North American Xyleborini north of Mexico: a review and key to genera and species (Coleoptera, Curculionidae, Scolytinae).

Bark and ambrosia beetles (Scolytinae) are the most successful group of invasive wood borers worldwide, and the most invasive among them are species in the tribe Xyleborini. This haplodiploid, highly inbred, fungus-farming group is represented by 30 non-native species in North America, of which at least five are serious pests. The few identification resources for Xyleborini that exist are becoming outdated due to new species arrivals and nomenclatural changes. Here we present a new comprehensive key to Xyleborini currently known from the continental United States. Compared to the previous key, the following species have been added to the North American fauna: Ambrosiodmus minor (Stebbing), Ambrosiophilus nodulosus (Eggers), Anisandrus maiche Kurentsov, Coptoborus pseudotenuis (Schedl), Cyclorhipidion fukiense (Eggers), Dryocoetoides reticulatus Atkinson, Dryoxylon onoharaense (Murayama), Euwallacea interjectus (Blandford), Xyleborinus andrewesi (Blandford), Xyleborinus artestriatus (Eichhoff), Xyleborinus octiesdentatus (Murayama), Xyleborus bispinatus Eichhoff, Xyleborus seriatus Blandford, Xyleborus spinulosus Blandford, and Xylosandrus amputatus (Blandford).

Introduction

Bark and ambrosia beetles (: ) are considered one of the most injurious groups of insects in native and planted forests (Raffa et al. 2015). The vast majority breeds in dead or dying tissues and do not have economic impact. However, some species attack living trees, seedlings, or seeds of commercial importance causing severe damage (Raffa et al. 2015). Scolytines are among the most commonly intercepted taxa at United States ports of entry. True bark beetles (phloeophagous species) are intercepted more often than ambrosia-feeding species; however, ambrosia beetles of the tribe represent half of the 60 non-native scolytines established in the United States (Haack and Rabaglia 2013).

The , with 1177 recognized species, is the most species-rich tribe within (Smith and Hulcr 2015). This tribe of ambrosia beetles also includes some of the most abundant and widely distributed species (Rabaglia et al. 2006). The combination of fungus-farming, wide host range, and arrhenotokous inbreeding (haplodiploidy) makes the one of the most successful groups of colonizers in the world (Atkinson et al. 1990, Smith and Hulcr 2015).

In the last decade, several exotic species of have successfully established in North America. The detection and control of both native and exotic species relies on a solid understanding of the systematics and identity of species. Since the last review of North American (Rabaglia et al. 2006), 15 additional non-native species have been recorded in North America and several nomenclatural changes have been made. The aim of this article is to review the species of occurring in continental North America, diagnose the new species for the region, and provide illustrated keys to genera and species.

Materials and methods

Specimens examined were from the cryo-preserved collection University of Florida Forest Entomology lab managed by JH (University of Florida, Gainesville, Florida, USA), the Florida State Collection of Arthropods (Gainesville, Florida, USA), and/or collected by the authors during various state, regional, and national surveys. Distribution records are as reported in Wood and Bright (1992), Atkinson (2017), recent publications, and unpublished data from the authors. Diagnostic characters used in the keys and notes are for the identification of genera and species occurring in North America and may not be useful for taxa occurring in other regions. Antennal club types are described as Hulcr et al. (2007). Interstria 1 is defined as the sutural interstria. Table 1 includes the complete list of species of occurring in continental North America.

Table 1.

List of species of occurring in continental North America north of Mexico.

Ambrosiodmus devexulus (Wood, 1978)	Xyleborinus artestriatus (Eichhoff, 1878)
Ambrosiodmus lecontei Hopkins, 1915	Xyleborinus attenuatus (Blandford, 1894)
Ambrosiodmus lewisi (Blandford, 1894)	Xyleborinus gracilis (Eichhoff, 1868)
Ambrosiodmus minor (Stebbing, 1909)	Xyleborinus octiesdentatus (Murayama, 1931)
Ambrosiodmus obliquus (LeConte, 1878)	Xyleborinus saxesenii (Ratzeburg, 1837)
Ambrosiodmus opimus (Wood, 1974)	Xyleborus affinis Eichhoff, 1868
Ambrosiodmus rubricollis (Eichhoff, 1875)	Xyleborus bispinatus Eichhoff, 1868
Ambrosiodmus tachygraphus (Zimmermann, 1868)	Xyleborus celsus Eichhoff, 1868
Ambrosiophilus atratus (Eichhoff, 1875)	Xyleborus ferrugineus (Fabricius, 1801)
Ambrosiophilus nodulosus (Eggers, 1941)	Xyleborus glabratus Eichhoff, 1877
Anisandrus dispar (Fabricius, 1792)	Xyleborus horridus Eichhoff, 1869
Anisandrus maiche Kurentsov, 1941	Xyleborus impressus Eichhoff, 1868
Anisandrus obesus (LeConte, 1868)	Xyleborus intrusus Blandford, 1898
Anisandrus sayi Hopkins, 1915	Xyleborus pfeilii (Ratzeburg, 1837)
Cnestus mutilatus (Blandford, 1894)	Xyleborus planicollis Zimmermann, 1868
Coptoborus pseudotenuis (Schedl, 1936)	Xyleborus pubescens Zimmermann, 1868
Cyclorhipidion bodoanum (Reitter, 1913)	Xyleborus seriatus Blandford, 1894
Cyclorhipidion fukiense (Eggers, 1941)	Xyleborus spinulosus Blandford, 1898
Cyclorhipidion pelliculosum (Eichhoff, 1878)	Xyleborus viduus Eichhoff, 1878
Dryocoetoides reticulatus Atkinson, 2009	Xyleborus volvulus (Fabricius, 1775)
Dryoxylon onoharaense (Murayama, 1934)	Xyleborus xylographus (Say, 1826)
Euwallacea fornicatus (Eichhoff, 1868)	Xylosandrus amputatus (Blandford, 1894)
Euwallacea interjectus (Blandford, 1894)	Xylosandrus compactus (Eichhoff, 1875)
Euwallacea similis (Ferrari, 1867)	Xylosandrus crassiusculus (Motschulsky, 1866)
Euwallacea validus (Eichhoff, 1875)	Xylosandrus curtulus (Eichhoff, 1869)
Theoborus ricini (Eggers, 1932)	Xylosandrus germanus (Blandford, 1894)
Xyleborinus andrewesi (Blandford, 1896)

Synonyms listed for each genus and species are cited from Alonso-Zarazaga and Lyal (2009), Alonso-Zarazaga et al. (2017), Beaver 2011, Bright (2014), and Wood and Bright (1992). References to original descriptions and synonymies are cited from Bright and Rabaglia (1999), Hulcr and Cognato (2009), Wood and Bright (1987, 1992), and Wood (1986). The type material collection information and repository correspond to Wood and Bright (1992). Abbreviations for location of type material are:

British Museum of Natural History, London;

Canadian National Collection of Insects, Ottawa;

Forest Research Institute, Dehradun;

Institut Royal des Sciences Naturelles de Belgique, Brussels;

Institute of Zoology at Moscow, Moscow;

Museum of Comparative Zoology, Cambridge, MA;

Natural History Museum Budapest, Budapest;

Naturhistorisches Museum Wien, Wien;

National Museum of Natural History, Washington, DC;

Senckenberg Deutsches Entomologisches Institut, Müncheberg;

Universitets Zoologisk Museum, Copenhagen;

Zoological Institute of the Russian Academy of Sciences, St. Petersburg;

Zoological Research Museum Alexander Koenig, Bonn; and

Zoologisches Institut und Zoologisches Museum, Hamburg.

Photographs were taken by JH and DG using an Olympus SZX16 stereomicroscope. Each image is a composite of up to 50 separate images taken with a Canon EOS Rebel T3i camera, and later stacked using the Helicon Focus software (v 6.0, Helicon Soft).

Antennal club types in . First row, types of antennae; second row, examples of variation; third row, rear face; fourth row, lateral view. Modified from Hulcr et al. (2007).

Systematics

Key to genera of female American north of Mexico

Hopkins, 1915

Motschulsky, 1863. Synonymy Wood 1966.

Nunberg, 1963. Synonymy Wood 1980.

Type species.

Zimmermann.

Species of differ from other members of the tribe by the asperities covering the entire surface of the pronotum.

(Wood, 1978)

Fig. 2

Figure 2.

Lateral and dorsal views of species. From top left, , , , and . Scale bar: 1.0 mm.

Wood, 1977. Preoccupied Schedl 1977.

Wood, 1978. Replacement name for

Schedl, 1979. Unnecessary replacement name.

Type material.

Holotype female; Homestead, FL; NMNH.

Distribution.

North America: Antilles, United States: Florida.

Notes.

This species is very similar to , but it is distinguished by its smaller size, lack of declivital granules, and interstriae 1 not elevated. It is only known from southern Florida, Puerto Rico, and the Dominican Republic.

Hopkins, 1915

Fig. 2

Eggers, 1931. Synonymy Wood 1972.

Holotype female; Keene, FL; USNM.

North America: Antilles, United States: Alabama, Florida, Louisiana, South Carolina, Texas; South America: Brazil.

In North America, this species is distinguished by the smaller size and the much deeper, coarser strial punctures compared to .

(Blandford, 1894)

Fig. 2

Blandford, 1894.

Strohmeyer, 1912. Synonymy Beaver and Liu 2010.

Eggers, 1923. Synonymy Schedl 1950.

Eggers, 1923. Synonymy

Syntypes female; Japan; BMNH.

Asia; North America (introduced): United States: Alabama, Georgia, Pennsylvania.

was first reported in North America from southeastern Pennsylvania (Hoebeke 1991). This non-native species is the largest in North America, and can be distinguished from by the tubercles on declivital interstriae 2, which are distinctly larger than those on other interstriae.

(Stebbing, 1909)

Fig. 2

Stebbing, 1909.

Hagedorn, 1910. Synonymy Schedl 1962.

Syntypes female; Assam: labeled Kochujan, printed as Goalpara Sal Forests; FRI.

Asia; North America (introduced): United States: Alabama, Florida, Georgia, Mississippi.

The first collection in North America of was in Florida in 2011 (Rabaglia and Okins 2011). Similar to but with tubercles on declivital interstriae 2 not distinctly larger than those on other interstriae.

(LeConte, 1878)

Fig. 3

Figure 3.

Lateral and dorsal views of species. From top left, , , , and . Scale bar: 1.0 mm.

LeConte, 1878.

Blandford, 1898. Synonymy Wood 1975.

Hopkins, 1915. Synonymy

Eggers, 1928. Synonymy Wood 1975.

Eggers, 1931. Synonymy Wood 1972.

Eggers, 1941. Synonymy Bright 1985.

Schedl, 1941. Synonymy Wood 1975.

Schedl, 1978. Synonymy

Holotype female; Enterprise, FL; MCZ.

Africa; Central America: Costa Rica, Guatemala, Honduras, Panama; North America: Antilles Islands, Mexico, United States: Alabama, Delaware, District of Columbia, Florida, Georgia, Louisiana, Maryland, Mississippi, North Carolina, South Carolina, Tennessee, Texas, Virginia; South America: Brazil, Colombia, Ecuador, Peru.

Similar to but with less prominent punctures.

(Wood, 1974)

Fig. 3

Wood, 1974.

Holotype female; Sebring, FL; NMNH.

North America: United States: Florida; South America: Brazil.

Similar to in North America, but interstriae 1 armed by several fine granules in .

(Eichhoff, 1875)

Fig. 3

Eichhoff, 1875.

Schedl, 1952. Synonymy

Schedl, 1975. Synonymy

Holotype Female; Japan; IRSNB.

Asia; Australia (introduced); Europe (introduced): Italy; North America (introduced): Mexico, United States: Alabama, Arkansas, Connecticut, Delaware, Florida, Georgia, Indiana, Louisiana, Maryland, Michigan, Mississippi, Missouri, North Carolina, Ohio, Oklahoma, Pennsylvania, South Carolina, Tennessee, Texas, Virginia.

This non-native species, first found in Maryland (Bright 1968), is now well-established and commonly found in the mid-Atlantic and southeastern states. This species is distinguished from other by the combination of the red color, the small size, and the equally granulate declivital interstriae.

(Zimmermann, 1868)

Fig. 3

Zimmermann, 1868.

Holotype female; North Carolina; MCZ.

North America: United States: Alabama, Arkansas, Delaware, District of Columbia, Florida, Georgia, Illinois, Indiana, Kentucky, Louisiana, Maryland, Mississippi, New Jersey, North Carolina, Ohio, Oklahoma, Pennsylvania, South Carolina, Tennessee, Texas, Virginia, West Virginia.

Widely distributed in the eastern United States. It is among the largest species of in North America.

Hulcr & Cognato, 2009

(Schedl).

Species of differ from other members of the tribe by the black and robust body combined with the absence of asperities on a flat pronotal disc, and the rounded edge of elytral declivity.

(Eichhoff, 1875)

Fig. 4

Figure 4.

Lateral and dorsal views of species. From top left, and . Scale bar: 1.0 mm.

Eichhoff, 1875.

Holotype female; Japan. ZMUH, lost.

Asia; North America (introduced): United States: Alabama, Delaware, Florida, Georgia, Kansas, Kentucky, Louisiana, Maine, Maryland, Michigan, Mississippi, Missouri, North Carolina, Ohio, Oklahoma, Pennsylvania, South Carolina, Tennessee, Texas, West Virginia; Oceania.

was first reported in eastern North America from Georgia, Maryland, Tennessee, Virginia and West Virginia (Atkinson et al. 1990). Differs from by the absence of tubercles on the declivity.

(Eggers, 1941)

Fig. 4

Eggers, 1941.

Schedl, 1957. Synonymy

Smith & Cognato, 2015. Synonymy

Holotype female; Fukien [Fujian Province, China]; ZMFK.

Asia; North America (introduced): United States: Georgia.

A recent introduction in the U.S. (Smith and Cognato 2015, Smith et al. 2017), is likely to expand its distribution. Differs from by its smaller size and by the presence of evenly spaced tubercles on the declivity.

Ferrari, 1867

(Fabricius).

Species of differ from other members of the tribe by the combination of serrations on the frontal edge of pronotum, a tuft of setae at the base of the pronotum, the contiguous procoxae, and an obliquely truncate antennal club with the first segment of club covering the entire posterior side.

(Fabricius, 1792)

Fig. 5

Figure 5.

Lateral and dorsal views of species. From top left, , , , and . Scale bar: 1.0 mm.

Fabricius, 1792.

Panzer, 1793. Synonymy Eichhoff 1878.

Panzer, 1793. Synonymy Hagedorn 1910.

Peck, 1817. Synonymy Hubbard 1897.

Sahlberg, 1834. Synonymy Eichhoff 1878.

Kolenati, 1846. Synonymy Ferrari 1867.

Reitter, 1913. Synonymy Mandelshtam 2001.

Drake, 1921. Synonymy Wood 1957.

Eggers 1922.

Eggers, 1937. Synonymy Schedl 1964.

Murayama, 1943. Synonymy Knížek 2011.

Syntypes female; Germaniae; UZMC.

Asia; Europe; North America (introduced): Canada: British Columbia, New Brunswick, Nova Scotia, Ontario; United States: California, District of Columbia, Idaho, Illinois, Indiana, Maine, Maryland, Massachusetts, Michigan, New Jersey, New York, North Carolina, Ohio, Oregon, Pennsylvania, Rhode Island, Utah, Virginia, Washington, West Virginia.

Representing the first non-native scolytine reported in North America (Rabaglia et al. 2006), was likely unintentionally introduced before 1817 (Wood 1977). Found across North America from southern Canada through northern United States. Similar to but larger.

Kurentsov, 1941

Fig. 5

Kurentsov, 1941.

Syntypes female; Ussuri, USSR [Russia]; IZL [ZIN], Leningrad [St. Petersburg].

Asia; North America (introduced): United States: Ohio, Pennsylvania, West Virginia, and Wisconsin.

was first reported in the US from Pennsylvania, Ohio, and West Virginia (Rabaglia et al. 2009). Similar to but smaller. This non-native species was originally reported from western Pennsylvania and eastern Ohio, but is becoming increasingly common in northeastern states.

(LeConte, 1868)

Fig. 5

LeConte, 1868.

Swaine, 1910. Synonymy Hopkins 1915.

Swaine, 1917. Synonymy Schedl 1964.

Lectotype female; Virginia; MCZ.

North America: Canada: New Brunswick, Ontario, Quebec; United States: Connecticut, Illinois, Kentucky, Massachusetts, Michigan, Minnesota, New Jersey, New York, Ohio, Virginia, West Virginia, Wisconsin.

Distinguished from other by the presence of a series of tubercles on the posterolateral margin of the declivity.

Hopkins, 1915

Fig. 5

Swaine, 1910. Synonymy Wood 1957.

Schedl, 1950. Synonymy Wood 1957.

Holotype female; Morgantown, WV; NMNH.

North America: Canada: New Brunswick, Ontario, Quebec; United States: Alabama, Connecticut, Delaware, District of Columbia, Georgia, Illinois, Indiana, Iowa, Kentucky, Maine, Maryland, Massachusetts, Michigan, Mississippi, Missouri, New Jersey, New York, North Carolina, Ohio, Pennsylvania, South Carolina, Tennessee, Texas, Virginia, West Virginia.

This is the most common species of in the northeastern U.S. Distinguished from other by the absence of significant sculpture on the elytral declivity. Wood (1957) synonymized with , but Swaine’s name is available and should have priority.

Sampson, 1911

Murayama, 1950. Synonymy Browne 1955.

Sampson

Species of differ from other members of the tribe by the truncate elytra, which are shorter than the pronotum.

(Blandford, 1894)

Fig. 6

Figure 6.

Lateral and dorsal view of . Scale bar" 1.0 mm.

Blandford, 1894.

Eggers, 1930. Synonymy

Schedl, 1939. Synonymy Kalshoven 1960.

Eggers, 1939. Synonymy

Holotype female; Japan; BMNH.

Asia; North America (introduced): United States: Florida, Georgia, Kentucky, Louisiana, Mississippi, Pennsylvania, South Carolina, Tennessee, Texas; Oceania.

was first collected in North America from Mississippi in 1999 (Schiefer and Bright 2004). This species is easily distinguished from other by the truncate and short elytra, with a circular declivity delimited by a distinct carina posteriorly and laterally.

Hopkins, 1915

Hopkins

Species of differ from other members of the tribe by the type 3 antennal club, the light brown or yellowish color, and the narrowed or acuminate elytral apex.

(Schedl, 1936)

Fig. 7

Figure 7.

Lateral and dorsal view of . Scale bar: 1.0 mm.

Schedl, 1936.

Schedl, 1948. Synonymy Wood 1976.

Holotype female; Brasilien; Schedl Collection in NHMW.

Central America: Costa Rica, Panama; North America: Mexico, United States: Florida; South America: Bolivia, Brazil, Ecuador, French Guiana, Peru, Venezuela.

was first documented in the US based on a reared specimen from southern Florida in 2004 (Atkinson et al. 2010). Common in South America, either introduced or naturally spread to Florida. Distinguished from other by the slightly impressed interstria 2, 1 and 3 with 3–5 small denticles, and by the elevated apical margin of interstriae 1 and 2.

Hagedorn, 1912

Hopkins, 1915. Synonymy

Nunberg, 1961. Synonymy

Hagedorn

Species of differ from other members of the tribe by being overall pubescent and covered with minute, dense, confused punctures.

(Reitter, 1913)

Fig. 8

Figure 8.

Lateral and dorsal views of species. From top left, , , and . Scale bar: 1.0 mm.

Reitter, 1913.

Kurentsov, 1948. Synonymy Mandelshtam 2001.

Wood, 1975. Synonymy Knížek 2011.

Syntypes female; Ostsibirien: Sotka-gora; NHMB.

Asia; North America (introduced): United States: Alabama, Arkansas, California, Delaware, Florida, Georgia, Kansas, Louisiana, Maryland, Michigan, Mississippi, Missouri, North Carolina, Ohio, Oklahoma, Oregon, South Carolina, Tennessee, Texas, Washington.

was first reported in the eastern US (Maryland, Delaware, South Carolina, and Arkansas) in 2000 (Vandenberg et al. 2000) but was originally known by its synonym Wood. Distinguished from other in North America by the size and the yellowish brown color.

(Eggers, 1941)

Fig. 8

Eggers, 1941.

Murayama, 1952. Synonymy

Schedl, 1953. Synonymy Beaver and Liu 2010.

Holotype female; Fukien [Fujian Province, China]; ZMFK.

Asia; North America (introduced): United States: Florida, Georgia.

This recently detected non-native species is very similar in general appearance to both and except for body length, with an intermediate size (Hoebeke et al. 2018).

(Eichhoff, 1878)

Fig. 8

Eichhoff, 1878.

Murayama, 1930. Synonymy Knížek 2011.

Kurenzov, 1948. Synonymy Knížek 2011.

Nunberg, 1956. Synonymy Knížek 2011.

Syntypes female; Japan; ZMUH, lost.

Asia; North America (introduced): United States: Delaware, Illinois, Kentucky, Maine, Maryland, Massachusetts, Missouri, New Jersey, North Carolina, Ohio, Pennsylvania, Rhode Island, Tennessee, Virginia.

was first documented in the US from Pennsylvania in 1987 and from Maryland in 1989 (Atkinson et al. 1990). Distinguished from other in North America by the larger size and the blackish brown color.

Hopkins, 1915

Hopkins.

Species of differ from other members of the tribe by the stick-like protibia, the posterior face of which is rugose.

Atkinson, 2009

Fig. 9

Figure 9.

Lateral and dorsal view of . Scale bar: 1.0 mm.

Holotype female; United States; NMNH.

North America: United States: Florida.

Distinguished from other by the clearly indicated punctures in the declivital striae, the uniseriate tubercles in the interstriae, and the dull declivity. Only known from south Florida (Atkinson 2009).

Bright & Rabaglia, 1999

Murayama.

Species of differ from other members of the tribe by the long body, the not impressed submentum, the narrow protibia with a few large teeth on outer margin, and by the deeply concave elytral declivity.

(Murayama, 1934)

Fig. 10

Figure 10.

Lateral and dorsal view of . Scale bar: 1.0 mm.

Murayama, 1934.

Bright & Rabaglia, 1999 (incorrect subsequent spelling).

: Alonso-Zarazaga & Lyal, 2009. Correction for

Lectotype female; Japan; NMNH.

Asia; North America (introduced): United States: Alabama, Arkansas, Delaware, Florida, Georgia, Louisiana, Maryland, Mississippi, North Carolina, Ohio, South Carolina, Tennessee, Texas, Virginia.

This is the only species of the genus. Bright and Rabaglia (1999) placed it in the based on tibial characters, but molecular analyses place it within the (Jordal et al. 2000, Jordal 2002). Distinguished by the obliquely truncate antennal club, the narrow protibiae with a few large teeth on outer margin, and by the distinctly concave, densely pubescent, and unarmed elytral declivity.

Hopkins, 1915

Blandford.

Species of differ from other members of the tribe by the costate and broad posterolateral edge of declivity. In most species the pronotum is subquadrate.

(Eichhoff, 1868)

Fig. 11

Figure 11.

Lateral and dorsal views of species. From top left, , , and . Scale bar: 1.0 mm.

Eichhoff, 1868.

Eggers, 1923. Synonymy Beeson 1930 (as variety).

Schedl, 1942. Synonymy

Schedl, 1951. Synonymy Wood,1989.

Schedl, 1951. Synonymy Beaver 1991.

Schedl, 1951. Synonymy

Syntypes: Ceylon; ZMUH, lost.

Africa; Asia; Central America (introduced): Costa Rica, Panama; North America (introduced): Mexico, United States: California, Florida, Hawaii; Oceania (introduced); South America (introduced): Brazil.

This species is a complex of several distinct genotypes, the most common of which are known as the Tea shot hole borer, Polyphagous shot hole borer, and the Kuroshio shot hole borer (Stouthamer et al. 2017). The different lineages are supported by rapidly evolving mitochondrial genes and more conserved nuclear gene regions. Although these potential different species display morphological differences, reliable morphological diagnosis has not been established (Chen et al. 2016).

(Blandford, 1894)

Fig. 11

Blandford, 1894.

Eggers, 1925. Synonymy Schedl 1958.

Holotype female; Japan, China [presumably syntypes]; BMNH.

Asia; North America (introduced): United States: Florida, Georgia, Hawaii, Kentucky, Louisiana, South Carolina, Texas, Virginia.

The first American occurrence of this species was in Louisiana in 1984, originally confused with (Cognato et al. 2015). Specimens from Asia can be larger in size, up to 3.8 mm long, overlapping with body size.

(Ferrari, 1867)

Fig. 11

Boheman, 1858. Synonymy Schedl 1960.

Ferrari, 1867.

Eichhoff, 1868. Synonymy Schedl 1959.

Eichhoff, 1876. Synonymy Schedl 1959.

Blandford, 1896. Synonymy Eggers 1929.

Schaufuss, 1897. Synonymy Schedl 1959.

Schaufuss, 1897. Synonymy Schedl 1959.

Schedl, 1936. Synonymy

Schedl, 1953. Synonymy

Holotype female; “Insula Keeling”. NHMW.

Africa; Asia; North America (introduced): United States: Texas; Oceania; South America (introduced): Brazil.

The designation of Motschulsky, 1863 as a synonym of this species (Mandelshtam and Nikitskij 2010) is not considered valid (Alonso-Zarazaga pers. comm.). Wood designated a specimen of as the lectotype of , not a specimen of (although they occurred on the same pin).

(Eichhoff, 1875)

Fig. 11

Eichhoff, 1875.

Syntypes female; Japan; IRSNB.

Asia; North America (introduced): Canada: Ontario; United States: Alabama, Delaware, Georgia, Kentucky, Maryland, Michigan, Mississippi, New Jersey, New York, North Carolina, Ohio, Pennsylvania, South Carolina, Tennessee, Texas, Virginia, West Virginia.

was first collected in the US from Nassau County, New York in 1975 (Wood 1977) and later from Pennsylvania in 1980 (Wood 1982) and Louisiana in 1984 (Chapin and Oliver 1986). This species is distinguished from in North America by the larger size, the absence of tubercles from the apical half of the interstriae 2, and by uneven and tuberculate declivital costae.

Hopkins, 1915

Hopkins.

Species of differ from other members of the tribe by the light-brown color, the type 3 antennal club, the pointed elytral declivity apex in dorsal view, and the smooth posterior face of protibia.

(Eggers, 1932)

Fig. 12

Figure 12.

Lateral and dorsal view of . Scale bar: 1.0 mm.

Eggers, 1932.

Schedl, 1954. Synonymy

Holotype female; “Congostaat”; NMNH.

Africa (introduced); Central America: Costa Rica; North America: Antilles, Mexico, United States: Florida; South America: Brazil, Colombia, Venezuela.

It is unclear if this species was introduced from South America or is native to North America. Distinguished by the light-brown color, the short and steep elytral declivity with stout and short interstrial setae, and the smooth posterior face of protibia.

Reitter, 1913

Ratzeburg.

Species of differ from other members of the tribe by the conical scutellum surrounded by setae.

(Blandford, 1896)

Fig. 13

Figure 13.

Lateral and dorsal views of species. From top left, , , and . Scale bar: 1.0 mm.

Blandford, 1896.

Schedl, 1970. Synonymy Beaver and Brownie 1978.

Bright, 1972. Synonymy Bright 1985.

Browne, 1984. Synonymy Beaver 1995.

Holotype female; India; BMHN.

Africa; Asia; North America (introduced): Antilles, United States: Florida, Hawaii; Oceania.

was first reported in the US from Lee County, Florida (Okins and Thomas 2010). Distinguished by the narrow, strongly convergent (as opposed to rounded) posterior margin of elytra.

(Eichhoff, 1878)

Fig. 13

Eichhoff, 1878.

Blandford, 1896. Synonymy Schedl 1958.

Schedl, 1953. Synonymy

Browne, 1978. Synonymy

Holotype female; ZMUH, lost.

Asia; North America (introduced): United States: Georgia, Texas; Oceania.

was reported for the first time in North America based on specimens from Georgia and Texas (Cognato et al. 2013). Distinguished by the wide denticles of interstriae 3 and the sulcate declivity.

(Blandford, 1894)

Fig. 13

Blandford, 1894.

Niisima, 1909. Synonymy Knížek 2011.

Holotype female; Nikko, Japan; BMNH.

Asia; Europe (introduced); North America (introduced): Canada: British Columbia, Nova Scotia, Ontario, Quebec; United States: Maine, Maryland, Michigan, New York, Oregon, Pennsylvania, Washington.

Similar to , but can be distinguished by the larger size and the pointed and hooked tubercles on the declivity (Holzschuh 1994, Hoebeke and Rabaglia 2007).

(Eichhoff, 1868)

Fig. 13

Eichhoff, 1868.

Eggers, 1941. Synonymy Bright 1985.

Schedl, 1954. Synonymy Bright 1985.

Schedl, 1981. Synonymy

Lectotype; Brasilia; NMNH.

Africa; Central America: Costa Rica, Honduras, Panama; North America: Mexico, United States: Florida, Louisiana, Missouri, North Carolina, South Carolina, Texas; South America: Argentina, Brazil, Colombia, Ecuador, Venezuela.

Distinguished from other by the blunt tubercles of declivital interstriae 3.

(Murayama, 1931)

Fig. 14

Figure 14.

Lateral and dorsal views of species. From top left, and . Scale bar: 1.0 mm.

Murayama, 1931.

Holotype; Kannanri, Korea; NMNH.

Asia; North America (introduced): Alabama, Louisiana, Mississippi, South Carolina.

was reported for the first time from North America based on specimens from Alabama and Louisiana (Rabaglia et al. 2010). Distinguished from other by the 4 pairs of long, pointed spines increasing in size towards apex, on interstriae 3.

(Ratzeburg, 1837)

Fig. 14

Ratzeburg, 1837.

Wollaston, 1854. Synonymy Eichhoff 1878.

Boieldieu, 1859. Synonymy Ferrari 1867.

Ferrari, 1867. Synonymy Eichhoff 1878.

Eichhoff, 1875. Synonymy Schedl 1964.

Rey, 1883. Synonymy Bedel 1888.

Blackburn, 1885. Synonymy Samuelson 1981.

Hopkins, 1915. Synonymy Wood 1962.

Hopkins, 1915. Synonymy Wood 1962.

Hopkins, 1915. Synonymy Wood 1962.

Hopkins, 1915. Synonymy Wood 1957.

Eggers, 1930. Synonymy

Swaine, 1934. Synonymy Wood 1957.

Swaine, 1934. Synonymy Wood 1957.

Schedl, 1937. Synonymy

Schedl, 1940. Synonymy

Eggers, 1944. Synonymy Schedl 1980.

Schedl, 1948. Synonymy Schedl 1964.

Schedl, 1948. Synonymy

Schedl, 1976. Synonymy

Schedl, 1980. Synonymy

Syntypes female; “Südlichen Deutschland”; type location is indicated as presumably at SDEI by Wood and Bright (2007), unconfirmed.

Africa (introduced); Asia, Europe (introduced), North America (introduced): Mexico, Canada: British Columbia, New Brunswick, Ontario, United States: Alabama, Arizona, Arkansas, California, Colorado, Connecticut, Delaware, Florida, Georgia, Hawaii, Illinois, Indiana, Iowa, Kansas, Kentucky, Louisiana, Maine, Maryland, Massachusetts, Michigan, Mississippi, Missouri, New Hampshire, New Jersey, New York, North Carolina, Ohio, Oregon, Pennsylvania, South Carolina, Tennessee, Texas, Utah, Virginia, Washington, West Virginia; Oceania (introduced); South America (introduced): Argentina, Brazil, Chile, Ecuador, Paraguay, Uruguay.

This species is widely distributed. Wood and Bright (1992) and most authors list this species as , but Holzschuh (1994) points out that Ratzeburg’s original description was , with the ii ending. The synonymy stated by Wood (1989) between Schedl, 1980 with , supported by Brockerhoff et al. (2003), Knížek (2011) and Alonso-Zarazaga et al. (2017), may be in error (Beaver pers. comm.).

Eichhoff, 1864

Dugès, 1887. Synonymy Hagedorn 1910.

Blandford, 1896. Synonymy Hagedorn 1910.

Broun, 1904. Synonymy Bain 1976.

Reitter, 1913. Synonymy Schedl 1934.

Hopkins, 1915. Synonymy Schedl 1952.

Schedl, 1934. Synonymy

Fabricius.

Species of differ from most (but not all) members of the tribe by the truncate antennal club, the first segment of which is corneous. Species of s. str. (Hulcr and Cognato 2013) have an inflated prosternal posterocoxal process. Some species currently placed in do not have this feature, but the proper genus placement of many such species is unclear.

Eichhoff, 1868

Fig. 15

Figure 15.

Lateral and dorsal views of species. From top left, , , and . Scale bar: 1.0 mm.

Eichhoff, 1878. Synonymy Schedl 1959.

Eichhoff, 1878. Synonymy Wood 1960.

Eichhoff, 1878. Synonymy Schedl 1959.

Hopkins, 1915. Synonymy Schedl 1959.

Eggers, 1933. Synonymy Schedl 1959.

Beeson, 1935. Synonymy Beaver 1991.

Eggers, 1943. Synonymy Schedl 1962.

Syntypes female; “America bor.”, Cuba; ZMUH, lost; 1 in NMNH.

Africa (introduced); Asia (introduced); Central America: Belize, Costa Rica, El Salvador, Guatemala, Honduras, Nicaragua, Panama; Europe (introduced), North America: Antilles, Canada: Quebec, Mexico, United States: Alabama, Arkansas, California, Delaware, District of Columbia, Florida, Georgia, Hawaii, Illinois, Kansas, Kentucky, Louisiana, Maryland, Massachusetts, Michigan, Mississippi, Missouri, New Jersey, New York, North Carolina, Ohio, Oklahoma, Pennsylvania, South Carolina, Tennessee, Texas, Virginia, West Virginia; Oceania (introduced); South America: Argentina, Bolivia, Brazil, Chile, Colombia, Ecuador, Fr. Guiana, Guyana, Paraguay, Peru, Suriname, Trinidad, Uruguay, Venezuela.

This widely distributed species can cause economic damage in moist lowland areas of the Neotropics. This species is distinguished by the broadly sloping shagreened declivity and the small denticles in interstriae 1 and 3.

Eichhoff, 1868

Fig. 15

Syntypes female; Brazil; IRSNB.

Central America: Belize, Costa Rica, Guatemala, Honduras, Panama; North America: Mexico, United States: Florida, Georgia, Louisiana, North Carolina, Texas; Oceania; South America: Argentina, Bolivia, Brazil, Colombia, Ecuador, Peru, Venezuela.

This species was removed from synonymy with by Kirkendall and Jordal (2006) and its taxonomic status is unclear.

Eichhoff, 1868

Fig. 15

LeConte, 1868. Synonymy Eichhoff 1878.

Syntypes female; “America boreali”. ZMUH, lost.

North America: Canada: Ontario, United States: Alabama, Arkansas, Connecticut, Delaware, District of Columbia, Florida, Georgia, Illinois, Indiana, Iowa, Kansas, Kentucky, Louisiana, Maryland, Michigan, Minnesota, Mississippi, Missouri, New Jersey, New York, North Carolina, Ohio, Oklahoma, Pennsylvania, South Carolina, Texas, Vermont, Virginia, West Virginia.

This species is distinguished by its large size and its two pairs of large pointed tubercles on declivital interstriae 1.

(Fabricius, 1801)

Fig. 15

Fabricius, 1801.

Wollaston, 1867. Synonymy Schedl 1960.

Eichhoff, 1868. Synonymy Schedl 1960.

Eichhoff, 1868. Synonymy Schedl 1957.

Zimmermann, 1868. Synonymy

Eichhoff, 1869. Synonymy Schedl 1960.

Sharp, 1885. Synonymy Schedl 1960.

Hagedorn, 1910. Synonymy Schedl 1960.

Hopkins, 1915. Synonymy

Hopkins, 1915. Synonymy Schedl 1960.

Beeson, 1929. Synonymy Schedl 1960.

Schedl, 1931. Synonymy Schedl 1960.

Eggers, 1932. Synonymy

Eggers, 1934. Synonymy Schedl 1960.

Beeson, 1940. Synonymy Beaver 1991.

Eggers, 1941. Synonymy Schedl 1960.

Schedl, 1948. Synonymy Schedl 1960.

Lectotype female; “America meridionali”; UZMC.

Africa (introduced); Asia; Central America: Belize, Costa Rica, El Salvador, Guatemala, Honduras, Nicaragua, Panama; North America: Antilles, Canada: Ontario, Mexico, United States: Alabama, Arizona, Arkansas, California, Delaware, District of Columbia, Florida, Georgia, Hawaii, Illinois, Indiana, Kansas, Kentucky, Louisiana, Maryland, Massachusetts, Michigan, Mississippi, Missouri, New Jersey, New York, North Carolina, Ohio, Oklahoma, Pennsylvania, South Carolina, Tennessee, Texas, Virginia, West Virginia; Oceania (introduced); South America: Argentina, Bolivia, Brazil, Chile, Colombia, Ecuador, Fr. Guiana, Guyana, Paraguay, Peru, Suriname, Trinidad, Uruguay, Venezuela.

This species was removed from synonymy with by Kirkendall and Jordal (2006). It is distinguished from by the smaller size, discal interstrial setae sparse or absent, and by its light orange to reddish brown color (Atkinson et al. 2013).

Eichhoff, 1877

Fig. 16

Figure 16.

Lateral and dorsal views of species. From top left, , , and . Scale bar: 1.0 mm.

Syntype female; Japan. IRSNB.

Asia; North America (introduced): United States: Alabama, Florida, Georgia, Mississippi, South Carolina.

In the US, was first detected in a survey trap near Port Wentworth, Georgia in 2002 (Rabaglia et al. 2006). The ambrosia fungus vectored by this species is responsible for the death of 300 million bay trees ( spp.) and other in the southeastern United States (Hughes et al. 2017). This species is distinguished by the dark color and the glabrous elytral disc and declivity with small granules in all interstriae decreasing in size toward apex.

Eichhoff, 1869

Fig. 16

Schedl, 1972. Synonymy

Lectotype female; Mexico. IRSNB.

Central America: Belize, El Salvador, Guatemala, Honduras, Panama; North America: Mexico, United States: Texas.

This species is distinguished by the presence of abundant, confused punctures and setae, which completely obscure declivital striae.

Eichhoff, 1868

Fig. 16

Lectotype female; “Amer. Bor.”; NMNH.

North America: United States: Alabama, Arkansas, Florida, Georgia, Illinois, Indiana, Louisiana, Massachusetts, Mississippi, Missouri, New Jersey, North Carolina, Ohio, Oklahoma, South Carolina, Tennessee, Texas, Virginia.

This species was removed from synonymy with by Rabaglia (2005). It is distinguished by the presence of three prominent tubercles on declivital interstriae 3, while interstriae 1 are unarmed.

Blandford, 1898

Fig. 16

Hopkins, 1915. Synonymy Wood 1972.

Hopkins, 1915. Synonymy

Hopkins, 1915. Synonymy Wood 1972.

Lectotype female; Guatemala; BMNH.

Central America: El Salvador, Guatemala, Honduras; North America: Antilles, Canada: British Columbia, Mexico, United States: Arizona, California, Colorado, District of Columbia, Georgia, Idaho, Maine, Maryland, Mississippi, Montana, New Jersey, New Mexico, North Carolina, Oregon, Pennsylvania, South Carolina, South Dakota, Utah, Virginia, West Virginia.

One of the few species of the genus restricted to conifers. Distinguished from other by the steep declivity which occupies the apical ¼ of the elytra, and broadly rounded posterolateral margin of the declivity. It is distinguished from by the larger declivital denticles and smaller, deeply impressed declivital strial punctures.

(Ratzeburg, 1837)

Fig. 17

Figure 17.

Lateral and dorsal views of species. From top left, , , and . Scale bar: 1.0 mm.

Ratzeburg, 1837.

Mulsant & Rey, 1856. Synonymy Hagedorn 1910.

Eichhoff, 1875. Synonymy Schedl 1963.

Blandford, 1894.Synonymy Schedl 1963.

Syntypes female; “im Lüneburgschen und in Bayern”; not located, if extant, probably in SDEI.

Africa; Asia; Europe; North America (introduced): Canada: British Columbia; United States: Maryland, Oregon, Pennsylvania, Washington; South America: Brazil.

was first detected in North America in Maryland in 1992 (Vandenberg et al. 2000) and in Oregon in 1997–98 (Mudge et al. 2001). Distinguished from by its larger size. Wood and Bright (1992) suggest that this species may be a synonym of .

Zimmermann, 1868

Fig. 17

Holotype female; Pennsylvania; MCZ.

North America: United States: Arkansas, Illinois, Indiana, Maryland, Michigan, Mississippi, Missouri, North Carolina, Pennsylvania, Texas, West Virginia.

Distinguished by the flattened anterior portion of pronotum.

Zimmermann, 1868

Fig. 17

Eichhoff, 1868. Erroneous identification.

Eichhoff, 1869. Synonymy Wood 1973.

Lectotype female; “southern states”, USA; MCZ.

Central America: El Salvador; North America: Antilles, Canada: Ontario; United States: Alabama, Arkansas, Delaware, District of Columbia, Florida, Georgia, Kentucky, Louisiana, Maryland, Michigan, Mississippi, Missouri, New Jersey, New York, North Carolina, Oklahoma, Pennsylvania, South Carolina, Tennessee, Texas, Virginia, West Virginia.

This species and are common in pines. It can be distinguished from by the larger, shallow strial punctures and the smaller declivital denticles.

Blandford, 1894

Fig. 17

Eggers, 1933. Synonymy Mandelshtam 2007.

Kurenzov, 1941. Synonymy

Kurenzov, 1941. Synonymy

Schedl, 1957. Synonymy Browne 1962.

Syntypes; Higo, Japan; BMNH.

Asia; North America (introduced): United States: Massachusetts.

First found in Massachusetts in 2005 and 2006 (Hoebeke and Rabaglia 2008), is distinguished from other by the distinctly impressed area adjacent to the scutellum and the alternating series of longer and shorter setae on the elytra (Hoebeke and Rabaglia 2008). Both Kurenzov and Kurenzov were listed as synonyms of by Wood and Bright (1992). Mandelshtam (2007) synonymised with , without mentioning Kurenzov’s subspecies.

Blandford, 1898

Fig. 18

Figure 18.

Lateral and dorsal views of species. From top left, , , and . Scale bar: 1.0 mm.

Eggers, 1934. Synonymy Wood 1979.

Schedl, 1934. Synonymy Wood 1966.

Schedl, 1935. Synonymy Wood 1979.

Lectotype female; San Geronimo, Guatemala; BMNH.

Central America: Costa Rica, Guatemala, Honduras; North America (introduced): Antilles Mexico, United States: Hawaii, Texas; South America: Argentina, Brazil, Colombia, Ecuador, Guyana, Peru, Venezuela.

, native to Central America and lowland Mexico, was first found in the US in Texas in 1994 (Atkinson and Riley 2013). It is distinguished by its unique declivity, which is armed by spine-like tubercles.

Eichhoff, 1878

Fig. 18

Syntypes female; uncertain: Brasilia or America septentrionali (USA). ZMUH, lost.

North America: United States: Alabama, Arkansas, Florida, Illinois, Indiana, Kansas, Maryland, Mississippi, Missouri, Oklahoma, Tennessee, Texas, West Virginia.

Distinguished by the impressed anterior portion of pronotum. Distinguished from by the impressed, shining, and tuberculate declivity.

(Fabricius, 1775)

Fig. 18

Fabricius, 1775.

Eichhoff, 1868. Synonymy Wood 1960.

Eichhoff, 1869. Synonymy Eggers 1929.

Eichhoff, 1869. Synonymy Wood 1960.

Eichhoff, 1878. Synonymy

Dugès, 1887. Synonymy Wood 1983.

Hopkins, 1915. Synonymy Schedl 1952.

Hopkins, 1915. Synonymy

Hopkins, 1915. Synonymy

Hopkins, 1915. Synonymy Wood 1972.

Eggers, 1920. Synonymy

Beeson, 1929. Synonymy

Schedl, 1948. Synonymy Wood 1972.

Schedl, 1950. Synonymy

Lectotype female; “America ligno Dom v. Rohr (presumably Cuba)”; UZMC.

Africa; Asia; Central America: Belize, Costa Rica, El Salvador, Guatemala, Honduras, Nicaragua, Panama; North America: Antilles, Mexico, United States: Florida, Hawaii; Oceania; South America: Argentina, Bolivia, Brazil, Colombia, Ecuador, Fr. Guiana, Guyana, Paraguay, Peru, Suriname, Trinidad, Uruguay, Venezuela.

Distinguished by the slightly convex to flattened declivity bearing prominent tubercles of varying sizes.

(Say, 1826)

Fig. 18

Say, 1826.

Eichhoff, 1868. Synonymy Eichhoff 1878.

Swaine, 1917. Synonymy Wood 1957.

Neotype female; North Carolina; CNCI.

Asia (introduced); North America: Antilles, Canada: British Columbia, Ontario, Quebec; United States: Arkansas, California, District of Columbia, Florida, Georgia, Illinois, Indiana, Iowa, Kansas, Kentucky, Maine, Maryland, Massachusetts, Michigan, Minnesota, Mississippi, Missouri, New Hampshire, New Jersey, New York, North Carolina, Ohio, Oklahoma, Oregon, Pennsylvania, South Carolina, Utah, Virginia, West Virginia, Wisconsin.

Distinguished by the lusterless and steep declivity, occupying no more than posterior 15% of elytra.

Reitter, 1913

Wood, 1980. Synonymy Wood 1984.

Blandford.

Species of differ from other members of the tribe by widely separated procoxae.

(Blandford, 1894)

Fig. 19

Figure 19.

Lateral and dorsal views of species. From top left, , , , and . Scale bar: 1.0 mm.

Blandford, 1894.

Eggers, 1926. Synonymy Beaver 2010.

Holotype female; Japan: Higo; BMNH.

Asia; North America (introduced): Florida, Georgia.

was first discovered in the US from Florida in 2010 (Cognato et al. 2011). Distinguished by the truncate elytral declivity with a carina forming a complete cirumdeclivital ring.

(Eichhoff, 1875)

Fig. 19

Eichhoff, 1875.

Hagedorn, 1912. Synonymy Murayama and Kalshoven 1962.

Syntypes female; Japan; ZMUH, lost. 1 syntype Schedl Collection NHMW.

Africa; Asia; North America (introduced): Antilles, United States: Alabama, Florida, Georgia, Hawaii, Illinois, Louisiana, Mississippi, North Carolina, Pennsylvania, Texas; Oceania (introduced); South America: Brazil, Fr. Guiana, Peru, Trinidad.

Commonly known as the black twig borer, was first collected in the US at Ft. Lauderdale, Florida in 1941 (Wood 1982). It attacks healthy twigs of living trees and shrubs in the southeastern United States. Distinguished by the small size, the black color, and the shining declivity.

(Motschulsky, 1866)

Fig. 19

Motschulsky, 1866.

Eichhoff, 1878. Synonymy Wood 1969.

Blandford, 1896. Synonymy Schedl 1959.

Niisima, 1909. Synonymy Choo 1983.

Stebbing, 1908. Synonymy Beeson 1915.

Hagedorn, 1908. Synonymy Eggers 1923.

Schedl, 1935. Synonymy Schedl 1959.

Schedl, 1936. Synonymy Schedl 1959.

Syntypes female; Ceylon; IZM.

Africa; Asia; Central America (introduced): Costa Rica, Guatemala, Panama;, North America (introduced): Antilles, Canada: Ontario; United States: Alabama, Delaware, Florida, Georgia, Hawaii, Indiana, Kentucky, Louisiana, Maryland, Michigan, Mississippi, Missouri, North Carolina, Ohio, Oklahoma, Oregon, South Carolina, Tennessee, Texas, Virginia; Oceania (introduced); South America (introduced): Argentina, Brazil, Fr. Guiana, Uruguay.

A widely introduced species around the globe, has spread in the US along the lower Piedmont region and coastal plain to North Carolina, Louisiana, Florida, and beyond (Atkinson et al. 2012). The first US record is based on a specimen collected in South Carolina in 1974 (Anderson 1974, as ). Distinguished by the confused declivital granules giving the declivity a dull appearance. Causes economic damage in nurseries and stored hardwood lumber (Smith and Hulcr 2015).

(Eichhoff, 1869)

Fig. 19

Eichhoff, 1869.

Hopkins, 1915. Synonymy

Eggers, 1941. Synonymy

Schedl, 1963. Synonymy Wood 1973.

Schedl, 1972. Synonymy Wood 1992.

Holotype female: Brazil; IRSNB.

Central America: Costa Rica, Guatemala, Honduras, Nicaragua, Panama; North America: Antilles, Mexico, United States: Florida; South America: Argentina, Bolivia, Brazil, Colombia, Venezuela.

This species is currently only known from central and southern Florida in the United States. Distinguished by the dark brown body, the small size, and the hairy and shagreened declivity.

(Blandford, 1894)

Fig. 19

Blandford, 1894.

Blandford, 1894. Synonymy Choo 1983.

Syntypes; Japan; BMNH.

Asia; Europe (introduced); North America (introduced): Canada: British Columbia, Ontario, Quebec; United States: Alabama, Connecticut, Delaware, Florida, Georgia, Hawaii, Illinois, Indiana, Kentucky, Maine, Maryland, Massachusetts, Michigan, Mississippi, Missouri, New Hampshire, New Jersey, New York, North Carolina, Ohio, Oregon, Pennsylvania, Rhode Island, South Carolina, Tennessee, Vermont, Virginia, West Virginia.

Originating from Asia, has now spread across much of North America, including the Northeast, South and Southeast, and the Pacific Northwest (Weber and McPherson 1982; LaBonte et al. 2005); it was first thought to have been introduced into the US in a Long Island area greenhouse in 1932 (Felt 1932). Distinguished by the black color and the lack of strial setae on declivity.

1	Body conspicuously long, 3.5 times as long as wide; protibiae narrow with five large teeth on outer margin; elytral declivity deeply concave and densely pubescent, declivital surface and lateral margins not armed	Dryoxylon
–	Body stout to slender, never 3.5 times as long as wide; protibiae broad and with more than 5 small denticles on the outer margin; elytral declivity usually not concave; if impressed, lateral margins armed with denticles	2
2	Scutellum minute, conical; base of elytra at suture notched, with abundant setae	Xyleborinus
–	Scutellum flat, shiny, its surface flush with adjacent elytra, or scutellum rounded, surrounded by a moderately deep impressed area at the base of elytra	3
3	Procoxae moderately to widely separated; intercoxal piece continuous, not longitudinally emarginate	4
–	Procoxae contiguous; intercoxal piece longitudinally emarginate	5
4	Elytra wider than long, shorter than pronotum, truncate; pronotum with lateral carina	Cnestus
–	Elytra never wider than long nor shorter than pronotum, usually not truncate; pronotum with lateral margins rounded	Xylosandrus
5	Pronotal asperities extending from apex to base	Ambrosiodmus
–	Pronotal asperities confined to apical half, basal half flat, shiny, or dull	6
6	Posterior face of antennal club with segments 2 and 3 at least partially visible (type 3) (Fig. 1)	7
–	Posterior face of antennal club with no sutures visible at or near apex (type 1 or 2) (Fig. 1)	12
7	Elytral punctures confused; elytral vestiture abundant and confused	Cyclorhipidion
–	Strial and interstrial punctures in rows; elytral vestiture confined to strial and interstrial rows	8
8	Elytra narrowly rounded at apex, sutural apex strongly emarginate (Fig. 7), body slender	Coptoborus
–	Elytral apex broadly rounded, sutural apex entire, body stout	9
9	Posterolateral margin of declivity costate and broad; pronotum subquadrate or rounded	Euwallacea
–	Posterolateral margin of declivity rounded, costa blunt; pronotum rounded, never quadrate	10
10	Protibia stick-like, posterior face rugose	Dryocoetoides
–	Protibia flattened, posterior face smooth	11
11	Color black; segment 2 of antennal club non-corneous or corneous only on anterior face	Ambrosiophilus
–	Color light-brown; segment 2 of antennal club corneous on both sides	Theoborus
12	Anterior margin of pronotum distinctly armed by several coarse serrations (flat teeth); body stout, < 2.2 times as long as wide	Anisandrus
–	Anterior margin of pronotum not armed by large serrations; if serrations present, they are not larger than asperities on anterior slope of pronotum; body more slender, > 2.3 times as long as wide	Xyleborus

1	Declivital interstriae 2 either unarmed or granules smaller than those on 1 or 3	2
–	Declivital interstriae 2 with tubercles as large as or larger than those on 1 or 3	3
2	Declivital interstriae 1 feebly elevated, usually as high as 3, 2 feebly sulcate, its granules as large as those on 1; discal interstriae 3 to 4 times as wide as striae; color reddish brown to black; slightly larger, length 2.0–2.4 mm	obliquus (LeConte)
–	Declivital interstriae 1 not elevated, declivital granules absent; elytral punctures larger, deeper; discal interstriae twice as wide as striae; color very dark brown to black; smaller, length 1.8–2.1 mm	devexulus (Wood)
3	Interstrial punctures on elytral disc strongly confused to irregularly biseriate, smooth to weakly granulate	4
–	Interstrial punctures on elytral disc weakly confused to uniseriate, finely granulate	5
4	Declivital interstriae all equally tuberculate, tubercles somewhat irregular in size, but those on 2 not distinctly larger than those on other interstriae; length 3.5 mm	minor (Stebbing)
–	Declivital interstriae 1 unarmed or bearing small granules, 2 strongly tuberculate; length 3.6–4.0 mm	lewisi (Blandford)
5	Declivital interstriae all equally granulate, granules somewhat irregular in size, but those on 2 not distinctly larger than those on other interstriae; 2.4–2.6 mm	rubricollis (Eichhoff)
–	Declivital interstriae 1 unarmed or bearing very small granules, 2 strongly granulate or tuberculate	6
6	Sutural area of declivity feebly impressed, interstriae 1 armed by several fine granules; rare; 2.4 mm	opimus (Wood)
–	Sutural area of declivity moderately to strongly impressed, interstriae 1 unarmed; longer than 2.4 mm	7
7	Strial punctures on disc coarse, deep; interstriae less than 1.5 times as wide as striae; reddish, slightly bicolored; smaller, 2.5–3.0 mm	lecontei Hopkins
–	Strial punctures on disc rather small, very shallow; interstriae more than 2 times as wide as striae; black; larger, 3.7–3.9 mm	tachygraphus (Zimmermann)

1	Body larger (length 3.0–3.2 mm); declivital striae 1 and interstriae 2 weakly impressed and finely granulate	atratus (Eichhoff)
–	Body smaller (length 2.4–2.7 mm); declivital striae 1 impressed and interstriae 2 convex, with evenly spaced tubercles from base to apex	nodulosus (Eggers)

1	Posterolateral costa on declivity armed by 3–5 distinct tubercles	obesus (LeConte)
–	Posterolateral costa on declivity, may appear undulating, but without distinct tubercles	2
2	Anterior margin of pronotum armed by 2–6 serrations, median pair conspicuously larger than the others; declivity evenly convex, granules few and small; body length 2.5–2.7 mm	sayi Hopkins
–	Anterior margin of pronotum armed by 6–8 subequal serrations; declivital interstriae 1 slightly to conspicuously raised, granules numerous; body length smaller than 2.5 mm or larger than 3.2 mm	3
3	Larger, body length 3.2–3.7 mm; declivital interstriae 1 slightly raised, 2 and 3 even; interstrial punctures on elytral disc confused to irregularly biseriate	dispar (Fabricius)
–	Smaller, body length 1.8–2.3 mm; declivital interstriae 1 raised, 2 impressed, 3 raised with numerous distinct granules; interstrial punctures on elytral disc uniseriate	maiche Kurentsov

1	Body length 1.82–2.16 mm; elytra pale yellowish brown; elytral declivity dull, almost flat, not impressed between interstriae 1 and 3; pronotum longer than wide; declivital strial punctures large, shallow, distinct, with interior surfaces reticulate, separated by less than their diameter; denticles on declivital interstriae 1 and 3 small and more or less uniform in size	bodoanum (Reitter)
–	Body length more than 2.40 mm; elytra chestnut brown to blackish brown; elytral declivity shining, impressed between interstriae 1 and 3; pronotum only slightly longer than wide or nearly quadrate; declivital strial punctures smaller and reticulate, generally separated at least by their diameter or slightly more; denticles on declivital interstriae 1 and 3 larger than others	2
2	Body length 2.45–2.76 mm; declivital interstriae 2 slightly impressed; elytra chestnut-brown; strial punctures and interstrial punctures on elytral declivity of equal size, confused	fukiense (Eggers)
–	Body length 3.07–3.36 mm; declivital interstriae 2 noticeably impressed; elytra blackish brown; strial punctures on elytral declivity clearly distinct and larger than interstrial punctures, distinctly seriate	pelliculosum (Eichhoff)

1	Body slender, length 1.8–2.5 mm; light reddish brown color; pronotum subquadrate from above; elytral declivity with striae 1 strongly diverging from suture on lower half, interstriae 1 with one to three small tubercles near base, and one large tubercle slightly below middle	similis (Ferrari)
–	Body stout; dark brown to black; pronotum subquadrate to subcircular; elytral declivity with striae parallel throughout, declivity without distinctive tubercles	2
2	Body length 1.9–2.3 mm; elytra 1.2 times as long as wide; pronotum subcircular anteriorly (not subquadrate), anterior margin procurved, coarsely serrate	fornicatus (Eichhoff)
–	Body length 3.4–3.8 mm; elytra at least 1.5 times as long as wide; pronotum more nearly subquadrate	3
3	Body narrower, 3.7–3.9 mm; elytra 2 times as long as wide; elytral declivity steeply sloped from summit to apex, surface dull, punctures in striae deep, interstriae 2 with tubercles mostly absent from the apical half	validus (Eichhoff)
–	Body stout, 3.4–3.6 mm; elytra 1.5 times as long as wide; elytra gradually sloped from the base to the apex, surface shiny, punctures in striae shallow, interstria 2 with tubercles present from the base to the apex	interjectus (Blandford)

1	Elytral apex strongly convergent	andrewesi (Blandford)
–	Elytral apex broadly rounded	2
2	Declivital interstriae 1 with small denticles; 1 and 3 equally, weakly elevated	3
–	Declivital interstriae 1 without denticles and not elevated	4
3	Denticles on declivital interstriae 1 and 3 larger, those on 3 pointed, spine-like, slightly incurved; denticles on ventrolateral area of the elytra large, sharply pointed, spine-like, curved slightly downwards and to the suture; declivital interstriae 2 flattened; 2.5–2.8 mm.	attenuatus (Blandford)
–	Denticles on declivital interstriae 1 and 3 smaller, obtusely pointed; denticles on ventrolateral areas of the elytra small, less pointed; declivital interstriae 2 slightly impressed; 2.0–2.4 mm.	saxesenii (Ratzeburg)
4	Declivity flattened, declivital interstriae 3 slightly elevated with 3 pairs of small tubercles, the pair near the posterior margin largest and often blunt; 1.6–1.9 mm.	gracilis (Eichhoff)
–	Declivity sulcate, interstriae 2 impressed, tubercles on interstriae 3 of equal size; longer than 2.0 mm.	5
5	Declivital interstriae 3 slightly elevated with 2–3 pairs of tubercles, with bases wider than their length; 2.0–2.5 mm.	artestriatus (Eichhoff)
–	Declivital interstriae 3 strongly elevated with 4 pairs of long, narrow, pointed spines increasing in size approaching posterior margin, 2.1–2.4 mm	octiesdentatus (Murayama)

1	Area adjacent to scutellum impressed; pronotum nearly as broad as long, posterolateral areas distinctly, strongly asperate; 1.9–2.5 mm.	seriatus Blandford
–	Area adjacent to scutellum not impressed, flush with elytral base; pronotum stout or elongate, posterolateral areas not asperate	2
2	Declivital striae completely obscured by abundant, confused punctures and setae; body slightly more stout, 2.3–2.6 times as long as wide; 3.8–4.2 mm	horridus Eichhoff
–	Declivital striae obviously indicated or not, never obscured as above; body slender, more than 2.6 times as long as wide	3
3	Tubercles on declivital interstriae 1 distinctly larger than tubercles on other interstriae	4
–	Tubercles on declivital interstriae 1 either similar in size to tubercles on other interstriae or absent (except at base or apex)	5
4	Elytral disc and declivity setose; all declivital interstriae armed by strong tubercles at base; declivital interstriae 1 armed by two very large pointed tubercles, declivital interstriae 3 armed by several smaller tubercles; declivity weakly sulcate; larger species, 3.6–4.5 mm	celsus Eichhoff
–	Elytral disc and declivity glabrous; all declivital interstriae armed by small granules, gradually decreasing in size toward apex; interstriae 1 near apex armed by one or two small tubercles; declivity flattened, convex at suture toward apex; smaller species, 2.0 mm.	glabratus Eichhoff
5	Tubercles on declivital interstriae 3 distinctly larger than tubercles on other interstriae; tubercles absent on interstriae 1 (one or two small denticles may be present at base or apex, but not on declivital face); declivity shallowly to strongly sulcate	6
–	Tubercles on declivital interstriae 3 not distinctly larger than those on other interstriae; tubercles present on interstriae 1; declivity flat to convex	10
6	Anterior portion of pronotum flattened, weakly sulcate; 2.0–2.5 mm.	viduus Eichhoff
–	Anterior portion of pronotum convex, normal	7
7	Apex of declivity at interstriae 3 armed by two prominent, elongate tubercles; declivital setae spatulate; 1.8–2.6 mm.	spinulosus Blandford
–	Apex of declivity at interstriae 3 unarmed; declivital setae hairlike	8
8	Declivital interstriae 1 unarmed, interstriae 3 with usually three prominent tubercles; declivity distinctly sulcate, interstriae 2 impressed, strial punctures less distinct; 2.0–2.5 mm	impressus Eichhoff
–	Declivital interstriae 1 armed by one or two small denticles at base, interstriae 3 with one prominent tubercle near middle of declivity (minor denticles may also be present); declivity flat to subsulcate, interstriae 2 not impressed, strial punctures distinct	9
9	Discal interstrial setae regularly spaced, numerous; larger, more robust species; color dark reddish brown; 2.8–3.2 mm.	bispinatus Eichhoff
–	Discal interstrial setae sparse or absent; smaller, more slender species; color light orange to reddish brown; 2.4–2.9 mm.	ferrugineus (Fabricius)
10	Surface of declivity opaque	11
–	Surface of declivity shining	13
11	Anterior portion of pronotum flattened, weakly sulcate; 2.3–2.4 mm	planicollis Zimmermann
–	Anterior portion of pronotum convex, normal	12
12	Declivity broadly sloping, occupying posterior 30-40% of elytra, shagreened; declivital denticles on interstriae 1 and 3 small but conspicuous; 2.0–2.7 mm.	affinis Eichhoff
–	Declivity steep, occupying posterior 15% of elytra; denticles on declivital interstriae 1 and 3 very small; 2.3–2.7 mm.	xylographus (Say)
13	Declivity steep, posterolateral margin rounded	14
–	Declivity broadly sloping, posterolateral margin subacute	15
14	Discal interstriae twice the width of striae; some declivital tubercles with height and basal width greater than the diameter of strial punctures; declivital strial punctures small, deep; 2.2–2.7 mm.	intrusus Blandford
–	Discal interstriae less than 1.5 times width of striae; some declivital tubercles with height and basal width less than the diameter of strial punctures; declivital strial punctures large, shallow; 2.3–2.7 mm.	pubescens Zimmermann
15	Color reddish brown; declivity flattened to slightly convex, interstriae 2 moderately impressed, interstriae 1 near apex less elevated; punctures of declivital striae 1 and 2 small, anterolateral margin of punctures not raised; smaller, 2.1–2.8 mm.	volvulus (Fabricius)
–	Color usually black; declivity flattened, interstriae 2 impressed, especially from middle of declivity, interstriae 1 near apex distinctly elevated; punctures of declivital striae 1 and 2 large, shallow, anterolateral margin of punctures produced into a short ridge; larger, 3.0–3.6 mm	pfeilii (Ratzeburg)

1	Elytral declivity sharply truncate, margin of declivity with posterolateral carina extending to suture, forming a complete cirumdeclivital ring; body length 2.7–2.9 mm	amputatus (Blandford)
–	Elytral declivity rounded or only obliquely truncate, margin of declivity with carina not extending beyond 7th interstriae	2
2	Declivity without punctures or carina, surface with dense and confused granules, dull; body length 3.0 mm.; reddish brown	crassiusculus (Motschulsky)
–	Declivital striae with punctures clearly impressed, in rows; declivital surface shining, granules in sparse rows when present; body length less than 2.7 mm; black or dark brown	3
3	Body length 2.0–2.3 mm; strial setae on declivity absent (only interstrial setae present); declivital striae at least weakly impressed, interstriae very slightly elevated; typically black	germanus (Blandford)
–	Body length 1.7 mm or less; strial setae on declivity present, at least one-third as long as those on interstriae; declivital striae not impressed, interstriae flat	4
4	Elytra almost evenly arched from middle of disc to apex; setae on pronotal disc more evenly distributed, slightly more abundant on a transverse row in median area at the base; posterior portion of pronotum shining; body black, length 1.6–1.7 mm.	compactus (Eichhoff)
–	Elytra more abruptly arched from base of declivity to middle of declivity; pronotal disc glabrous except for a dense, median tuft of setae extending from base about half distance to summit; posterior portion of pronotum mostly reticulate; body dark brown, length 1.6–1.8 mm.	curtulus (Eichhoff)