Background: Familial hypercholesterolemia is a human monogenic disease caused by populationspecific mutations in the low density lipoprotein (LDL) receptor gene. Despite thirteen different mutations of the LDL receptor gene were reported from Russia prior to 2003, the whole spectrum of disease-causing gene alterations in this country is poorly known and requires further investigation provided by the current study.
G197del is the most prevalent LDL receptor (LDLR) mutation causing familial hypercholesterolemia (FH) in Ashkenazi Jew (AJ) individuals. The purpose of this study was to determine the origin, age, and population distribution of G197del, as well as to explore environmental and genetic effects on disease expression. Index cases from Israel (n=46), South Africa (n=24), Russia (n=7), The Netherlands (n=1), and the United States (n=1) were enlisted. All trace their ancestry to Lithuania. A highly conserved haplotype (D19S221:104-D19S865:208-D19S413:74) was identified in G197del chromosomes, suggesting the occurrence of a common founder. When two methods were used for analysis of linkage disequilibrium (LD) between flanking polymorphic markers and the disease locus and for the study of the decay of LD over time, the estimated age of the deletion was found to be 20 +/- 7 generations (the 95% confidence interval is 15-26 generations), so that the most recent common ancestor of the mutation-bearing chromosomes would date to the 14th century. This corresponds with the founding of the Jewish community of Lithuania (1338 a.d.), as well as with the great demographic expansion of AJ individuals in eastern Europe, which followed this settlement. The penetrance of mutation-linked severe hypercholesterolemia is high (94% of heterozygotes have a baseline concentration of LDL cholesterol (LDL-C) that is >160 mg/dl), and no significant differences in the mean baseline lipid level of G197del carriers from different countries were found. Polymorphisms of apolipoprotein E and of scavenger-receptor class B type I were observed to have minor effects on the plasma lipid profile. With respect to determinative genetic influences on the biochemical phenotype, there is no evidence that could support the possibility of a selective evolutionary metabolic advantage. Therefore, the founder effect in a rapidly expanding population from a limited number of families remains a simple, parsimonious hypothesis explaining the spread of G197del-LDLR-linked FH in AJ individuals.
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 CryphalusErichson, 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 EidophelusEichhoff, 1876 (=Scolytogenes Eichhoff, 1878 syn. nov.; PtilopodiusHopkins, 1915syn. nov.; ErnoporicusBerger, 1917syn. nov.; CryphalogenesWood, 1980syn. nov.); ErnoporusThomson, 1859 (=ErnocladiusWood, 1980syn. nov.; AllothenemusBright and Torres, 2006syn. nov.); Hemicryphalus Schedl, 1963; and ProcryphalusHopkins, 1915. Trypophloeini Nüsslin, 1911 stat. res. includes the genera Afrocosmoderes Johnson and Jordal gen. nov.; AtomothenemusBright, 2019; Cosmoderes Eichhoff, 1878 (=AllernoporusKurentsov, 1941syn. nov.); HypothenemusWestwood, 1834 (=PeriocryphalusWood, 1971syn. nov.); MacrocryphalusNobuchi, 1981stat. res.; Microcosmoderes Johnson and Jordal gen. nov.; MicrosomusBright, 2019; PygmaeoborusBright, 2019; and TrypophloeusFairmaire, 1864. Xyloterini LeConte, 1876 is maintained, containing Indocryphalus Eggers, 1939; TrypodendronStephens, 1830 and XyloterinusSwaine, 1918. AcorthylusBrèthes, 1922, CryptocarenusEggers, 1937, Neocryphus Nunberg, 1956, Stegomerus Wood, 1967, and TrypolepisBright, 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 basihirtusBeeson, 1929comb. 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 mangiferaeStebbing, 1914comb. 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 montanusSchedl, 1974syn. nov.); Cryphalus pellicius Johnson nom. nov. (=Hypocryphalus pilifer Schedl, 1979 syn. nov.); Cryphalus punctistriatulus Johnson nom. nov. (=Cryphalus striatulusBrowne, 1981syn. 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 densepilosusSchedl, 1943syn. nov.); Cryphalus storckiellae Johnson nom. nov. (=Cryphalus striatusBrowne, 1974syn. 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 minutusBright, 1992syn. 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 aterNobuchi, 1975syn. nov.); Eidophelus parvulus Johnson nom. nov. (=Cryphalus parvus Browne, 1984 syn. nov.); Eidophelus rhododendri Johnson nom. nov. (=Hemicryphalus squamosusBright, 1992syn. nov.); Eidophelus schedli Johnson nom. nov. (=Cryphalomorphus ceylonicus Schedl, 1959 syn. nov.); Eidophelus yinae Johnson nom. nov. (=Scolytogenes venustusYin, 2001syn. 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 browneiBeaver, 1991syn. nov.); Macrocryphalus oblongusNobuchi, 1981stat. res. (=Hypothenemus nobuchiiKnížek, 2011syn. 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 horridusEichhoff, 1878a (=Cryphalus horridusEichhoff, 1878bsyn. nov); Cryphalus numidicusEichhoff, 1878a (=Cryphalus numidicusEichhoff, 1878bsyn. nov); Cryphalus submuricatusEichhoff, 1878a (=Cryphalus submuricatusEichhoff, 1878bsyn. nov); Eidophelus aspericollis (Eichhoff, 1878a) (=Eidophelus aspericollisEichhoff, 1878bsyn. nov); Hypothenemus arundinis (Eichhoff, 1878a) (=Hypothenemus arundinisEichhoff, 1878bsyn. nov); Hypothenemus birmanus (Eichhoff, 1878a) (=Hypothenemus birmanusEichhoff, 1878bsyn. nov); Hypothenemus fuscicollis (Eichhoff, 1878a) (=Hypothenemus fuscicollisEichhoff, 1878bsyn. nov); Hypothenemus rotundicollis (Eichhoff, 1878a) (=Hypothenemus rotundicollisEichhoff, 1878bsyn. nov). Subjective species-level changes are minimal. The following synonymies are proposed: Cryphalus papuanus (Schedl, 1973) (=Ernoporus antennariusSchedl, 1974syn. nov.); Eidophelus concentralis (Schedl, 1975) (=Margadillius concentralis Schedl, 1975 syn. nov.). A neotype for Periocryphalus sobrinus Wood, 1974 and its replacement name Hypothenemus marginatusnom. nov. is designated at USNM due to the holotype being lost and replaced with a different species.
Cylindrobrotus pectinatusgen. et sp.n., a new scolytine species from Cretaceous Lebanese amber, is described. A new tribe, Cylindrobrotini trib.n., is proposed for this unique species, which demonstrates an unusual combination of some archaic and many advanced characters. This finding suggests that the Scolytinae became a distinct lineage of Curculionoidea from the Lower Cretaceous. Fossil records are reviewed, and some remarks on the origin and taxonomic position of bark and ambrosia beetles are made. Some comments on the various phylogenetic interpretations of the last 30 years are given, particularly in respect of their correspondence with the fossil record. The early appearance of Scolytinae in the fossil record before other Curculionidae (which appeared in the Upper Cretaceous) can be used as evidence against the hypothesis of bark beetles as offspring of weevils. The question of the taxonomic rank of bark beetles (separate subfamily or family) and their placement among other groups of the superfamily remains unsolved.
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