Scolytinae

Female Phloeosinus dentatus, copyright N. Wright.


Belongs within: Curculionidae.
Contains: Xyloterini, Micracini, Bothrosternini, Scolytus, Diamerini, Scolytodes, Phloeotribus, Hylurgops, Tomicini, Polygraphini, Crypturgini, Pityophthorina, Corthylina, Cryphalus, Xyloctonini, Hypoborini, Ips, Hypothenemus, Scolytogenes, Cyrtogenius, Xyleborus.

The Scolytinae, bark beetles, are a group of small weevils that live and feed in the subcortical tissues of plants. They are characterised by a reduced rostrum, geniculate antennae with a distinct, compact club, and tibiae marginally armed by spines or socketed denticles (Wood 1986). The majority of species live in galleries in the phloem layer of their hosts but some feed on other tissues such as xylem or seeds. Some species feed on ambrosia fungi growing on host xylem. Many bark beetles are monogamous with a single mating pair living in a gallery; eggs are laid within the gallery and the parents guard the burrow entrance while the larvae develop. Other species exhibit various modes of polygyny.

Wood (1986) recognised the Scolytidae as a separate family from the Curculionidae, divided between two subfamilies Scolytinae and Hylesininae on the basis of features such as form of the pronotum and base of the elytra, but more recent studies have not supported monophyly of the latter two groups. Tribes of the Scolytinae include the Scolytini, members of which are most diverse in the Americas and lack armature on the lateral margins of the pro- and metatibiae except for a single, apical, spinelike process curving towards and extending beyond the process of the inner apical angle (Wood 1986).

Characters (from Wood 1986, as Scolytidae): Adult with head narrower than pronotum; eyes nearly flat against head, often emarginate to divided; pregular sutures present and reinforced internally by massive apodemal ridges; rostrum short to nonexistent; hypostomal area small, posterior mandibular condyle more anterior in position; axis of mandibular hinge oblique to transverse, cutting edge on mesal margin; pronotum often asperate, almost never with constriction near middle; when visible, scutellum usually flush with elytral surface, often flattened; metepisternum stout to elongate, anterior margin usually procurved, anteroventral angle displaced slightly caudad and distinctly extended ventrad; mesepimeron flat, oblique; mesepisternum smaller, declivous cephalad; tibiae armed on apical and/or lateral margins either by spines or socketed denticles, apical mucro formed by mesal element; tarsal segment 1 about equal in length to 2 or 3; male spiculum gastrale present. Larva with clypeus distinct, separate from frons; frontal sutures never reaching articulating membrane of mandibles.

<==Scolytinae [Hylesininae, Scolytarii] D09
    |--Scolytini [Camptoceridae] GC-P14
    |    |--Scolytus W86
    |    |--Scolytopsis Blandford 1896 W86
    |    |    `--*S. puncticollis W86
    |    |--Camptocerus Latreille 1829 W86
    |    |    |--*C. aeneipennis [=Hylesinus aeneipennis] W86
    |    |    `--C. auricomus W86
    |    `--Cnemonyx Eichhoff 1868 (see below for synonymy) W86
    |         |--*C. galeritus W86
    |         |--C. barbatus [=*Minulus barbatus] W86
    |         |--C. flavicornis [=*Loganius flavicornis] W86
    |         |--C. hylurgoides [=*Coptodryas hylurgoides, *Coptosomus hylurgoides] W86
    |         |--C. jucundus [=*Ceratolepis jucundus] W86
    |         `--C. panamensis W86
    `--+--+--+--Diamerini GC-P14
       |  |  `--+--Hexacolini [Ctenophoridae, Ctenophorini, Hexacolidae, Problechilidae] GC-P14
       |  |     |    |--Scolytodes W86
       |  |     |    |--Gymnochilus Eichhoff 1867 [=Problechilus Eichhoff 1878; incl. Meringopalpus Hagedorn 1905] W86
       |  |     |    |    `--*G. zonatus [=*Problechilus zonatus; incl. *Meringopalpus fallax] W86
       |  |     |    |--Pycnarthrum Eichhoff 1878 (see below for synonymy) W86
       |  |     |    |    `--P. hispidum (see below for synonymy) W86
       |  |     |    `--Microborus Blandford 1897 [incl. Pseudocrypturgus Eggers 1919] W86
       |  |     |         `--*M. boops [incl. *Pseudocrypturgus camerunus] W86
       |  |     `--Phloeotribini [Phloeotribidae, Phthorophloeides] GC-P14
       |  |          |--Phloeotribus W86
       |  |          `--Aricerus Blandford 1894 [incl. Hylesinosoma Lea 1910] W86
       |  |               |--*A. chapuisi W86
       |  |               `--A. eichhoffi [incl. Hylesinus fici, *Hylesinosoma fici] W86
       |  `--+--Hylesinini (see below for synonymy) GC-P14
       |     |    |--Hylurgops W86
       |     |    |--Hylastinus Bedel 1888 W86
       |     |    |    `--*H. obscurus [=Ips obscurus] W86
       |     |    |--Kissophagus Chapuis 1869 W86
       |     |    |    `--*K. hederae [=Hylesinus hederae] W86
       |     |    |--Pteleobius Bedel 1888 W86
       |     |    |    `--*P. vittatus [=Bostrichus vittatus] W86
       |     |    |--Cryptocurus Schedl 1957 [incl. Hyloperus Browne 1970] W86
       |     |    |    `--*C. spinipennis [incl. *Hyloperus bicornis]
       |     |    |--Neopteleobius Nobuchi 1971 W86
       |     |    |    `--*N. scutulatus [=Hylesinus scutulatus] W86
       |     |    |--Scierus LeConte 1876 W86
       |     |    |    `--*S. annectens W86
       |     |    |--Hylurgus Latreille 1807 M94
       |     |    |    `--*H. ligniperda (Fabricius 1775) [=Hylesinus ligniperda] M94
       |     |    |--Dendrosinus Chapuis 1869 W86
       |     |    |    `--*D. globosus [=Hyulesinus globosus] W86
       |     |    |--Hyleops Schedl 1938 W86
       |     |    |    `--*H. glabratus W86
       |     |    |--Pseudochramesus Blackman 1939 W86
       |     |    |    `--*P. acuteclavatus [=Chramesus acuteclavatus] W86
       |     |    |--Phloeosinopsoides Schedl 1964 [=Phloeosinopsis Schedl 1964 non Schedl 1936] W86
       |     |    |    `--*P. triseriatus [=*Phloeosinopsis triseriatus] W86
       |     |    |--Phloeocranus Schedl 1942 [incl. Diamerides Browne 1949] W86
       |     |    |    `--*P. bruchoides [incl. *Diamerides litseae] W86
       |     |    |--Phloeoditica Schedl 1962 W86
       |     |    |    `--*P. curta [=Kissophagus curtus] W86
       |     |    |--Carphotoreus Wood 1973 W86
       |     |    |    `--*C. alni [=Chaetophloeus alni] W86
       |     |    |--Catenophorus Nunberg 1956 W86
       |     |    |    `--*C. congonus W86
       |     |    |--Phloeosinites Hagedorn 1907 W86
       |     |    |    `--*P. rehi W86
       |     |    |--Phloeoborus Erichson 1836 [incl. Phloeotrupes Erichson 1836] W86
       |     |    |    |--*P. rudis W86
       |     |    |    `--P. grandis [=*Phloeotrupes grandis] W86
       |     |    |--Dactylipalpus Chapuis 1869 (see below for synonymy) W86
       |     |    |    |--D. transversus [=*Dactylopselaphus transversus] W86
       |     |    |    `--D. cicatricosus [=*Ethadopselaphus cicatricosus] W86
       |     |    |--Alniphagus Swaine 1918 [incl. Hylastinoides Spessivitzev 1919] W86
       |     |    |    |--*A. aspericollis [=Hylesinus aspericollis] W86
       |     |    |    `--A. alni [=Hylastes alni, *Hylastinoides alni] W86
       |     |    |--Rhopalopselion Hagedorn 1909 [incl. Hapalogenius Hagedorn 1912] W86
       |     |    |    |--*R. bituberculatum W86
       |     |    |    `--R. globosus [=*Hapalogenius globosus] W86
       |     |    |--Ficicis Lea 1910 [incl. Ficiphagus Murayama 1958] W86
       |     |    |    |--*F. varians W86
       |     |    |    `--‘Hylesinus’ porcatus Chap. 1869 W86, M87 [incl. *Ficiphagus goliathoides W86]
       |     |    |--Pachycotes Sharp 1877 M94
       |     |    |    |--P. peregrinus M94 [=Hylesinus peregrinus W86; incl. *P. ventralis Sharp 1877 M94]
       |     |    |    `--P. australis Schedl 1938 M94
       |     |    |--Hylesinus Fabricius 1801 [incl. Leperisinus Reitter 1913] W86
       |     |    |    |--*H. crenatus W86
       |     |    |    |--H. antipodius W86
       |     |    |    `--H. varius W86 (see below for synonymy)
       |     |    |--Cladoctonus Strohmeyer 1911 (see below for synonymy) W86
       |     |    |    |--*C. affinis W86
       |     |    |    |--C. banosus [=*Hoplites banosus, *Hoplitontus banosus] W86
       |     |    |    `--C. interruptus [=Hoplites interruptus; incl. *Hoplitophthorus sentosus] W86
       |     |    |--Olonthogaster Motschulsky 1866 (see below for synonymy) W86
       |     |    |    |--*O. nitidicollis [=*Holonthogaster nitidicollis] W86
       |     |    |    |--O. asper [=*Hyledius asper] W86
       |     |    |    |--O. spinifer [=Phloeosinus spinifer; incl. *Phloeosinopsis armatus] W86
       |     |    |    `--O. sumatranus [=*Hylurgulus sumatranus] W86
       |     |    |--Chramesus LeConte 1868 (see below for synonymy) W86
       |     |    |    |--*C. hickoriae W86
       |     |    |    |--C. annectans [=*Prochramesus annectans] W86
       |     |    |    |--C. incomptus W86
       |     |    |    |--C. quadridens W86
       |     |    |    |--C. rotundatus [incl. Dendrosinus bonnairei, *Thaumasinulus bonnairei] W86
       |     |    |    `--C. tuberculatus [=*Rhopalopleurus tuberculatus] W86
       |     |    |--Hylesinopsis Eggers 1920 (see below for synonymy) W86
       |     |    |    |--*H. dubius W86
       |     |    |    |--H. africanus [=*Pseudophloeotribus africanus] W86
       |     |    |    |--H. baphiae [=*Glochicopterus baphiae] W86
       |     |    |    |--H. brinckei [=Metahylesinus brinckei, *Hapalophloeus brinckei] W86
       |     |    |    |--H. endroedyi [=*Hemihylesinus endroedyi] W86
       |     |    |    |--H. joveri [=*Trypographus joveri] W86
       |     |    |    |--H. planicollis [=*Chilodendron planicolle] W86
       |     |    |    `--H. togonus [=*Pseudohylesinus togonus, *Metahylesinus togonus] W86
       |     |    `--Phloeosinus Chapuis 1873 M94
       |     |         |--*P. dentatus [=Hylurgus dentatus] M94
       |     |         |--P. bicolor W86
       |     |         |--P. canadensis M94
       |     |         |--P. cristatus W86
       |     |         |--P. cupressi Hopkins 1903 M94
       |     |         |--P. perlatus B14
       |     |         |--P. punctatus W86
       |     |         `--P. thujae [=Hylesinus thujae] W86
       |     `--+--Hylastes Erichson 1836 M94 [Hylastini GC-P14, Hylastinides]
       |        |    |--*H. ater (Paykull 1800) [=Bostrichus ater, Hylesinus ater] M94
       |        |    |--H. brunneus RW02
       |        |    |--H. cunicularius RW02
       |        |    |--H. glabratus C01
       |        |    `--H. nigrinus W86
       |        `--Tomicini GC-P14
       `--+--+--Polygraphini GC-P14
          |  `--Crypturgini GC-P14
          `--+--+--Corthylini [Amphicranidae, Araptidae, Corthyli, Pityophthoridae] GC-P14
             |  |    |--Pityophthorina W86
             |  |    `--Corthylina W86
             |  `--+--Cryphalus GC-P14
             |     `--Xyloctonini GC-P14
             `--+--Hypoborini GC-P14
                `--+--+--Premnobius Eichhoff 1878 W86 [incl. Premnophilus Browne 1962 W86; Premnobiini GC-P14]
                   |  |    |--*P. cavipennis W86
                   |  |    `--P. joveri [=Xyleborus joveri, *Premnophilus joveri] W86
                   |  `--Ipini [Eidophelinae, Ernoporinae, Trypophloeinae] GC-P14
                   |       |--Ips W86
                   |       |--Hypothenemus W86
                   |       |--Scolytogenes W86
                   |       |--Dendrochilus Schedl 1957 W86
                   |       |    `--*D. strombosiopsis W86
                   |       |--Cryphalities Cockerell 1917 W86
                   |       |    `--*C. rugosissimus Cockerell 1917 W86
                   |       |--Periocryphalus Wood 1971 W86
                   |       |    `--*P. pullus W86
                   |       |--Trischidias Hopkins 1915 W86
                   |       |    `--*T. georgiae W86
                   |       |--Neocryphus Nunberg 1956 W86
                   |       |    `--*N. argentinensis W86
                   |       |--Stegomerus Wood 1967 W86
                   |       |    `--*S. vulgaris W86
                   |       |--Allernoporus Kurentzov 1941 W86
                   |       |    `--*A. euonymi W86
                   |       |--Coriacephilus Schedl 1939 W86
                   |       |    `--*C. coriaceus [=Stephanoderes coriaceus] W86
                   |       |--Ernocladius Wood 1980 W86
                   |       |    `--*E. corpulentus [=Cryphalus corpulentus] W86
                   |       |--Cryphalogenes Wood 1980 W86
                   |       |    `--*C. euphorbiae W86
                   |       |--Hemicryphalus Schedl 1963 W86
                   |       |    `--*H. argutus [=Eidophelus argutus] W86
                   |       |--Margadillius Hopkins 1915 W86
                   |       |    `--*M. margadilaonis W86
                   |       |--Ptilopodius Hopkins 1915 W86
                   |       |    `--*P. stephegynis W86
                   |       |--Xyleborinus Reitter 1913 M94
                   |       |    `--*X. saxeseni (Ratzeburn 1837) [=Bostrichus saxeseni] M94
                   |       |--Pityokteines Fuchs 1911 [incl. Orthotomides Wood 1951] W86
                   |       |    |--*P. curvidens [=Ips curvidens] W86
                   |       |    `--P. lasiocarpi [=Orthotomicus lasiocarpi, *Orthotomides lasiocarpi] W86
                   |       |--Orthotomicus Ferrari 1867 [=Neotomicus Fuchs 1911, Onthotomicus] W86
                   |       |    |--*O. laricis [=Bostrichus laricis, *Neotomicus laricis] W86
                   |       |    `--O. erosus JKW03
                   |       |--Trypophloeus Fairmaire 1868 [=Glyptoderus Eichhoff 1878 non LaFerté Sénectère 1851] W86
                   |       |    |--*T. binodulus [=Bostrichus binodulus, *Glyptoderus binodulus] W86
                   |       |    `--T. populi [=Cryphalus populi] W86
                   |       |--Eidophelus Eichhoff 1875 [incl. Phellodendrophagus Krivolutskaya 1958] W86
                   |       |    |--*E. imitans W86
                   |       |    `--E. elegans [=*Phellodendrophagus elegans] W86
                   |       |--Procryphalus Hopkins 1915 W86
                   |       |    |--P. mucronatus [=Cryphalus mucronatus; incl. *P. populi] W86
                   |       |    `--P. utahensis W86
                   |       |--Cryptocarenus Eggers 1937 [incl. Tachyderes Blackman 1943] W86
                   |       |    |--*C. diadematus W86
                   |       |    `--C. seriatus [incl. *Tachyderes floridensis] W86
                   |       |--Acorthylus Brèthes 1922 [incl. Phacrylus Schedl 1938] W86
                   |       |    |--*A. asperatus W86
                   |       |    `--A. bosqui [=*Phacrylus bosqui] W86
                   |       |--Stephanopodius Schedl 1941 (see below for synonymy) W86
                   |       |    |--*S. dispar [=Stephanoderes dispar] W86
                   |       |    `--S. ghanaensis (see below for synonymy) W86
                   |       |--Pityogenes Bedel 1888 [incl. Eggersia Lebedev 1926, Pityoceragenes Balachowsky 1947] W86
                   |       |    |--*P. chalcographus [=Dermestes chalcographus] W86
                   |       |    |--P. bidentatus [=Bostrichus bidentatus, *Eggersia bidentata] W86
                   |       |    `--P. quadridens [=Bostrichus quadridens, *Pityoceragenes quadridens] W86
                   |       |--Ernoporus Thomson 1859 (see below for synonymy) W86
                   |       |    |--*E. tiliae [=Bostrichus tiliae; incl. Cryphalus lederi, *Cryphalops lederi] W86
                   |       |    |--E. concentralis [=*Euptilius concentralis] w86
                   |       |    `--E. melodori [=*Stephanorhopalus melodori] W86
                   |       |--Ernoporicus Berger 1917 (see below for synonymy) W86
                   |       |    |--*E. spessivtzevi W86
                   |       |    |--E. caucasicus [=Ernoporus caucasicus, *Ernopocerus caucasicus] W86
                   |       |    `--E. semenovi [=*Eocryphalus semenovi] W86
                   |       |--Cosmoderes Eichhoff 1878 (see below for synonymy) W86
                   |       |    |--*C. monilicollis [incl. *Vitaderes luffa (n. n.)] W86
                   |       |    |--C. granulicollis [=*Dendriops granulicollis] W86
                   |       |    `--C. setistriatus [=Cryphalus setistriatus, *Erioschidias setistriatus] W86
                   |       |--Hypocryphalus Hopkins 1915 [incl. Dacryphalus Hopkins 1915] W86
                   |       |    |--*H. rotundus W86
                   |       |    |--H. mangiferae W86
                   |       |    `--H. obesus [=*Dacryphalus obesus] W86
                   |       |--Acanthotomicus Blandford 1894 [incl. Isophthorus Schedl 1938, Mimips Eggers 1932] W86
                   |       |    |--*A. spinosus W86
                   |       |    |--A. mimicus W86
                   |       |    |--A. pilosus [=Ips pilosus, *Mimips pilosus] W86
                   |       |    `--A. quadrituberculatus [=*Isophthorus quadrituberculatus] W86
                   |       `--Coccotrypes Eichhoff 1879 M94 (see below for synonymy)
                   |            |--*C. dactyliperda [=Bostrichus dactyliperda] W86
                   |            |--C. advena [incl. *Thamnurgides persicae] W86
                   |            |--C. carpophagus (Hornung 1842) M94 (see below for synonymy)
                   |            |--C. rhizophorae [=*Spermatoplex rhizophorae, *Dendrurgus rhizophorae] W86
                   |            `--C. sannio [=*Poecilips sannio] W86
                   `--+--Dryocoetini [=Dryocoetoideae, Taphrorychini, Thamnurginae] GC-P14
                      |    |--Cyrtogenius W86
                      |    |--Toxophthorus Wood 1986 [=Toxophorus Eggers 1920 (preoc.)] W86
                      |    |    `--*T. africanus (Eggers 1920) [=*Toxophorus africanus] W86
                      |    |--Taphramites Schedl 1947 W86
                      |    |    `--*T. gnathotrichus Schedl 1947 W86
                      |    |--Thamnurgus Eichhoff 1864 W86
                      |    |    `--*T. euphorbiae [=Bostrichus euphorbiae] W86
                      |    |--Chiloxylon Schedl 1959 W86
                      |    |    `--*C. rufulus W86
                      |    |--Dactylotrypes Eggers 1927 W86
                      |    |    `--D. longicollis [=Xyloterus longicollis; incl. *D. uyttenboogaarti] W86
                      |    |--Dendrocranulus Schedl 1937 W86
                      |    |    `--*D. tardus W86
                      |    |--Lymantor Lovendal 1889 W86
                      |    |    `--‘Tomicus’ coryli [incl. *Lymantor sepicola] W86
                      |    |--Taphronurgus Reitter 1913 W86
                      |    |    `--*T. exul [=Thamnurgus exul] W86
                      |    |--Dolurgocleptes Schedl 1965 W86
                      |    |    `--*D. malgassicus W86
                      |    |--Dendrographus Schedl 1964 [incl. Protopityophthorus Schedl 1973] W86
                      |    |    `--*D. pygmaeus [=Pelicerus pygmaeus; incl. *Protopityophthorus durus] W86
                      |    |--Peridryocoetes Wood 1984 W86
                      |    |    `--*P. nitens [=Ozodendron nitens] W86
                      |    |--Ozopemon Hagedorn 1908 W86
                      |    |    `--*O. regius W86
                      |    |--Dryocoetiops Schedl 1957 W86
                      |    |    |--*D. laevis [=Ozopemon laevis] W86
                      |    |    `--D. coffeae W86
                      |    |--Triotemnus Wollaston 1864 [incl. Cladoctoproctus Schedl 1975] W86
                      |    |    |--*T. subretusus W86
                      |    |    `--T. scrafa [=*Cladoctoproctus scrafa] W86
                      |    |--Dryocoetes Eichhoff 1864 (see below for synonymy) W86
                      |    |    |--*D. autographus [=Bostrichus autographus, *Anodius autographus] W86
                      |    |    |--D. confusus W86
                      |    |    `--D. villosus [=Bostrichus villosus, *Dryocoetinus villosus] W86
                      |    |--Tiarophorus Schreiner 1882 (see below for synonymy) W86
                      |    |    |--*T. elongatus W86
                      |    |    |--T. camerunus [=*Hypaspistes camerunus, *Orthaspistes camerunus] W86
                      |    |    `--T. scrutator [=Thamnurgus scrutator, *Pseudothamnurgus scrutator] W86
                      |    |--Xylocleptes Ferrari 1867 [incl. Hylonius Nunberg 1973, Xestips Hagedorn 1912] W86
                      |    |    |--*X. bispinus [=Bostrichus bispinus] W86
                      |    |    |--X. brunneus [=*Hylonius brunneus] W86
                      |    |    `--X. marginatus [=*Xestips marginatus] W86
                      |    `--Taphrorychus Eichhoff 1878 (see below for synonymy) W86
                      |         |--*T. bicolor [=Bostrichus bicolor] W86
                      |         |--T. machnovskii [=Hypothenemus machnovskii, *Saliciphilus machnovskii] W86
                      |         |--T. mikuniyamensis [=*Pseudopoecilips mikuniyamensis] W86
                      |         `--T. primitus [=*Taphroterus primitus] W86
                      `--Xyleborini [Webbinae, Xylebori] GC-P14
                           |--Australoplatypus incompertus GE05
                           |--Xyleborus W86
                           |--Xyleborites Wickham 1913 W86
                           |    `--*X. longipennis Wickham 1913 W86
                           |--Sampsonius Eggers 1935 W86
                           |    `--*S. sexdentatus W86
                           |--Schedlia Browne 1950 W86
                           |    `--*S. sumatrana [=Xyleborus sumatranus] W86
                           |--Dryocoetoides Hopkins 1915 W86
                           |    `--D. capucinus [=Xyleborus capucinus; incl. *D. guatemalensis] W86
                           |--Leptoxyleborus Wood 1980 W86
                           |    `--*L. sordicauda [=Phloeotrogus sordicauda] W86
                           |--Theoborus Hopkins 1915 W86
                           |    `--*T. theobromae W86
                           |--Euwallacea Hopkins 1915 W86
                           |    `--*E. wallacei [=Xyleborus wallacei] W86
                           |--Taurodemus Wood 1980 W86
                           |    `--*T. sharpi [=Xyleborus sharpi] W86
                           |--Mesoscolytus Broun 1904 W86
                           |    `--*M. inurbanus W86
                           |--Hadrodemius Wood 1980 W86
                           |    `--*H. globus [=Xyleborus globus] W86
                           |--Taphrodasus Wood 1980 W86
                           |    `--*T. percorthylus [=Xyleborus percorthylus] W86
                           |--Cryptoxyleborus Schedl 1937 W86
                           |    `--*C. naevus W86
                           |--Cnestus Sampson 1911 [incl. Tosaxyleborus Murayama 1950] W86
                           |    |--*C. magnus W86
                           |    `--C. murayamai [incl. *Tosaxyleborus pallidipennis] W86
                           |--Coptoborus Hopkins 1915 [incl. Streptocranus Schedl 1939] W86
                           |    |--C. vespatorius [=Xyleborus vespatorius; incl. *C. emarginatus] W86
                           |    `--C. mirabilis [=*Streptocranus mirabilis] W86
                           |--Arixyleborus Hopkins 1915 [incl. Xyleboricus Eggers 1923] W86
                           |    |--*A. rugosipes W86
                           |    `--A. canaliculatus [=*Xyleboricus canaliculatus] W86
                           |--Terminalinus Hopkins 1915 [incl. Kelantanius Nunberg 1961] W86
                           |    |--*T. terminaliae W86
                           |    `--T. punctatopilosus [=Xyleborus punctatopilosus, *Kelantanius punctatopilosus] W86
                           |--Eccoptopterus Motschulsky 1863 (see below for synonymy) W86
                           |    |--E. spinosus [=Scolytus spinosus; incl. *Eccoptopterus sexspinosus] W86
                           |    `--E. gracilipes [=*Platydactylus gracilipes, *Eurydactylus gracilipes] W86
                           |--Ambrosiodmus Hopkins 1915 (see below for synonymy) W86
                           |    |--*A. tachygraphus [=Xyleborus tachygraphus] W86
                           |    |--A. obliquecaudata [=*Phloeotrogus oliquecaudata] W86
                           |    `--A. obliquus (see below for synonymy) W86
                           |--Amasa Lea 1894 [incl. Anaxyleborus Wood 1980, Pseudoxyleborus Eggers 1930] W86
                           |    |--*A. thoracicus W86
                           |    |--*Pseudoxyleborus’ beesoni W86
                           |    `--A. truncatus (Erichs. 1842) W86, M87 (see below for synonymy)
                           |--Coptodryas Hopkins 1915 (see below for synonymy) W86
                           |    |--*C. confusa W86
                           |    |--C. pullus [=Xyleborus pullus; incl. *Adryocoetes nitidus (n. n.)] W86
                           |    `--C. theae [=Xyleborus theae, *Microperus theae] W86
                           |--Webbia Hopkins 1915 (see below for synonymy) W86
                           |    |--*W. dipterocarpi W86
                           |    |--W. pabo W86
                           |    |--W. subuculae [=*Prowebbia subuculae] W86
                           |    `--W. trepanicauda [=Xyleborus trepanicauda, *Pseudowebbia trepanicauda] W86
                           |--Xylosandrus Reitter 1913 [incl. Apoxyleborus Wood 1980] W86
                           |    |--*X. morigerus [=Xyleborus morigerus] W86
                           |    |--X. compactus MHG04
                           |    |--X. mancus [=Xyleborus mancus, *Apoxyleborus mancus] W86
                           |    `--X. retusus W86
                           `--Anisandrus Ferrari 1867 GC-P14, W86
                                `--*A. dispar W86, GC-P14 [=Xyleborus (Apate) dispar W86]

Scolytinae incertae sedis:
  Bufonus Eggers 1919 W86
    `--*B. obscurus Eggers 1919 W86
  Cactopinus Schwarz 1899 [incl. Cactopinorus Bright 1967; Cactopinini] W86
    |--*C. hubbardi Schwarz 1899 B38
    |--C. cactophthorus [=*Cactopinorus cactophthorus] W86
    |--C. depressus W86
    |--C. desertus W86
    |--C. koebelei Blackman 1938 B38
    |--C. pini Blackman 1938 B38
    `--C. rhois Blackman 1938 B38
  Scolytoplatypus Schaufuss 1893 (see below for synonymy) W86
    |--*S. permirus W86
    |--S. congonus W86
    |--S. mikado [=*Strophionocerus mikado, *Taeniocerus mikado] W86
    |--S. papuanus W86
    `--S. tycon [=*Spongocerus tycon] W86
  Carphodicticini W86
    |--Dendrodicticus Schedl 1958 W86
    |    `--*D. argentiniae W86
    |--Carphodicticus Wood 1971 W86
    |    `--*C. cristatus W86
    `--Craniodicticus Blandford 1895 W86
         `--*C. mucronatus W86
  Xyloterini W86
  Micracini W86
  Phrixosoma Blandford 1897 (see below for synonymy) W86
    |--*P. rude W86
    |--P. magna W86
    |--P. psaltes [=*Bothryperus psaltes] W86
    |--P. quadrioculatum [=*Neohylesinus quadrioculatus] W86
    `--P. striatum [=*Sphaerosinus striatus] W86
  Hyorrhynchini [Hyorrhynchinae, Sueinae] W86
    |--Sueus Murayama 1951 [incl. Neohyorrhynchus Schedl 1962] W86
    |    `--S. niisimai (see below for synonymy) W86
    |--Hyorrhynchus Blandford 1894 W86
    |    `--*H. lewisi W86
    `--Pseudohyorrhynchus Murayama 1950 W86
         `--*P. wadai W86
  Bothrosternini W86

Amasa truncatus (Erichs. 1842) W86, M87 [=Tomicus truncatus W86, *Anaxyleborus truncatus W86]

Ambrosiodmus Hopkins 1915 [incl. Browneia Nunberg 1963, Phloeotrogus Motschulsky 1863 (nom. rej.)] W86

Ambrosiodmus obliquus [=Pityophthorus obliquus; incl. Xyleborus illepidus, *Browneia illepida] W86

Chramesus LeConte 1868 [incl. Prochramesus Wood 1956, Rhopalopleurus Chapuis 1869, Thaumasinulus Reitter 1913] W86

Cladoctonus Strohmeyer 1911 [incl. Hoplites Eggers 1923 nec Koch 1869 nec Neumayr 1875, Hoplitontus Wood 1961, Hoplitophthorus Wood 1961] W86

Cnemonyx Eichhoff 1868 [incl. Ceratolepis Chapuis 1869, Coptodryas Schedl 1948 non Hopkins 1915, Coptosomus Schedl 1952, Loganius Chapuis 1869, Minulus Eggers 1912] W86

Coccotrypes Eichhoff 1879 M94 [incl. Cryphaloides Formenek 1908 W86, Dendrurgus Eggers 1923 W86, Hyphaene Hagedorn 1913 (n. n.) W86, Poecilips Schaufuss 1897 W86, Spermatoplex Hopkins 1915 W86, Thamnurgides Hopkins 1915 W86]

Coccotrypes carpophagus (Hornung 1842) M94 [=Bostrichus carpophagus M94; incl. *Cryphaloides donisthorpei W86, *Hyphaene guineensis (n. n.) W86]

Coptodryas Hopkins 1915 [incl. Adryocoetes Eggers in Schedl 1952 (n. n.), Microperus Wood 1980] W86

Cosmoderes Eichhoff 1878 [incl. Dendriops Schedl 1953, Erioschidias Schedl 1938, Vitaderes Beeson 1941 (n. n.)] W86

Dactylipalpus Chapuis 1869 [=Dactylopselaphus Gemminger & Harold 1872; incl. Ethadopselaphus Blandford 1896] W86

Dryocoetes Eichhoff 1864 [=Anodius Motschulsky 1860 (nom. rej.); incl. Dryocoetinus Balachowsky 1949] W86

Eccoptopterus Motschulsky 1863 [incl. Eurydactylus Hagedorn 1909 non LaFerté Sénectère 1851, Platydactylus Eichhoff 1886] W86

Ernoporicus Berger 1917 [incl. Eocryphalus Kurentzov 1941, Ernopocerus Balachowsky 1949] W86

Ernoporus Thomson 1859 [incl. Cryphalops Reitter 1889, Euptilius Schedl 1940, Stephanorhopalus Hopkins 1915] W86

Hylesinini [Alniphagini, Dactylipalpi, Hylesinen, Phloeobori, Phloeosinides, Phloeosinini, Phloeotrupides] GC-P14

Hylesinopsis Eggers 1920 [incl. Chilodendron Schedl 1953, Glochicopterus Schedl 1954, Hapalophloeus Schedl 1966, Hemihylesinus Schedl 1967, Metahylesinus Eggers 1922, Pseudohylesinus Eggers 1919 non Swaine 1917, Pseudophloeotribus Eggers 1933, Trypographus Schedl 1950] W86

Hylesinus varius W86 [=Bostrichus varius W86; incl. B. fraxini W86, Hylesinus fraxini L86, *Leperisinus fraxini W86]

Olonthogaster Motschulsky 1866 [=Holonthogaster Gemminger & Harold 1872; incl. Hyledius Sampson 1921, Hylurgulus Eggers 1927, Phloeosinopsis Schedl 1936] W86

Phrixosoma Blandford 1897 [incl. Bothryperus Hagedorn 1909, Neohylesinus Eggers 1920, Sphaerosinus Eggers 1929; Phrixosomini] W86

Pycnarthrum Eichhoff 1878 [incl. Monebius Hopkins 1914, Nemobius Chapuis 1869 (preoc.), Nomebius Navas 1915] W86

Pycnarthrum hispidum [=Hypoborus hispidus; incl. *P. gracile, *Nemobius lambottei, *Monebius lambottei, *Nomebius lambottei] W86

Scolytoplatypus Schaufuss 1893 [incl. Spongocerus Blandford 1893, Strophionocerus Sampson 1921, Taeniocerus Blandford 1893; Scolytoplatypini, Scolytoplatypodini, Spongocerinae, Taeniocerini] W86

Stephanopodius Schedl 1941 [incl. Cryphalomimetes Browne 1963, Cryphalomimus Browne 1962 non Eggers 1927] W86

Stephanopodius ghanaensis [=Hypocryphalus ghanaensis, *Cryphalomimetes ghanaensis, *Cryphalomimus ghanaensis] W86

Sueus niisimai [=Hyorrhynchus niisimai, *Neohyorrhynchus niisimai; incl. *Sueus sphaerotrypoides] W86

Taphrorychus Eichhoff 1878 [incl. Pseudopoecilips Murayama 1957, Saliciphilus Sokanovskii 1954, Taphroterus Schedl 1965] W86

Tiarophorus Schreiner 1882 [incl. Hypaspistes Hagedorn 1908 (preoc.), Orthaspistes Hagedorn 1909, Pseudothamnurgus Eggers 1912] W86

Webbia Hopkins 1915 [incl. Prowebbia Browne 1962, Pseudowebbia Browne 1961, Xelyborus Schedl 1939 (n. n.)] W86

*Type species of generic name indicated

REFERENCES

[B38] Blackman, M. W. 1938. New species of Cactopinus Schwarz (Coleoptera: Scolytidae). Proceedings of the Entomological Society of Washington 40 (6): 151–157.

[B14] Bouchard, P. (ed.) 2014. The Book of Beetles: A lifesize guide to six hundred of nature's gems. Ivy Press: Lewes (United Kingdom).

[C01] Csiki, E. 1901. Bogarak [Coleopteren]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 75–120. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[GC-P14] Gillett, C. P. D. T., A. Crampton-Platt, M. J. T. N. Timmermans, B. H. Jordal, B. C. Emerson & A. P. Vogler. 2014. Bulk de novo mitogenome assembly from pooled total DNA elucidates the phylogeny of weevils (Coleoptera: Curculionoidea). Molecular Biology and Evolution 31 (8): 2223–2237.

[GE05] Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

[JKW03] Jacobs, K., T. Kirisits & M. J. Wingfield. 2003. Taxonomic re-evaluation of three related species of Graphium, based on morphology, ecology and phylogeny. Mycologia 95 (4): 714–727.

[L86] Lindquist, E. E. 1986. The world genera of Tarsonemidae (Acari: Heterostigmata): a morphological, phylogenetic, and systematic revision, with a reclassification of family-group taxa in the Heterostigmata. Memoirs of the Entomological Society of Canada 118 (S136): 1–517.

[M87] Masters, G. 1887. Catalogue of the described Coleoptera of Australia. Part VI. Proceedings of the Linnean Society of New South Wales, series 2, 1 (4): 979–1036.

[M94] May, B. M. 1994. An introduction to the immature stages of Australian Curculionoidea. In: Zimmerman, E. C. Australian Weevils (Coleoptera: Curculionoidea) vol. 2. Brentidae, Eurhynchidae, Apionidae and a chapter on immature stages by Brenda May pp. 365–728. CSIRO Australia.

[MHG04] Maynard, G. V., J. G. Hamilton & J. F. Grimshaw. 2004. Quarantine—phytosanitary, sanitary and incursion management: an Australian entomological perspective. Australian Journal of Entomology 43 (3): 318–328.

[RW02] Reay, S. D., & P. J. Walsh. 2002. Relative attractiveness of some volatiles to the introduced pine bark beetles, Hylastes ater and Hylurgus ligniperda (Curculionidae: Scolytinae). New Zealand Entomologist 25: 51–56.

[W86] Wood, S. L. 1986. A reclassification of the genera of Scolytidae (Coleoptera). Great Basin Naturalist Memoirs 10: 1–126.

Last updated: 19 September 2018.

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