| Abstrakt: |
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 Nüsslin, 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 Nüsslin, 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 Brèthes, 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 Knížek, 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. [ABSTRACT FROM AUTHOR] |
