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Helotidae is a small and morphologically uniform family in Nitiduloidea. In this study, we report an unusual form of helotids, represented by Lobatihelota lescheni Li, Liu & Cai gen. nov., sp. nov. and L. iridescens Li, Liu & Cai sp. nov. from mid-Cretaceous Burmese amber. Lobatihelota is unique within the family in having a leg morphology typical of some Nitidulidae and Kateretidae (tibia widened distally, tarsomeres 1–3 bilobed, tarsomere 4 shortened). Additionally, new diagnostic characters are suggested for Trihelota from the same deposit (prosternum and mesoventrite with paired carinae, metanepisterna short, metacoxae meeting elytra laterally), and the morphological divergence of representative genera of the superfamilies Erotyloidea, Nitiduloidea and Cucujoidea is visualised using a phylomorphospace approach. ZooBank: urn:lsid:zoobank.org:pub:FE0E4D6A-8157-4E14-9240-7B87F285AEA5 Helotidae is a small family in Nitiduloidea. We report an unusual form of helotids, represented by Lobatihelota lescheni gen. nov., sp. nov. and L. iridescens sp. nov., from mid-Cretaceous Burmese amber. Lobatihelota is unique within the family in having a leg morphology typical of some Nitidulidae and Kateretidae. Additionally, new diagnostic characters are suggested for Trihelota from the same deposit, and the morphological divergence of representative genera of Erotyloidea, Nitiduloidea and Cucujoidea is visualised using a phylomorphospace approach. [ABSTRACT FROM AUTHOR]

Simple Summary: The family Helotidae represents a unique and primitive group of Cucujoidea, with important implications in understanding the phylogeny of beetles. To better understand the characteristics of the helotid mitochondrial (mt) genome and the evolution of Cucujoidea, we sequenced and compared the first recorded Helotidae mt genomes to reveal their characteristics and reconstruct the phylogenetic relationships of 13 basal families of this group. Phylogenetic analysis of the mt genomes indicated the positions of seven families within Cucujoidea but did not statistically support the presence of the Erotylid series and the Nitidulid series as distinct groups in this superfamily. In the phylogenetic results, Helotidae and Protocucujidae are sister groups. This study provides a new phylogenetic hypothesis regarding the basal relations of Cucujoidea. Helotid beetles are commonly found in places where sap flows from tree trunks and in crevices in bark. The Helotidae family is a rare and primitive group of Cucujoidea. To date, no complete mitochondrial (mt) genome has been sequenced for this family. To better understand the characteristics of the mt genome and the evolution of Cucujoidea, we sequenced and annotated the complete mt genomes of Helota thoracica (Ritsema, 1895) and Helota yehi Lee, 2017 using next-generation sequencing. These are the first record of Helotidae mt genomes. The RNA secondary structures of both species were also predicted in this study. The mt genomes of H. thoracica and H. yehi are circular, with total lengths of 16,112 bp and 16,401 bp, respectively. After comparing the mt genomes of H. thoracica and H. yehi, we observed the gene arrangement, codon usage patterns, base content, and RNA secondary structures of both species to be similar, which has also been noted in other Coleoptera insects. The nucleotide sequence of the coding regions and the control region has small differences. The phylogenetic analysis indicated that Helotidae and Protocucujidae are sister groups and revealed the relationship between seven families; however, the validity of the two series (Erotylid series and Nitidulid series) as larger groups in the superfamily was not supported. The mt phylogenomic relationships have strong statistical support. Therefore, the division of Cucujoidea into series should be re-examined. Our results will provide a better understanding of the mt genome and phylogeny of Helotidae and Cucujoidea and will provide valuable molecular markers for further genetic studies. [ABSTRACT FROM AUTHOR]

The material of family Helotidae collected over 2008-2014 in the area of Mount Phu Pane, in the small part of northeastern Laos, was studied and the results are given. Twenty-two species of this family were recognized, including two species new to science described herein: Helota luciae sp. nov. (the Helota thibetana species group) and Helota catharinae sp. nov. (the Helota vigorsii species group). An updated key for three mentioned species groups sensu Lee (2010) including the newly described species is provided. Figures of protibiae, genitalia, internal sacs, eighths abdominal tergites, and fifth abdominal ventrites are given for all new species. New records from Laos are presented for the two new species only. [ABSTRACT FROM AUTHOR]

A new genus belonging to the cucujoid family Helotidae, Trihelota gen. nov., is described from mid-Cretaceous (ca. 99 million years old) amber from Kachin State, northern Myanmar, containing a single species T. fulvata sp. nov. The minute fossil was documented using fl uorescence microscopy and X-ray microtomography to reveal fi ne morphological details. Trihelota is characterised by its transverse head, antennal club about 1.5 times longer than wide, pronotum almost parallel-sided, and elytra not costate. Notably, each elytron possesses three coloured spots, a character unique within Helotidae. The new genus considerably expands the known morphological disparity of Cretaceous helotids. A key to extinct and extant genera of the family Helotidae is provided along with a checklist of fossil species. [ABSTRACT FROM AUTHOR]

Abstract Three new species of Helotidae, Palaeohelota parva gen. et sp. nov. from the Lower Cretaceous of Yixian Formation, Burmahelota pengweii gen. et sp. nov. and Metahelotella monochromata sp. nov. from the Upper Cretaceous of Northern Myanmar are described, with the first taxon representing the oldest fossil record of the family Helotidae. The diversity of Helotidae during the Cretaceous suggests the family originated and diversified at least in the Early Cretaceous. Laodiscis Lin, 1971 described in Helotidae from the Paleocene of China is here removed from Helotidae and transferred to Coleoptera, family incertae sedis. [ABSTRACT FROM AUTHOR]

Members of the culta species group of the genus Neohelota Ohta are revised. Ten species are recognized, including a new species described herein, N. barclayi sp. nov., from China, India, Laos, Nepal, Thailand, and Vietnam. The following synonymies are proposed: Helota klapperichi Mader, 1955, as a junior synonym of Neohelota chinensis (Mader, 1955); Helota taiwana Ohta, 1929, as a junior synonym of Neohelota boulei (Ritsema, 1915); Helota gestroi Ritsema, 1891, Helota sharpi Ritsema, 1915, Helota similis Lee and Satô, 2006, as junior synonyms of Neohelota pusilla (Oberthür, 1883); Helota notata Ritsema, 1891, and Helota severini Ritsma, 1893, as junior synonyms of Neohelota cula (Olliff, 1883); Helota hopei Ritsema, 1915, as a junior synonym of Neohelota fryi ( Ritsema, 1894); Helota olliffii Ritsema, 1899, as a junior synonym of Neohelota boysii (Ritsema, 1889); Helota bretaudeaui Ritsema, 1894, as a junior synonym of Neohelota affinis (Ritsema, 1891 ); Helota jordanii Ritsema, 1899, and Helota indicator Ritsema, 1905, as junior synonyms of Neohelota dubia (Ritsema, 1891). Lectotypes are designated for Helota chinensis Mader, 1955, H klapperichi, Helota boulei Ritsema, 1915, Helota consanguinea Ritsema, 1915, H. sharpi, H. notata, H. severini, H. olliffii, H. bretaudeaui, and H. jordanii. A key for the culta species group is provided. Illustrations of diagnostic characters are provided for all species. [ABSTRACT FROM AUTHOR]

The attenuata species group of Neoiielota Ohta is revised. Six species are recognized, including one described herein, N. vietnamensis sp. nov., from Vietnam and Laos. Helota moutonii Ritsema, 1905 is recognized as a junior synonym oiNeohelota renati (Ritsema, 1905), Helota bhutanensis Chujo, 1975, as a junior synonym of N. durelii (Ritsema, 1905), and H. sonani Ohta, 1929 as a junior synonym of N. letvisi (Ritsema, 1915). A lecotype is designated for Helota attenuata Ritsema, 1894. A key for the attenuata species group is provided. Figures of protibiae, genitalia, internal sacs, eighth abdominal tergites, and fifth abdominal ventrites are given for all species. [ABSTRACT FROM AUTHOR]

The candezei species group of Neohelota Ohta is revised. Six species are recognized, including one described herein, Neohelota jakli sp. nov., from Indonesia (Sumatra). Helota claudi Nguyen-Phung, 1985 is placed as junior synonym of Neohelota candezei (Ritsema, 1899), and Helot jocelynae Nguyen-Phung, 1985 as a junior synonym of Neohelota montana (Ohta, 1929). Lectotypes are designated for Helota krugeri Ritsema, 1900; Helota margarethae Ritsema, 1900; and Helota mari Ritsema, 1899. A key for the candezei species group is provided. Figures of protibiae, genitalia, internal sacs, eighth abdominal tergites, and fifth abdominal ventrites are given for all species. [ABSTRACT FROM AUTHOR]

Species of the genus Neohelota are classified into six species groups, and 60 species are recognized. Taxonomy of the laevigata and helleri species groups is presented in detail. Two new species are described: N. viklundi sp. nov. from Myanmar and hr. satoi sp. nov. from Laos. A lectotype is designated for N. tibialis Ritsema. Protibiae, genitalia, aedeagal internal sacs, eighth abdominal tergites, and fifth abdominal ventrites are illustrated for all species. [ABSTRACT FROM AUTHOR]

The species belonging to the genus Metahelotella Kirejtshuk are revised. Six species are recognized, including two described herein, M. schawalleri sp. nov. from Vietnam and M. sprecherae sp. nov. from China (Shaanxi Province). Helotafulvitarsis Ritsema and H. difficilis Ritsema are placed as junior synonyms of M. semifulva Ritsema. Lectotypes are designated for H. fulvitarsis Ritsema, H. difficilis Ritsema, and H. immaculata Ritsema. A key and diagnosis of the species belonging to the genus Metahelotella Kirejtshuk are provided. Figures of protibiae, genitalia, internal sacs, eighth abdominal tergites, and fifth abdominal ventrites are given for all species. [ABSTRACT FROM AUTHOR]

The thibetana species group of Helota MacLeay is revised. Five species are considered to be valid. The following synonyms are proposed: Helota sinensis Olliff, 1883 (=H. tonkinensis Ritsema, 1905a); and H. thibetana Westwood, 1842 (=H. verrucosa Ritsema, 1894; H. donckieri Ritsema, 1907; H. westwoodii Ritsema, 1907). Lectotypes are designated for Helota sinensis Olliff, H. tonkinensis Ritsema, H. donckieri Ritsema, and H. westwoodii Ritsema. A key for the thibetana species group is provided. Figures of the protibia, penis, parameres, internal sac, and eighth abdominal tergite are given for each species. A checklist of the 23 valid species and subspecies of the genus Helota MacLeay is provided. [ABSTRACT FROM AUTHOR]

The vigorsii species group of Helota MacLeay is revised. Ten species and one subspecies are recognized, including a new species H. brancuccii described, from Laos and Thailand. The following new synonyms are proposed: H. vandepolli Ritsema, 1891 (=Helota fruhstorferi Ritsema, 1905; H. lugubris Ritsema, 1914); H. servillei Hope, 1840 (=Helota oberthueri Ritsema, 1889); H. scintillans Olliff, 1884 (=Helota pasteuri Ritsema, 1893); Helota vigorsii vigorsii MacLeay, 1825 (=Helota vigorsii sumatrensis Ritsema, 1909). Lectotypes are designated for Helota oberthueri Ritsema, H. feai Ritsema, H. vigorsii vigorsii MacLeay, H. vigorsii sumatrensis Ritsema, H. vigorsii borneensis Ritsema, H. fairmairei Ritsema, and H. longipes Ritsema. A key and diagnosis for the Helota vigorsii MacLeay species group are provided. Figures of protibia, penis, parameres, internal sac, eighth abdominal tergite are given for all species. [ABSTRACT FROM AUTHOR]

The gemmata species group of Helota MacLeay is revised. Seven species are recognized, including two described herein, H. schuhi sp. nov. from China (Hubei Province) and H. ohbayashii sp. nov. from Laos and Thailand. H. distincta Ritsema and H. ritsemana Heller are placed as junior synonyms of H. fidviventris Kolbe; H. caudata Ritsema is placed as a junior synoym of H. gorhami Olliff. Lectotypes are designated for H. kolbei Ritsema, H. ventralis Ritsema, H. gemmata Gorham, H. fidviventris Kolbe, H. ritsemana Heller, H. distincta Ritsema, and H. gorhami Olliff. A key for the gemmata species group is provided. Figures of protibia, penis, parameres, internal sac, eighth abdominal tergite are presented for all species. [ABSTRACT FROM AUTHOR]