Classification and taxonomy of the nitidulid-group of families (Coleoptera: Cucujoidea): comments on morphology, bionomics, phylogeny and methodology, with a key to species of the subgenus Myothorax Murray, 1864 of the genus Carpophilus Stephens, 1830 (Nitidulidae: Carpophilinae)

The paper provides an overview of the author’s methodological opinion, which became the basis for developing the system and phylogeny of the nitidulid-group of families. The proposals for changing the system and phylogeny of these families are discussed, including the justification for using the taxon Nitiduloidea. The most reliable evidence of ancient divergences of these families can be the structure of the genitalia of both sexes, and therefore, in a number of cases, fossils without exposed genitalia can be arranged to a certain supraspecies group only tentatively. According to the structure of the genitalia of both sexes, two subgroups of families are distinguished: kateretid-subgroup (Apophisandridae, Boganiidae, Kateretidae, Parandrexidae, and Smicripidae) and nitidulid-subgroup (Helotidae, Monotomidae, and Nitidulidae), which reveal significant antiquity and many parallelisms in structure and bionomy and seem to have diverged earlier than the Middle Jurassic. The family Nitidulidae is also clearly divided into two phyletic lineages based on the structure of the male genitalia: nitiduline-lineage (Cillaeinae, Cryptarchinae, Cybocephalinae, Maynipeplinae, and Nitidulinae) and carpophiline-lineage (Amphicrossinae, Calonecrinae, Carpophilinae, and Epuraeinae). Recent proposals for changing the system and phylogeny are discussed. These do not take into account the fundamental features discussed in this paper. They were obtained after preliminary comparisons of some sequencing that require careful additional checks and clarifications. There are also disscussed the independent appearance of trophic interactions with the plant generative organs in representatives of many groups of the considered families; misinterpretations of these interactions have often led to hasty and insufficiently substantiated conclusions, including serious errors in constructing classifications and phylogenetic proposals. The latter methodological defects require verification, which can be achieved by the principle (method) of multiple parallelisms. This principle harmoniously complements the concept of integrated taxonomy and phylogenomics, providing the latter with an objective basis and a method of verification. The possible role of participation of representatives of the considered families in pollination of gymnosperms and angiosperms in the past, starting from the Middle Jurassic and up to the present day, are discussed. Using the method of multiple parallelisms, it has been shown that if the nitidulid-group of families has a common origin, then the kateretid-subgroup should be considered as mainly Mesozoic with some genera represented in the modern biota, then the nitidulid-subgroup should be considered, despite the antiquity of its origin, as prosperous in the Cenozoic, having given rise to a large number of modern forms with imaginal or complete anthophagy. At the same time, modern nitidulids, which usually live on dioecious palms, exhibit greatest convergent similarity of many structures with those of Mesozoic apophisandrids.
An overview of the dimidiatus-group of species of the subgenus Myothorax Murray, 1864 of the genus Carpophilus Stephens, 1830 is presented. This group includes widespread food pests that cause significant economic damages. Identification of these species has so far been an almost insoluble problem for pest control specialists. A key to these pests facilitating their identification is developed. New taxa are proposed: subfamily Vetunitidulinae subfam. n. (Apophisandridae, type genus Vetunitidula Zhao, Engel, Huang et Cai, 2025), tribe Plesiogethini trib. n. (Cybocephalinae, type genus Plesiogethes Zaitsev, Vasilenko et Perkovsky, 2025) and the genus Mesohelotopsis gen. n. (Helotidae, type species Metahelotella monochromata Liu, Ślipiński, Ren et Pang, 2019), as a result a new combination is established: Mesohelotopsis monochromata (Liu, Ślipiński, Ren et Pang, 2019), comb. n. The following new species are described: Carpophilus (Myothorax) assignatus sp. n. (Malaysia: Kalimantan, Sabah; Indonesia: Sulawesi), C. (M.) fumatoides sp. n. (India: Karnataka; Laos), C. (M.) generosus sp. n. (Indonesia: Java, Bali), and C. (M.) languescens sp. n. (Cambodia). In addition, new synonymy is introduced for the following species: Epuraeinae: Epuraea (Epuraea) excisicollis Reitter, 1872 = E. (E.) dolosa Kirejtshuk, 1995, syn. n.; Carpophilinae: Carpophilus (Ecnomorphus) plagiatipennis (Motschulsky, 1858) = C. (Ecnomorphus) jahari Dasgupta et Pal, 2019, syn. n., Carpophilus (Myothorax) contegens (Walker, 1858) = C. (M.) maculatus Murray, 1864, syn. n. = C. (M.) vittiger var. nigritus Murray, 1864, syn. n. = C. (M.) vittiger var. testaceus Murray, 1864, syn. n. = C. (M.) vittiger var. dilutus Murray, 1864, syn. n., non Colastus dilutus Motschulsky, 1858, Carpophilus (Myothorax) fusciceps Grouvelle, 1913 = C. (M.) scotti Grouvelle, 1913, syn. n., Carpophilus (Myothorax) lewisi Reitter, 1884 = C. (M.) signatus Grouvelle, 1908, syn. n. = C. (M.) signatus var. ornatus Grouvelle, 1908, syn. n. = C. (M.) subcalvus Kirejtshuk, 1984, syn. n., Carpophilus (Myothorax) schioedtei Murray, 1864 = C. (M.) pallescens Murray, 1864, syn. n. = C. (M.) vittiger var. limbalis Murray, 1864, syn. n., Carpophilus (Myothorax) pilipennis Macleay, 1873 = C. (M.) davidsoni Dobson, 1952, syn. n.; Nitidulinae: Aethina (Aethina) aeneipennis Reitter, 1873 = A. (A.) zhizhuaca Chen et Huang, 2024, syn. n., Atarphia quadripunctata Reitter, 1884 = A. cincta Jelínek et Hájek, 2012, syn. n., Physoronia wajdelota (Wankowicz, 1869) = P. japonica Reitter, 1873, syn. n., Pocadius nobilis Reitter, 1873 = P. fasciatus Cline, 2008, syn. n. = P. okinawaensis Cline, 2008, syn. n. = P. tenebrosus Chen et Huang, 2020, syn. n. = P. zhangjiajieensis Chen et Huang, 2020, syn. n.; Cryptarchinae: Glischrochilus (Glischrochilus) quadripunctatus (Linnaeus, 1758) = G. (G.) tremulae Clayhills, Audisio et Cline, 2016, syn. n.; Cybocephalinae: Cybocephalus bicinctus Kirejtshuk, 1988 = C. chlorocapitis Hisamatsu, 2013, syn. n. Besides, lectotypes are designated for Aethina suturalis Reitter, 1884, Carpophilus lewisi Reitter, 1884, C. ochropterus Boheman, 1851, C. pilosellus Motschulsky, 1858, C. (Ecnomorphus) nigricans Grouvelle, 1897, C. (Eidocolastus) bosschae Grouvelle, 1892, C. (Myothorax) vittiger var. limbalis Murray, 1864, C. (M.) luridus Murray, 1864, C. (M.) maculatus Murray, 1864, C. (M.) vittiger var. nigritus Murray, 1864, C. (M.) nepos Murray, 1864, C. (M.) notatus Murray, 1864, C. (M.) pallescens Murray, 1864, C. (M.) vittiger var. robustus Murray, 1864, C. (M.) schioedtei Murray, 1864, C. (M.) vittiger var. testaceus Murray, 1864, C. (M.) vittiger Murray, 1864, Colastus dilutus Motschulsky, 1858, non Carpophilus (Myothorax) vittiger var. dilutus Murray, 1864, Colastus plagiatipennis Motschulsky, 1858, Nitidula contegens Walker, 1858, Nitidula hemiptera Fabricius, 1792, non Dermestes hemipterus Linnaeus, 1758. The name Carpophilus (Myothorax) robustus Murray, 1864, stat. n. should be used as valid for a separate species, which was originally proposed to designate a variety of C. (M.) vittiger, however, according to the studied lectotypes, the latter should be recognized as a junior synonym of C. (M.) contegens.