Revision of the Bark Beetle Genera Within the Former Cryphalini (Curculionidae: Scolytinae) Author Johnson, Andrew J. School of Forest Resources and Conservation, University of Florida, Gainesville, FL 32611, andrewjavanjohnson@gmail.com Author Hulcr, Jiri School of Forest Resources and Conservation, University of Florida, Gainesville, FL 32611, Author Knížek, Miloš Department of Forest Protection Service, Forestry and Game Management Research Institute, Strnady, Jíloviště, Praha 5, Zbraslav CZ- 15600, Czechia, Author Atkinson, Thomas H. Texas Natural History Collections, Integrative Biology, University of Texas at Austin, Austin, TX 78712, Author Mandelshtam, Michail Yu. Department of Forest Protection, Wood Science and Game Management, Saint-Petersburg State Forest Technical University named after S. M. Kirov, Institutskii per., 5, 194021 Saint-Petersburg, Russia, Author Smith, Sarah M. Department of Entomology, Michigan State University, East Lansing, MI 48824, Author Cognato, Anthony I. Department of Entomology, Michigan State University, East Lansing, MI 48824, Author Park, Sangwook Research Institute of Forest Insect Diversity, Namyangju 12113, South Korea, Author Li, You School of Forest Resources and Conservation, University of Florida, Gainesville, FL 32611, Author Jordal, Bjarte H. University Museum of Bergen, University of Bergen, P. O. 7800, 5020 Bergen, and text Insect Systematics and Diversity 2020 2020-05-31 4 3 1 1 81 journal article 22148 10.1093/isd/ixaa002 be792f86-9e77-414c-9e2f-767704ff704e 2399-3421 3826789 Ernoporus Thomson, 1859: 147 ( Figs. 27–29 ) Synonymy = Cryphalops Reitter, 1889: 94 . = Stephanorhopalus Hopkins, 1915: 35 . = Euptilius Schedl, 1940c: 589 . = Ernocladius Wood, 1980: 93 syn. nov. = Allothenemus Bright and Torres, 2006: 400 syn. nov. Type of genus Apate tiliae Panzer, 1793 . Diagnosis This genus can be diagnosed by the combination of the entire eye, by the pronotum with concentric rows of asperities, by the distinct summit, and by the proventriculus which has anteriorly pointing spines posterior to the masticatory brush. Female Robust body shape, less than 2.1 times as long as wide. Eye oval shaped. Antennae with three or four funicle segments. Antennal club in most species with sutures straight to profoundly procurved. Pronotum with two or more marginal asperities. Asperities on the pronotum always arranged in concentric rows, leading to a distinct summit. Sides of pronotum rounded, sometimes with a weakly visible carina extending slightly from the posterior edge. Elytra with scale-like interstrial bristles. Apex of elytra vertical. Proventriculus with a simple, short apical plate. Crop spines hair-like. An area of anteriorly pointing spine-shaped or spatula-shaped setae is present beyond the masticatory brush. Male Similar to females, except for a more produced anterior margin of the pronotum, typically with feather-like setae on the protarsi and two or more spines on the posteroventral margin of the seventh tergite. Penis apodemes much shorter than penis body, fused at apex. Tegmen open dorsally, much narrower than penis apodemes. Tegminal apodemes absent. Spiculum gastrale thicker than penis apodemes, with a fork. Basal sclerites sometimes visible. Sometimes large hook-like end plates are present. Distribution Asia, Europe, Africa, Caribbean (likely introduced), Australia (introduced). Remarks Nineteen species known. Ernoporus are most easily recognized by the stout appearance with concentric asperities on the pronotum, though a few members of Eidophelus also share this feature, as well as Acorthylus and Neocryphus . The tuft of setae posterior to the proventriculus has not been observed in any other Ernoporini, but is present in other genera such as Acorthylus and at least some Stegomerus species. Several species were previously placed in the genus Euptilius , presented separately in Fig. 25 . These are characterized by antennal club with profoundly procurved sutures and extensive split setae on the hypomeron. No fresh specimens were available for study and no genetic information exists. However, based on the external morphology as well as a dissection of the proventriculus, there was no justification for resurrecting this genus based on the information available ( Wood 1980 ). All the characters which are distinctive are present in other Ernoporus species (procurved antennal sutures, split setae on the pronotum), although to a much lesser extent. Members of this species group have a very similar appearance to Neocryphus , but differ in the antennal club and the eye shape. The genera Ernocladius and Allothenemus closely match the characters for this genus. Ernocladius was described as distinct from Ernoporus based on the interstrial ground vestiture being absent on the elytral disc and sparse or absent on the declivity, by the antennal funicle with only two segments, and by the procurved sutures of the antennal club ( Wood 1980 ). This combination of characters is not consistent within the groups, and is not stable across other genera. Allothenemus was described based on the unique morphology compared to former Cryphalini from the Americas, but also matches the diagnostic characters of Ernocladius and closely matches some Asian species. It is likely that the species represent non-native species based on the distribution and the close similarity to Margadillius minor Schedl, 1942 . Erioschidias imitatrix Schedl, 1977 is transferred from Cosmoderes , supported by internal and external morphology and molecular phylogenetics. Several species formerly in the genus Margadillius have been moved here. The previous misclassification is unsurprising, since a specimen in Schedl’s collection (NHMW) labeled as a homeotype (which is usually specimen that was directly compared to and that matches the primary type ) of Margadillius margadilaonis , was in fact a misidentified member of Ernoporus . Type material examined Holotype of Euptilius armatus Browne, 1981 ( BMNH ) ; holotype and paratype of Ernoporus concentralis Eggers, 1936 ( BMNH ) ; allotypes (x2) of Cryphalus corpulentus Sampson, 1919 ( BMNH ) ; holotype of Erioschidias imitatrix Schedl, 1977 ( NHMW ) ; holotype of Ernoporus japonicus Nobuchi, 1966 ( ITLJ ) ; syntypes (x2) of Margadillius minor Schedl, 1942 ( NHMW and BMNH ) ; holotype of Margadillius parvulus Eggers, 1943 ( NHMW ) ; paratype of Euptilius thailandicus Schedl, 1967 ( NHMW ) ; holotype of Euptilius tuberculatus Browne, 1981 ( BMNH ) . Included species Ernoporus acanthopanaxi ( Niisima, 1913: 4 ) ( Cryphalus ) . Ernoporus armatus ( Browne, 1981: 129 ) ( Euptilius ) . Ernoporus concentralis Eggers, 1936c: 629 . Ernoporus corpulentus ( Sampson, 1919: 113 ) ( Cryphalus ) comb. nov. [ Ernocladius ]. = Margadillius corpulentus sundri Schedl, 1969: 48 (syn: Wood, 1989 ). Ernoporus dispar ( Schedl, 1972f: 49 ) ( Cryphalops ) . Ernoporus exquisitus ( Bright, 2019: 105 ) ( Allothenemus ) comb. nov. [ Allothenemus ]. Ernoporus guiboutiae ( Schedl, 1957b: 53 ) ( Miocryphalus ) comb. nov. [ Ernocladius ]. Ernoporus imitatrix ( Schedl, 1977c: 499 ) ( Erioschidias ) comb. nov. [ Cosmoderes ]. Ernoporus inermis ( Schedl, 1939b: 343 ) ( Stephanorhopalus ) . Ernoporus japonicus Nobuchi, 1966b: 52 . Ernoporus melodori ( Hopkins, 1915: 36 ) ( Stephanorhopalus ) . Ernoporus minor ( Schedl, 1942a: 176 ) ( Margadillius ) comb. nov. [ Margadillius ]. Ernoporus minutus ( Bright and Torres, 2006: 400 ) ( Allothenemus ) comb. nov. [ Allothenemus ]. Ernoporus parvulus ( Eggers, 1943b: 75 ) ( Margadillius ) comb. nov. [ Margadillius ]. Ernoporus quadridens ( Schedl, 1971a: 284 ) ( Cryphalops ) . Ernoporus shimanensis Murayama, 1953: 36 . Ernoporus thailandicus ( Schedl, 1967a: 127 ) ( Euptilius ) . Ernoporus tiliae ( Panzer, 1793: 14 ) ( Apate ) . = Cryphalus ratzeburgi Ferrari, 1867: 11 . = Cryphalus lederi Reitter, 1889: 93 . = Ernoporus eggersi Kurentsov, 1941: 155 (syn: Sokanovskiy, 1954 ). = Ernoporus starki Eggers, 1942: 31 (syn: Schedl, 1952e ). Ernoporus tuberculatus ( Browne, 1981: 128 ) ( Euptilius ) .