Cryphalus Erichson, 1836: 61

Johnson, Andrew J., Hulcr, Jiri, Knížek, Miloš, Atkinson, Thomas H., Mandelshtam, Michail Yu., Smith, Sarah M., Cognato, Anthony I., Park, Sangwook, Li, You & Jordal, Bjarte H., 2020, Revision of the Bark Beetle Genera Within the Former Cryphalini (Curculionidae: Scolytinae), Insect Systematics and Diversity 4 (3), No. 1, pp. 1-81 : 16-24

Cryphalus Erichson, 1836: 61 ( Figs. 10–12 )

Synonymy

= Pseudocryphalus Ferrari, 1869: 252 .

= Taenioglyptes Bedel, 1888: 398 .

= Cryptarthrum Blandford, 1896a: 200 .

= Allarthrum Hagedorn, 1912c: 355 .

= Dacryphalus Hopkins, 1915: 42 syn. nov.

= Ericryphalus Hopkins, 1915: 38 .

= Hypocryphalus Hopkins, 1915: 41 syn. nov.

= Margadillius Hopkins, 1915: 37 syn. nov.

= Piperius Hopkins, 1915: 39 .

= Toenioglyptes Balachowsky, 1949: 205 (unavailable name). = Ernocryphalus Murayama, 1958: 934 .

= Acryphalus Tsai and Li, 1963: 604 (unavailable name). = Gugocryphelus Tsai and Li, 1963: 603 (unavailable name). = Jugocryphalus Tsai and Li, 1963: 604 .

Type of genus

Bostrichus asperatus Gyllenhal, 1813 .

Diagnosis

Cryphalus can be distinguished from similar genera by the combination of emarginated eyes, sutures on the antennae clearly visible but without a septum, by the weakly bilobed third tarsal segments, the proventriculus with a large apical plate, and the aedeagus that has a tegmen sclerotized completing a ring, usually with two tegminal apodemes.

Female

Eye emarginated. Antennae with three to five funicle segments. Antennal club with three visible procurved sutures, occasionally recurved, never septate. Frons convex, simple, sometimes with weak impression above epistoma, and sometimes with aciculations converging at epistoma. Pronotum with two or more marginal asperities, the margin produced in some species. Hypomeron with bifurcating setae (rare exceptions). Base of elytra sometimes with a slightly elevated ridge, straight to recurved. Angle of apex of declivity shallow, with the apex of each elytron obtuse (typically exposing part of the pygidium). Ground vestiture dense in most species (rarely sparse or absent), which can be hair-like, dagger-like or with tridentate scales. Metatibia with lateral denticles distributed over at least the apical quarter. Mesocoxae not contiguous, usually of a similar distance to distance between metacoxae or wider. Third tarsal segment weakly bilobed. Proventriculus with large apical plate, typically with sutural teeth and much larger apical teeth.

Male

Similar to female except sixth ventrite usually visible, frons with a transverse ridge, occasionally with enlarged setae on the protibiae and protarsi, and modified vestiture in the gular region. Penis apodemes typically as long as or longer than penis body, and separated or weakly fused at apex. Tegmen dorsally fused to form a ring. Tegmen with two short tegminal apodemes (absent in some species). Spiculum gastrale long, of a similar thickness to the penis apodemes, and without a fork. Basal sclerites not visible.

Distribution

Africa, Asia, Europe, Oceania, North America, Central America (introduced), South America (introduced).

Remarks

In total, 252 species known. Almost all species possess many split setae on the hypomeron which can easily separate species in this genus from most similar bark beetles. The presence, number, and arrangement of sutural teeth on the apical plate of the proventriculus have been extensively used to diagnose Chinese species (see Tsai and Li (1963)).

Many species are described. It is likely that several of the species names should be synonymized, but many types are in a poor condition with few external diagnostic characters known. The genus was previously split into three genera, including Hypocryphalus and Margadillius based primarily on the number of funicular segments. This is, however, a rapidly evolving character which may vary within a species and has a poor phylogenetic signal. The large number of species, particularly those described by Schedl, create many secondary homonyms. Where appropriate, these have been corrected.

Tsai and Li (1963) proposed three subgenera, Cryphalus s. str., Acryphalus , and Jugocryphalus . The latter two are listed as unavailable names, based on an apparent lack of type designation (Alonso- Zarazaga and Lyal 2009). However, the types are indicated in the original description of the subgenera (‘ ỄĦḡ ’ after the name, translating to ‘ type of subgenus’), with Cryphalus lipingensis the type of Acryphalus , and Cryphalus fulvus as the type of Jugocryphalus . Among the species described in the subgenera, the descriptions may represent monophyletic clades.

Where known, all species are monogamous with cave-like galleries feeding under bark.

Type material examined

Allotype of Ernoporus antennarius Schedl, 1974 (ANIC) ; paratype of Ernoporus antennarius Schedl, 1974 (NHMW) ; holotype of Taenioglyptes aquilonius Nobuchi, 1975 (ITLJ) ; holotype of Cryphalus araucariae Schedl, 1970 (ANIC) ; holotype of Taenioglyptes artestriatus Browne, 1970 (BMNH) ; holotype of Ericryphalus artocarpus Schedl, 1939 (BMNH) ; holotype and paratype of Cryphalus asperulus Schedl, 1948 (BMNH) ; paratype of Cryphalus asperulus Schedl, 1948 (BMNH) ; holotype of Cryphalus ater Browne, 1984 (BMNH) ; holotype of Cryphalus basihirtus Beeson, 1929 (BMNH) ; holotype of Taenioglyptes bicarinatus Nobuchi, 1975 (ITLJ) ; holotype of Cryphalomorphus bicolor Browne, 1983 (BMNH) ; holotype of Hypocryphalus bidentatus Browne, 1980 (BMNH) ; holotype of Cryphalus brasiliensis Schedl, 1976 (NHMW) ; paratype of Ptilopodius brevis Browne, 1970 (BMNH) ; allotype of Cryphalus brevisetosus Schedl, 1943 (NHMW) ; lectotype of Cryphalus brevisetosus Schedl, 1943 (NHMW) ; holotype of Cryphalus brimblecombei Schedl, 1948 (BMNH) ; holotype of Cryphalus brunneus Browne, 1984 (BMNH) ; holotype of Cryphalus buloloensis Browne, 1984 (BMNH) ; paratype of Cryphalus buloloensis Browne, 1984 (BMNH) ; holotype of Margadillius carinatus Browne, 1980 (BMNH) ; paratype of Cryphalus carpini Berger, 1917 (ZIN) ; holotype of Cryphalus cylindricus Browne, 1980 (BMNH) ; holotype of Cryphalus cylindrus Browne, 1984 (BMNH) ; holotype of Dacryphalus cylindrus Browne, 1950 (BMNH) ; paratype of Cryphalus cylindrus Browne, 1984 (BMNH) ; holotype of Cryphalus dilutus Eichhoff, 1878 (NHMW) ; holotype of Cryphalus dipterocarpi Wood, 1989 (USNM) ; holotype of Taenioglyptes dissimilis Nobuchi, 1975 (ITLJ) ; holotype and paratype of Cryphalus diversicolor Browne, 1984 (BMNH) ; holotype of Cryphalus exiguus Blandford, 1894 (BMNH) ; holotype of Euptilius exiguus Browne, 1984 (BMNH) ; holotype of Cryphalus felis Wood, 1989 (USNM) ; holotype of Hypocryphalus fici Browne, 1986 (BMNH) ; holotype of Hypocryphalus froggatti Nunberg, 1961 (BMNH) ; holotype of Cryphalus fuliginosus Blandford, 1895 (BMNH) ; holotype of Cryphalus fulmineus Wood, 1989 (USNM) ; holotype of Margadillius fulvus Browne, 1984 (BMNH) ; cotype of Cryphalus garciniae Nobuchi, 1959 (ITLJ) ; holotype of Cryphalus grayi Schedl, 1968 (ANIC) ; holotype of Hypothenemus griseus Blackburn, 1885 (BMNH) ; holotype of Cryphalus helopioides Schedl, 1953 (BMNH) ; holotype of Taenioglyptes hirsutus Nobuchi, 1975 (ITLJ) ; holotype of Cryphalus inops Eichhoff, 1872 (RBINS) ; holotype of Taenioglyptes kagoshimensis Nobuchi, 1975 (ITLJ) ; holotype and paratype of Cryphalus kesiyae Browne, 1975 (BMNH) ; paralectotype of Cryphalus kurenzovi Stark, 1936 (ZIN) ; holotype of Hypocryphalus laevis Browne, 1980 (BMNH) ; holotype and paratype of Cryphalus laevis Browne, 1984 (BMNH) ; type of Cryphalus laricis Niisima, 1909 (ITLJ) ; holotype and paratype of Hypocryphalus laticollis Browne, 1974 (BMNH) ; holotype of Cryphalus longior Browne, 1984 (BMNH) ; paratype of Cryphalus longior Browne, 1984 (BMNH) ; holotype and two paratypes of Taenioglyptes longipennis Browne, 1970 (BMNH) ; holotype of Hypocryphalus longipilis Browne, 1981 (BMNH) ; holotype of Taenioglyptes longisetosus Nobuchi, 1975 (ITLJ) ; holotype and paratype of Margadillius magnus Browne, 1984 (BMNH) ; holotype of Hypocryphalus malayensis Schedl, 1942 (BMNH) ; syntype of Cryphalus mandschuricus Eggers, 1929 (ITLJ) ; paralectotype and lectotype of Cryphalus mangiferae Stebbing, 1914 (BMNH) ; holotype of Margadillius margadilaonis Hopkins, 1915 (USNM) ; holotype of Taenioglyptes meridionalis Nobuchi, 1975 (ITLJ) ; Two paralectotypes of Cryphalus minimus Eggers, 1927 (NHMW, BMNH) ; holotype of Hypocryphalus minutus Browne, 1980 (BMNH) ; holotype of Cryphalus mollis Schedl, 1955 (BMNH) ; holotype of Cryphalus montanus Nobuchi, 1964 (ITLJ) ; paratype of Hypocryphalus montanus Schedl, 1974 (NHMW) ; holotype and paratype of Cryphalus negrosensis Browne, 1979 (BMNH) ; holotype of Cryphalus nitens Browne, 1980 (BMNH) ; holotype and paratype of Cryphalus nitidipennis Browne, 1984 (BMNH) ; paratype of Cryphalus nothofagi Browne, 1984 (BMNH) ; holotype of Margadillius papuanus Schedl, 1973 (ANIC) ; paratype of Margadillius papuanus Schedl, 1973 (NHMW) ; lectotype of Cryphalus perminimus Schedl, 1942 (NHMW ; holotype of Cryphalus pilosulus Browne, 1980 (BMNH) ; holotype of Taenioglyptes pulchellus Nobuchi, 1975 (ITLJ) ; holotype and paratype of Cryphalus punctatus Browne, 1984 (BMNH) ; holotype of Margadillius quadrituberculatus Schedl, 1963 (NHMW) ; holotype of Hypocryphalus reflexus Browne, 1980 (BMNH) ; syntypes of Cryphalus rhusii Niisima, 1909 (ITLJ) ; syntypes (6) of Cryphalus robustus Eichhoff, 1872 (RBINS(5), BMNH(1)) ; holotype of Hypocryphalus rotundus Hopkins, 1915 (USNM) ; holotype of Ericryphalus rugosus Schedl, 1958 (BMNH) ; holotype of Cryphalus samoensis Beeson, 1929 (BMNH) ; syntype of Cryphalus sandakanensis Schedl, 1937 (BMNH) ; holotype of Cryphalus sawadai Nobuchi and Takahashi, 1965 (ITLJ) ; holotype of Cryphalus scabripennis Schedl, 1972 (BMNH) ; holotype of Ericryphalus securus Schedl, 1940 (BMNH) ; holotype of Taenioglyptes sordidus Nobuchi, 1975 (ITLJ) ; holotype of Cryphalomorphus striatulus Browne, 1978 (BMNH) ; holotype of Cryphalus striatulus Browne, 1981 (BMNH) ; holotype of Cryphalus striatus Browne, 1974 (BMNH) ; lectotype and 3 paralectotypes Cryphalus swezeyi Schedl, 1942 (BMNH(2), NHMW(2)) ; syntype of Cryphalus swezeyi Schedl, 1942 (BMNH) ; holotype of Hypothenemus sylvicola Perkins, 1900 (BMNH) ; holotype of Cryphalus terminaliae Browne, 1980 (BMNH) ; holotype of Margadillius terminaliae Browne, 1984 (BMNH) ; syntype of Cryphalus trypanus Sampson, 1914 (BMNH) ; holotype of Cryphalus vestitus Blandford, 1895 (BMNH) ; holotype and paratype of Cryphalus vitiensis Browne, 1974 (BMNH) ; holotype of Cryptarthrum walkeri Blandford, 1896 (BMNH) .

Included species

Cryphalus abbreviatus Schedl, 1943: 35 .

Cryphalus aciculatus ( Schedl, 1939b: 341) (Hypocryphalus) comb. nov. [ Hypocryphalus ].

Cryphalus afiamalus ( Schedl, 1951d: 148) (Hypocryphalus) comb. nov. [ Hypocryphalus ].

Cryphalus amplicollis Johnson nom. nov.

= Cryphalus laticollis Browne, 1984b: 288 syn. nov. (secondary homonym).

Remarks: A replacement name is proposed for Cryphalus laticollis Browne, 1984b because the name is now occupied by Hypocryphalus laticollis Browne, 1974 . The replacement name is an adjective formed from ampli (= large) and collis (=collar).

Cryphalus angustior Eggers, 1927b: 300 comb. res. [ Hypocryphalus ]. Cryphalus aquilonius ( Nobuchi, 1975: 52) (Taenioglyptes) .

Cryphalus araucariae Schedl, 1970a: 214 .

Cryphalus armatus Schedl, 1974: 459 .

Cryphalus artestriatus ( Browne, 1970: 553) (Taenioglyptes) .

Cryphalus artocarpus ( Schedl, 1939b: 432) (Ericryphalus) .

= Cryphalus artocarpus Schedl, 1958d: 498 (syn: Schedl, 1975d). = Cryphalus brownei Wood, 1992b: 79 (unnecessary replacement name).

Cryphalus asper ( Broun, 1881: 742) (Tomicus) comb. nov. [ Hypocryphalus ].

Cryphalus asperatus ( Gyllenhal, 1813: 368) (Bostrichus) .

= Bostrichus abietis Ratzeburg, 1837: 163 (syn: Wood, 1972b).

Cryphalus asperulus Schedl, 1948b: 26 .

Cryphalus ater Browne, 1984d: 91 .

Cryphalus babai Murayama, 1961: 29 .

Cryphalus bakeri ( Eggers, 1927c: 77) (Stephanoderes) comb. nov. [ Hypocryphalus ].

Cryphalus balanopselaphus Eggers, 1920: 21 .

Cryphalus basihirtus Beeson, 1929: 227 comb. nov. [ Hypocryphalus ]. Cryphalus bellus Schedl, 1957b: 47 .

Cryphalus bicarinatus ( Nobuchi, 1975: 53) (Taenioglyptes) .

Cryphalus bicolor ( Browne, 1983b: 69) (Cryphalomorphus) .

Cryphalus bidentatus ( Browne, 1980b: 383) (Hypocryphalus) comb. nov. [ Hypocryphalus ].

Cryphalus boettcheri Schedl, 1943: 38 .

Cryphalus brasiliensis Schedl, 1976: 65 .

Cryphalus brevior ( Schedl, 1943: 40) (Hypocryphalus) comb. nov. [ Hypocryphalus ].

Cryphalus brevipilosus Schedl, 1942c: 173 .

Cryphalus brimblecombei Schedl, 1948b: 26 .

Cryphalus brunneus Browne, 1984d: 93 .

Cryphalus buloloensis Browne, 1984c: 67 .

Cryphalus capucinicollis Schedl, 1950b: 47 .

Cryphalus capucinoides Eggers, 1939b: 4 .

Cryphalus capucinomorphus Schedl, 1950b: 48 .

Cryphalus capucinus Schedl, 1938c: 425 .

Cryphalus carinatus ( Browne, 1980b: 382) (Margadillius) comb. nov. [ Margadillius ].

Cryphalus carpini Berger, 1917: 234 .

Cryphalus carpinivorus Murayama, 1930: 14 .

Cryphalus chamaecipariae Niisima, 1910: 10 .

Cryphalus chinlingensis Tsai and Li, 1963: 612 .

Cryphalus ciliatipes Blandford, 1896b: 242 .

Cryphalus cinereotestaceus ( Motschulsky, 1866: 403) (Hypoborus) . Cryphalus compactus Lea, 1910: 139 .

Cryphalus confusus ( Hopkins, 1915: 38) (Margadillius) comb. nov. [ Margadillius ].

Cryphalus constrictus Schedl, 1942c: 174 .

= Cryphalus perminimus Schedl, 1942d: 13 (syn: Schedl, 1958e). = Hypocryphalus constrictus Schedl, 1942d: 22 .

= Hypocryphalus froggatti Nunberg, 1961: 611 (syn: Schedl, 1962c).

Cryphalus corpulentus ( Schedl, 1942d: 22) (Hypocryphalus) comb. nov. [ Hypocryphalus ].

Cryphalus coryli Stark, 1936: 144 .

Cryphalus cryptomeriae Niisima, 1908: 91 .

Cryphalus cylindricus Browne, 1980b: 384 .

Cryphalus cylindripennis ( Schedl, 1959a: 483) (Hypocryphalus) comb. nov. [ Hypocryphalus ].

Cryphalus cylindrus ( Browne, 1950: 647) (Dacryphalus) comb. nov. [ Hypocryphalus ].

Cryphalus densepilosus ( Schedl, 1942d: 21) (Hypocryphalus) comb. nov. [ Hypocryphalus ].

Cryphalus dilutus Eichhoff, 1878a: 384 comb. res. [ Hypocryphalus ]. = Cryphalus dilutus Eichhoff, 1878b: 490 (syn: Johnson et al., 2017).

Cryphalus dipterocarpi Wood, 1989: 179 .

Cryphalus discrepans ( Schedl, 1965a: 58) (Hypocryphalus) comb. nov. [ Hypocryphalus ].

Cryphalusdiscretus Eichhoff,1878a: 385 comb. res. [ Hypocryphalus ]. = Cryphalus discretus Eichhoff, 1878b: 490 (syn: Johnson et al., 2017).

= Cryphalus scabricollis Eichhoff, 1878b: 491 (syn: Wood, 1989). = Cryphalus brevisetosus Schedl, 1943: 36 (syn: Wood, 1989).

Cryphalus dissimilis ( Nobuchi, 1975: 57) (Taenioglyptes) .

Cryphalus diversicolor Browne, 1984d: 90 .

Cryphalus dorsalis ( Motschulsky, 1866: 403) (Hypoborus) .

= Hylesinus sericeus Motschulsky, 1866: 402 (syn: Wood, 1969). = Hypoborus nebulosus Motschulsky, 1866: 403 (syn: Wood, 1969).

= Cryphalus indicus Eichhoff, 1878a: 384 syn. nov.

= Cryphalus indicus Eichhoff, 1878b: 489 .

Remarks: No formal synonymy has been proposed for this species name. Wood (1969) indicated that they are probably the same, and Wood and Bright (1992) listed the synonyms as above, choosing Cryphalus dorsalis as the valid name with dorsalis getting priority chosen by the first reviser. The nomenclature is being treated as valid, but the subjective synonymy is uncertain (see Mandelshtam and Nikitsky 2005).

Cryphalus dubiosus Schedl, 1963c: 66 .

Cryphalus duplosquamosus Schedl, 1942d: 15 .

Cryphalus eggersi Johnson nom. nov.

= Cryphalus confusus Eggers, 1927b: 395 syn. nov. (secondary homonym).

Remarks: A replacement name is proposed for Cryphalus confusus Eggers, 1927b , because the combination is already occupied following the transfer of Margadillius confusus Hopkins, 1915 to Cryphalus . The replacement name is a noun in the genitive case, honoring the original author.

Cryphalus elaboratus Schedl, 1950b: 51 .

Cryphalus elongatus Schedl, 1962f: 105 .

Cryphalus eriobotryae Johnson, 2019: 3 .

Cryphalus erraticus Schedl, 1979c: 96 .

Remarks: The original and subsequent spelling is erraticum as an adjective, which should be the masculine erraticus .

Cryphalus erythrinae ( Hopkins, 1915: 38) (Margadillius) comb. nov. [ Margadillius ].

Cryphalus exiguus Blandford, 1894b: 82 .

Cryphalus felis Wood, 1989: 180 .

Cryphalus fici ( Browne, 1986: 90) (Hypocryphalus) comb. nov. [ Hypocryphalus ].

Cryphalus ficivorus Murayama, 1958: 933 .

Cryphalus formosanus Schedl, 1942a: 175 .

Cryphalus fugax Schedl, 1973b: 87 .

Cryphalus fuliginosus Blandford, 1895: 319 .

Cryphalus fulmineus Wood, 1989: 180 .

Cryphalus fulvus Niisima, 1908: 92 .

= Cryphalus pini Eggers, 1921: 39 (syn: Eggers, 1923b).

Cryphalus furukawai Murayama, 1934a: 59 .

Cryphalus fuscus Johnson nom. nov.

= Cryphalus cylindrus Browne, 1984d: 92 syn. nov. (secondary homonym).

Remarks: A replacement name is proposed for Cryphalus cylindrus Browne, 1984d following the transfer of Dacryphalus cylindrus Browne, 1950 from Hypocryphalus . The replacement name is an adjective, meaning either brown or indistinct.

Cryphalus garambaensis Nunberg, 1965: 25 .

Cryphalus garciniae Nobuchi, 1959: 24 .

Cryphalus giganteus Schedl, 1950a: 22 .

Cryphalus gigas Schedl, 1975b: 218 .

Cryphalus glabratus ( Schedl, 1959a: 484) (Hypocryphalus) comb. nov. [ Hypocryphalus ].

Cryphalus gracilis Johnson nom. nov.

= Cryphalus laevis Browne, 1984b: 288 syn. nov. (secondary homonym).

Remarks: A replacement name is proposed for Cryphalus laevis Browne, 1984b because the name is occupied by Hypocryphalus laevis Browne, 1980a . The replacement name is an adjective meaning slender or graceful.

Cryphalus granulatus ( Schedl, 1942a: 176) (Hypocryphalus) comb. nov. [ Hypocryphalus ].

Cryphalus helopioides Schedl, 1953b: 295 .

Cryphalus hirsutus ( Nobuchi, 1975: 52) (Taenioglyptes) .

Cryphalus horridus Eichhoff, 1878a: 384 .

= Cryphalus horridus Eichhoff, 1878b: 488 syn. nov.

Cryphalus imitans ( Schedl, 1951d: 148) (Hypocryphalus) comb. nov. [ Hypocryphalus ].

Cryphalus infimus Schedl, 1972c: 287 .

Cryphalus intermedius Ferrari, 1867: 79 .

Cryphalus interponens ( Schedl, 1953a: 126) (Hypocryphalus) comb. nov. [ Hypocryphalus ].

Cryphalus jeholensis Murayama, 1939: 143 .

Cryphalus jezoensis Inouye and Nobuchi, 1957: 49 .

Cryphalus juglansi Niisima, 1913: 3 .

Cryphalus kagoshimensis ( Nobuchi, 1975: 56) (Taenioglyptes) .

Cryphalus kalambanganus ( Schedl, 1943: 39) (Hypocryphalus) comb. nov. [ Hypocryphalus ].

Cryphalus kesiyae Browne, 1975: 288 .

Cryphalus kivuensis Schedl, 1957b: 48 .

Cryphalus kolbei ( Hagedorn, 1912c: 355) (Allarthrum) .

Cryphalus kurenzovi Stark, 1936: 150 .

= Cryphalus punctulatus Eggers, 1942: 29 (syn: Pfeffer, 1944).

= Cryphalus ussuriensis Eggers, 1942: 29 (syn: Schedl, 1952e).

Cryphalus kyotoensis Nobuchi, 1966a: 53 .

Cryphalus laevis ( Browne, 1980a: 374) (Hypocryphalus) comb. nov. [ Hypocryphalus ].

Cryphalus laricis Niisima, 1909: 142 .

Cryphalus laticollis ( Browne, 1974: 67) (Hypocryphalus) comb. nov. [ Hypocryphalus ].

Cryphalus latus Eggers, 1929a: 10 .

= Cryphalus premayaensis Murayama, 1943: 97 (syn: Mandelshtam et al., 2007).

Cryphalus lepocrinus Tsai and Li, 1963: 619 .

Cryphalus lipingensis Tsai and Li, 1959: 90 .

Cryphalus longior Browne, 1984d: 91 .

Cryphalus longipennis ( Browne, 1970: 552) (Taenioglyptes) comb. nov. [ Hypocryphalus ].

Cryphalus longipilis ( Browne, 1981: 128) (Hypocryphalus) comb. nov. [ Hypocryphalus ].

Cryphalus longipilus Schedl, 1943: 34 .

Cryphalus longisetosus ( Nobuchi, 1975: 57) (Taenioglyptes) .

Cryphalus longus ( Eggers, 1926b: 136) (Ernoporus) .

= Cryphalus alni Krivolutskaya, 1958: 145 (syn: Mandelshtam, 2002).

Cryphalus luteus Johnson nom. nov. [ Margadillius ].

= Margadillius fulvus Browne, 1984b: 289 syn. nov. (secondary homonym).

Remarks: A replacement name is proposed for Margadillius fulvus Browne, 1984b after its transfer to Cryphalus , because the name is already occupied by Cryphalus fulvus Niisima, 1908 . The name is an adjective, meaning either brown or yellow, maintaining the original meaning.

Cryphalus magnus ( Browne, 1984d: 88) (Margadillius) comb. nov. [ Margadillius ].

Cryphalus major Stebbing, 1903: 270 .

= Cryphalus morinda Stebbing, 1903: 265 (syn: Wood, 1989).

Cryphalus malayensis ( Schedl, 1942a: 176) (Hypocryphalus) comb. nov. [ Hypocryphalus ].

Cryphalus malloti Schedl, 1943: 37 .

Cryphalus malus Niisima, 1909: 144 .

= Cryphalus padi Krivolutskaya, 1958: 144 (syn: Mandelshtam, 2002).

Cryphalus mandschuricus Eggers, 1929b: 10 .

Cryphalus mangiferae Stebbing, 1914: 542 comb. res. [ Hypocryphalus ] (nomen protectum).

= Cryphalus inops Eichhoff, 1872a: 131 (syn: Wood, 1982) (nomen oblitum).

= Hypothenemus griseus Blackburn, 1885: 194 (syn: Wood, 1982) (nomen oblitum).

= Hypocryphalus mangiferae Eggers, 1928: 85 (syn: Eggers, 1930).

= Cryphalus subcylindricus Schedl, 1942d: 16 (syn: Schedl, 1958e).

= Cryphalus mimicus Schedl, 1942d: 17 (syn: Kalshoven, 1958).

= Hypocryphalus opacus Schedl, 1942d: 20 .

Cryphalus margadilaonis ( Hopkins, 1915: 38) (Margadillius) comb. nov. [ Margadillius ].

Cryphalus markangensis Tsai and Li, 1963: 618 .

Cryphalus massonianus Tsai and Li, 1963: 613 .

Cryphalus meridionalis ( Nobuchi, 1975: 55) (Taenioglyptes) .

Cryphalus mindoroensis ( Schedl, 1943: 39) (Hypocryphalus) comb. nov. [ Hypocryphalus ].

Cryphalus minimus Eggers, 1927c: 76 .

Cryphalus minor ( Schedl, 1943: 40) (Hypocryphalus) comb. nov. [ Hypocryphalus ].

Cryphalus minusculus Johnson nom. nov. [ Hypocryphalus ].

= Hypocryphalus minutus Browne, 1980c: 495 syn. nov. (secondary homonym).

Remarks: A replacement name is proposed for Hypocryphalus minutus Browne, 1980c following its transfer to Cryphalus . The name is occupied by Margadillius minutus Hopkins, 1915 . The replacement name is an adjective, and maintains the original meaning, referencing the very small size.

Cryphalus minutus ( Hopkins, 1915: 37) (Margadillius) comb. nov. [ Margadillius ].

Cryphalus miyalopiceus Tsai and Li, 1963: 608 .

Cryphalus mollis Schedl, 1955a: 288 comb. res. [ Hypocryphalus ].

= Hypocryphalus tongaensis Schedl, 1979c: 104 (syn: Beaver, 1987). Cryphalus montanus Nobuchi, 1964: 129 .

Cryphalus moorei ( Schedl, 1964b: 247) (Hypocryphalus) comb. nov. [ Hypocryphalus ].

Cryphalus neglectus Schedl, 1962d: 106 .

Cryphalus negrosensis Browne, 1979: 85 .

Cryphalus niger Schedl, 1942c: 172 .

Cryphalus nigericus Browne, 1973: 289 .

Cryphalus nigricans Schedl, 1943: 35 .

Cryphalus nigrosetosus ( Schedl, 1948b: 27) (Hypocryphalus) comb. nov. [ Hypocryphalus ].

Cryphalus niponensis Inouye and Nobuchi, 1957: 51 .

Cryphalus nitens Browne, 1980c: 494 .

Cryphalus nitidicollis ( Schedl, 1975b: 219) (Hypocryphalus) comb. nov. [ Hypocryphalus ].

Cryphalus nitidipennis Browne, 1984d: 89 .

Cryphalus nothofagi Browne, 1984c: 68 .

Cryphalus numidicus Eichhoff, 1878a: 385 .

= Cryphalus numidicus Eichhoff, 1878b: 487 syn. nov.

Cryphalus nyalubombeae Schedl, 1957b: 49 .

Cryphalus obesus ( Hopkins, 1915: 42) (Dacryphalus) .

Cryphalus oblongus Niisima, 1910: 9 .

Cryphalus obscurus ( Hopkins, 1915: 41) (Hypocryphalus) comb. nov. [ Hypocryphalus ].

Cryphalus orientalis Eggers, 1911: 122 .

Cryphalus ovalicollis ( Schedl, 1942c: 177) (Hypocryphalus) comb. nov. [ Hypocryphalus ].

Cryphalus ozopemoides Johnson nom. nov. [ Hypocryphalus ].

= Hypocryphalus montanus Schedl, 1974: 460 syn. nov. (secondary homonym).

Remarks: A replacement name is proposed for Hypocryphalus montanus Schedl, 1974 following its transfer to Cryphalus , because the name is already occupied by Cryphalus montanus Nobuchi, 1964 . The replacement name is an adjective, refenceing the gross similarity between this species and the genus Ozopemon Hagedorn, 1910 , for which females have a very similar shape and overall appearance.

Cryphalus paganus Eichhoff, 1878b: 129 .

Cryphalus palawanus Schedl, 1942a: 174 .

Cryphalus pallidus Eichhoff, 1872a: 131 .

Cryphalus papuanus ( Schedl, 1973b: 87) (Margadillius) comb. nov. [ Margadillius ].

= Ernoporus antennarius Schedl, 1974: 461 syn. nov.

Cryphalus parvulus Niisima, 1910: 8 .

Cryphalus pellicius Johnson nom. nov. [ Hypocryphalus ].

= Hypocryphalus pilifer Schedl, 1979c: 98 syn. nov. (secondary homonym).

Remarks: A replacement name is proposed for Hypocryphalus pilifer Schedl, 1979c following transfer to the genus Cryphalus , because the name is already occupied by Cryphalus pilifer Eggers, 1927b . The replacement name is an adjective, following the descriptive meaning of the name, meaning furry.

Cryphalus pexus Schedl, 1979b: 104 .

Cryphalus piceae ( Ratzeburg, 1837: 163) (Bostrichus) .

= Cryphalus hattorii Kôno, 1938: 67 (syn: Inouye and Nobuchi, 1957).

= Cryphalus subdepressus Eggers, 1940d: 37 (syn: Wood, 1992a). Cryphalus piceus Eggers, 1926b: 136 .

Cryphalus pilifer Eggers, 1927b: 394 .

Cryphalus piliger ( Schedl, 1975b: 219) (Hypocryphalus) comb. nov. [ Hypocryphalus ].

Cryphalus pilosellus Erichson, 1842: 212 .

Cryphalus pilosulus Browne, 1980b: 385 .

Cryphalus pilosus Tsai and Li, 1963: 606 .

Cryphalus planicollis Schedl, 1978b: 74 .

Cryphalus polynesiae ( Schedl, 1979c: 105) (Hypocryphalus) comb. nov. [ Hypocryphalus ].

Cryphalus procerus Schedl, 1953b: 296 .

Cryphalus pruni Eggers, 1929b: 11 .

Cryphalus pseudochinlingensis Tsai and Li, 1963: 610 .

Cryphalus pseudotabulaeformis Tsai and Li, 1963: 613 .

Cryphalus puberulus Schedl, 1942c: 171 .

Cryphalus pubescens Hopkins, 1915: 39 .

= Cryphalus subconcentralis Hopkins, 1915: 40 (syn: Wood, 1954).

Cryphalus pulchellus ( Nobuchi, 1975: 54) (Taenioglyptes) .

Cryphalus punctatostriatus Schedl, 1942a: 175 .

Cryphalus punctatus Browne, 1984d: 66 .

Cryphalus punctipennis Schedl, 1942c: 169 .

Cryphalus punctistriatulus Johnson nom. nov.

= Cryphalus striatulus Browne, 1981: 127 syn. nov. (secondary homonym).

Remarks: A replacement name is proposed for Cryphalus striatulus Browne, 1981 because the name is occupied, following the revised status of Cryphalomorphus striatulus Browne, 1978 . The new is formed from the diminutive of the adjective punctatus meaning punctured, and s striatus meaning with lines or rows.

Cryphalus pusillimus Schedl, 1942c: 171 .

Cryphalus pusillus Schedl, 1943: 38 .

Cryphalus quadrituberculatus ( Schedl, 1963c: 64) (Margadillius) comb. nov. [ Margadillius ].

Cryphalus redikorzevi Berger, 1917: 232 .

Cryphalus reflexus ( Browne, 1980b: 383) (Hypocryphalus) comb. nov. [ Hypocryphalus ].

Cryphalus resiniferi Schedl, 1943: 36 .

Cryphalus rhusii Niisima, 1909: 145 .

= Cryphalus kurilensis Krivolutskaya, 1968: 53 (syn: Mandelshtam, 2006).

Cryphalus robustus Eichhoff,1872a: 131 comb. res. [ Hypocryphalus ]. Cryphalus rotundus ( Hopkins, 1915: 41) (Hypocryphalus) comb. nov. [ Hypocryphalus ].

Cryphalus rubentis Hopkins, 1915: 40 .

Cryphalus ruficollis Hopkins, 1915: 40 .

= Cryphalus fraseri Hopkins, 1915: 40 (syn: Wood, 1989).

= Cryphalus approximatus Hopkins, 1915: 41 (syn: Wood, 1954). = Cryphalus balsameus Hopkins, 1915: 41 (syn: Wood, 1954).

= Cryphalus amabilis Chamberlin, 1917: 321 (syn: Wood, 1954). = Cryphalus grandis Chamberlin, 1917: 323 (syn: Wood, 1954).

= Cryphalus canadensis Chamberlin, 1918: 88 (syn: Wood, 1954). = Cryphalus mainensis Blackman, 1922b: 126 (syn: Wood, 1954). = Cryphalus ruficollis coloradensis Wood, 1954: 1008 (syn: Wood, 1977c).

Cryphalus rufopilosus Schedl, 1942a: 174 .

Cryphalus rugosus ( Schedl, 1958b: 102) (Ericryphalus) .

Cryphalus saltuarius Weise, 1891: 336 (nomen protectum).

= Bostrichus asperatus Ratzeburg, 1837: 163 (unavailable name). = Cryphalus scriba Gozis, 1886: 31 (nomen oblitum).

Cryphalus samoensis Beeson, 1929: 224 .

Cryphalus sandakanensis Schedl, 1937b: 548 comb. res. [ Hypocryphalus ].

= Hypocryphalus maculatus Browne, 1962a: 303 (syn: Wood, 1989).

Cryphalus sarawakensis Schedl, 1972h: 199 .

Cryphalus sawadai Nobuchi and Takahashi, 1965: 3 .

Cryphalus scabripennis Schedl, 1972h: 199 .

Cryphalus schedli Johnson nom. nov. [ Hypocryphalus ].

= Hypocryphalus formosanus Schedl, 1952b: 62 syn. nov. (secondary homonym).

Remarks: A replacement name is proposed for Hypocryphalus formosanus Schedl, 1952b because the name is occupied by Cryphalus formosanus Schedl, 1942a . The replacement name is a noun in the genitive case, honoring the original author. The types of both species were unavailable for study.

Cryphalus scopiger Berger, 1917: 228 .

Cryphalus securus ( Schedl, 1940a: 436) (Ericryphalus) .

Cryphalus sejugatus Schedl, 1965a: 54 .

Cryphalus sichotensis Kurentsov, 1941: 152 .

Cryphalus sidneyanus ( Nördlinger, 1856: 75) (Bostrichus) .

Cryphalus silvanus Schedl, 1951d: 145 .

Cryphalus similis ( Eggers, 1922: 168) (Stephanoderes) .

Cryphalus simplex ( Schedl, 1939b: 341) (Ericryphalus) .

Cryphalus sinoabietis Tsai and Li, 1963: 606 .

⊇ Cryphalus sinoabietis opienensis Tsai and Li, 1963: 607 .

Cryphalus solomonensis Johnson nom. nov.

= Margadillius terminaliae Browne, 1984c: 450 syn. nov. (secondary homonym).

Remarks: A replacement name is proposed for Margadillius terminaliae Browne, 1984c because the name is already occupied by Cryphalus terminaliae Browne, 1980b . The replacement name is a toponym derived from the type locality.

Cryphalus sordidus ( Nobuchi, 1975: 51) (Taenioglyptes) .

Cryphalus sparsepilosus Schedl, 1942c: 172 .

Cryphalus spathulatus ( Schedl, 1938b: 49) (Hypocryphalus) comb. nov. [ Hypocryphalus ].

Cryphalus spissepilosus Johnson nom. nov.

= Cryphalus densepilosus Schedl, 1943: 36 syn. nov. (secondary homonym).

Remarks: A replacement name is proposed for Cryphalus densepilosus Schedl, 1943 because the name is now occupied by Hypocryphalus densepilosus Schedl, 1942d . The replacement name is formed from the adverb spisse (= densely) and the adjective pilosus (= hairy).

Cryphalus squameus Schedl, 1943: 38 .

Cryphalus squamulosus Strohmeyer, 1911: 20 .

Cryphalus storckiellae Johnson nom. nov.

= Cryphalus striatus Browne, 1974: 66 syn. nov. (secondary homonym).

Remarks: A replacement name is proposed for Cryphalus striatus Browne, 1974 because the name is now occupied by Hypocryphalus striatus Hopkins, 1915 . The replacement name is a noun in the genitive case, named after the host plant Storckiella .

Cryphalus striatulus ( Browne, 1978: 576) (Cryphalomorphus) comb. nov. [ Hypothenemus ].

= Hypothenemus brownei Beaver, 1991: 53 syn. nov. (unnecessary replacement name).

Remarks: Cryphalus striatulus ( Browne, 1978) is transferred to Cryphalus based on the external diagnostic characters visible. The replacement name is no longer necessary.

Cryphalus striatus ( Hopkins, 1915: 41) (Hypocryphalus) comb. nov. [ Hypocryphalus ].

Cryphalus strigilatus Wichmann, 1914: 137 .

Cryphalus strigipennis Schedl, 1950b: 50 .

Cryphalus strohmeyeri Stebbing, 1914: 538 .

= Cryphalus indicus Stebbing, 1902: 403 (syn: Stebbing, 1914) (Permanently invalid name due to primary homonymy).

Cryphalus subcompactus Lea, 1910: 140 .

Cryphalus subgranulatus Schedl, 1943: 37 .

Cryphalus submuricatus Eichhoff, 1878a: 385 .

= Cryphalus submuricatus Eichhoff, 1878b: 492 syn. nov.

= Cryphalus tuberculatus Schedl, 1943: 37 syn. nov.

Cryphalus substriatus Schedl, 1942a: 174 .

Cryphalus subtuberculatus Schedl, 1942c: 168 .

Cryphalus subvestitus Schedl, 1959a: 482 .

Cryphalus sumatranus ( Schedl, 1939c: 38) (Dacryphalus) comb. nov. [ Hypocryphalus ].

Cryphalus sundaensis Schedl, 1942d: 14 .

Cryphalus sylvicola ( Perkins, 1900: 181) (Hypothenemus) . = Ericryphalus henshawi Hopkins, 1915: 38 (syn: Schedl, 1941b). = Piperius pini Hopkins, 1915: 39 (syn: Schedl, 1963c). = Cryphalus swezeyi Schedl, 1942b: 147 (syn: Wood, 1960). = Cryphalus dimorphus Schedl, 1950b: 38 (syn: Beaver, 1991). = Cryphalus sylvicola obliquus Schedl, 1950b: 48 .

Cryphalus szechuanensis Tsai and Li, 1963: 614 .

⊇ Cryphalus szechuanensis tehchangensis Tsai and Li, 1963: 615 . Cryphalus tabulaeformis Tsai and Li, 1963: 609 .

⊇ Cryphalus tabulaeformis chienzhuangensis Tsai and Li, 1963: 610 .

Cryphalus taiwanus Schedl, 1970c: 359 .

Cryphalus takahashii Johnson nom. nov. [ Ernoporus ]

= Euptilius exiguus Browne, 1984c: 451 syn. nov. (secondary homonym).

Remarks: A replacement name is proposed for Euptilius exiguus Browne, 1984c because the name is occupied by Cryphalus exiguus Blandford, 1894b . The replacement name is a noun in the genitive case, honoring the original collector, K. Takahashi.

Cryphalus tenuis Schedl, 1942d: 16 .

Cryphalus terminaliae Browne, 1980b: 384 .

Cryphalus tetricus ( Schedl, 1940a: 437) (Ericryphalus) .

= Cryphalus papuanus Schedl, 1942c: 170 (syn: Schedl, 1979c). = Cryphalus grayi Schedl, 1968: 265 (syn: Schedl, 1979c). = Ptilopodius brevis Browne, 1970: 556 (syn: Schedl, 1972j). Cryphalus theobromae Nunberg, 1956b: 208 (replacement name). = Cryphalus horridus Graham, 1908: 113 (syn: Nunberg, 1956b) (permanently invalid name due to primary homonymy).

Cryphalus triangularis ( Schedl, 1975a: 351) (Hypocryphalus) comb. nov. [ Hypocryphalus ].

Cryphalus trypanoides Beeson, 1935a: 106 .

Cryphalus trypanus Sampson, 1914: 383 .

Cryphalus tutuilaensis ( Schedl, 1951d: 147) (Hypocryphalus) comb. nov. [ Hypocryphalus ].

Cryphalus uapouensis ( Beeson, 1935a: 107) (Ericryphalus) . Cryphalus upoluensis Schedl, 1951d: 144 .

Cryphalus variolosus Schedl, 1950b: 49 .

Cryphalus vestitus Blandford, 1895: 318 .

Cryphalus viburni Stark, 1936: 151 .

= Cryphalus viburni Eggers, 1942: 30 (syn: Pfeffer, 1944). Cryphalus vitiensis Browne, 1974: 67 .

Cryphalus walkeri ( Blandford, 1896a: 200) (Cryptarthrum) . = Coccotrypes hagedorni Eggers, 1908: 216 (syn: Schedl, 1963c). = Cryphalus javanus Schedl, 1954a: 148 (syn: Schedl, 1958e). Cryphalus wapleri Eichhoff, 1872a: 131 .

= Cryphalus mekeoi Schedl, 1972e: 61 (syn: Schedl, 1979d). Cryphalus yamaguchii Inouye and Nobuchi, 1957: 55 .

Cryphalus zimmermani Schedl, 1950b: 51 .

Figures

Figure 10. Images of Cryphalus spp.: Dorsal and lateral photographs of A) C. asperatus, B) C. bicolor, C) C. dorsalis, D) C. kesiyae, E) C. amplicollis, F) C. meridionalis.

Figure 11. Images of Cryphalus spp.: Dorsal and lateral photographs of A) C. ozopemoides, B) C. nr. piceus. Eye, antennae, and frons of C) C. kesiyae and D) C. mangiferae.

Figure 12. Images of Cryphalus spp.: Aedeagus of A) Cryphalus asperatus, B) C. keysiae, C) C. mangiferae. Proventriculus of D) C. asperatus, E) C. dorsalis, F) C. mangiferae, and G) C. ozopemoides.