Xylosandrus compactus (shot-hole borer)
- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- History of Introduction and Spread
- Risk of Introduction
- Hosts/Species Affected
- Host Plants and Other Plants Affected
- Growth Stages
- List of Symptoms/Signs
- Species Vectored
- Biology and Ecology
- Natural enemies
- Notes on Natural Enemies
- Means of Movement and Dispersal
- Plant Trade
- Wood Packaging
- Impact Summary
- Impact: Biodiversity
- Threatened Species
- Risk and Impact Factors
- Prevention and Control
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Xylosandrus compactus (Eichhoff, 1875)
Preferred Common Name
- shot-hole borer
Other Scientific Names
- Xyleborus compactus Eichhoff
- Xyleborus morstatti Hagedorn, 1912
- Xylosandrus morstatti (Hagedorn)
International Common Names
- English: black coffee borer; black coffee twig borer; black twig borer; tea stem borer
- French: scolyte des rameaux du caféier; scolyte noir des rameaux; scolyte noir du caféier
Local Common Names
- Germany: Bohrer, Schwarzer Kaffeezweig-; Schwarzer Zweigbohrer an Kaffee
- Netherlands: takkenboeboek; zwarte takkenboeboek
Summary of InvasivenessTop of page X. compactus should be considered a high-risk quarantine pest; most of the species in Xylosandrus and related genera should be considered potential quarantine pests. This is because members of the tribe Xyleborini (Xylosandrus plus related genera) are all inbreeding, with the males generally mating with their sisters within the parental gallery system before dispersal. Thus the introduction of only a few mated females may lead to the establishment of an active population if suitable host plants can be found and environmental conditions are satisfactory. Even suitable host plants may not be a limiting factor because the adult beetle does not actually feed on the plant material but uses it as a medium for growing the fungus, which is the larval food. Any woody material of suitable moisture content and size may be all that is required. A very wide range of host plants has been recorded for many species of Xylosandrus. The direct risk of establishment of species of Xylosandrus into tropical and subtropical areas should be considered extremely serious.
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Metazoa
- Phylum: Arthropoda
- Subphylum: Uniramia
- Class: Insecta
- Order: Coleoptera
- Family: Scolytidae
- Genus: Xylosandrus
- Species: Xylosandrus compactus
Notes on Taxonomy and NomenclatureTop of page Xylosandrus compactus was described by Eichhoff in 1875 in the genus Xyleborus. Xylosandrus was described by Reitter in 1913, and X. compactus was transferred to Xylosandrus by Nunberg (1959) and Browne (1963). Xyleborus morstatti was recognized as a synonym of X. compactus by Murayama and Kalshoven (1962). No other synonyms have been recognized.
DescriptionTop of page Eggs
The egg of X. compactus is about 0.3 mm wide and 0.5 mm long. It is white and ovoid with a smooth surface (Hara and Beardsley, 1979). The incubation period varies from 3 to 5 days with over 80% of eggs hatching after 4 days (Hara and Beardsley, 1979).
The mature larva is about 2.0 mm long. The body is creamy white with a pale-brown head. It has no legs. The mean head width of final-instar larvae is about 0.36 mm (Ngoan et al., 1976). A detailed description of the larva has not been published.
The pupae are illustrated by Hara and Beardsley (1979). The body of the pupa is creamy white and exarate. It is about the same length as the adult.
Bright (1968) provided a brief description of the female and male of X. compactus. The adult females are dark brown to almost black, 1.4-1.9 mm long and about two times longer than wide. The front of the head is convex, with a weak transverse impression just above the mouthparts. The antennal funicle is five-segmented, and the antennal club is obliquely truncate, about 1.2 times longer than wide. The pronotum, viewed from above, is subcircular. The anterior margin of the pronotum is broadly rounded, with 6-8 (sometimes 10) distinct, equal-sized serrations. The anterior half of the pronotum is finely asperate whereas the posterior portion is smooth with distinct, shallow punctures. The elytra are 1.1 times longer than wide, convex and steeply declivitous posteriorly. The strial punctures on the elytra are distinctly impressed, about equal in size to those between the striae. Each interstria bears a row of long setae, these are about two times longer than the interstrial width. The steeply convex, posterior portion of the elytra is similar to the remaining portion of the elytra.
The small, wingless males are about 0.8-1.1 mm long and two times longer than wide. The pronotum is narrowly rounded in front without serrations. The anterior portion of the pronotum is flattened and slightly concave in the median portion and the asperities are very low, almost obsolete.The elytral striae and interstrial are irregularly punctured.
A partial list of illustrations of the adult is given in Wood and Bright (1992) and Bright and Skidmore (1997).
Wood (1982) provided a key to species of Xylosandrus found in North and Central America, including X. compactus.
DistributionTop of page In addition to the records given here, there are unpublished records from Brunei Darussalam, Christmas Island and Malaysia (Sarawak) (RA Beaver, Chiangmai, Thailand, personal communication, 2004). The record of X. compactus from New Zealand (Wood and Bright, 1992; CABI/EPPO, 1997) must refer to intercepted specimens not seen by the New Zealand Quarantine Service, or be incorrect. Brockerhoff et al. (2003) noted that there is no breeding population in New Zealand and they do not include X. compactus in their list of Scolytidae intercepted in the country.
Distribution TableTop of page
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/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Cambodia||Present||Native||Not invasive||Waterhouse, 1993; CABI/EPPO and, 1997; EPPO, 2013|
|China||Present||Native||Not invasive||CABI/EPPO and, 1997; EPPO, 2013|
|-Guangdong||Present||Native||Not invasive||CABI/EPPO and, 1997; EPPO, 2013|
|-Guizhou||Present||Native||Not invasive||CABI/EPPO and, 1997; EPPO, 2013|
|-Hainan||Present||Native||Not invasive||CABI/EPPO and, 1997; EPPO, 2013|
|-Hunan||Present||Native||Not invasive||CABI/EPPO and, 1997; EPPO, 2013|
|East Timor||Restricted distribution||IPPC, 2016|
|India||Present||Native||Not invasive||CABI/EPPO and, 1997; EPPO, 2013|
|-Gujarat||Present||Native||Not invasive||CABI/EPPO and, 1997; EPPO, 2013|
|-Karnataka||Present||Native||Not invasive||CABI/EPPO and, 1997; EPPO, 2013|
|-Kerala||Present||Native||Not invasive||CABI/EPPO and, 1997; EPPO, 2013|
|-Madhya Pradesh||Present||Native||Not invasive||CABI/EPPO and, 1997; EPPO, 2013|
|-Maharashtra||Present||Native||Not invasive||CABI/EPPO and, 1997; EPPO, 2013|
|-Tamil Nadu||Present||Native||Not invasive||CABI/EPPO and, 1997; EPPO, 2013|
|Indonesia||Present||Native||Not invasive||Waterhouse, 1993; CABI/EPPO and, 1997; EPPO, 2013|
|-Irian Jaya||Present||Native||Not invasive||CABI/EPPO and, 1997; EPPO, 2013|
|-Java||Present||Native||Not invasive||CABI/EPPO and, 1997; EPPO, 2013|
|-Kalimantan||Present||Native||Not invasive||CABI/EPPO and, 1997; EPPO, 2013|
|-Sulawesi||Present||Native||Not invasive||CABI/EPPO and, 1997; EPPO, 2013|
|-Sumatra||Present||Native||Not invasive||CABI/EPPO and, 1997; EPPO, 2013|
|Japan||Present||Native||Not invasive||CABI/EPPO and, 1997; EPPO, 2013|
|-Hokkaido||Present||Native||Not invasive||CABI/EPPO and, 1997; EPPO, 2013|
|-Honshu||Present||Native||Not invasive||CABI/EPPO and, 1997; EPPO, 2013|
|-Kyushu||Present||Native||Not invasive||CABI/EPPO and, 1997; EPPO, 2013|
|-Ryukyu Archipelago||Present||Native||Not invasive||CABI/EPPO and, 1997; EPPO, 2013|
|-Shikoku||Present||Native||Not invasive||CABI/EPPO and, 1997; EPPO, 2013|
|Laos||Present||Native||Not invasive||CABI/EPPO and, 1997; EPPO, 2013|
|Malaysia||Present||Native||Not invasive||Waterhouse, 1993; CABI/EPPO and, 1997; EPPO, 2013|
|-Peninsular Malaysia||Present||Native||Not invasive||CABI/EPPO and, 1997; EPPO, 2013|
|-Sabah||Present||Native||Not invasive||CABI/EPPO and, 1997; EPPO, 2013|
|Myanmar||Present||Native||Not invasive||Waterhouse, 1993; CABI/EPPO and, 1997; EPPO, 2013|
|Philippines||Present||Native||Not invasive||Wood and Bright, 1992; CABI/EPPO and, 1997; EPPO, 2013|
|Singapore||Present||Native||Not invasive||CABI/EPPO and, 1997; EPPO, 2013|
|Sri Lanka||Present||Native||Not invasive||CABI/EPPO and, 1997; EPPO, 2013|
|Taiwan||Present||Native||Not invasive||Wood and Bright, 1992; CABI/EPPO and, 1997; EPPO, 2013|
|Thailand||Present||Native||Not invasive||Wood and Bright, 1992; CABI/EPPO and, 1997; EPPO, 2013|
|Vietnam||Present||Native||Not invasive||Waterhouse, 1993; CABI/EPPO and, 1997; EPPO, 2013|
|Benin||Present||Introduced||Invasive||CABI/EPPO and, 1997; EPPO, 2013|
|Cameroon||Present||Introduced||Invasive||CABI/EPPO and, 1997; EPPO, 2013|
|Central African Republic||Present||Introduced||Invasive||CABI/EPPO and, 1997; EPPO, 2013|
|Comoros||Present||Introduced||Invasive||Wood and Bright, 1992; CABI/EPPO and, 1997; EPPO, 2013|
|Congo||Present||Introduced||Invasive||CABI/EPPO and, 1997; EPPO, 2013|
|Congo Democratic Republic||Present||EPPO, 2013|
|Côte d'Ivoire||Present||Introduced||Invasive||CABI/EPPO and, 1997; EPPO, 2013|
|Equatorial Guinea||Present||EPPO, 2013|
|Gabon||Present||Introduced||Invasive||CABI/EPPO and, 1997; EPPO, 2013|
|Ghana||Present||Introduced||Invasive||CABI/EPPO and, 1997; EPPO, 2013|
|Guinea||Present||Introduced||Invasive||CABI/EPPO and, 1997; EPPO, 2013|
|Guinea-Bissau||Present||Introduced||Invasive||CABI/EPPO and, 1997; EPPO, 2013|
|Kenya||Present||Introduced||Invasive||CABI/EPPO and, 1997; EPPO, 2013|
|Liberia||Present||Introduced||Invasive||CABI/EPPO and, 1997; EPPO, 2013|
|Madagascar||Present||Introduced||Invasive||CABI/EPPO and, 1997; EPPO, 2013|
|Mauritania||Present||Introduced||Invasive||Wood and Bright, 1992; CABI/EPPO and, 1997; EPPO, 2013|
|Mauritius||Present||Introduced||Invasive||CABI/EPPO and, 1997; EPPO, 2013|
|Nigeria||Present||Introduced||Invasive||CABI/EPPO and, 1997; EPPO, 2013|
|Réunion||Present||Introduced||Invasive||CABI/EPPO and, 1997; EPPO, 2013|
|Senegal||Present||Introduced||Invasive||Wood and Bright, 1992; CABI/EPPO and, 1997; EPPO, 2013|
|Seychelles||Present||Introduced||Invasive||CABI/EPPO and, 1997; EPPO, 2013|
|Sierra Leone||Present||Introduced||Invasive||CABI/EPPO and, 1997; EPPO, 2013|
|South Africa||Present||Introduced||Invasive||Wood and Bright, 1992; EPPO, 2013|
|Tanzania||Present||Introduced||Invasive||CABI/EPPO and, 1997; EPPO, 2013|
|Togo||Present||Introduced||Invasive||CABI/EPPO and, 1997; EPPO, 2013|
|Uganda||Present||Introduced||Invasive||CABI/EPPO and, 1997; EPPO, 2013|
|Zimbabwe||Present||Introduced||Invasive||CABI/EPPO and, 1997; EPPO, 2013|
|USA||Restricted distribution||EPPO, 2013|
|-Alabama||Present||Introduced||Invasive||CABI/EPPO and, 1997; EPPO, 2013|
|-Florida||Present||Introduced||Invasive||Wood, 1977; CABI/EPPO and, 1997; EPPO, 2013|
|-Georgia||Present||Introduced||Invasive||Wood, 1977; CABI/EPPO and, 1997; EPPO, 2013|
|-Hawaii||Present||Introduced||Invasive||CABI/EPPO and, 1997; EPPO, 2013|
|-Louisiana||Present||Introduced||Invasive||CABI/EPPO and, 1997; EPPO, 2013|
|-Mississippi||Present||Introduced||Invasive||Wood, 1977; Wood and Bright, 1992; CABI/EPPO and, 1997; EPPO, 2013|
|-South Carolina||Present||Introduced||Invasive||CABI/EPPO and, 1997; EPPO, 2013|
|-Texas||Present||Introduced||Invasive||Wood and Bright, 1992; CABI/EPPO and, 1997; EPPO, 2013|
Central America and Caribbean
|British Virgin Islands||Present||Introduced||Invasive||CABI/EPPO and, 1997; EPPO, 2013|
|Cuba||Present||Introduced||Invasive||Bright, 1985; Wood, 1977; CABI/EPPO and, 1997; EPPO, 2013|
|Curaçao||Present||Introduced||Invasive||Vazquez and Monteagudo, 1988|
|Netherlands Antilles||Present||Introduced||Invasive||CABI/EPPO and, 1997; EPPO, 2013|
|Puerto Rico||Present||Introduced||Invasive||Franqui, 1991; CABI/EPPO and, 1997; EPPO, 2013|
|United States Virgin Islands||Present||Introduced||Invasive||Bright, 1985; CABI/EPPO and, 1997; EPPO, 2013|
|Brazil||Present||Introduced||Invasive||Wood, 1980; CABI/EPPO and, 1997; EPPO, 2013|
|-Amazonas||Present||Introduced||Invasive||Abreu et al., 2001; Wood, 1980|
|-Goias||Present||Oliveira et al., 2008|
|Italy||Present||Francardi et al., 2012; Garonna et al., 2012; Pennacchio et al., 2012; EPPO, 2013|
|American Samoa||Present||EPPO, 2013|
|Fiji||Present||Introduced||Invasive||CABI/EPPO and, 1997; EPPO, 2013|
|New Zealand||Absent, unreliable record||Wood and Bright, 1992; CABI/EPPO and, 1997; Brockerhoff et al., 2003; EPPO, 2013|
|Papua New Guinea||Present||Introduced||Invasive||CABI/EPPO and, 1997; EPPO, 2013|
|Samoa||Present||Introduced||Invasive||CABI/EPPO and, 1997; EPPO, 2013|
|Solomon Islands||Present||Introduced||Invasive||Bigger, 1985; EPPO, 2013|
History of Introduction and SpreadTop of page X. compactus was probably unintentionally introduced to the Afrotropical region from the Oriental region hundreds of years ago by early traders, and is now widespread. It was accidentally introduced into the Americas before the middle of the twentieth century. Wood (1977) indicated that X. compactus was present in Florida in 1941, in Cuba in 1958, in Mississippi in 1968 and in Georgia ca 1975. Since then it has spread further in the USA and has been accidentally introduced to other Caribbean countries. It was collected in Brazil (Amazonas) in 1979 (Wood, 1980) and again in the same province more recently (Abreu et al., 2001), but has apparently not yet spread to other countries of South America. It was intercepted in Hawaii in 1931, but not found again until 1961 in Oahu, by which time it was established and spreading (Samuelson, 1981). According to Samuelson (1981), it spread to the islands of Kauai by 1962, Hawaii by 1966, Maui by 1968, Molokai by 1974 and Lanai by 1975.
Risk of IntroductionTop of page Two other species of Xylosandrus, Xylosandrus crassiusculus and Xylosandrus morigerus, with similar habits to X. compactus, have become important pests of tree crops, ornamental and native trees in tropical and subtropical areas where they have been introduced. The risk of introduction for X. compactus must be considered high, most probably in the twigs and small branches of imported plants. Once established, such species are difficult to eradicate and are likely to spread with the movement of infested plants, as well as by normal dispersal of the adults. X. compactus is listed as a quarantine pest in New Zealand, but apparently not elsewhere.
Hosts/Species AffectedTop of page Over 225 species of plants, belonging to 62 families, are susceptible to X. compactus (Ngoan et al., 1976). Browne (1961) remarked that X. compactus does not appear to be highly host specific in its natural mixed-forest habitat, and it is only when it finds special conditions of concentrated cultivation that it tends to be a pest. Only a selection of the recorded hosts is given here. Further host lists can be found in Schedl (1963), Brader (1964), Hara and Beardsley (1979) and Wood and Bright (1992).
The main economic host of X. compactus is coffee (especially Coffea canephora robusta, also Coffea arabica). In Japan, X. compactus is a pest of tea (Kaneko et al., 1965). X. compactus is also a pest of avocado and cocoa in South-East Asia and elsewhere (Kalshoven, 1958; Browne, 1961; Beaver, 1976; Waterhouse, 1997; Nair, 2000; Matsumoto, 2002). In India, X. compactus is reported as infesting and killing the seedlings and saplings of Khaya grandifoliola, and Khaya senegalensis, shade trees in coffee plantations (Meshram et al., 1993); in Africa, Erythrina sp. and Melia azedarach (Le Pelley, 1968). Attacks on seedlings and young plantations of a variety of forest trees can be severe (Browne, 1968; Intachat and Kirton, 1997). In addition to the large range of dicotyledonous trees and shrubs, it will sometimes attack both monocotyledonous plants, such as orchids and gingers, and conifers (Hara and Beardsley, 1979). Its attacks can also endanger rare native trees (Ziegler, 2001, 2002).
Host Plants and Other Plants AffectedTop of page
|Acacia auriculiformis (northern black wattle)||Fabaceae||Other|
|Acacia mangium (brown salwood)||Fabaceae||Other|
|Annona muricata (soursop)||Annonaceae||Main|
|Annona squamosa (sugar apple)||Annonaceae||Other|
|Aucoumea klaineana (okoume)||Burseraceae||Other|
|Caesalpinia kavaiensis||Fabaceae||Wild host|
|Camellia sinensis (tea)||Theaceae||Main|
|Cedrela odorata (Spanish cedar)||Meliaceae||Other|
|Cinnamomum verum (cinnamon)||Lauraceae||Other|
|Coffea arabica (arabica coffee)||Rubiaceae||Main|
|Coffea canephora (robusta coffee)||Rubiaceae||Main|
|Colubrina oppositifolia||Rhamnaceae||Wild host|
|Cornus florida (Flowering dogwood)||Cornaceae||Other|
|Dalbergia (rosewoods)||Fabaceae||Wild host|
|Entandrophragma utile (ogipogo-mahogany)||Meliaceae||Other|
|Erythrina abyssinica (red hot poker tree)||Fabaceae||Other|
|Eusideroxylon zwageri (billian)||Lauraceae||Wild host|
|Hevea brasiliensis (rubber)||Euphorbiaceae||Other|
|Khaya grandifoliola (big-leaf mahogany)||Meliaceae||Other|
|Khaya ivorensis (African mahogany)||Meliaceae||Other|
|Khaya senegalensis (dry zone mahogany)||Meliaceae||Other|
|Laurus nobilis (sweet bay)||Lauraceae||Other|
|Leucaena leucocephala (leucaena)||Fabaceae||Other|
|Macadamia integrifolia (macadamia nut)||Proteaceae||Other|
|Mangifera indica (mango)||Anacardiaceae||Other|
|Melia azedarach (Chinaberry)||Meliaceae||Other|
|Ochroma pyramidale (balsa)||Bombacaceae||Other|
|Persea americana (avocado)||Lauraceae||Other|
|Pometia pinnata (fijian longan)||Sapindaceae||Other|
|Punica granatum (pomegranate)||Punicaceae||Other|
|Swietenia macrophylla (big leaved mahogany)||Meliaceae||Main|
|Swietenia mahagoni (Cuban mahogany)||Meliaceae||Other|
|Theobroma cacao (cocoa)||Sterculiaceae||Other|
|Toona ciliata (toon)||Meliaceae||Other|
Growth StagesTop of page Flowering stage, Fruiting stage, Seedling stage, Vegetative growing stage
SymptomsTop of page X. compactus bores into the current year's twigs, killing them in a few weeks or causing them to break from the weight of the crop. The pest weakens and retards the fruiting of young plants and makes the replacement of trees very difficult. The typical host symptoms that characterize X. compactus infestation are necrosis of the leaves and stem extending from the entrance hole distally to the end of the branch. Flagging of branches occurs about 5-7 days after initial tunnelling and gallery formation. Wilting of twigs and branches usually becomes evident within weeks of infestation. The entrance holes are small (0.8 mm diameter) and are located on the underside of branches. Cankers, 10-210 mm long, are commonly seen around the attacked areas of larger twigs and branches (Dixon and Woodruff, 1982). A whitish pile of dust from boring may be seen at each hole.
X. compactus is one of the few species of ambrosia beetles that can attack and kill live twigs and branches. Most of the other species of ambrosia beetles primarily attack newly felled, stressed, dead or dying trees and shrubs. Apparently, the pathogenic action of the ambrosia fungus, Fusarium solani to the host plant enables X. compactus to attack live plants. The pathogenic action of F. solani to woody host plants has been proven by pure culture isolates of F. solani from discoloured vascular tissues of a large number of host species (Dixon and Woodruff, 1982).
List of Symptoms/SignsTop of page
|Leaves / necrotic areas|
|Leaves / wilting|
|Stems / internal feeding|
|Stems / necrosis|
|Whole plant / plant dead; dieback|
Biology and EcologyTop of page Several studies of the biology of X. compactus have been made. Browne (1961) reviewed the biology of X. compactus in South-East Asia, under the name Xylosandrus morstatti. Brader (1964) studied the biology of the pest when attacking coffee in West Africa, with particular reference to the effects of climatic factors, and relationships to the associated ambrosia fungus, and to the host plant. Kaneko (1965) and Kaneko et al. (1965) studied the biology of X. compactus in tea plants in Japan. Entwistle (1972) gives a detailed review of earlier work. Dixon and Woodruff (1982) summarized the biology and ecology of X. compactus in Florida, USA. Hara and Beardsley (1979) reported on the biology of X. compactus in Hawaii, USA. Beaver (1988) studied the biology and host associations of X. compactus in the Seychelles. Wood and Bright (1992) give several hundred references relating to the biology, habits, taxonomy and control of X. compactus. Bright and Skidmore (1997, 2002) give more recent references.
Only the adult females initiate the attack on the host plants. X. compactus is mainly a borer of seedlings, shoots and small twigs, but it will also breed in cut branches and poles up to a diameter of about 6 cm, rarely in larger material (Browne, 1961). Attacks in the tap root of seedlings have been noted in West Africa (Entwistle, 1972), but such attacks are more likely to be caused by the related species, Xylosandrus morigerus. On cocoa seedlings in Nigeria, attacks were most abundant 20-40 cm above ground level, and on stems of 6-10 mm diameter (Entwistle, 1972). The female constructs an entrance tunnel into the pith or wood of the host to a depth of 1-3 cm. The tunnel system consists of a simple or bifurcated entrance tunnel, and a longitudinal chamber or irregular tunnel where a loose cluster of eggs is deposited (Browne, 1961; Entwistle, 1972). One or more females may occupy a twig or branch. Generally, there is only one female if the twig diameter is less than 7 mm, but up to 20 females may be found on branches of diameter 8-22 mm.
Entwistle (1964) and Takenouchi and Tagaki (1967) report arrhenotokous parthenogenesis in X. compactus. Unmated females produce an all-male brood, but such broods are rare (Brader, 1964; Entwistle, 1972). Hara and Beardsley (1979) found seven all-male broods out of 416 examined.
The size of the brood varies considerably. Browne (1961) found that in peninsular Malaysia, broods rarely exceeded 10 individuals, and in the Seychelles, the largest brood that was observed by Brown (1954) included two eggs and seven larvae. Chevalier (1931) reported broods of 30-50 individuals in tropical Africa. In one gallery system in Fiji, 26 individuals of all instars were found (Lever, 1938). Entwistle (1972) found a mean of 12.3 offspring in field-collected galleries, but the number could occasionally exceed 60. The larvae feed on an ambrosia fungus growing on the walls of the gallery.
The pupation and mating of brood adults occurs in the infested material; the (usually) single male in each gallery mating with his sisters. The brood adults emerge through the entrance holes made by the parent beetles. In tropical Africa, Lavabre (1958, 1959) found that oviposition began 7-8 days after the parent female began her gallery. The egg stage lasted 4-5 days, larval development took 11 days, 7 days were spent in the pupal stage and the teneral adults remained in the gallery system for another 6 days before emerging. Thus about 37 days were required from the time the female first began boring into the branch until sexual maturity of the next generation. Ngoan et al. (1976) found that approximately 28 days (at 25°C) were required for development from egg to adult. According to Ngoan et al. (1976) and Hara and Beardsley (1979), there are two larval instars. The ratio of females to males varies, but is usually approximately 9:1 (Entwistle, 1972; Hara and Beardsley, 1979).
In Japan, there are normally two generations per year, and adult females overwinter, but in most parts of the range, breeding is continuous, with overlapping generations, so that the species is active at all times, and in all stages of development (Browne, 1968).
Natural enemiesTop of page
|Natural enemy||Type||Life stages||Specificity||References||Biological control in||Biological control on|
|Tetrastichus sp. nr. xylebororum||Parasite||Larvae|
Notes on Natural EnemiesTop of page The immature stages have few natural enemies. The female parent normally remains in the gallery entrance whilst the immature stages are developing, preventing the entry of potential predators and parasitoids. Provided that the female remains alive and the growth of the ambrosia fungus on which the larvae feed is satisfactory, mortality of the immature stages is likely to be very low. Most mortality is probably during the dispersal of the adults and during gallery establishment.
Several studies of the natural enemies of X. compactus have been made in India. Sreedharan et al. (1992) reported that larvae of the clerid coleopteran Callimerus sp. were found in over 4% of the gallery systems of X. compactus. Although the predator feeds on all stages of the scolytid, it prefers feeding on the larvae. In India, Eupelmus sp. was found in nearly 8% of the branches of robusta coffee examined (Balakrishnan et al., 1989). The larvae of this eupelmid act as predators when several ambrosia beetle larvae are available. The incidence of parasitism ranged from 1.3% in January to nearly 21% in September. In Java, the eulophid Tetrastichus xylebororum parasitizes both this species and Xylosandrus morigerus (Le Pelley, 1968), a related species recorded from India (Dhanam et al., 1992), and a further species of Tetrastichus from Hawaii (Tenbrink and Hara, 1994). Le Pelley (1968) mentioned an undescribed bethylid ectoparasitoid of X. compactus larvae, and the bethylid, Prorops nasuta, a well-known parasite of the coffee berry borer, Hypothenemus hampei, has been recorded attacking X. compactus in West Africa (Brader, 1964). However, parasitism is not normally an important cause of mortality in Xylosandrus species.
One species of entomopathogenic fungus, Beauveria bassiana, was found infecting X. compactus in India (Balakrishnan et al., 1994) and has also been recorded in West Africa (Brader, 1964).
During gallery establishment, the adults are frequently attacked by ants (Lavabre, 1962; Brader, 1964). Brader (1964) noted attacks by Oecophylla longinoda in Côte d'Ivoire, and similar attacks by Oecophylla smaragdina occur in South-East Asia. Lizards and clerid beetles prey on the adults of ambrosia beetles, such as Xylosandrus as the latter attempt to bore into the host tree.
Means of Movement and DispersalTop of page Natural Dispersal
The adult females fly readily, and flight is one the main means of movement and dispersal to previously uninfected areas. Entwistle (1972) found that adult females dispersed at least 200 m, and it is likely that dispersal over several kilometres is possible, especially if wind-aided. However, of more importance for long distance movement is the transport of infested seedlings, saplings or cut branches.
The female has a mycangium, a pouch used to carry spores of the ambrosia fungus on which both adult and larvae feed, opening between the pronotum and mesonotum, and extending below the pronotum (Beaver, 1989). The ambrosia fungus of X. compactus has been variously identified by different researchers. Brader (1964) described it as Ambrosiella xylebori in Côte d'Ivoire. Bhat and Sreedharan (1988) agreed, but Muthappa and Venkatasubbaiah (1981) suggested that in India, the ambrosia fungus is Ambrosiella macrospora. In North America, the fungus has been identified as Fusarium solani (Ngoan et al., 1976; Hara and Beardsley, 1979). It is possible that A. xylebori is a pleomorphic form of F. solani (Hara and Beardsley, 1979). The records of Cladosporium cladosporioides and Penicillium pallidum [Geosmithia putterillii] (Brown, 1954) from X. compactus galleries in the Seychelles are of secondary saprophytic fungi (Schedl, 1963). Ambrosiella spp. are not known to be pathogenic, although they do cause staining of the wood around the gallery systems. However, F. solani is well known as a plant pathogen, and its pathogenicity to host plants of X. compactus has been confirmed (Hara and Beardsley, 1979; Dixon and Woodruff, 1983). Entwistle (1972) noted that fungal attack always follows gallery formation. In West Africa, the fungi Botryodiplodia theobromae [Lasiodiplodia theobromae] and Calonectria rigidiuscula [Nectria rigidiuscula] (the perfect stage of Fusarium decemcellulare), both of which are wound parasites of weak pathogenicity, are involved (Entwistle, 1972).
Plant TradeTop of page
|Plant parts liable to carry the pest in trade/transport||Pest stages||Borne internally||Borne externally||Visibility of pest or symptoms|
|Bark||adults||Yes||Pest or symptoms usually visible to the naked eye|
|Seedlings/Micropropagated plants||adults; eggs; larvae; pupae||Yes||Pest or symptoms not visible to the naked eye but usually visible under light microscope|
|Stems (above ground)/Shoots/Trunks/Branches||adults; eggs; larvae; pupae||Yes||Pest or symptoms not visible to the naked eye but usually visible under light microscope|
|Wood||adults; eggs; larvae; pupae||Yes||Pest or symptoms not visible to the naked eye but usually visible under light microscope|
|Plant parts not known to carry the pest in trade/transport|
|Fruits (inc. pods)|
|Growing medium accompanying plants|
|True seeds (inc. grain)|
Wood PackagingTop of page
|Wood Packaging not known to carry the pest in trade/transport|
|Loose wood packing material|
|Processed or treated wood|
|Solid wood packing material with bark|
|Solid wood packing material without bark|
Impact SummaryTop of page
|Fisheries / aquaculture||None|
ImpactTop of page X. compactus is a serious pest of shrubs and trees. It causes extensive damage to coffee and cocoa throughout tropical Africa, Indonesia, southern India and the West Indies. There is no recent objective assessment of crop losses caused by X. compactus. In India, Ramesh (1987) observed that losses due to X. compactus were 21% on 45-year-old coffee plants and 23.5% on young plants. Infestation rates of 60-70% in African mahogany were reported by Meshram et al. (1993) in India. Lavabre (1958, 1959) reported losses of about 20% of the coffee crop in Cameroon. In Japan, X. compactus is a major pest of tea causing extensive dieback (Kaneko et al., 1965). In China, Yan et al. (2001) recorded an attack rate of 78% on the main stems of young chesnut trees.
Impact: BiodiversityTop of page In Hawaii, X. compactus attacks several rare and threatened native trees, including Colubrina oppositifolia (Ziegler, 2001) and Caesalpinia kavaiensis (Ziegler, 2002), providing an additional threat to their survival. Similar threats to rare native trees may occur elsewhere in the range of the beetle as a result of its very wide host range.
Threatened SpeciesTop of page
|Threatened Species||Conservation Status||Where Threatened||Mechanism||References||Notes|
|Cyanea calycina||CR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered); USA ESA listing as endangered species USA ESA listing as endangered species||Hawaii||Herbivory/grazing/browsing||US Fish and Wildlife Service, 2012|
|Cyanea lanceolata (lanceleaf cyanea)||USA ESA listing as endangered species USA ESA listing as endangered species||Hawaii||Herbivory/grazing/browsing||US Fish and Wildlife Service, 2012|
|Doryopteris takeuchii (Takeuch's lipfern)||NatureServe NatureServe; USA ESA listing as endangered species USA ESA listing as endangered species||Hawaii||Herbivory/grazing/browsing||US Fish and Wildlife Service, 2012|
|Flueggea neowawraea (mehamehame)||CR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered); USA ESA listing as endangered species USA ESA listing as endangered species||Hawaii||Pathogenic||,|
|Melicope christophersenii||EN (IUCN red list: Endangered) EN (IUCN red list: Endangered); NatureServe NatureServe; USA ESA listing as endangered species USA ESA listing as endangered species||Hawaii||Pathogenic||US Fish and Wildlife Service, 2012|
|Melicope hiiakae||NatureServe NatureServe; USA ESA listing as endangered species USA ESA listing as endangered species||Hawaii||Poisoning||US Fish and Wildlife Service, 2012|
|Melicope makahae||EN (IUCN red list: Endangered) EN (IUCN red list: Endangered); NatureServe NatureServe; USA ESA listing as endangered species USA ESA listing as endangered species||Hawaii||Pollen swamping||US Fish and Wildlife Service, 2012|
|Plantago princeps||NatureServe NatureServe; USA ESA listing as endangered species USA ESA listing as endangered species||Hawaii||Pest and disease transmission||US Fish and Wildlife Service, 2010|
|Santalum freycinetianum var. lanaiense||No Details||Hawaii||Herbivory/grazing/browsing||US Fish and Wildlife Service, 2011|
|Schiedea nuttallii||CR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered); USA ESA listing as endangered species USA ESA listing as endangered species||Hawaii||Pest and disease transmission||US Fish and Wildlife Service, 1999; US Fish and Wildlife Service, 2009|
|Serianthes nelsonii||Guam; Northern Mariana Islands||Pest and disease transmission||US Fish and Wildlife Service, 1994|
Risk and Impact FactorsTop of page Impact mechanisms
- Pest and disease transmission
- Pollen swamping
Prevention and ControlTop of page
The decision to use chemical control is influenced by environmental concerns and the difficulties of applying chemicals to the concealed habitats in which X. compactus feeds. Mangold et al. (1977) reported that when chlorpyrifos was applied with a hand sprayer to individual twigs of flowering dogwood in Florida, USA, there was 77% mortality of all stages of the beetle. In subsequent field studies, hydraulic sprays of chlorpyrifos killed 83-92% of all beetle stages per infested twig. Bambara (2003) suggested the use of chlorpyrifos, permethrin or bifenthrin. Yan et al. (2001) used quinalphos or chlorypyrifos plus cypermethrin mixed with yellow soil and painted it on the main stem of young chestnut trees, and reported good control.
Material that is infested with X. compactus should be pruned and destroyed. Where practicable, this is perhaps the most effective method of control, but it may not be economic (Le Pelley, 1968). Practices that promote tree vigour and health will aid recovery from beetle damage (Dixon and Woodruff, 1982; Bambara, 2003). Entwistle (1972) noted the attraction of X. compactus to cocoa seedlings with overhead shade, and with a growing ground cover, but he pointed out that shade may be essential for seedling establishment, and that its removal may also render the plant more susceptible to other pests. In Malaysia, Anuar (1986) found that when growing robusta and Liberian coffee under shaded and unshaded conditions, only robusta showed damage caused by X. compactus. The frequency and severity of damage was significantly higher on shaded than on unshaded trees.
X. compactus is "singularly free from attack by parasites and predators" (Entwistle, 1972). In Africa, there are no effective parasites of X. compactus (Brader, 1964). Parasites are known in Indonesia and Le Pelley (1968) suggested that they have "an appreciable effect from time to time". The entomopathogenic fungus, Beauveria bassiana, causes some mortality in X. compactus and its potential usefulness is being investigated (Balakrishnan et al., 1994). However, biological control methods seem unlikely to be effective for X. compactus.
ReferencesTop of page
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Distribution MapsTop of page
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