Xylosandrus crassiusculus (Asian ambrosia beetle)

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Adult

Xylosandrus crassiusculus (Asian ambrosia beetle); lateral view. USA. VA. Albemarle Co., Charlottesville, 20 May 2005, from funnel trap; det. R.J. Rabaglia, 2007.

©Pest & Diseases Image Library (PaDIL)/Bugwood.org - CC BY 3.0 AU

Identity

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Preferred Scientific Name

Xylosandrus crassiusculus (Motschulsky)

Preferred Common Name

Asian ambrosia beetle

Other Scientific Names

Dryocoetes bengalensis Stebbing
Xyleborus bengalensis Stebbing
Xyleborus crassiusculus (Motschulsky)
Xyleborus declivigranulatus Schedl
Xyleborus ebriosus Niisima
Xyleborus mascarenus Hagedorn
Xyleborus okoumeensis Schedl
Xyleborus semigranosus Blandford
Xyleborus semiopacus Eichhoff
Xylosandrus semigranosus (Blandford)
Xylosandrus semiopacus (Eichhoff)

International Common Names

English: granulate ambrosia beetle

EPPO code

XYLBCR (Xyleborus crassiusculus)
XYLBEB (Xyleborus ebriosus)

Summary of Invasiveness

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X. crassiusculus should be considered a high-risk quarantine pest; most of the species in Xyleborus and related genera should be considered potential quarantine pests. This reflects the fact that since members of the tribe Xyleborini (Xyleborus plus related genera) are all parthenogenetic, the introduction of only a few individuals (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 since 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 density may be all that is required. A very wide range of host plants have been recorded for many species of Xyleborus and related genera. The direct risk of establishment of populations of species of Xyleborus into tropical and sub-tropical areas should be considered extremely serious.

Taxonomic Tree

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Domain: Eukaryota
Kingdom: Metazoa
Phylum: Arthropoda
Subphylum: Uniramia
Class: Insecta
Order: Coleoptera
Family: Scolytidae
Genus: Xylosandrus
Species: Xylosandrus crassiusculus

Notes on Taxonomy and Nomenclature

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Many species previously classified in the genus Xyleborus have now been transferred into other genera such as Ambrosiodmus, Euwallacea, Xylosandrus and Xyleborinus, including X. crassiusculus. These species are all ambrosia beetles. A number of species within the Xyleborini, the tribe in which Xyleborus and related genera are placed, can be considered potential pests to agriculture and forestry; X. crassiusculus is one of the more important species.

Description

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Adult Female

Length 2.2-2.5 mm. Frons weakly convex, with a distinct median line, surface coarsely granulate, sparsely punctate. Antennal club solid on posterior face, no sutures present. Pronotum about as long as wide; sides weakly arcuate, anterior margin narrowly rounded, with 8 or 9 weak serrations. Elytra 1.2-1.3 times longer than wide, apex broadly rounded. Elytral declivity abrupt, convex, surface opaque with dense, confused granules and rows of long stout setae.

Immature Stages

The egg and pupa have not been described. The larvae of X. crassiusculus and Xylosandrus discolor are briefly described and keyed by Gardner (1934). Gardner (1934) describes the mature larva (as Xyleborus semigranosus) as follows: length about 3.5 mm. Head pale, slightly wider than long. Labrum with strong posterior extension. Epipharyngeal setae very small. Maxillary palp with apical segment only slightly longer than wide. Labial palps separated by about width of a basal segment, apical segment globular, not longer than wide. Abdominal terga with two distinct folds separated by an extremely narrow subdivision. Spiracles with combined width of air-tubes equal to diameter of atrium. Skin rather densely covered with micro-asperities.

Too little is known of the larvae of other species of Xylosandrus to be sure whether the description is adequate to separate X. crassiusculus larvae from those of related species. Gardner (1934) distinguishes the species from Xylosandrus discolor by the micro-asperate (versus smooth) skin.

Distribution

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There are unpublished records from Brunei Darussalam, Christmas Island (Indian Ocean), Reunion, South Africa (RA Beaver, Chiangmai, Thailand, personal communication, 2004). Browne (1968) reported this species from Fiji, however, this record is incorrect and this species has not yet been recorded from the Fiji islands.

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.

Last updated: 25 Feb 2021

Continent/Country/Region	Distribution	Origin	Invasive	Reference	Notes
Africa
Cameroon	Present	Introduced		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
Congo, Democratic Republic of the	Present	Introduced		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
Côte d'Ivoire	Present	Introduced		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
Equatorial Guinea	Present	Introduced		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
Gabon	Present			EPPO (2020)
Ghana	Present	Introduced		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
Kenya	Present	Introduced		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
Madagascar	Present	Introduced		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
Mauritania	Present	Introduced		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
Mauritius	Present	Introduced		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
Nigeria	Present	Introduced		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
Seychelles	Present	Introduced		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
Sierra Leone	Present	Introduced		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
Tanzania	Present	Introduced		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
Asia
Bhutan	Present	Native		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
China	Present	Native		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
-Fujian	Present	Native		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
-Hunan	Present	Native		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
-Sichuan	Present	Native		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
-Tibet	Present	Native		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
-Yunnan	Present	Native		Yin et al. (1984) CABI and EPPO (2009) EPPO (2020)
Hong Kong	Present	Native		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
India	Present	Native		Wood and Bright (1992) Keshavareddy et al. (2008) CABI and EPPO (2009) Prathapan and Hiremath (2018) EPPO (2020)
-Andaman and Nicobar Islands	Present	Native		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
-Assam	Present	Native		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
-Himachal Pradesh	Present	Native		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
-Karnataka	Present	Native		Sreedharan et al. (1991) Keshavareddy et al. (2008) CABI and EPPO (2009) EPPO (2020)
-Kerala	Present			CABI and EPPO (2009) Prathapan and Hiremath (2018) EPPO (2020)
-Madhya Pradesh	Present	Native		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
-Maharashtra	Present	Native		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
-Tamil Nadu	Present	Native		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
-Uttar Pradesh	Present	Native		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
-West Bengal	Present	Native		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
Indonesia	Present	Native		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
-Irian Jaya	Present	Native		Schedl (1940) CABI and EPPO (2009) EPPO (2020)
-Java	Present	Native		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
-Lesser Sunda Islands	Present			EPPO (2018)
-Maluku Islands	Present	Native		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
-Sulawesi	Present	Native		Kalshoven (1959) CABI and EPPO (2009) EPPO (2020)
-Sumatra	Present	Native		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
Israel	Absent, Unconfirmed presence record(s)			EPPO (2020)
Japan	Present	Native		Wood and Bright (1992) CABI and EPPO (2009) Ito and Kajimura (2009) EPPO (2020)
-Bonin Islands	Present	Native		Wood and Bright (1992) CABI and EPPO (2009)
-Hokkaido	Present	Native		Murayama (1953) CABI and EPPO (2009) Ito and Kajimura (2009) EPPO (2020)
-Honshu	Present	Native		Murayama (1953) CABI and EPPO (2009) EPPO (2020)
-Kyushu	Present	Native		Murayama (1953) CABI and EPPO (2009) EPPO (2020)
-Ryukyu Islands	Present			Ito and Kajimura (2009)
-Shikoku	Present	Native		Murayama (1953) CABI and EPPO (2009) EPPO (2020)
Malaysia	Present	Native		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
-Peninsular Malaysia	Present	Native		Browne (1961) CABI and EPPO (2009) EPPO (2020)
-Sabah	Present	Native		Chey VunKhen (2002) CABI and EPPO (2009) EPPO (2020)
-Sarawak	Present	Native		Schedl (1964) CABI and EPPO (2009) EPPO (2020)
Myanmar	Present	Native		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
Nepal	Present	Native		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
North Korea	Present	Native		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
Pakistan	Present			Khuhro et al. (2005) CABI and EPPO (2009) EPPO (2020)
Philippines	Present	Native		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
South Korea	Present	Native		Choo et al. (1983) CABI and EPPO (2009) EPPO (2020) CABI (Undated)
Sri Lanka	Present	Native		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
Taiwan	Present	Native		MURAYAMA (1934) Eggers (1939) CABI and EPPO (2009) EPPO (2020)
Thailand	Present	Native		Beaver and Browne (1975) CABI and EPPO (2009) EPPO (2020)
Vietnam	Present	Native		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
Europe
Belgium	Absent			EPPO (2020)
France	Present, Localized			EPPO (2020)
Italy	Present	Introduced		Pennacchio et al. (2003) CABI and EPPO (2009) Francardi et al. (2017) EPPO (2020)
Lithuania	Absent, Confirmed absent by survey			EPPO (2020)
Slovenia	Present, Transient under eradication			EPPO (2020) Kavčič (2018) Kavčič (2019)
Spain	Present, Localized			EPPO (2020)
North America
Canada	Present, Few occurrences			EPPO (2020)
-Ontario	Present, Few occurrences			EPPO (2020)
Costa Rica	Present			CABI and EPPO (2009) EPPO (2020)
Guatemala	Present, Localized			EPPO (2020)
Honduras	Present			EPPO (2020)
Panama	Present			CABI and EPPO (2009) EPPO (2020)
United States	Present	Introduced	Invasive	Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
-Alabama	Present			Horn and Horn (2006) CABI and EPPO (2009) EPPO (2020)
-Arkansas	Present			EPPO (2020) CABI and EPPO (2009)
-Delaware	Present			CABI (Undated) CABI and EPPO (2009) EPPO (2020)	Original citation: Rabaglia and Valenti (2003)
-Florida	Present	Introduced	Invasive	Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
-Georgia	Present	Introduced	Invasive	Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
-Hawaii	Present	Introduced	Invasive	Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
-Indiana	Present	Introduced	Invasive	Pierce et al. (2005) CABI and EPPO (2009) EPPO (2020)
-Kansas	Present			EPPO (2020)
-Louisiana	Present	Introduced	Invasive	Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
-Maryland	Present	Introduced	Invasive	Bambara and Casey (2003) CABI and EPPO (2009) EPPO (2020)
-Michigan	Present			CABI and EPPO (2009) EPPO (2020)
-Mississippi	Present	Introduced	Invasive	Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
-Missouri	Present			CABI and EPPO (2009) EPPO (2020)
-Nebraska	Present			CABI and EPPO (2009) EPPO (2020)
-New Jersey	Present			CABI and EPPO (2009) EPPO (2020)
-New York	Present			CABI and EPPO (2009) EPPO (2020)
-North Carolina	Present	Introduced	Invasive	Wood and Bright (1992) Bambara and Casey (2003) CABI and EPPO (2009) EPPO (2020)
-Ohio	Present	Introduced		Lightle et al. (2007) CABI and EPPO (2009) EPPO (2020)
-Oklahoma	Present	Introduced	Invasive	Bambara and Casey (2003) CABI and EPPO (2009) EPPO (2020)
-Oregon	Present	Introduced		CABI (Undated) CABI and EPPO (2009) EPPO (2020)	Original citation: LaBonte et al. (2005)
-South Carolina	Present	Introduced	Invasive	Wood and Bright (1992) Bambara and Casey (2003) CABI and EPPO (2009) EPPO (2020)
-Tennessee	Present	Introduced	Invasive	Oliver and Mannion (2001) Bright and Skidmore (2002) CABI and EPPO (2009) EPPO (2020)
-Texas	Present	Introduced	Invasive	Wood and Bright (1992) Bambara and Casey (2003) CABI and EPPO (2009) EPPO (2020)
-Virginia	Present	Introduced	Invasive	Bambara and Casey (2003) CABI and EPPO (2009) EPPO (2020)
-Washington	Present	Introduced		CABI (Undated) CABI and EPPO (2009) EPPO (2020)	Original citation: LaBonte et al. (2005)
Oceania
Australia	Present			CABI (Undated a) EPPO (2020)	Present based on regional distribution.
-Queensland	Present, Localized			IPPC (2017) EPPO (2020)
New Caledonia	Present	Introduced		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
New Zealand	Present, Localized			EPPO (2020) Brockerhoff et al. (2003) CABI and EPPO (2009)
Palau	Present	Introduced		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
Papua New Guinea	Present	Native		Wood and Bright (1992) CABI and EPPO (2009) EPPO (2020)
Samoa	Present	Introduced	Invasive	Beeson (1929) Beaver (1976) CABI and EPPO (2009) EPPO (2020)
South America
Argentina	Present, Localized			EPPO (2020)
Brazil	Present, Localized			EPPO (2020)
-Amapa	Present			EPPO (2020)
-Pernambuco	Present			EPPO (2020)
-Rio de Janeiro	Present			EPPO (2020)
-Sao Paulo	Present			EPPO (2020)
French Guiana	Present, Localized			EPPO (2020)
Uruguay	Present, Localized			EPPO (2020)

History of Introduction and Spread

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X. crassiusculus was probably introduced to the Afrotropical region from the Oriental region hundreds of years ago by early traders. It has become one of the commonest ambrosia beetles in the rain forest (Schedl, 1963) in both East and West Africa. In North America, it was first discovered near Charleston, South Carolina in 1974 (Anderson, 1974). From there it spread to North Carolina (Hunt, 1979), Louisiana and Florida (Chapin and Oliver, 1986; Deyrup and Atkinson, 1987), and to Mississippi and Texas (Atkinson et al., 1991). Most recently, it has been reported from Tennessee (Oliver and Mannion, 2001). The species is now well-established in south-eastern USA, and may be expected to spread further where climatic conditions are suitable. X.crassiusculus was intercepted in Canada in 1997, but has not established there (Krcmar-Nozic et al., 2000). It seems likely that climatic conditions are too harsh there for the species. In the Hawaiian islands, the species was first found on Hawaii in 1950, and became established on most of the windward islands during the 1950s (Samuelson, 1981). There is no information on when it was introduced to other Pacific Islands. In all cases, the introduction has been accidental.

Risk of Introduction

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Two other species of Xylosandrus, Xylosandrus compactus and Xylosandrus morigerus, with similar habits to X. crassiusculus, 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. crassiusculus must be considered high, most probably in the twigs and small branches of imported plants, although, because it can also breed in fresh timber, other pathways are also possible. 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. crassiusculus is listed as a quarantine pest in New Zealand, but apparently not elsewhere. This should be remedied.

Habitat List

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Category	Sub-Category	Habitat	Presence	Status
Terrestrial

Hosts/Species Affected

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Members of Xyleborus and the related genera Ambrosiodmus, Euwallacea, Xyleborinus and Xylosandrus are all ambrosia beetles that feed and breed in a variety of forest trees and shrubs. Depending on the species, they may be found in small branches and seedlings to large logs. All are potentially damaging to agriculture and/or forestry under suitable conditions. Many species, previously considered of only minor importance, may become important pests in agriculture and forestry as a result of the continuing destruction of natural forests and the expansion of forest and tree crop plantations, agroforestry and agriculture.

X. crassiusculus occurs in a very wide variety of host plants (e.g. Kalshoven, 1959; Browne, 1961; Schedl, 1963; Beaver, 1976; Samuelson, 1981). Schedl (1963) lists 94 species in 28 families in Africa, and 63 species in 34 families outside Africa, and many more species have since been added to this list (Wood and Bright, 1992). It is evident that almost any broad-leaved tree or sapling can be attacked, although the species has not been recorded from conifers. It is particularly important as a pest of crop and ornamental trees. Its attacks are sometimes primary on apparently healthy hosts. It has been recorded killing saplings of forest trees shortly after transplanting. Given the great range of host trees attacked, and the differences between geographical areas, it is not possible to distinguish 'main host' trees from 'other host' trees (see List of hosts). It may be expected that almost any non-coniferous crop, plantation or ornamental tree in a particular area can be attacked. The Host list in this datasheet contains a selection of hosts only.

Host Plants and Other Plants Affected

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Plant name	Family	Context
Acacia koa (koa)	Fabaceae	Wild host
Acacia mangium (brown salwood)	Fabaceae	Other
Albizia	Fabaceae	Other
Albizia lebbeck (Indian siris)	Fabaceae	Other
Artocarpus integer (champedak)	Moraceae	Other
Asimina triloba (Pawpaw-apple)	Annonaceae	Other
Aucoumea klaineana (okoume)	Burseraceae	Other
Bauhinia alba	Fabaceae	Other
Calamus	Arecaceae	Other
Carya illinoinensis (pecan)	Juglandaceae	Other
Castanea mollissima (hairy chestnut)	Fagaceae	Other
Ceiba pentandra (kapok)	Bombacaceae	Other
Cercis canadensis (eastern redbud)	Fabaceae	Other
Cinchona	Rubiaceae	Other
Cinnamomum camphora (camphor laurel)	Lauraceae	Other
Cinnamomum verum (cinnamon)	Lauraceae	Other
Coffea (coffee)	Rubiaceae	Other
Dalbergia sissoo	Fabaceae	Other
Deckenia nobilis	Arecaceae	Wild host
Delonix regia (flamboyant)	Fabaceae	Other
Diospyros kaki (persimmon)	Ebenaceae	Other
Elaeis guineensis (African oil palm)	Arecaceae	Other
Eucalyptus camaldulensis (red gum)	Myrtaceae	Other
Eucalyptus robusta (swamp mahogany)	Myrtaceae	Other
Ficus racemosa (cluster tree)	Moraceae	Other
Grevillea robusta (silky oak)	Proteaceae	Other
Hevea brasiliensis (rubber)	Euphorbiaceae	Other
Khaya ivorensis (African mahogany)	Meliaceae	Other
Laburnum	Fabaceae	Other
Leucaena leucocephala (leucaena)	Fabaceae	Other
Liquidambar styraciflua (Sweet gum)	Hamamelidaceae	Other
Litchi sinensis	Sapindaceae	Other
Macadamia ternifolia (Queensland nut)	Proteaceae	Other
Magnolia	Magnoliaceae	Other
Mangifera indica (mango)	Anacardiaceae	Other
Metrosideros collina	Myrtaceae	Wild host
Milicia excelsa (African teak)	Moraceae	Other
Mimosa (sensitive plants)	Fabaceae	Other
Myristica fragrans (nutmeg)	Myristicaceae	Other
Persea americana (avocado)	Lauraceae	Other
Prunus (stone fruit)	Rosaceae	Other
Prunus persica (peach)	Rosaceae	Other
Saccharum officinarum (sugarcane)	Poaceae	Other
Shorea	Dipterocarpaceae	Wild host
Syzygium aromaticum (clove)	Myrtaceae	Other
Syzygium cumini (black plum)	Myrtaceae	Other
Tectona grandis (teak)	Lamiaceae	Other
Theobroma cacao (cocoa)	Malvaceae	Other
Toona ciliata (toon)	Meliaceae	Other
Ulmus (elms)	Ulmaceae	Wild host

Growth Stages

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Flowering stage, Fruiting stage, Post-harvest, Seedling stage, Vegetative growing stage

Symptoms

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Attacked plants may show signs of wilting, branch die-back, shoot breakage, chronic debilitation, sun-scorch or a general decline in vigour.

List of Symptoms/Signs

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Sign	Life Stages	Type
Growing point / dieback
Stems / lodging; broken stems
Whole plant / wilt

Biology and Ecology

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The important pest species in the genus Xyleborus and the related genera Xylosandrus, Xyleborinus and Euwallacea are all ambrosia beetles in the Xyleborini, a tribe with a social organization of extreme polygamy. The sexual dimorphism is strongly developed, and the ratio of females to males is high. Some species infest small twigs and shoots, others are found in larger branches and poles, while others are found in large timber; others may breed in material of almost any size. In general, most species bore through the bark and into the wood where an enlarged chamber of varying size and shape is constructed. The tunnels into the wood are highly variable in depth and shape, depending on the species involved in the construction. Generally only unhealthy or newly fallen material is infested, but some species are capable of attacking host plants following only a slight set-back, for example, transplanting or temporarily unfavourable conditions such as drought or mechanical injury. A few species have become aggressive under certain conditions, and have thereby attained the status of important pests.

All species of Xyleborus and the related genera are closely associated with ambrosial fungi. Some of these fungi are phytopathogenic and all species of Xyleborus and related genera should be considered to be possible vectors of plant disease.

Some details of the biology of X. crassiusculus are given by Beeson (1930), Browne (1961), Schedl (1963) and Beaver (1976, 1988). The species is known to prefer fresh, moist wood (Beeson, 1930; Beaver, 1988), and attack-densities are usually higher on wood in the shade than in the sun, and higher on the lower side of logs. Stems of fairly small diameter (2.5 - 8 cm) are usually attacked, but sometimes larger logs. The gallery sytem is somewhat variable depending on the size of the stem. In large stems, it branches several times in one transverse plane, and may penetrate 5 cm or more. In small stems, there are fewer branches and one or more may extend along the axis of the stem. In the palm rachis, the galleries run more irregularly through the fibrous tissues. Brood sizes up to 65 have been recorded in the Congo, and up to 100 in Ghana (Schedl, 1963), but usually range between about 10 and 40 (Beaver, 1988). In the tropics, breeding is continuous throughout the year, with overlapping generations, so that the species is present at all times and in all stages of development (Browne, 1968). In south-eastern USA, beetles are active from the beginning of March until autumn, and the life cycle takes about 55 days (Bambara and Casey, 2003).

Notes on Natural Enemies

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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. Schedl (1962) records a species of the curculionid genus Scolytoproctus (Scolytoproctus schaumi) which acts as a nest parasite of X. crassiusculus in the Congo. The female Scolytoproctus forces its way into the beetle gallery, and lays its eggs near the gallery entrance. It is unclear, however, whether the ambrosia beetle is killed by the invader, and whether the ambrosia beetle brood continues to develop normally. Most mortality is probably during the dispersal of the adults, and during gallery establishment. The adults of ambrosia beetles are predated by lizards, clerid beetles and ants as they attempt to bore into the host tree. The adults will also fail to oviposit if the ambrosia fungus fails to establish in the gallery.

Means of Movement and Dispersal

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Natural Dispersal

Adult females fly readily, and flight is one of the main means of movement and dispersal to previously uninfected areas. Of more importance for long distance movement, however, is the transport of infested seedlings, saplings or cut branches. X. crassiusculus usually attacks stems of small diameter (not more than 5 cm diameter), but is sometimes found in larger timber, especially if fresh. Hence it may also be transported in crates or other packing material.

Vector Transmission

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. crassiusculus is a species of Ambrosiella (Kinuura, 1995; Dute et al., 2002). Ambrosiella spp. are not pathogenic, although they do cause staining of the wood around the gallery systems. 'Contamination ' of the mycangia by the spores of pathogenic fungi is possible. Spores of pathogenic fungi can also be transported on the cuticle of the beetle, although their chance of survival there is much less than in the mycangial pouch. There is some evidence for the transmission of wilt fungi by X. crassiusculus (Davis and Dute, 1997). The species has been reported to vector the sap-stain fungus, Botryodiplodia theobromae, into shade trees (Grevillea robusta) in coffee plantations in India (Sreedharan et al., 1991).

Plant Trade

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Plant parts liable to carry the pest in trade/transport	Pest stages	Borne internally	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
Bulbs/Tubers/Corms/Rhizomes
Flowers/Inflorescences/Cones/Calyx
Fruits (inc. pods)
Growing medium accompanying plants
Leaves
Roots
True seeds (inc. grain)

Wood Packaging

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Wood Packaging liable to carry the pest in trade/transport	Timber type	Used as packing
Loose wood packing material		No
Solid wood packing material with bark	Fresh sapwood	Yes
Solid wood packing material without bark	Fresh sapwood	Yes

Wood Packaging not known to carry the pest in trade/transport
Non-wood
Processed or treated wood

Impact Summary

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Category	Impact
Animal/plant collections	Negative
Animal/plant products	None
Biodiversity (generally)	None
Crop production	Negative
Environment (generally)	None
Fisheries / aquaculture	None
Forestry production	Negative
Human health	None
Livestock production	None
Native fauna	None
Native flora	Negative
Rare/protected species	Negative
Tourism	None
Trade/international relations	None
Transport/travel	None

Impact

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Xylosandrus species are known pests of various forest and agricultural plants, and have the potential to transmit pathogenic fungi to their host plants. X. crassiusculus has been recorded killing saplings of forest trees shortly after transplanting. Browne (1968) notes 'devastating' attacks on newly formed plantations of Aucoumea klaineana and Khaya ivoriensis in Ghana. It has also been found in apparently healthy Cinchona trees in Java, Indonesia (Kalshoven, 1959). No known stress factors could be associated with primary attacks on peach orchards in South Carolina, USA (Kovach and Gorsuch, 1985). Atkinson et al. (2000) note large numbers of attacks in Florida, USA on Shumard oak saplings which showed no other symptoms of stress, disease or attack by other insects, and consider that the beetles caused the death of the trees. Atkinson et al. (2000) also noted isolated attacks on large Drake elm saplings. The attacks did not kill the tree directly, but large cankers developed at the site of the attacks, and these sometimes resulted in the death of the trees by girdling. The species can breed successfully in newly sawn timber (Browne, 1961).

Impact: Biodiversity

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Samuelson (1981) notes that in Hawaii, X. crassiusculus has spread into native forest environments, where its hosts include the important endemic trees, Acacia koa and Metrosideros collina. Maeto et al. (1999) noted a large influx of the species from oil palm plantations, where it breeds in fallen palm leaf stalks, into lowland rain forest in Peninsular Malaysia.

Detection and Inspection

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Some success in the detection of the beetle has been obtained by using traps baited with ethanol placed in and around port facilities where infested material may be stored, and around nurseries with plants susceptible to attack. A simple type of trap is described by Bambara et al. (2002). Visual inspection of suspected infested material is required to detect the presence of ambrosia beetles. Infestations are most easily detected by the presence of entry holes made by the attacking beetles, and the presence of frass produced during gallery construction. In X. crassiusculus, the frass is pushed out in the form of a compact cylinder, which may reach a length of 3 to 4 cm before it breaks off, and forms a useful recognition character for the presence of attacks.

Prevention and Control

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Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.

When Xylosandrus species are detected in plant material, it is necessary to immediately destroy all of the infested material. When they are detected in traps, plant material in the vicinity of the trap should be inspected, with special attention directed towards imported woody products such as crating, dunnage and lumber milling scraps. If an active infestation is detected, chemical control using insecticides is not generally effective since the adult beetles bore deep into the host material. The following insecticides were found to be effective against a species of Euwallacea, destructive to tea: fenvalerate, deltamethrin, quinalphos and cypermethrin (Muraleedharan, 1995); these insecticides may also be effective against other ambrosia beetles. Bambara and Casey (2003) suggest the use of permethrin, but multiple treatments may be required during a season. They suggest the use of some attacked trees as trap trees, which need to be removed and burned before the life cycle of the beetle (about 55 days in North Carolina, USA) is completed. The concealed habitats in which these species feed and reproduce, the difficulties and high costs of insecticide application, and environmental concerns, all limit the effectiveness of chemical control. The use of radiation to kill, sterilize or inhibit the emergence of beetles in cut timber (Yoshida et al., 1975), is unlikely to be practical.

References

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Anderson DM, 1974. First record of Xyleborus semiopacus in the continental United States (Coleoptera, Scolytidae). Cooperative Economic Insect Report, 24(45/48):863-864

Atkinson TH, Foltz JL, Wilkinson RC, Mizell RF, 2000. Xylosandrus crassiusculus (Motschulsky) (Insecta: Coleoptera: Scolytidae) Asian ambrosia beetle, granulate ambrosia beetle. Florida Department of Plant Industry Entomology Circular, 310. http://creatures.ifas.ufl.edu/trees/asian_ambrosia_beetle.htm

Atkinson TH, Rabaglia RJ, Peck SB, Foltz JL, 1991. New records of Scolytidae and Platypodidae (Coleoptera) from the United States and the Bahamas. Coleopterists Bulletin, 45(2):152-164

Bambara S, Casey C, 2003. The Asian ambrosia beetle. North Carolina Cooperative Extension Service. http://www.ces.ncsu.edu/depts/ent/notes/O&T/trees/note111/note111.html

Bambara S, Stephan D, Reeves E, 2002. Asian ambrosia beetle trapping. North Carolina Cooperative Extension Service. http://www.ces.ncsu.edu/depts/ent/notes/O&T/trees/note122/note122.html

Beaver RA, 1976. The biology of Samoan bark and ambrosia beetles (Coleoptera, Scolytidae and Platypodidae). Bulletin of Entomological Research, 65(4):531-548

Beaver RA, 1988. Biological studies on ambrosia beetles of the Seychelles (Col., Scolytidae and Platypodidae). Journal of Applied Entomology, 105(1):62-73

Beaver RA, 1989. Insect-fungus relationships in the bark and ambrosia beetles. Insect-fungus interactions. 14th Symposium of the Royal Entomological Society of London in collaboration with the British Mycological Society [edited by Wilding, N.; Collins, N.M.; Hammond, P.M.; Webber, J.F.] London, UK; Academic Press, 121-143

Beaver RA, Browne FG, 1975. The Scolytidae and Platypodidae (Coleoptera) of Thailand. A checklist with biological and zoogeographical notes. Oriental Insects, 9(3):283-311

Beeson CFC, 1929. Platypodidae and Scolytidae. Insects of Samoa, 4:217-248

Beeson CFC, 1930. The biology of the genus Xyleborus, with more new species. Indian Forest Records, 14:209-272

Bright DE, Skidmore RE, 2002. A catalogue of Scolytidae and Platypodidae (Coleoptera), Supplement 2 (1995-1999). Ottawa, Canada: NRC Research Press, 523 pp

Brockerhoff EG, Knizek M, Bain J, 2003. Checklist of Indigenous and Adventive Bark and Ambrosia Beetles (Curculionidae: Scolytinae and Platypodinae) of New Zealand and Interceptions of Exotic Species (1952-2000). New Zealand Entomologist, 26:29-44

Browne FG, 1961. The biology of Malayan Scolytidae and Platypodidae. Malayan Forest Records, 22:1-255

Browne FG, 1968. Pests and diseases of forest plantation trees: an annotated list of the principal species occurring in the British Commonwealth. Oxford, UK: Clarendon Press

CABI/EPPO, 2009. Xylosandrus crassiusculus. [Distribution map]. Distribution Maps of Plant Pests, No.December. Wallingford, UK: CABI, Map 731

Chapin JB, Oliver AD, 1986. New records for Xylosandrus and Xyleborus species (Coleoptera: Scolytidae). Proceedings of the Entomological Society of Washington, 88(4):680-683

Chey VunKhen, 2002. Major insect pests and their management in forest plantations in Sabah, Malaysia. FORSPA Publication, No.30:19-23; 9 ref

Choo HY, Woo KS, Nobuchi A, 1983. A list of the bark and ambrosia beetles injurious to fruit and flowering trees from Korea (Coleoptera: Scolytidae). Korean Journal of Plant Protection, 22(3):171-173

Davis MA, Dute RR, 1997. Fungal associates of the Asian ambrosia beetle, Xylosandrus crassiusculus. Southern Nursery Association Research Conference, 42:106-112

Deyrup MA, Atkinson TH, 1987. New distribution records of Scolytidae from Indiana and Florida. Great Lakes Entomologist, 20:67-68

Dute RR, Miller ME, Davis MA, Woods FM, McLean KS, 2002. Effects of ambrosia beetle attack on Cercis canadensis. IAWA Journal, 23(2):143-160; 36 ref

Eggers H, 1939. Japanische Borkenkäfer II. Arbeiten über Morphologische und Taxonomische Entomologie, Berlin-Dahlem, 6:114-123

EPPO, 2014. PQR database. Paris, France: European and Mediterranean Plant Protection Organization. http://www.eppo.int/DATABASES/pqr/pqr.htm

EPPO, 2018. EPPO Global Database (available online). https://gd.eppo.int

Flechtmann CAH, Atkinson TH, 2016. First records of Xylosandrus crassiusculus (Motschulsky) (Coleoptera: Curculionidae: Scolytinae) from South America, with notes on its distribution and spread in the new world. Coleopterists Bulletin, 70(1):79-83. http://www.bioone.org/loi/cole

GARDNER JCM, 1934, August 15th. Immature Stages of Indian Coleoptera (15) (Scolytidae). Indian Forest Records, 20(pt. 8):17 pp

Horn S, Horn GN, 2006. New host record for the Asian ambrosia beetle, Xylosandrus crassiusculus (Motschulsky) (Coleoptera: Curculionidae). Journal of Entomological Science, 41(1):90-91

Hunt TN, 1979. A scolytid beetle (Xylosandrus crassiusculus) - North Carolina. USDA, Cooperative Plant Pest Report, 4:237

IPPC, 2017. Detection of Xylosandrus crassiculus (Granulate ambrosia beetle) in Queensland. IPPC Official Pest Report, No. AUS-79/1. Rome, Italy: FAO. https://www.ippc.int/

Kalshoven LGE, 1959. Studies on the biology of Indonesian Scolytoidea. 4. Data on the habits of Scolytidae. Second part. Tijdschrift voor Entomologie, 102:135-173

Kanzaki N, Giblin-Davis RM, Gonzalez R, Duncan R, Carrillo D, 2015. Description of Ruehmaphelenchus juliaen. sp. (Tylenchina: Aphelenchoididae) isolated from an ambrosia beetle, xylosandrus crassiusculus (Motschulsky), from South Florida. Nematology, 17(6):639-653. http://booksandjournals.brillonline.com/content/journals/10.1163/15685411-00002896

Khuhro RD, Nizamani SM, Abbasi QD, Rahoo GM, Jiskani MM, 2005. Asian ambrosia beetle: a new insect pest in Sindh Pakistan. Pakistan Journal of Agriculture, Agricultural Engineering, Veterinary Sciences, 21(1):44

Kinuura H, 1995. Symbiotic fungi associated with ambrosia beetles. JARQ, Japan Agricultural Research Quarterly, 29(1):57-63

Kovach J, Gorsuch CS, 1985. Survey of ambrosia beetle species infesting South Carolina peach orchards and a taxonomic key for the most common species. Journal of Agricultural Entomology, 2(3):238-247

Krcmar-Nozic E, Wilson B, Arthur L, 2000. The potential impacts of exotic forest pests in North America: a synthesis of research. Information Report - Pacific Forestry Centre, Canadian Forest Service, No. BC-X-387:ix + 33 pp.; Many ref

LaBonte JR, Mudge AD, Johnson KJR, 2005. Nonindigenous woodboring Coleoptera (Cerambycidae, Curculionidae: Scolytinae) new to Oregon and Washington, 1999-2002: consequences of the intracontinental movement of raw wood products and solid wood packing materials. Proceedings of the Entomological Society of Washington, 107(3):554-564. http://apt.allenpress.com/aptonline/?request=get-archive&issn=0013-8797

LaBonte JR, Mudge AD, Johnson KJR, 2005. Nonindigenous woodboring Coleoptera (Cerambycidae, Curculionidae: Scolytinae) new to Oregon and Washington, 1999-2002: consequences of the intracontinental movement of raw wood products and solid wood packing materials. Proceedings of the Entomological Society of Washington, 107:554-564

Lightle DM, Gandhi KJK, Cognato AI, Mosley BJ, Nielsen DG, Herms DA, 2007. New reports of exotic and native ambrosia and bark beetle species (Coleoptera: Curculionidae: Scolytinae) from Ohio. Great Lakes Entomologist, 40(3/4):194-200

Maeto K, Fukuyama K, Kirton LG, 1999. Edge effects on ambrosia beetle assemblages in a lowland rain forest, bordering oil palm plantations, in Peninsular Malaysia. Journal of Tropical Forest Science, 11(3):537-547; 28 ref

Muraleedharan N, 1995. Strategies for the management of shot-hole borer. Planters' Chronicle, January:23-24

Murayama J, 1934. On the Ipidae (Coleoptera) from Formosa with special reference to their food plants. Journal of the Society of Tropical Agriculture, Taihoku Imperial University, 6:505-512

Murayama J, 1953. The insect fauna of Mt. Ishizuchi and Omogo Valley, Iyo, Japan. The Scolytidae and Platypodidae (Coleoptera). Transactions of the Shikoku Entomological Society, 3:144-166

Oliver JB, Mannion CM, 2001. Ambrosia beetle (Coleoptera: Scolytidae) species attacking chestnut and captured in ethanol-baited traps in middle Tennessee. Environmental Entomology, 30(5):909-918; [Available online at http://www.entsoc.org/pubs/ee/eetocs]; 42 ref

Pelley RH le, 1968. Pests of Coffee. London and Harlow, UK: Longmans, Green and Co Ltd

Pennacchio F, Roversi PF, Francardi V, Gatti E, 2003. Xylosandrus crassiusculus (Motschulsky) a bark beetle new to Europe (Coleoptera Scolytidae). Redia, 86:77-80

Pierce CMF, Gibb TJ, Waltz RD, 2005. Insects and other arthropods of economic importance in Indiana in 2004. Proceedings of the Indiana Academy of Science, 114(2):105-110. http://www.indianaacademyofscience.org

Prathapan, K. D., Hiremath, S. R., 2018. Post-flood outbreak of Xylosandrus crassiusculus and Diuncus corpulentus (Coleoptera: Curculionidae: Scolytinae: Xyleborini) on tree spices in Kerala. Journal of Spices and Aromatic Crops, 27(2), 161-166.

Rabaglia RJ, Valenti MA, 2003. Annotated list of the bark and ambrosia beetles (Coleoptera: Scolytidae) of Delaware, with new distributional records. Proceedings of the Entomological Society of Washington, 105(2):312-319

Rabaglia RJ, Valenti MA, 2003. Annotated list of the bark and ambrosia beetles (Coleoptera: Scolytidae) of Delaware, with new distributional records. Proceedings of the Entomological Society of Washington, 105:312-319

Samuelson GA, 1981. A synopsis of Hawaiian Xyleborini (Coleoptera: Scolytidae). Pacific Insects, 23(1/2):50-92

Schedl KE, 1940. Scolytidae and Platypodidae. 61.contribution. Annals and Magazine of Natural History, series 11, 5:433-442

Schedl KE, 1962. Forstentomologische Beiträge aus dem ehemaligen Belgisch-Congo. Familie Curculionidae. Zeitschrift für Angewandte Entomologie, 50:255-289

Schedl KE, 1963. Scolytidae und Platypodidae Afrikas, Band II. Revista de Entomologia de Moçambique, 5 (1962):1-594

Schedl KE, 1964. Neue und interessante Scolytoidea von den Sunda-Inseln, Neu Guinea und Australien. Tijdschrift voor Entomologie, 107:297-306

Sreedharan K, Balakrishnan MM, Samuel SD, Bhat PK, 1991. A note on the association of wood boring beetles and a fungus with the death of silver oak trees on coffee plantations. Journal of Coffee Research, 21(2):145-148

Wood SL, Bright DE, 1992. A catalog of Scolytidae and Platypodidae (Coleoptera), Part 2: Taxonomic Index Volume A. Great Basin Naturalist Memoirs, 13:1-833

Yin HF, Huang FS, Li ZL, 1984. Coleoptera: Scolytidae. Economic Insect Fauna of China, Fasc. 29. Beijing, China: Science Press, 205 pp

Yoshida T, Fukami J-I, Fukunaga K, Matsuyama A, 1975. Control of the harmful insects in timbers by irradiation: doses required for kill, sterilization and inhibition of emergence in three species of ambrosia beetles (Xyleborini) in Japan. Japanese Journal of Applied Entomology and Zoology, 19:193-202

Distribution References

Bambara S, Casey C, 2003. The Asian ambrosia beetle. In: North Carolina Cooperative Extension Service, http://www.ces.ncsu.edu/depts/ent/notes/O&T/trees/note111/note111.html

Beaver R A, 1976. The biology of Samoan bark and ambrosia beetles (Coleoptera, Scolytidae and Platypodidae). Bulletin of Entomological Research. 65 (4), 531-548. DOI:10.1017/S0007485300006210

Beaver R A, Browne F G, 1975. The Scolytidae and Platypodidae (Coleoptera) of Thailand. A checklist with biological and zoogeographical notes. Oriental Insects. 9 (3), 283-311.

Beeson C F C, 1929. Platypodidae and Scolytidae. Insects of Samoa. 217-248.

Bright DE, Skidmore RE, 2002. A catalogue of Scolytidae and Platypodidae (Coleoptera), Supplement 2 (1995-1999., Ottawa, Canada: NRC Research Press. 523 pp.

Brockerhoff EG, Knizek M, Bain J, 2003. Checklist of Indigenous and Adventive Bark and Ambrosia Beetles (Curculionidae: Scolytinae and Platypodinae) of New Zealand and Interceptions of Exotic Species (1952-2000). In: New Zealand Entomologist, 26 29-44.

Browne F G, 1961. The biology of Malayan Scolytidae and Platypodidae. Malayan Forest Records. xi + 255.

CABI, EPPO, 2009. Xylosandrus crassiusculus. [Distribution map]. In: Distribution Maps of Plant Pests, Wallingford, UK: CABI. Map 731. DOI:10.1079/DMPP/20093321028

CABI, Undated. Compendium record. Wallingford, UK: CABI

CABI, Undated a. CABI Compendium: Status inferred from regional distribution. Wallingford, UK: CABI

CABI, Undated b. CABI Compendium: Status as determined by CABI editor. Wallingford, UK: CABI

Chey VunKhen, 2002. Major insect pests and their management in forest plantations in Sabah, Malaysia. In: FORSPA Publication. [ed. by Hutacharern C, Napompeth B, Allard G, Wylie F R]. Bangkok, Thailand: FAO Regional Office for Asia and the Pacific. 19-23.

Choo H Y, Woo K S, Nobuchi A, 1983. A list of the bark and ambrosia beetles injurious to fruit and flowering trees from Korea (Coleoptera: Scolytidae). Korean Journal of Plant Protection. 22 (3), 171-173.

Eggers H, 1939. (Japanische Borkenkäfer II. Arbeiten über Morphologische und Taxonomische Entomologie)., 6 Berlin-Dahlem, 114-123.

EPPO, 2018. EPPO Global database. In: EPPO Global database. Paris, France: EPPO. https://gd.eppo.int/

EPPO, 2020. EPPO Global database. In: EPPO Global database, Paris, France: EPPO. https://gd.eppo.int/

Francardi V, Noal A, Francescato S, Pinto R, Bruni A, Loffredi L, Bucini D, Guarnieri D, Bellantuono M, Esposito N, Nuccitelli L, Binazzi F, Vitale S, Giambattista G di, Roversi P F, Pennacchio F, 2017. Coexistence of Xylosandrus crassiusculus (Motschulsky) and X. compactus (Eichhoff) (Coleoptera Curculionidae Scolytinae) in the National Park of Circeo (Lazio, Italy). Redia. 149-155. http://www.redia.it/images/stories/Online%20first%20articles/19%20Francardi%20Noal%20et%20al__.pdf

Horn S, Horn G N, 2006. New host record for the Asian ambrosia beetle, Xylosandrus crassiusculus (Motschulsky) (Coleoptera: Curculionidae). Journal of Entomological Science. 41 (1), 90-91.

IPPC, 2017. Detection of Xylosandrus crassiculus (Granulate ambrosia beetle) in Queensland. In: IPPC Official Pest Report, No. AUS-79/1, Rome, Italy: FAO. https://www.ippc.int/

Ito M, Kajimura H, 2009. Phylogeography of an ambrosia beetle, Xylosandrus crassiusculus (Motschulsky) (Coleoptera: Curculionidae: Scolytinae), in Japan. Applied Entomology and Zoology. 44 (4), 549-559. DOI:10.1303/aez.2009.549

Kalshoven LGE, 1959. Studies on the biology of Indonesian Scolytoidea. 4. Data on the habits of Scolytidae. Second part. In: Tijdschrift voor Entomologie, 102 135-173.

Kavčič A, 2018. First record of the Asian ambrosia beetle, Xylosandrus crassiusculus (Motschulsky) (Coleoptera: Curculionidae, Scolytinae), in Slovenia. Zootaxa. 4483 (1), 191-193. DOI:10.11646/zootaxa.4483.1.9

Kavčič A, 2019. Finding of a new bark beetle species in Slovenia. (Najdba nove vrste podlubnika v Sloveniji.). In: Zbornik predavanj in referatov, 14. slovensko posvetovanje o varstvu rastlin z mednarodno udeležbo, 5.-6. marec 2019, Maribor, Slovenija [14. slovensko posvetovanje o varstvu rastlin z mednarodno udelezbo, 5.-6. marec 2019, Maribor, Slovenija.], Ljubljana, Slovenia: Društvo za varstvo rastlin Slovenije. 106-112.

Keshavareddy G, Abraham Verghese, Kumar H R M, 2008. Spatial distribution of the damage of shot hole borer (Coleoptera: Scolytidae) on grapes. Current Biotica. 2 (1), 18-31. http://www.currentbiotica.com/Images/Journals-IssueII/CB_2_1_3f.pdf

Khuhro R D, Nizamani S M, Abbasi Q D, Rahoo G M, Jiskani M M, 2005. Asian ambrosia beetle: a new insect pest in Sindh Pakistan. Pakistan Journal of Agriculture, Agricultural Engineering, Veterinary Sciences. 21 (1), 44.

Lightle D M, Gandhi K J K, Cognato A I, Mosley B J, Nielsen D G, Herms D A, 2007. New reports of exotic and native ambrosia and bark beetle species (Coleoptera: Curculionidae: Scolytinae) from Ohio. Great Lakes Entomologist. 40 (3/4), 194-200.

MURAYAMA J, 1934. On the Ipidae (Coleoptera) from Formosa with special References to their Food Plants. Journal of the Society of Tropical Agriculture. 6 (3), 505-512.

Murayama J, 1953. The insect fauna of Mt. Ishizuchi and Omogo Valley, Iyo, Japan. The Scolytidae and Platypodidae (Coleoptera). Transactions of the Shikoku Entomological Society. 144-166.

Oliver J B, Mannion C M, 2001. Ambrosia beetle (Coleoptera: Scolytidae) species attacking chestnut and captured in ethanol-baited traps in middle Tennessee. Environmental Entomology. 30 (5), 909-918. DOI:10.1603/0046-225X-30.5.909

Pennacchio F, Roversi P F, Francardi V, Gatti E, 2003. Xylosandrus crassiusculus (Motschulsky) a bark beetle new to Europe (Coleoptera Scolytidae). Redia. 77-80.

Pierce C M F, Gibb T J, Waltz R D, 2005. Insects and other arthropods of economic importance in Indiana in 2004. Proceedings of the Indiana Academy of Science. 114 (2), 105-110. http://www.indianaacademyofscience.org

Prathapan K D, Hiremath S R, 2018. Post-flood outbreak of Xylosandrus crassiusculus and Diuncus corpulentus (Coleoptera: Curculionidae: Scolytinae: Xyleborini) on tree spices in Kerala. Journal of Spices and Aromatic Crops. 27 (2), 161-166.

Schedl K E, 1940. Scolytidae and Platypodidae. 61.contribution. Annals and Magazine of Natural History, series 11. 433-442.

Schedl KE, 1964. (Neue und interessante Scolytoidea von den Sunda-Inseln, Neu Guinea und Australien). In: Tijdschrift voor Entomologie, 107 297-306.

Sreedharan K, Balakrishnan M M, Samuel S D, Bhat P K, 1991. A note on the association of wood boring beetles and a fungus with the death of silver oak trees on coffee plantations. Journal of Coffee Research. 21 (2), 145-148.

Wood SL, Bright DE, 1992. A catalog of Scolytidae and Platypodidae (Coleoptera), Part 2: Taxonomic Index Volume A. In: Great Basin Naturalist Memoirs, 13 1-833.

Yin HF, Huang FS, Li ZL, 1984. Coleoptera: Scolytidae. In: Economic Insect Fauna of China Fasc. 29, Beijing, China: Science Press. 205 pp.

Links to Websites

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Website	URL	Comment
GISD/IASPMR: Invasive Alien Species Pathway Management Resource and DAISIE European Invasive Alien Species Gateway	https://doi.org/10.5061/dryad.m93f6	Data source for updated system data added to species habitat list.

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Xylosandrus crassiusculus (Asian ambrosia beetle)

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