Revision Of The Bark Beetle Genera Within The Former Cryphalini (Curculionidae: Scolytinae)

Cryphalini Lindemann, 1877 are a speciose group of mostly miniscule beetles. The tribe Cryphalini is reviewed here which resulted in taxonomic and nomenclatural changes. This revision follows a recent molecular phylogenomic re-analysis focused on the tribe and related scolytine taxa. The analysis demonstrated that the tribe is polyphyletic, as found in other molecular phylogenies. To ensure monophyletic classification, we present a revision of the former tribe with two tribes resurrected, one described, and several genera transferred to other existing tribes. Additionally, extensive generic synonymy, and new combinations are presented. A key, photographs, and illustrations are provided to enable an accurate determination of genera. The revised Cryphalini contains only Cryphalus Erichson, 1836 (=Hypocryphalus Hopkins, 1915 syn. nov.; Margadillius Hopkins, 1915 syn. nov.). Coriacephilini Johnson trib. nov. contains only Coriacephilus Schedl, 1939. Ernoporini Nusslin, 1911 stat. res. contains Eidophelus Eichhoff, 1876 (=Scolytogenes Eichhoff, 1878 syn. nov.; Ptilopodius Hopkins, 1915 syn. nov.; Ernoporicus Berger, 1917 syn. nov.; Cryphalogenes Wood, 1980 syn. nov.); Ernoporus Thomson, 1859 (=Ernocladius Wood, 1980 syn. nov.; Allothenemus Bright and Torres, 2006 syn. nov.); Hemicryphalus Schedl, 1963; and Procryphalus Hopkins, 1915. Trypophloeini Nusslin, 1911 stat. res. includes the genera Afrocosmoderes Johnson and Jordal gen. nov.; Atomothenemus Bright, 2019; Cosmoderes Eichhoff, 1878 (=Allernoporus Kurentsov, 1941 syn. nov.); Hypothenemus Westwood, 1834 (=Periocryphalus Wood, 1971 syn. nov.); Macrocryphalus Nobuchi, 1981 stat. res.; Microcosmoderes Johnson and Jordal gen. nov.; Microsomus Bright, 2019; Pygmaeoborus Bright, 2019; and Trypophloeus Fairmaire, 1864. Xyloterini LeConte, 1876 is maintained, containing Indocryphalus Eggers, 1939; Trypodendron Stephens, 1830 and Xyloterinus Swaine, 1918. Acorthylus Brethes, 1922, Cryptocarenus Eggers, 1937, Neocryphus Nunberg, 1956, Stegomerus Wood, 1967, and Trypolepis Bright, 2019 are transferred to Corthylini LeConte, 1876. Stephanopodius Schedl, 1963 is transferred to Xyloctonini Eichhoff, 1878. As a consequence of generic synonymy, the following new or resurrected combinations are proposed: Cosmoderes euonymi (Kurentsov, 1941) comb. nov.; Cryphalus aciculatus (Schedl, 1939) comb. nov.; Cryphalus afiamalus (Schedl, 1951) comb. nov.; Cryphalus angustior Eggers, 1927 comb. res.; Cryphalus asper (Broun, 1881) comb. nov.; Cryphalus bakeri (Eggers, 1927) comb. nov.; Cryphalus basihirtus Beeson, 1929 comb. nov.; Cryphalus bidentatus (Browne, 1980) comb. nov.; Cryphalus brevior (Schedl, 1943) comb. nov.; Cryphalus carinatus (Browne, 1980) comb. nov.; Cryphalus confusus (Hopkins, 1915) comb. nov.; Cryphalus corpulentus (Schedl, 1942) comb. nov.; Cryphalus cylindripennis (Schedl, 1959) comb. nov.; Cryphalus cylindrus (Browne, 1950) comb. nov.; Cryphalus densepilosus (Schedl, 1942) comb. nov.; Cryphalus dilutus Eichhoff, 1878 comb. res.; Cryphalus discrepans (Schedl, 1965) comb. nov.; Cryphalus discretus Eichhoff, 1878 comb. res.; Cryphalus erythrinae (Hopkins, 1915) comb. nov.; Cryphalus fici (Browne, 1986) comb. nov.; Cryphalus glabratus (Schedl, 1959) comb. nov.; Cryphalus granulatus (Schedl, 1942) comb. nov.; Cryphalus imitans (Schedl, 1951) comb. nov.; Cryphalus interponens (Schedl, 1953) comb. nov.; Cryphalus kalambanganus (Schedl, 1943) comb. nov.; Cryphalus laevis (Browne, 1980) comb. nov.; Cryphalus laticollis (Browne, 1974) comb. nov.; Cryphalus longipennis (Browne, 1970) comb. nov.; Cryphalus longipilis (Browne, 1981) comb. nov.; Cryphalus magnus (Browne, 1984) comb. nov.; Cryphalus malayensis (Schedl, 1942) comb. nov.; Cryphalus mangiferae Stebbing, 1914 comb. res.; Cryphalus margadilaonis (Hopkins, 1915) comb. nov.; Cryphalus mindoroensis (Schedl, 1943) comb. nov.; Cryphalus minor (Schedl, 1943) comb. nov.; Cryphalus minutus (Hopkins, 1915) comb. nov.; Cryphalus mollis Schedl, 1955 comb. res.; Cryphalus moorei (Schedl, 1964) comb. nov.; Cryphalus nigrosetosus (Schedl, 1948) comb. nov.; Cryphalus nitidicollis (Schedl, 1975) comb. nov.; Cryphalus obscurus (Hopkins, 1915) comb. nov.; Cryphalus ovalicollis (Schedl, 1942) comb. nov.; Cryphalus papuanus (Schedl, 1973) comb. nov.; Cryphalus piliger (Schedl, 1975) comb. nov.; Cryphalus polynesiae (Schedl, 1979) comb. nov.; Cryphalus quadrituberculatus (Schedl, 1963) comb. nov.; Cryphalus reflexus (Browne, 1980) comb. nov.; Cryphalus robustus Eichhoff, 1872 comb. res.; Cryphalus rotundus (Hopkins, 1915) comb. nov.; Cryphalus sandakanensis Schedl, 1937 comb. res.; Cryphalus spathulatus (Schedl, 1938) comb. nov.; Cryphalus striatulus (Browne, 1978) comb. nov.; Cryphalus striatus (Hopkins, 1915) comb. nov.; Cryphalus sumatranus (Schedl, 1939) comb. nov.; Cryphalus triangularis (Schedl, 1975) comb. nov.; Cryphalus tutuilaensis (Schedl, 1951) comb. nov.; Eidophelus absonus (Schedl, 1975) comb. nov.; Eidophelus afer (Schedl, 1970) comb. nov.; Eidophelus africanus (Schedl, 1977) comb. nov.; Eidophelus aitutakii (Beaver and Maddison, 1990) comb. nov.; Eidophelus alniphagus (Nobuchi, 1975) comb. nov.; Eidophelus alternans (Schedl, 1975) comb. nov.; Eidophelus amanicus (Eggers, 1919) comb. nov.; Eidophelus ankius (Schedl, 1979) comb. nov.; Eidophelus apicalis (Schedl, 1971) comb. nov.; Eidophelus approximatus (Schedl, 1975) comb. nov.; Eidophelus aspericollis (Eichhoff, 1878) comb. nov.; Eidophelus ater (Eggers, 1923) comb. nov.; Eidophelus australis (Schedl, 1942) comb. nov.; Eidophelus badius (Nobuchi, 1975) comb. nov.; Eidophelus bambusae (Browne, 1983) comb. nov.; Eidophelus bangensis (Eggers, 1927) comb. nov.; Eidophelus basilaris (Wood, 1960) comb. nov.; Eidophelus birosimensis (Murayama, 1958) comb. nov.; Eidophelus braderi (Browne, 1965) comb. nov.; Eidophelus brimblecombei (Schedl, 1972) comb. nov.; Eidophelus buruensis (Eggers, 1926) comb. nov.; Eidophelus camelliae (Nobuchi, 1975) comb. nov.; Eidophelus candidus (Nobuchi, 1975) comb. nov.; Eidophelus capucinus (Schedl, 1971) comb. nov.; Eidophelus caucasicus (Lindemann, 1877) comb. nov.; Eidophelus ceylonicus (Schedl, 1959) comb. nov.; Eidophelus cicatricosus (Schedl, 1942) comb. nov.; Eidophelus coccotrypanoides (Schedl, 1939) comb. nov.; Eidophelus communis (Schaufuss, 1891) comb. nov.; Eidophelus confragosus (Sampson, 1914) comb. nov.; Eidophelus corni (Kurentsov, 1941) comb. nov.; Eidophelus corpulentus (Schedl, 1965) comb. nov.; Eidophelus corrugatus (Schedl, 1950) comb. nov.; Eidophelus creber (Schedl, 1975) comb. nov.; Eidophelus crenatus (Sampson, 1914) comb. nov.; Eidophelus cylindricus (Schedl, 1959) comb. nov.; Eidophelus darwini (Eichhoff, 1878) comb. nov.; Eidophelus devius (Schedl, 1975) comb. nov.; Eidophelus dubiosus (Wood, 1960) comb. nov.; Eidophelus eggersi (Schedl, 1962) comb. nov.; Eidophelus euphorbiae (Wood, 1980) comb. nov.; Eidophelus excellens (Schedl, 1979) comb. nov.; Eidophelus exiguus (Wood, 1980) comb. nov.; Eidophelus exilis (Yin, 2001) comb. nov.; Eidophelus eximius (Schedl, 1942) comb. nov.; Eidophelus expers (Blandford, 1894) comb. nov.; Eidophelus fagi (Fabricius, 1798) comb. nov.; Eidophelus fijianus (Schedl, 1950) comb. nov.; Eidophelus formosanus (Browne, 1981) comb. nov.; Eidophelus fugax (Schedl, 1975) comb. nov.; Eidophelus fujisanus (Nobuchi, 1975) comb. nov.; Eidophelus fulgens (Schedl, 1975) comb. nov.; Eidophelus fulgidus (Schedl, 1975) comb. nov.; Eidophelus fulvipennis (Nobuchi, 1975) comb. nov.; Eidophelus ghanaensis (Schedl, 1977) comb. nov.; Eidophelus glabratus (Yin, 2001) comb. nov.; Eidophelus gracilis (Schedl, 1950) comb. nov.; Eidophelus granulatus (Wood, 1960) comb. nov.; Eidophelus grobleri (Schedl, 1962) comb. nov.; Eidophelus hirtus (Wood, 1974) comb. nov.; Eidophelus hobohmi (Schedl, 1955) comb. nov.; Eidophelus hylesinopsis (Schedl, 1975) comb. nov.; Eidophelus incultus (Yin, 2001) comb. nov.; Eidophelus indicus (Wood, 1989) comb. nov.; Eidophelus insularis (Nobuchi, 1975) comb. nov.; Eidophelus insularum (Krivolutskaya, 1968) comb. nov.; Eidophelus jalappae (Letzner, 1849) comb. nov.; Eidophelus javanus (Schedl, 1942) comb. nov.; Eidophelus kanawhae (Hopkins, 1915) comb. nov.; Eidophelus landolphiae (Schedl, 1961) comb. nov.; Eidophelus leprosulus (Browne, 1974) comb. nov.; Eidophelus longipennis (Eggers, 1936) comb. nov.; Eidophelus magnocularis (Yin, 2001) comb. nov.; Eidophelus marquesanus (Beeson, 1935) comb. nov.; Eidophelus mauritianus (Schedl, 1965) comb. nov.; Eidophelus micans (Eggers, 1927) comb. nov.; Eidophelus minor (Eggers, 1927) comb. nov.; Eidophelus minutissimus (Schedl, 1943) comb. nov.; Eidophelus mus (Schedl, 1975) comb. nov.; Eidophelus nanulus (Wood, 1960) comb. nov.; Eidophelus nigellatus (Schedl, 1950) comb. nov.; Eidophelus nubilus (Wood, 1960) comb. nov.; Eidophelus ocularis (Schedl, 1965) comb. nov.; Eidophelus onyanganus (Schedl, 1941) comb. nov.; Eidophelus opacus (Schedl, 1959) comb. nov.; Eidophelus pacificus (Schedl, 1941) comb. nov.; Eidophelus papuanus (Schedl, 1974) comb. nov.; Eidophelus papuensis (Wood, 1989) comb. nov.; Eidophelus paradoxus (Wood, 1992) comb. nov.; Eidophelus parvus (Hopkins, 1915) comb. nov.; Eidophelus pityophthorinus (Schedl, 1943) comb. nov.; Eidophelus pleiocarpae (Schedl, 1957) comb. nov.; Eidophelus polisquamosus (Yin, 2001) comb. nov.; Eidophelus praeda (Browne, 1978) comb. nov.; Eidophelus puerarae (Choo and Woo, 1989) comb. nov.; Eidophelus pumilionides (Schedl, 1977) comb. nov.; Eidophelus pumilus (Wood, 1960) comb. nov.; Eidophelus punctatulus (Nobuchi, 1976) comb. nov.; Eidophelus punctatus (Schedl, 1951) comb. nov.; Eidophelus puncticollis (Schedl, 1950) comb. nov.; Eidophelus pygmaeolus (Schedl, 1971) comb. nov.; Eidophelus quadridens (Browne, 1983) comb. nov.; Eidophelus ramosus (Beeson, 1935) comb. nov.; Eidophelus robustus (Schedl, 1955) comb. nov.; Eidophelus rugosus (Schedl, 1943) comb. nov.; Eidophelus rusticus (Wood, 1974) comb. nov.; Eidophelus semenovi (Kurentsov, 1941) comb. nov.; Eidophelus separandus (Schedl, 1965) comb. nov.; Eidophelus setifer (Wood, 1974) comb. nov.; Eidophelus sodalis (Schedl, 1965) comb. nov.; Eidophelus spessivtzevi (Berger, 1917) comb. nov.; Eidophelus spirostachius (Schedl, 1958) comb. nov.; Eidophelus splendens (Schedl, 1975) comb. nov.; Eidophelus squamatilis (Schedl, 1977) comb. nov.; Eidophelus squamosus (Schedl, 1942) comb. nov.; Eidophelus squamulosus (Eggers, 1936) comb. nov.; Eidophelus stephegynis (Hopkins, 1915) comb. nov.; Eidophelus takahashii (Nobuchi, 1975) comb. nov.; Eidophelus tarawai (Beaver, 1990) comb. nov.; Eidophelus tonsus (Schedl, 1969) comb. nov.; Eidophelus tricolor (Lea, 1910) comb. nov.; Eidophelus trucis (Wood, 1974) comb. nov.; Eidophelus uncatus (Schedl, 1971) comb. nov.; Eidophelus usagaricus (Eggers, 1922) comb. nov.; Eidophelus varius (Schedl, 1975) comb. nov.; Eidophelus venustus (Schedl, 1953) comb. nov.; Eidophelus yunnanensis (Yin, 2001) comb. nov.; Eidophelus zachvatkini (Krivolutskaya, 1958) comb. nov.; Ernoporus corpulentus (Sampson, 1919) comb. nov.; Ernoporus exquisitus (Bright, 2019) comb. nov.; Ernoporus guiboutiae (Schedl, 1957) comb. nov.; Ernoporus minutus (Bright and Torres, 2006) comb. nov.; Hypothenemus attenuatus (Eggers, 1935) comb. nov.; Hypothenemus loranthus (Schedl, 1942) comb. nov.; Hypothenemus novateutonicus (Schedl, 1951) comb. nov.; Hypothenemus pullus (Wood, 1971) comb. nov. Following assessment of diagnostic characters, the following species were transferred to a different genus: Afrocosmoderes madagascariensis Schedl, 1961 comb. nov.; Afrocosmoderes caplandicus (Schedl, 1965) comb. nov.; Afrocosmoderes grobleri (Schedl, 1961) comb. nov.; Afrocosmoderes niger (Schedl, 1961) comb. nov.; Afrocosmoderes pellitus (Schedl, 1953) comb. nov.; Afrocosmoderes pennatus (Schedl, 1953) comb. nov.; Eidophelus concentralis (Schedl, 1975) comb. nov.; Eidophelus inermis (Browne, 1984) comb. nov.; Eidophelus insignis (Browne, 1984) comb. nov.; Eidophelus kinabaluensis (Bright, 1992) comb. nov.; Eidophelus philippinensis (Schedl, 1967) comb. nov.; Eidophelus podocarpi (Bright, 1992) comb. nov.; Ernoporus imitatrix (Schedl, 1977) comb. nov.; Ernoporus minor (Schedl, 1942) comb. nov.; Ernoporus parvulus (Eggers, 1943) comb. nov.; Indocryphalus sericeus (Schedl, 1942) comb. nov.; Macrocryphalus elongatus (Schedl, 1965) comb. nov.; Macrocryphalus punctipennis (Schedl, 1965) comb. nov.; Microcosmoderes shoreae (Schedl, 1953) comb. nov.; Stegomerus parvatis (Wood, 1974) comb. nov.; Stephanopodius dubiosus (Schedl, 1970) comb. nov. Twenty-nine secondary homonyms were created following genus synonymy, and are designated replacement names: Afrocosmoderes schedli Johnson nom. nov. (=Euptilius madagascariensis Schedl, 1963 syn. nov.); Cryphalus amplicollis Johnson nom. nov. (=Cryphalus laticollis Browne, 1984 syn. nov.); Cryphalus eggersi Johnson nom. nov. (=Cryphalus confusus Eggers, 1927 syn. nov.); Cryphalus fuscus Johnson nom. nov. (=Cryphalus cylindrus Browne, 1984 syn. nov.); Cryphalus gracilis Johnson nom. nov. (=Cryphalus laevis Browne, 1984 syn. nov.); Cryphalus luteus Johnson nom. nov. (=Margadillius fulvus Browne, 1984 syn. nov.); Cryphalus minusculus Johnson nom. nov. (=Hypocryphalus minutus Browne, 1980 syn. nov.); Cryphalus ozopemoides Johnson nom. nov. (=Hypocryphalus montanus Schedl, 1974 syn. nov.); Cryphalus pellicius Johnson nom. nov. (=Hypocryphalus pilifer Schedl, 1979 syn. nov.); Cryphalus punctistriatulus Johnson nom. nov. (=Cryphalus striatulus Browne, 1981 syn. nov.); Cryphalus schedli Johnson nom. nov. (=Hypocryphalus formosanus Schedl, 1952 syn. nov.); Cryphalus solomonensis Johnson nom. nov. (=Margadillius terminaliae Browne, 1984 syn. nov.); Cryphalus spissepilosus Johnson nom. nov. (=Cryphalus densepilosus Schedl, 1943 syn. nov.); Cryphalus storckiellae Johnson nom. nov. (=Cryphalus striatus Browne, 1974 syn. nov.); Cryphalus takahashii Johnson nom. nov. (=Euptilius exiguus Browne, 1984 syn. nov.); Eidophelus alstoniae Johnson nom. nov. (=Chiloxylon sumatranus Schedl, 1970 syn. nov.); Eidophelus brighti Johnson nom. nov. (=Hemicryphalus minutus Bright, 1992 syn. nov.); Eidophelus brownei Johnson nom. nov. (=Euptilius papuanus Browne, 1983 syn. nov.); Eidophelus furvus Johnson nom. nov.(=Cryphalophilus ater Schedl, 1972 syn. nov.); Eidophelus levis Johnson nom. nov. (=Eidophelus gracilis Browne, 1984 syn. nov.); Eidophelus lucidus Johnson nom. nov. (=Lepicerinus pacificus Schedl, 1959 syn. nov.); Eidophelus minusculus Johnson nom. nov. (=Eidophelus minutissimus Schedl, 1962 syn. nov.); Eidophelus niger Johnson nom. nov. (=Ernoporicus ater Nobuchi, 1975 syn. nov.); Eidophelus parvulus Johnson nom. nov. (=Cryphalus parvus Browne, 1984 syn. nov.); Eidophelus rhododendri Johnson nom. nov. (=Hemicryphalus squamosus Bright, 1992 syn. nov.); Eidophelus schedli Johnson nom. nov. (=Cryphalomorphus ceylonicus Schedl, 1959 syn. nov.); Eidophelus yinae Johnson nom. nov. (=Scolytogenes venustus Yin, 2001 syn. nov.); Hypothenemus marginatus Johnson nom. nov. (=Periocryphalus sobrinus Wood, 1974 syn. nov.); Hypothenemus squamosulus Johnson nom. nov. (=Ptilopodius squamosus Schedl, 1953 syn. nov.). Two replacement names are now unnecessary: Cryphalus striatulus (Browne, 1978) stat. res. (=Hypothenemus brownei Beaver, 1991 syn. nov.); Macrocryphalus oblongus Nobuchi, 1981 stat. res. (=Hypothenemus nobuchii Knizek, 2011 syn. nov.). We also acknowledge the original description of several species by Eichhoff, 1878a which have been widely referenced as a later description (Eichhoff, 1878b). The following taxonomic changes are provided to acknowledge the changes: Cryphalus horridus Eichhoff, 1878a (=Cryphalus horridus Eichhoff, 1878b syn. nov); Cryphalus numidicus Eichhoff, 1878a (=Cryphalus numidicus Eichhoff, 1878b syn. nov); Cryphalus submuricatus Eichhoff, 1878a (=Cryphalus submuricatus Eichhoff, 1878b syn. nov); Eidophelus aspericollis (Eichhoff, 1878a) (=Eidophelus aspericollis Eichhoff, 1878b syn. nov); Hypothenemus arundinis (Eichhoff, 1878a) (=Hypothenemus arundinis Eichhoff, 1878b syn. nov); Hypothenemus birmanus (Eichhoff, 1878a) (=Hypothenemus birmanus Eichhoff, 1878b syn. nov); Hypothenemus fuscicollis (Eichhoff, 1878a) (=Hypothenemus fuscicollis Eichhoff, 1878b syn. nov); Hypothenemus rotundicollis (Eichhoff, 1878a) (=Hypothenemus rotundicollis Eichhoff, 1878b syn. nov). Subjective species-level changes are minimal.The following synonymies are proposed: Cryphalus papuanus (Schedl, 1973) (=Ernoporus antennarius Schedl, 1974 syn. nov.); Eidophelus concentralis (Schedl, 1975) (=Margadillius concentralis Schedl, 1975 syn. nov.). A neotype for Periocryphalus sobrinus Wood, 1974 and its replacement name Hypothenemus marginatus nom. nov. is designated at USNM due to the holotype being lost and replaced with a different species.