2 urn:lsid:arphahub.com:pub:45048D35-BB1D-5CE8-9668-537E44BD4C7E urn:lsid:zoobank.org:pub:91BD42D4-90F1-4B45-9350-EEF175B1727A ZooKeys ZK 1313-2989 1313-2970 Pensoft Publishers 10.3897/zookeys.1126.90233 90233 Research Article Animalia Arthropoda Coleoptera Elateroidea Hexapoda Insecta Invertebrata Polyphaga Rhagophthalmidae Catalogues and Checklists Faunistics & Distribution Nomenclature Systematics Afghanistan Asia Central Asia China China Seas Far East Indian Sub-Continent and Himalayas Indonesia Japan Laos Malaysia Myanmar Philippines Singapore Taiwan Thailand Vietnam Rhagophthalmidae Olivier, 1907 (Coleoptera, Elateroidea): described genera and species, current problems, and prospects for the bioluminescent and paedomorphic beetle lineage Kundrata Robin robin.kundrata@upol.cz https://orcid.org/0000-0001-9397-1030 1 Conceptualization Writing - original draft Writing - review and editing Data curation Formal analysis Funding acquisition Investigation Project administration Resources Supervision Validation Visualization Hoffmannova Johana https://orcid.org/0000-0003-0216-6031 1 Writing - review and editing Data curation Formal analysis Funding acquisition Investigation Resources Hinson Kevin R. https://orcid.org/0000-0003-3111-4513 2 Writing - review and editing Data curation Formal analysis Investigation Resources Validation Keller Oliver https://orcid.org/0000-0001-5067-3316 3 Writing - review and editing Data curation Formal analysis Investigation Resources Validation Packova Gabriela https://orcid.org/0000-0001-7949-619X 1 Writing - review and editing Data curation Formal analysis Funding acquisition Investigation Resources Visualization Department of Zoology, Faculty of Science, Palacky University, 17. listopadu 50, 77900, Olomouc, Czech Republic Palacky University Olomouc Czech Republic EpiLogic GmbH Agrarbiologische Forschung und Beratung, Hohenbachernstr. 19–21, 85354, Freising, Germany EpiLogic GmbH Agrarbiologische Forschung und Beratung Freising Germany Florida State Collection of Arthropods, Florida Department of Agriculture and Consumer Services, P.O. Box 147100, Gainesville, FL, 32614-7100, USA Florida Department of Agriculture and Consumer Services Gainesville United States of America

Corresponding author: Robin Kundrata (robin.kundrata@upol.cz)

Academic editor: Vinicius S. Ferreira

 2022 01 11 2022 1126 55 130 22EA20E1-A925-5729-AAA6-1C4099DDD686 0ABE7C8D-BD9C-44ED-89D3-CACB78D12AB9 7550811 12 07 2022 12 09 2022 Robin Kundrata, Johana Hoffmannova, Kevin R. Hinson, Oliver Keller, Gabriela Packova This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. http://zoobank.org/0ABE7C8D-BD9C-44ED-89D3-CACB78D12AB9

Rhagophthalmidae are a small beetle family known from the eastern Palaearctic and Oriental realms. Rhagophthalmidae are closely related to railroad worms (Phengodidae) and fireflies (Lampyridae) with which they share highly modified paedomorphic females and the ability to emit light. Currently, Rhagophthalmidae include 66 species classified in the following 12 genera: Bicladodrilus Pic, 1921 (two spp.), Bicladum Pic, 1921 (two spp.), Dioptoma Pascoe, 1860 (two spp.), Diplocladon Gorham, 1883 (two spp.), Dodecatoma Westwood, 1849 (eight spp.), Falsophrixothrix Pic, 1937 (six spp.), Haplocladon Gorham, 1883 (two spp.), Menghuoius Kawashima, 2000 (three spp.), Mimoochotyra Pic, 1937 (one sp.), Monodrilus Pic, 1921 (two spp. in two subgenera), Pseudothilmanus Pic, 1918 (two spp.), and Rhagophthalmus Motschulsky, 1854 (34 spp.). The replacement name Haplocladon gorhami Kundrata, nom. nov. is proposed for Diplocladon hasseltii Gorham, 1883b (described in subgenus Haplocladon) which is preoccupied by Diplocladon hasseltii Gorham, 1883a. The genus Reductodrilus Pic, 1943 is tentatively placed in Lampyridae: Ototretinae. Lectotypes are designated for Pseudothilmanus alatus Pic, 1918 and P. marginalis Pic, 1918. Interestingly, in the eastern part of their distribution, Rhagophthalmidae have remained within the boundaries of the Sunda Shelf and the Philippines demarcated by the Wallace Line, which separates the Oriental and Australasian realms. This study is intended to be a first step towards a comprehensive revision of the group on both genus and species levels. Additionally, critical problems and prospects for rhagophthalmid research are briefly discussed.

 Catalogue classification Drilidae Lampyridae neoteny Oriental Region Phengodidae IGA of the Faculty of Science, UP Olomouc Citation

Kundrata R, Hoffmannova J, Hinson KR, Keller O, Packova G (2022) Rhagophthalmidae Olivier, 1907 (Coleoptera, Elateroidea): described genera and species, current problems, and prospects for the bioluminescent and paedomorphic beetle lineage. ZooKeys 1126: 55–130. https://doi.org/10.3897/zookeys.1126.90233

 Introduction

Rhagophthalmidae are a small elateroid family distributed in South, East, and Southeast Asia (Wittmer 1979; Kawashima et al. 2010; Kundrata and Bocak 2011a; Kazantsev 2012). Soft-bodied males are capable of flight, whereas all known females are strongly paedomorphic and remain larva-like as adults (Fig. 1). Predaceous larvae occur in soil and leaf litter where they feed on millipedes. Both larvae and adults are bioluminescent, although the biology and ecology of most species are unknown (Li and Liang 2008; Kawashima et al. 2010). Rhagophthalmidae have a convoluted history of classification. Most genera were originally placed either in Lampyridae or the widely defined Drilidae (currently Drilini in Elateridae: Agrypninae; Kundrata and Bocak 2011b). The separate family Rhagophthalmidae was proposed by Olivier (1907, 1910) for genera which had antennae with 12 antennomeres and more or less emarginate eyes. However, since their erection, the composition and classification of Rhagophthalmidae have varied greatly, and various authors have recognized 3–11 genera in the group. At various times, the majority of Rhagophthalmidae have been considered either a subgroup of Lampyridae (e.g., McDermott 1964, 1966) or Phengodidae (Crowson 1972; Lawrence and Newton 1995; Bocak 2007), or a separate family close to one of the two above-mentioned families (Olivier 1910; Winkler 1925; Wittmer and Ohba 1994). Recent phylogenomic approaches suggest Rhagophthalmidae are sister to Phengodidae, and both are closely related to Lampyridae, Sinopyrophoridae, and Elateridae (Zhang et al. 2018; Douglas et al. 2021; Kusy et al. 2021; Cai et al. 2022).

10.3897/zookeys.1126.90233.figure1 DD030532-6271-5CC0-A267-5EEEEF2720AA

Morphology of RhagophthalmidaeA habitus of Dioptoma adamsii from Sri Lanka (syntype of D. greeni), female, NHMUK, dorsal view B head of Dioptoma adamsii from Sri Lanka, male, NHMUK, frontal view C head and pronotum of Diplocladon hasseltii hasseltii from Indonesia, male, SDEI, dorsal view D habitus of Falsophrixothrix sp. from Indonesia, male, NHMUK, dorsal view E head and pronotum of Rhagophthalmus sp. from China, male, first author’s collection, lateral view F habitus of Rhagophthalmus sp. from China, male, first author’s collection, dorsal view. Scale bars: 5.0 mm (A, F); 1.0 mm (B, E); 2.0 mm (C, D).

https://binary.pensoft.net/fig/764389

The early history of Rhagophthalmidae systematic research dates back to 1849, when Westwood (1849) described Dodecatoma Westwood, 1849 based on a single species from India. Motschulsky (1854) then described Rhagophthalmus Motschulsky, 1854 based on one species from China, and Pascoe (1860, 1862) added Dioptoma Pascoe, 1860 and Ochotyra Pascoe, 1862 from Bangladesh and India, respectively. While Dodecatoma was placed in the widely delimited Drilidae, the remaining genera were classified in Lampyridae (e.g., Gemminger 1869). Gorham (1883a, b) described Diplocladon Gorham, 1883 and its subgenus Haplocladon Gorham, 1883, both from Indonesia, and placed them in Drilidae.

Several new species of Rhagophthalmus from Southeast Asia, India, and China were added by Olivier (1885) and Fairmaire (1889, 1896, 1899). Gorham (1895) described the second species of Dodecatoma from India, and classified Dioptoma, Diplocladon, Dodecatoma, Haplocladon (originally as a subgenus), and Ochotyra in Drilinae. He later described a second species of Haplocladon, which was collected in India (Gorham 1903). Olivier (1907) erected the family Rhagophthalmidae for Dioptoma, Ochotyra, and Rhagophthalmus. In 1910, he provided the first catalogues for Rhagophthalmidae and Drilidae (Olivier 1910), with the latter including Diplocladon (with Haplocladon as a synonym) and Dodecatoma. Jakobson (1911) included many soft-bodied groups, including “Rhagophthalmini”, in his “Cantharididae”. Olivier (1912) revised Rhagophthalmus and recognized 12 species, five of which were newly described from China and Sri Lanka. Gahan in Morice (1913) reported a new species of Dioptoma from Sri Lanka.

Many new taxa currently belonging to Rhagophthalmidae were then described by the French coleopterist Maurice Pic, a person famous for his usually short and uninformative descriptions (e.g., Villiers 1958; Bezděk and Regalin 2015). Pic described the following taxa from Asia: one new species of Dioptoma and four species of Rhagophthalmus from India, Sri Lanka, China, and Indochina (Pic 1916, 1917, 1925a, b); genus Pseudothilmanus Pic, 1918, with its monotypic subgenus Drilothilmanus Pic, 1918 from northern India (Pic 1918); genus Bicladodrilus Pic, 1921, with two species from the Philippines and Vietnam (Pic 1921a, 1923); genus Bicladum Pic, 1921, with two species from Borneo and Sumatra (Pic 1921b, 1930a); a new variety and a new species of Dodecatoma from Indonesia and the Philippines, respectively (Pic 1921b, 1924); a new variety of Diplocladon from Indonesia (Pic 1921b); a new genus Monodrilus Pic, 1921 from Indonesia (Pic 1921b) and subsequently the monotypic subgenus Dodecatomorpha Pic, 1928 from Vietnam (Pic 1928); and a monotypic Mimoochotyra Pic, 1937 from Indonesia. Pic (1937) also erected Falsophrixothrix Pic, 1937 for two species from Indonesia, one of which was already described by Pic in the genus Phrixothrix Olivier, 1909 (currently in Phengodidae; Pic 1914).

Later, Wittmer (1939, 1944) added another three species from Indonesia and Singapore to Falsophrixothrix, with one being new and two transferred from Phrixothrix (Olivier 1911; Pic 1921a). Wittmer (1944) published a comprehensive catalogue of genera and species in Drilidae in which he listed many genera that are currently in Rhagophthalmidae, i.e., Bicladodrilus, Bicladum (as Bicladon [sic!]), Diplocladon (with Haplocladon as a synonym), Dodecatoma, Falsophrixothrix, Mimoochotyra, Monodrilus, and Pseudothilmanus. Pic (1951) described an additional species of Falsophrixothrix from Vietnam. In his major works on Lampyridae, McDermott (1964, 1966) included Dioptoma, Mimoochotyra (as Mimochotyra [sic!]), Ochotyra, and Rhagophthalmus in the subfamily Rhagophthalminae.

Crowson (1972) redefined Drilidae to include only a few core genera. Although Crowson excluded the majority of genera from Drilidae, he did not suggest any family placement for many, which left them in an uncertain position. Crowson (1972) also redefined Phengodidae by including Cydistus Bourgeois, 1885 as well as genera which are currently in Rhagophthalmidae, i.e., Dioptoma, Diplocladon, Falsophrixothrix, and Rhagophthalmus. Lawrence and Newton (1995) distinguished the subfamily Rhagophthalminae within Phengodidae, and included the genera Cydistus, Dioptoma, Diplocladon, Dodecatoma, Falsophrixothrix, Mimoochotyra (as Mimochotrya [sic!]), Ochotyra (as Ochotrya [sic!]), and Rhagophthalmus. Other major works on Rhagophthalmidae were those by Walter Wittmer, who described three new species of Dodecatoma from Afghanistan, India, and Nepal (Wittmer 1979, 1995), synonymized Ochotyra with Rhagophthalmus (Wittmer and Ohba 1994), and described eight new species of Rhagophthalmus from China, Japan, and Myanmar (Wittmer and Ohba 1994; Wittmer 1997).

Kawashima (1998) described the morphology of a larviform adult female of Rhagophthalmus. He also erected Menghuoius Kawashima, 2000 for two Chinese species originally classified in Rhagophthalmus, and later described the third species of that genus from Myanmar (Kawashima 2000, 2002). Kawashima and Satô (2001) described three species of Rhagophthalmus from Myanmar, Taiwan, and Thailand, and Kawashima and Sugaya (2003) added an additional new species from Taiwan. Branham and Wenzel (2003) studied the evolution of bioluminescence in the soft-bodied elateroids (i.e., “cantharoids”) and confirmed that Rhagophthalmus is closely related to Dioptoma and Diplocladon. Li and Liang (2008) described the morphology of a larviform adult female of Diplocladon from China. Li et al. (2008a) described two new species of Rhagophthalmus from China, provided information on the morphology and distribution for several other species, and provided a distribution map for all species in China and surrounding regions.

In the Rhagophthalmidae chapter of the Handbook of Zoology, Kawashima et al. (2010) included only Dioptoma, Diplocladon, Dodecatoma, Menghuoius, Mimoochotyra (as Mimochotyra [sic!]), and Rhagophthalmus. Kundrata and Bocak (2011a) revised the long-neglected genus Pseudothilmanus (with its subgenus Drilothilmanus, which they synonymized with Pseudothilmanus), added it to Rhagophthalmidae, and also listed Bicladodrilus, Bicladum (as Bicladon [sic!]), Dioptoma, Diplocladon, Dodecatoma, Falsophrixothrix, Mimoochotyra (as Mimochotyra [sic!]), Monodrilus, Reductodrilus, and Rhagophthalmus (including Menghuoius and Ochotyra). Ho et al. (2012) described two new species of Rhagophthalmus from Taiwan. Kazantsev (2012) described two species of Dodecatoma from India and Nepal. Most recently, Yiu (2017) described a new species of Diplocladon and a new species of Rhagophthalmus from Hong Kong. Roza (2020) added information on the morphology and distribution of Pseudothilmanus.

Besides research on the diversity, systematics, and morphology of Rhagophthalmidae, many studies in the 21st century have focused on their bioluminescence (Ohmiya et al. 2000; Ohba 2004a; Chen et al. 2010; Oba et al. 2011; Oba 2015; Liu et al. 2020) and embryogenesis (Kobayashi et al. 2001, 2002, 2003). Additionally, the rapid development of molecular phylogenetic methods in the last decades has enabled scientists to test the phylogenetic placement of Rhagophthalmidae within Elateroidea using one or several markers (e.g., Suzuki 1997; Bocakova et al. 2007; Sagegami-Oba et al. 2007; Stanger-Hall et al. 2007; Kundrata and Bocak 2011b; Kundrata et al. 2014; McKenna et al. 2015), mitogenomes (Li et al. 2007; Amaral et al. 2016; Chen et al. 2019), or a phylogenomic approach (Zhang et al. 2018; Amaral et al. 2019; Douglas et al. 2021; Kusy et al. 2021; Cai et al. 2022).

Despite the long history of rhagophthalmid systematic research, we lack a comprehensive study which would summarize all relevant information of all genera and species in the group. Therefore, in this study, we provide an annotated catalogue of genera and species of Rhagophthalmidae, including information on their synonyms, type material, distribution, and bibliography. We believe this study will serve as a robust framework for subsequent taxonomic revisions of all genera in addition to studies devoted to diversity, evolution, nature conservation, and ecology of the group.

 Materials and methods

Names of family-, genus-, and species-group taxa are given with the name of the author, and the year and page of publication. The page given is the page where the taxon name and description are printed. The year and page given for the incorrect subsequent spellings are the first year and page in which they are used. Incorrect subsequent spellings not in prevailing usage are unavailable (ICZN 1999, Art. 33.3). Complete data and comments for genus-group names are presented with the lowest-rank name, i.e., subgenus rather than genus, since these criteria follow the Principle of Coordination (ICZN 1999, Art. 36.1 and 43.1).

We provide the type species for each genus-group name, including information on its designation. We follow Recommendation 73F of the Code (ICZN 1999) and provide lectotype designations to fix the species identity for two species of Pseudothilmanus Pic, 1918. These species were originally described based on an unknown number of specimens, then redescribed (Kundrata and Bocak 2011a) under the assumption that the original descriptions were based only on holotypes. We do not provide lectotype designations for species in other genera, as they must first be revised in detail. Under each name, the currently valid name is listed first, followed by synonyms in chronological order.

Misspellings and unavailable names are followed by a colon “:”. We list all relevant references known to us for all genera, as well as for the family Rhagophthalmidae, particularly those that include information on systematics, classification, phylogeny, biology, and ecology. Since PhD or any other student theses are not officially published in the sense of the Code (ICZN 1999), we list only the relevant works (i.e., Ho 2002; Jeng 2008; Roza 2022) at the end of the Literature sections under each genus and species. Dates of publications and exact bibliographic references (especially problematic ones, often not cited uniformly by researchers) are taken from the following comprehensive general works: Chandler (2000); Bouchard et al. (2011); Bousquet (2016); and Evenhuis (2020). For the date of publication of F. P. Pascoe’s description of the genus Dioptoma (Pascoe 1860), we follow Evenhuis (2020).

Type depositories

ESRIEndemic Species Research Institute, JiJi, Nantou, Taiwan, Taiwan

ICM Insect Center, Moscow, Russia

KNHMZKunming Natural History Museum of Zoology, Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming, China

MNHNMuseum National d’Histoire Naturelle, Paris, France

MSNGMuseo Civico di Storia Naturale, Genova, Italy

MZB Bogor Zoology Museum, Bogor, Indonesia

NHMBNaturhistorisches Museum, Basel, Switzerland

NHMUKNatural History Museum, London, The United Kingdom

NKME Naturkundemuseum Erfurt, Germany

NMNSDepartment of Entomology, National Taiwan University, Taiwan

NTU Department of Entomology, National Taiwan University, Taipei, Taiwan

NWU Nagoya Women’s University, Nagoya, Japan

PCIK collection of I. Kawashima, Yokosuka-shi, Kanagawa, Japan

RMNHNaturalis Biodiversity Center, Leiden, The Netherlands

SMNHSwedish Museum of Natural History, Stockholm, Sweden

SMNSStaatliches Museum für Naturkunde, Stuttgart, Germany

TARITaiwan Agricultural Research Institute, Taichung, Taiwan

TLES Insect Museum, Tai Lung Experimental Station, Hong Kong, China

YCMYokosuka City Museum, Yokosuka, Japan

ZMM Zoological Museum of M.V. Lomonosov State University, Moscow, Russia

 Systematics Animalia Coleoptera Rhagophthalmidae D27F3E3C-1F04-5C21-94A8-DF877076DE15 Rhagophthalmidae Olivier, 1907 Rhagophthalmidae E. Olivier, 1907: 63. Type genus. Rhagophthalmus Motschulsky, 1854. Rhagophtalmidae : Junk 1912: 24 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Rhagophthalmidae : Blair in Gahan 1925: vi [unavailable name, incorrect subsequent spelling not in prevailing usage]. Rhagopthalmidae: Harvey 1952: 389 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Rhagophthalidae: Ohba 1998: 2 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Rhagophthammidae: Suzuki and Kobayashi 2009: 31 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Literature.

Olivier (1907: 1, 63): catalogue; Lefroy (1909: 327): catalogue; Olivier (1910: 3): catalogue; Jakobson (1911: 662, 687): catalogue [as Rhagophthalmini]; Junk (1912: 24): bibliography [as Rhagophtalmidae [sic!]]; Olivier (1912: 467): revision of Rhagophthalmus; Blair (1915a: 411): bioluminescence; Pic (1923: 25): catalogue; Gahan (1925: vi): remark [as Phagophthalmidae [sic!]; attributed to KG Blair]; Handlirsch (1925: 589): catalogue [as Rhagophthalmini]; Winkler (1925: 522): catalogue; Ridley (1934: 58): larval biology and morphology; Pic (1937: 137): genus description; Harvey (1952: 389, 450): remark, bioluminescence [also as Rhagopthalmidae [sic!], also as Rhagopthalminae [sic!]]; Brues et al. (1954: 565): classification; Crowson (1955: 68): remark, morphology [as Rhagophthalminae]; Harvey (1955: 19): checklist, bioluminescence; Raj (1957: 788): larval biology; McDermott (1964: 49): revision [as Rhagophthalminae]; McDermott (1966: preface (unnumbered),121): catalogue, distribution [as Rhagophthalminae]; Mikšić and Mikšić (1966: 31): remark [as Rhagophthalminae]; Nakane (1968: 3): remark [as Rhagophthalminae]; Crowson (1972: 50): classification, morphology [as Rhagophthalminae]; McElroy et al. (1974: 415): remark [as Rhagopthalmidae [sic!]]; Lawrence (1982: 512): remark; Haneda (1985: 167): bioluminescence [as Rhagopthalmidae [sic!]]; Herring (1987: 158): checklist [as Rhagophthalminae]; LeSage (1991: 424): remark [also as Rhagophthalminae]; Wittmer and Ohba (1994: 341): taxonomy, biology; Lawrence and Newton (1995: 857): catalogue, review [as Rhagophthalminae]; Chen and Ho (1996: 46): distribution; Ohba et al. (1996a: 1): morphology, biology; Ohba (1997a: 5): checklist; Ohba (1997c: 51): breeding; Suzuki (1997: 11, 38): phylogeny, biology [also as Rhagophthalminae]; Wittmer (1997: 257): species descriptions; Chen and Ho (1998: 34): bioluminescence; Ohba (1998: 2): biology [also as Rhagophthalidae [sic!]]; Costa et al. (1999: 22): remark [as Rhagophthalminae]; Goto and Kawashima (2000: 141): distribution; Jeng et al. (2000: 316): remark; Kawashima (2000: 131): genus description; Kim et al. (2000: 214): molecular phylogeny; Ohmiya et al. (2000: 32): luciferase; Branham and Wenzel (2001: 565): phylogeny [also as Rhagophthalminae and Rhagopthalmidae [sic!]]; Kawashima and Satô (2001: 423): species descriptions; Kobayashi et al. (2001: 1): embryogenesis, morphology [also as Rhagophthalminae]; Hua (2002: 71): catalogue; Kawashima (2002: 487): species description; Kobayashi et al. (2002: 1): embryogenesis, morphology [also as Rhagophthalminae]; Branham and Wenzel (2003: 3): phylogeny; Chen (2003: 52): morphology, bioluminescence; Hayashi and Suzuki (2003: 4): biology, morphology, phylogeny, figure of mating; Kawashima and Sugaya (2003: 353): species description; Kawashima et al. (2003: 255): catalogue; Kobayashi et al. (2003: 19): embryogenesis, morphology; DeCock (2004: 341): bioluminescence; Ohba (2004a: 225): bioluminescence, biology; Lau and Meyer-Rochow (2006: 19): eye morphology; Li et al. (2006: 817): molecular phylogeny; Arnoldi et al. (2007: 2): molecular phylogeny, remark; Bocak (2007: 224): catalogue [as Rhagophthalminae]; Bocakova et al. (2007: 477): molecular phylogeny [also as Rhagophthalminae]; Hunt et al. (2007: 1915): molecular phylogeny; Li et al. (2007: 197): mitochondrial genome, phylogeny [also as Rhagophthalminae]; Sagegami-Oba et al. (2007: 110): molecular phylogeny [also as Rhagophthalminae]; Stanger-Hall et al. (2007: 38): molecular phylogeny; Bocak et al. (2008: 2021): molecular phylogeny; Li and Liang (2008: 109): female morphology; Li et al. (2008a: 259): species descriptions, distribution [also as Rhagophthalminae]; Li et al. (2008b: 494): review [also as Rhagophthalminae]; Bogahawatta et al. (2009: 5): distributional remark [as Rhagophthalminae]; Levkanicova and Bocak (2009: 212): molecular phylogeny; Suzuki and Kobayashi (2009: 30): embryogenesis [also as Rhagophthalminae and Rhagophthammidae [sic!]]; Chen et al. (2010: 196): biology, bioluminescence; Kawashima et al. (2010: 135): book chapter [also as Rhagophthalminae]; Lawrence et al. (2010a: 5): classification; Lawrence et al. (2010b: 165): remark; Bouchard et al. (2011: 326): family-group names catalogue; Kundrata and Bocak (2011a: 57): revision of Pseudothilmanus; Kundrata and Bocak (2011b: 364): molecular phylogeny [also as Rhagophthalminae]; Lawrence et al. (2011: 7): phylogeny; Oba et al. (2011: 775): biology, bioluminescence [also as Rhagophthalminae]; Yiu (2011a: 14): remark; Yiu (2011b: 20): bioluminescence, larva; Amaral et al. (2012: 1262): luciferase, phylogeny [as Rhagophthalminae]; Ho et al. (2012: 1): species descriptions; Johnson et al. (2012: 178): ICZN case; Kazantsev (2012: 349): species descriptions; Timmermans and Vogler (2012: 299): remark, molecular phylogeny; Kundrata et al. (2013: 201): molecular phylogeny; Yiu (2013: 101): remark, bioluminescence; Amaral et al. (2014: 415): molecular phylogeny; Bocak et al. (2014: 103): molecular phylogeny; Hosoe et al. (2014: 331): biology; ICZN (2014: 195): ICZN case; Kundrata et al. (2014: 163): molecular phylogeny; Li et al. (2015: 269): catalogue; Martin et al. (2015: 516): molecular phylogeny; McKenna et al. (2015: 843): molecular phylogeny [also as Rhagopthalmidae [sic!]]; Oba (2015: 99): bioluminescence; Amaral et al. (2016: 255): molecular phylogeny; Bocak et al. (2016: 2): molecular phylogeny; Kundrata et al. (2016: 293): molecular phylogeny; Lawrence (2016: 17): classification; Wijekoon et al. (2016: 69): checklist [also as Rhagophthalminae]; Amaral et al. (2017a: 674): mitogenome, phylogeny; Kundrata et al. (2017: 153): molecular phylogeny; Martin et al. (2017: 564): phylogeny; Wang et al. (2017: 1): phylogeny; Yiu (2017: 60): species descriptions, key; Bocak et al. (2018: 2): molecular phylogeny; Fallon et al. (2018: 2, 96): genomes, bioluminiscence; Kusy et al. (2018a: 5): molecular phylogeny; Kusy et al. (2018b: 2): molecular phylogeny; Tan (2018: 127, 135): distribution, photographs; Zhang et al. (2018: 3): molecular phylogeny; Amaral et al. (2019: 283): molecular phylogeny [also as Rhagophtalmidae [sic!]]; Chen et al. (2019: 4): molecular phylogeny; Jeng (2019: 8): biofluorescence, biology; Kundrata et al. (2019: 1259): molecular phylogeny; Martin et al. (2019: 2): molecular phylogeny [also as Rhagophthalminae]; McKenna et al. (2019: 4): molecular phylogeny; Liu et al. (2020: 46): luciferase, phylogeny [also as Rhagophthalminae]; Rosa et al. (2020: 7): molecular phylogeny; Roza (2020: 421): morphology, distribution; Zhang et al. (2020: 1): molecular phylogeny, bioluminescence; Douglas et al. (2021: 2): molecular phylogeny; Ge et al. (2021: 3): mitogenomic phylogeny; Kusy et al. (2021: 111): molecular phylogeny; Li et al. (2021a: 5): remark; Li et al. (2021b: 1): phylogeny, distribution, morphology; Seri and Rahman (2021: 715): remark; Cai et al. (2022: 6): molecular phylogeny; Ge et al. (2022: 2): mitogenomic phylogeny; Powell et al. (2022: 1): molecular phylogeny, bioluminescence [also as Rhagophtalmidae [sic!]]. In addition to the aforementioned literature, Rhagophthalmidae were mentioned in some student works, e.g., PhD theses by Ho (2002), Jeng (2008), and Roza (2022).

 Remarks.

As defined here, Rhagophthalmidae include 12 genera (one of them with two subgenera) and 66 species distributed primarily in East, South, and Southeast Asia, with a few species found on the border of South and Central Asia (i.e., Afghanistan). Males can be recognized by antennae with 12 antennomeres, with antennomere III longer than antennomere II. In cases where the antennae are serrate or pectinate, antennomere III is not simple, i.e., the serration or rami begin on antennomere III. Females are more (e.g., Diplocladon or Haplocladon; see Remarks under these genera) or less (e.g., Rhagophthalmus) larviform (for more information, see Kawashima et al. 2010). Known larvae are predators of millipedes, similar to larvae of the closely related Phengodidae. Although Rhagophthalmidae were credited by McDermott (1966) to “Olivier, 1902”, we found no evidence of the publication to which McDermott referred, similar to Lawrence and Newton (1995: 858).

 Animalia Coleoptera Rhagophthalmidae B0185C71-CABA-51E4-964E-32CACCB767FD Bicladodrilus Pic, 1921 Bicladodrilus Pic, 1921a: 15. Gender: masculine. Type species. Bicladodrilus bakeri Pic, 1921; by monotypy. Bieladodrilus : Pic 1923: 62 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Bicalodrilus : Pic 1930b: 320 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Literature.

Pic (1921a: 15): original description; Pic (1923: 62): species description [as Bieladodrilus [sic!]]; Pic (1930b: 320): remark [as Bicalodrilus [sic!]], key; Wittmer (1941: 197): catalogue, distribution; Wittmer (1944: 211): catalogue; Bocakova et al. (2007: 484): molecular phylogeny; Hunt et al. (2007: suppl.): molecular phylogeny; Bocak et al. (2008: 2019): molecular phylogeny; Levkanicova and Bocak (2009: 214): molecular phylogeny; Costa and Zaragoza-Caballero (2010: 134): remark; Kawashima et al. (2010: 139): book chapter; Kundrata and Bocak (2011a: 57): remark; Kundrata and Bocak (2011b: 370): molecular phylogeny; Kundrata et al. (2013: 202): molecular phylogeny; Kundrata et al. (2014: 167): molecular phylogeny; Bocak et al. (2016: suppl.): molecular phylogeny; Kovalev and Kirejtshuk (2016: 205): remark; Kundrata et al. (2016: 296): molecular phylogeny; Bocak et al. (2018: 4): molecular phylogeny; Kundrata et al. (2019: 1263): molecular phylogeny; Liu et al. (2020: 46): remark. In addition to the aforementioned literature, this genus was included in PhD theses by Jeng (2008) and Roza (2022).

 Remarks.

This genus currently contains two described species from the Philippines and Vietnam, respectively. The generic assignment of a specimen reported as “Bicladodrilus sp.” from China, which was used in the molecular phylogenetic analyses by Bocakova et al. (2007), Bocak et al. (2008, 2018), Levkanicova and Bocak (2009), and other studies, needs a careful re-examination. Bicladodrilus is similar to Bicladum and Diplocladon in having strongly bipectinate antennae and long elytra. This generic complex is in need of revision.

 Animalia Coleoptera Rhagophthalmidae 0BD39A61-BB56-51D6-993C-E1CC29B97B1B Bicladodrilus bakeri Pic, 1921 Bicladodrilus bakeri Pic, 1921a: 15. Type depository.

Described based on an unknown number of specimens. Syntype, male (MNHN).

 Type locality.

Philippines: Mindanao.

 Distribution.

Philippines.

 Literature.

Pic (1921b: 15): original description; Pic (1923: 63): comparison with B. laticollis Pic, 1923; Wittmer (1941: 197): catalogue, distribution; Wittmer (1944: 211): catalogue.

 Animalia Coleoptera Rhagophthalmidae BC39FB9B-2F7E-5312-BA3E-E71FDBA0BD26 Bicladodrilus laticollis Pic, 1923 Bieladodrilus [sic!] laticollis Pic, 1923: 62. Type depository.

Described based on an unknown number of specimens. Syntype, male (MNHN).

 Type locality.

Vietnam: Lào Cai [Tonkin: Lao-Kay].

 Distribution.

Vietnam.

 Literature.

Pic (1923: 62): original description; Wittmer (1944: 211): catalogue.

 Animalia Coleoptera Rhagophthalmidae 76BB5109-16CE-560A-925C-0AD76EC685EE Bicladum Pic, 1921 Bicladum Pic, 1921b: 12. Gender: neuter. Type species. Bicladum multipunctatum Pic, 1921; by monotypy. Bicladon : Pic 1930a: 2 [unavailable name, incorrect subsequent spelling]. Literature.

Pic (1921b: 12): original description; Pic (1921a: 15): comparison with Bicladodrilus; Pic (1930a: 2): species description [as Bicladon [sic!]]; Pic (1930b: 320, 321): remark, key [as Bicladon [sic!]]; Wittmer (1944: 211): catalogue [as Bicladon [sic!]]; Lawrence et al. (2010b: 175): remark [as Bicladon [sic!]]; Kundrata and Bocak (2011a: 57): remark [as Bicladon [sic!]]; Janisova and Bocakova (2013: 3): remark [as Bicladon [sic!]]; Kovalev and Kirejtshuk (2016: 205): remark [as Bicladon [sic!]]. In addition to the aforementioned literature, this genus was included in a PhD thesis by Jeng (2008).

 Remarks.

This genus currently contains two described species from Borneo and Sumatra, respectively. It is similar to Bicladodrilus and Diplocladon in having strongly bipectinate antennae and long elytra. This generic complex is in need of revision.

 Animalia Coleoptera Rhagophthalmidae 31AB7D66-0507-513D-9AAD-818B41A21FD3 Bicladum mjobergi Pic, 1930 Bicladon [sic!] mjöbergi [sic!] Pic, 1930a: 2, 4. Type depositories.

Described based on an unknown number of specimens. One syntype, male (MNHN), two syntypes, males (labelled as “Typus” and “Paratypus”) (SMNH).

 Type locality.

Indonesia: Sumatra, Medan.

 Distribution.

Indonesia (Sumatra).

 Literature.

Pic (1930a: 2, 4): original description; Wittmer (1944: 211): catalogue [as Bicladon [sic!]].

 Remarks.

Pic (1930a: 5) also reported an unnamed variety of B. mjobergi based on a specimen from Tjinta Radja. This specimen is deposited in SMNH and bears the label “Typus”; however, based on Article 72.4.1. of the Code (ICZN 1999) it should not be considered a part of the type series.

 Animalia Coleoptera Rhagophthalmidae 651CD9BF-32F0-595E-B93C-6167BA93EF71 Bicladum multipunctatum Pic, 1921 Bicladum multipunctatum Pic, 1921b: 12. Type depository.

Described based on an unknown number of specimens. Syntype, male (MNHN).

 Type locality.

Borneo (without any further data).

 Distribution.

Borneo (probably northern region).

 Literature.

Pic (1921b: 12): original description; Pic (1930a: 5): comparison with B. mjobergi; Wittmer (1944: 211): catalogue [as Bicladon [sic!]].

 Animalia Coleoptera Rhagophthalmidae 53D1C3FD-5D9F-5AE5-8197-1E79BE98F47E Dioptoma Pascoe, 1860 Fig. 1A, B Dioptoma Pascoe, 1860: 118. Gender: feminine. Type species. Dioptoma adamsii Pascoe, 1860; by monotypy. Diaptoma : Wijekoon et al. 2016: 70 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Literature.

Pascoe (1860: 118): original description, drawings of male habitus, head, and antenna; Pascoe (1862: 323): comparison with Ochotyra; Gerstaecker (1863: 409): remark; Gemminger (1869: 1647): catalogue; Gorham (1880: 66): remark; Gorham (1881: 63): remark; Olivier (1885: 372): remark; Gorham (1890: 550): catalogue; Gorham (1895: 309): redescription; Sharp (1899: 251): remark; Gorham (1903: 330): distributional note; Olivier (1907: 63): catalogue; Gahan (1908a: xlviii): remark; Gahan (1908b: 205): remark; Olivier (1910: 1): catalogue; Olivier (1912: 467): remark; Morice (1913: cxviii): introduction of a new species attributed to Gahan; Green (1913: 718): male and female morphology, bioluminescence, drawing of male habitus; McDermott (1914: 304): remark; Blair (1915a: 413): bioluminescence; Blair (1915b: 191): bioluminescence; Blair (1915c: 37): bioluminescence, morphology; Gravely (1915: 502): remark; Bugnion (1916: 83): remark; Pic (1916: 8): species description; Lucas (1920: 241): catalogue; Bugnion (1929: 4): remark; Brues (1941: 41): remark; Harvey (1952: 392): remark; Harvey (1955: 19): checklist, bioluminescence; Bess (1956: 25): remark; McDermott (1964: 50): revision; McDermott (1966: 122): catalogue; Mikšić and Mikšić (1966: 32): remark; Lloyd (1971: 101): remark, drawing of male habitus with distribution of luminous organs; Crowson (1972: 52): remark; Paulus (1975: 78): remark; Herring (1978: 471): checklist; Lloyd (1978: 252): remark, drawing of male habitus with distribution of luminous organs; Lloyd (1979: 302): remark; Ohba (1980: 14): remark; Crowson (1981: 314): remark, drawing of male habitus with distribution of luminous organs; Sivinski (1981: 168): remark; Lloyd (1983: 136): remark, bioluminescence; Hoffmann (1984: 230): remark; Herring (1987: 158): checklist; Cicero (1988: 148): remark; Viviani and Bechara (1993: 615): remark; Wittmer and Ohba (1994: 342): remark; Lawrence and Newton (1995: 857): catalogue, remark; Branham (1996: 18): remark; Ohba et al. (1996a: 17): remark; Viviani and Bechara (1997: 389): remark; Sivinski et al. (1998: 29): remark; Kawashima (2000: 131): remark; Branham and Wenzel (2001: 566): phylogeny; O’Keefe (2002: 182): remark; Branham and Wenzel (2003: 5): phylogeny; Li et al. (2008a: 259): remark; Li et al. (2008b: 495): review; Li and Liang (2008: 111): remark; Bogahawatta et al. (2009: 1): remark; Suzuki and Kobayashi (2009: 30): remark; Chen et al. (2010: 196): remark; Kawashima et al. (2010: 135): book chapter; Kundrata and Bocak (2011a: 57): remark; Oba et al. (2011: 777): remark; Wijekoon et al. (2016: 70): checklist [as Diaptoma [sic!]]; Liu et al. (2020: 46): remark. In addition to the aforementioned literature, this genus was included in PhD theses by Jeng (2008) and Roza (2022).

 Remarks.

This genus currently contains two described species from Bangladesh, India, and Sri Lanka. Males are characterized by short antennae and deeply emarginate eyes, each with a smaller upper portion and a larger lower portion (Fig. 1B). Regarding the gender of Dioptoma, Pascoe (1860) treated it as feminine and since the name is not a Greek noun, here we follow Pascoe’s decision.

 Animalia Coleoptera Rhagophthalmidae 355E0479-9A71-59C9-9A06-4C7CD9069AC1 Dioptoma adamsii Pascoe, 1860 Fig. 1A, B Dioptoma adamsii Pascoe, 1860: 118. Dioptoma adamsi : Gemminger 1869: 1647 [unavailable name, incorrect subsequent spelling]. Dioptoma greeni Gahan in Morice 1913: cxviii. Synonymized with D. adamsii (as a variety) by McDermott (1966: 122). McDermott (1966) attributed the name D. greeni to Gahan (1908a: xlviii); however, there is no such name in that publication, and we believe that this name first appeared in 1913. Dioptoma ademsi : Bogahawatta et al. 2009: 1 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Type depository.

Holotype of D. adamsii, male (NHMUK). 25 syntypes of D. greeni (eight males from Maskeliya, eight males and four females from Dikoya, four males and one female from Bogawantalawa; Fig. 1A) (NHMUK).

 Type locality of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Dioptoma">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="adamsii">adamsii</tp:taxon-name-part></tp:taxon-name></italic>.

Bangladesh: Dhaka [“India: Dacca”]. Type localities of D. greeni. Sri Lanka: Bogawantalawa, Dikoya, and Maskeliya.

 Distribution.

Bangladesh, India (Karnataka, Kerala, Tamil Nadu, Uttarakhand), Sri Lanka.

 Literature.

Pascoe (1860: 118): original description, drawings of male habitus, head, and antenna; Gemminger (1869: 1647): catalogue [as D. adamsi [sic!]]; Gorham (1880: 66): remark [as D. adamsi [sic!]]; Olivier (1885: 372): remark [as D. adamsi [sic!]]; Gorham (1890: 550): catalogue [as D. adamsi [sic!]]; Gorham (1895: 310): redescription, distributional note [as D. adamsi [sic!]]; Sharp (1899: 251): remark [as D. adamsi [sic!]]; Gorham (1903: 330): distributional note [as D. adamsi [sic!]]; Olivier (1910: 1): catalogue [as D. adamsi [sic!]]; Morice (1913: cxviii): original description of D. greeni (attributed to Gahan), remarks on D. adamsi [sic!]; Green (1913: 718): male and female morphology, bioluminescence, drawing of male habitus [as D. adamsi [sic!]]; McDermott (1914: 304): remark; Blair (1915a: 413): bioluminescence [as D. adamsi [sic!]]; Blair (1915b: 191): bioluminescence [as D. adamsi [sic!]]; Blair (1915c: 37): bioluminescence, morphology [as D. adamsi [sic!]]; Gravely (1915: 502): remark [as D. adamsi [sic!]]; Bugnion (1916: 96): remark [as D. adamsi [sic!]]; Pic (1916: 8): comparison with D. atripennis Pic, 1916 [as D. adamsi [sic!]]; Lucas (1920: 241): catalogue [as D. adamsi [sic!]]; Brues (1941: 41): remark [as D. adamsi [sic!]]; Harvey (1952: 450): remark, bioluminescence [as D. adamsi [sic!]]; McDermott (1964: 50): redescription [as D. adamsi [sic!]]; McDermott (1966: 122): catalogue, synonymization of D. greeni with D. adamsi [sic!]; Lloyd (1971: 101): remark, drawing of male habitus with distribution of luminous organs [as D. adamsi [sic!]]; Lloyd (1978: 252): remark, drawing of male habitus with distribution of luminous organs [as D. adamsi [sic!]]; Lloyd (1979: 302): remark [as D. adamsi [sic!]]; Crowson (1981: 314): remark, drawing of male habitus with distribution of luminous organs [as D. adamsi [sic!]]; Sivinski (1981: 168): remark [as D. adamsi [sic!]]; Lloyd (1983: 136): remark, bioluminescence [as D. adamsi [sic!]]; Hoffmann (1984: 230): remark [as D. adamsi [sic!]]; Branham (1996: 18): remark [as D. adamsi [sic!]]; Ohba et al. (1996a: 17): remark; Sivinski et al. (1998: 29): remark [as D. adamsi [sic!]]; Kawashima (2000: 131): remark; Branham and Wenzel (2001: 567): phylogeny [as D. adamsi [sic!]]; Branham and Wenzel (2003: 5): phylogeny [as D. adamsi [sic!]]; Li et al. (2008b: 496): review [also as D. adamsi [sic!]]; Bogahawatta et al. (2009: 1): remark [both D. ademsi [sic!] and D. greeni]; Kawashima et al. (2010: 135): book chapter [as D. adamsi [sic!]]; Wijekoon et al. (2016: 70): catalogue [both Diaptoma adamsi [sic!] and D. greeni]. In addition to the aforementioned literature, this species was included in PhD theses by Jeng (2008) and Roza (2022).

 Remarks.

This species was referred to as “adamsi” in the majority of publications. The original spelling “adamsii” was used only by McDermott (1914), Ohba et al. (1996a), and Kawashima (2000). However, following Article 33.4. of the Code (ICZN 1999), the original spelling should be maintained. It should be noted that the current concept of D. adamsii may include several species.

 Animalia Coleoptera Rhagophthalmidae DB60CEF4-783B-5564-80FC-4CF0FF6A6ED3 Dioptoma atripennis Pic, 1916 Dioptoma atripennis Pic, 1916: 8. Type depository.

Described based on an unknown number of specimens. Two syntypes, males (MNHN).

 Type locality.

India: Tamil Nadu, Madurai [Madura].

 Distribution.

India (Tamil Nadu).

 Literature.

Pic (1916: 8): original description; McDermott (1966: 122): catalogue; Li et al. (2008b: 496): review.

 Animalia Coleoptera Rhagophthalmidae 09D151D2-3825-57C6-A32F-819651C145ED Diplocladon Gorham, 1883 Fig. 1C Diplocladon Gorham, 1883a: 5. Gender: neuter. Type species. Diplocladon hasseltii Gorham, 1883, by monotypy. Diplocadum : Pic 1921b: 12 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Diplocladum : Pic 1928: 86 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Diplocadon : Viviani and Bechara 1993: 615 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Diploclodon : Tan 2018: 135 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Literature.

Gorham (1883a: 5): original description; Gorham (1883b: 249, 250): comparison with Haplocladon; Gorham (1883c: 597): remark; Gorham (1887: 76): catalogue, redescription; Waterhouse (1890: 25): remark, figure of male habitus; Gorham (1895: 310): remark; Olivier (1910: 8): catalogue; Lucas (1920: 243): catalogue; Rüschkamp (1920: 386): distributional note; Pic (1921b: 12): comparison with Bicladum and Monodrilus [as Diplocadum [sic!]]; Pic (1928: 86): remark [as Diplocadum [sic!]]; Pic (1930a: 2): distributional note [as Diplocladum [sic!]]; Pic (1930b: 320): remark, key; Ridley (1934: 60): larval biology and morphology; Wittmer (1944: 211): catalogue; Haneda (1950: 2): bioluminescence; Harvey (1952: 451): bioluminescence, drawings of female habitus with position of luminous organs, photographs of male and female habitus; Crowson (1955: 68, 171): remark; Haneda (1955: 364): remark, bioluminescence; Harvey (1955: 19): checklist, bioluminescence; Harvey (1957: 554): remark; McDermott (1964: 50): remark; Nakane (1968: 3): remark; Lloyd (1971: 101): remark, drawing of female habitus with luminous organs; Crowson (1972: 52): remark; Paulus (1972: 49): remark; Halverson et al. (1973: 1332): biology, bioluminescence; McElroy et al. (1974: 417): remark; Paulus (1975: 78): remark; Case and Strause (1978: 332): remark; Herring (1978: 471): checklist; Lloyd (1978: 252): remark, drawing of female habitus with distribution of luminous organs; Ohba (1980: 14): remark; Crowson (1981: 314): remark, drawing of female habitus with distribution of luminous organs; Sivinski (1981: 168): remark; Lloyd (1983: 136): remark, bioluminescence; Hoffmann (1984: 229): remark; Haneda (1985: 167): bioluminescence; Herring (1987: 157): checklist; Cicero (1988: 148): remark; De Keyzer (1989: 54): remark; Viviani and Bechara (1993: 615): remark [as Diplocadon [sic!]]; Wittmer and Ohba (1994: 350): remark; Lawrence and Newton (1995: 857): catalogue, remark; Branham (1996: 18): remark; Ohba et al. (1996a: 13): remark; Ohba et al. (1996b: 30): remark; Ohba (1997a: 17): remark; Viviani and Bechara (1997: 389): remark [as Diplocadon [sic!]]; Branham and Wenzel (2001: 566): phylogeny; O‘Keefe (2002: 182): remark; Branham and Wenzel (2003: 3): remark; Li and Liang (2008: 109): remark, female description; Li et al. (2008b: 495): review; Suzuki and Kobayashi (2009: 31): remark; Kawashima et al. (2010: 135): book chapter, figures of male and female habitus, and details of female abdominal segments; Kundrata and Bocak (2011a: 57): remark; Oba et al. (2011: 777): remark; Yiu (2012: 30): catalogue, figures of habitus; Yiu (2013: 113): remark, biology; Kovalev and Kirejtshuk (2016: 205): remark; Yiu (2017: 64): description of a new species, identification key; Tan (2018: 135): possible larva, distribution, figures of larval habitus and bioluminescence [also as Diploclodon]; Liu et al. (2020: 46): remark; Lawrence et al. (2021: 456): wing morphology; Li et al. (2021b: 4): remark; Seri and Rahman (2021: 721): remark. In addition to the aforementioned literature, this genus was included in PhD theses by Jeng (2008) and Roza (2022).

 Remarks.

See more information on Haplocladon, which was once considered a subgenus of Diplocladon (Gorham 1883b) or even its synonym (e.g., Wittmer 1944), under the genus name Haplocladon below. Some authors who mentioned Diplocladon were actually probably referring to Haplocladon (for more details, see Remarks under D. hasseltii). Diplocladon currently contains two described species, one from China and one from Indonesia. It is similar to Bicladodrilus and Bicladum in having strongly bipectinate antennae (Fig. 1C) and long elytra. This generic complex is in need of revision.

 Animalia Coleoptera Rhagophthalmidae BE33CF86-A240-5027-AE5B-84EC4C7ED7A1 Diplocladon atripenne Yiu, 2017 Diplocladon atripennis [sic!] Yiu, 2017: 64. Type depository.

Holotype, male (TLES). Paratype, male (TLES).

 Type locality.

China: Hong Kong, Lantau, Wo Tin (22.27351°N, 113.98819°E).

 Distribution.

China (Hong Kong).

 Literature.

Yiu (2017: 64): original description, figures of male habitus, pregenital segments and genitalia.

 Animalia Coleoptera Rhagophthalmidae 775AC322-4459-51C5-8356-43D6D4CC8FE2 Diplocladon hasseltii hasseltii Gorham, 1883 Fig. 1C Diplocladon hasseltii Gorham, 1883a: 6. Diplocladon hasselti : Olivier 1910: 8 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Type depository.

Described based on two specimens (Gorham 1887). One syntype, male (RMNH); one syntype, male (MNHN).

 Type locality.

Indonesia: Sumatra, Boenga mas (Palembang).

 Distribution.

Indonesia (Sumatra, Java).

 Literature.

Gorham (1883a: 6): original description; Gorham (1887: 76): catalogue, redescription; Waterhouse (1890: 25): remark, figure of male habitus; Olivier (1910: 8): catalogue [as D. hasselti [sic!]]; Lucas (1920: 243): catalogue [as D. hasselti [sic!]]; Ridley (1934: 60): larval biology and morphology [as D. hasselti [sic!]]; Wittmer (1944: 211): catalogue [as D. hasselti [sic!]]; Haneda (1950: 2): bioluminescence, drawings of adult male and female, and position of luminous organs; Harvey (1952: 451): bioluminescence, drawings of female habitus with position of luminous organs, photographs of male and female habitus; Haneda (1955: 364): remark, bioluminescence; Lloyd (1971: 101): remark, drawing of female habitus with luminous organs [as D. hasselti [sic!]]; Lloyd (1978: 252): remark, drawing of female habitus with distribution of luminous organs [as D. hasselti [sic!]]; Crowson (1981: 314): remark, drawing of female habitus with distribution of luminous organs [as D. hasselti [sic!]]; Sivinski (1981: 168): remark [as D. hasselti [sic!]]; Lloyd (1983: 136): remark, bioluminescence [as D. hasselti [sic!]]; Hoffmann (1984: 229): remark [as D. hasselti [sic!]]; Haneda (1985: 167): bioluminescence, drawings of adult male and female, and position of luminous organs [as D. hasselti [sic!]]; De Keyzer (1989: 54): remark [as D. hasselti [sic!]]; Wittmer and Ohba (1994: 350): remark [as D. hasselti [sic!]]; Branham (1996: 18): remark [as D. hasselti [sic!]]; Ohba et al. (1996a: 13): remark; Ohba et al. (1996b: 30): remark; Ohba (1997a: 17): remark; Li and Liang (2008: 109): remark; Kawashima et al. (2010: 135): book chapter, figures of male and female habitus, and details of female abdominal segments [as D. hasselti [sic!]]; Yiu (2017: 64): comparison with D. atripennis; Lawrence et al. (2021: 456): wing morphology, figure of hind wing [as D. hasselti [sic!]]. In addition to the aforementioned literature, this species was included in PhD theses by Jeng (2008) and Roza (2022).

 Remarks.

Based on the available figures, adults of both sexes which were reported by Haneda (1950) from Singapore, and repeatedly mentioned in subsequent studies (e.g., Harvey 1952; Haneda 1955, 1985; Lloyd 1971, 1978; Crowson 1981; Kawashima et al. 2010), are probably members of Haplocladon. We are aware of several Haplocladon specimens from Singapore (deposited in NHMUK) but no Diplocladon are known from that area.

 Animalia Coleoptera Rhagophthalmidae 66186E05-2202-5317-A540-5D6307EE2409 Diplocladon hasseltii testaceum Pic, 1921 Diplocadum [sic!] hasselti [sic!] var. testaceum Pic, 1921b: 12. Diplocladum [sic!] hasselti [sic!] var. testaceus [sic!]: Pic 1930a: 2. Type depository.

Described based on an unknown number of specimens. No type specimen found in MNHN by RK.

 Type locality.

Indonesia: Sumatra.

 Distribution.

Indonesia (Sumatra).

 Literature.

Pic (1921a: 12): original description [as a variety of Diplocadum [sic!] hasselti [sic!]]; Pic (1930b: 2): distributional note; Wittmer (1944: 211): catalogue.

 Remarks.

The name “testaceum” is deemed to be subspecific according to Article 45.6.4. of the Code (ICZN 1999).

 Animalia Coleoptera Rhagophthalmidae 7523762A-0AAD-572B-B413-AF186157C9EC Dodecatoma Westwood, 1849 Dodecatoma Westwood, 1849: 1. Gender: feminine. Type species. Dodecatoma bicolor Westwood, 1849, by monotypy. Dodecatomax : Crowson 1955: 171 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Literature.

Westwood (1849: 1): original description, drawings of male habitus, head, mouthparts, antenna, and leg; Schaum (1850: 165): morphology, remark; Lacordaire (1857: 377): catalogue, redescription; Motschulsky (1861: 134): comparison with Pachytarsus Motschulsky, 1861; Gemminger (1869: 1686): catalogue; Gorham (1895: 309): species description, remark; Olivier (1910: 8): catalogue; Fowler (1912: 138): remark; Lucas (1920: 246): catalogue; Rüschkamp (1920: 386): distributional note; Pic (1921b: 12): species description; Pic (1924: 713): species description, remark; Pic (1930b: 321): remark; Wittmer (1941: 197): catalogue; Wittmer (1944: 211): catalogue; Harvey (1952: 392): remark; Crowson (1955: 68, 171): remark [also as Dodecatomax [sic!]]; Goidanich (1957: 565): remark; McDermott (1964: 50): remark; Paulus (1972: 49): remark; Wittmer (1979: 89): species description, drawing of male antenna; Lawrence and Newton (1995: 857): catalogue, remark; Wittmer (1995: 110): species descriptions; Bocak (2007: 225): catalogue; Li et al. (2008b: 495): review; Kawashima et al. (2010: 135): book chapter; Kundrata and Bocak (2011a: 58): remark; Oba et al. (2011: 777): remark; Kazantsev (2012: 349): descriptions of new species, identification key; Johnson et al. (2012: 178): ICZN case; ICZN (2014: 195): ICZN case; Kovalev and Kirejtshuk (2016: 205): remark; Liu et al. (2020: 46): remark; Lawrence et al. (2021: 456): wing morphology. In addition to the aforementioned literature, this genus was included in PhD theses by Jeng (2008) and Roza (2022).

 Remarks.

Dodecatoma currently contains eight described species from Afghanistan, India, Nepal, Indonesia, and the Philippines. This genus is in need of revision; taxa from Southeast Asia should be removed from Dodecatoma, and the generic assignment of the species with serrate antennae described recently by Kazantsev (2012) needs careful re-examination (the remaining species of Dodecatoma, including the type species, have pectinate antennae).

 Animalia Coleoptera Rhagophthalmidae E8B995FD-B4FD-5C72-B0DF-1950C9E6E19B Dodecatoma bicolor Westwood, 1849 Dodecatoma bicolor Westwood, 1849: 1. Type depository.

Described based on an unknown number of specimens. Syntype, male (OUMNH).

 Type locality.

India: Deccan Plateau (without further details; “North India” on the label of the syntype in OUMNH).

 Distribution.

India (Karnataka, Maharashtra).

 Literature.

Westwood (1849: 1): original description, drawings of male habitus and body parts; Schaum (1850: 165): morphology, remark; Gemminger (1869: 1686): catalogue; Gorham (1895: 309): distributional note; Olivier (1910: 8): catalogue; Lucas (1920: 246): catalogue; Pic (1924: 714): comparison with D. testaceiceps Pic, 1924; Pic (1930b: 321): remark; Wittmer (1944: 211): catalogue; Lawrence and Newton (1995: 858): catalogue, remark; Bocak (2007: 225): catalogue; Johnson et al. (2012: 178): ICZN case; ICZN (2014: 195): ICZN case; Lawrence et al. (2021: 456): wing morphology, figure of hind wing. In addition to the aforementioned literature, this species was included in PhD theses by Jeng (2008) and Roza (2022).

 Animalia Coleoptera Rhagophthalmidae 4B374DE5-B946-5D8B-95B0-DFC9FB3DB09F Dodecatoma fuscicornis fuscicornis Gorham, 1895 Dodecatoma fuscicornis Gorham, 1895: 309. Type depository.

Described based on “several examples” (Gorham 1895: 309). Three syntypes, males (NHMUK). Several specimens from Belgaum deposited in MNHN are potentially syntypes (RK pers. obs.).

 Type locality.

India: Karnataka, Belgaum.

 Distribution.

India (Karnataka).

 Literature.

Gorham (1895: 309): original description; Olivier (1910: 8): catalogue; Wittmer (1944: 211): catalogue; Wittmer (1979: 90): comparison with other species; Johnson et al. (2012: 179): ICZN case.

 Animalia Coleoptera Rhagophthalmidae 661DC4F8-8805-52CF-B0ED-22B10DE19E3F Dodecatoma fuscicornis testaceicornis Pic, 1921 Dodecatoma fuscicornis var. testaceicornis Pic, 1921b: 12. Type depository.

Described based on an unknown number of specimens. Syntype, male (MNHN).

 Type locality.

Indonesia: Java.

 Distribution.

Indonesia (Java).

 Literature.

Pic (1921b: 12): original description; Wittmer (1944: 212): catalogue.

 Remarks.

The name “testaceicornis” is deemed to be subspecific according to Art. 45.6.4. of the Code (ICZN 1999). This taxon is not morphologically similar to D. fuscicornis Gorham, 1895 nor to any other species of Dodecatoma.

 Animalia Coleoptera Rhagophthalmidae 330A1435-D4AC-5FE7-935B-AB09FC2839CC Dodecatoma gracilis Wittmer, 1995 Dodecatoma gracilis Wittmer, 1995: 110. Type depository.

Holotype, male (NHMB). One paratype, male (NHMB).

 Type locality.

Nepal: near Simra Abhabar, 200 m.

 Distribution.

Nepal.

 Literature.

Wittmer (1995: 110): original description, figures of male antenna and genitalia; Bocak (2007: 225): catalogue; Johnson et al. (2012: 179): ICZN case; Kazantsev (2012: 349): comparison with D. saluki and D. schmidti, identification key.

 Animalia Coleoptera Rhagophthalmidae 289E71E3-C000-5BC8-A15F-DC5C23F3E277 Dodecatoma parvicornis Wittmer, 1979 Dodecatoma parvicornis Wittmer, 1979: 89. Type depository.

Holotype, male (NHMB). Two paratypes, males (NHMB).

 Type locality.

Afghanistan: Nuristan, Baschgultal.

 Distribution.

Afghanistan, Pakistan.

 Literature.

Wittmer (1979: 89): original description, drawing of antenna; Bocak (2007: 225): catalogue; Johnson et al. (2012: 179): ICZN case.

 Animalia Coleoptera Rhagophthalmidae C9E0D1A8-DCC2-5BAA-8FA8-525AE1DB9477 Dodecatoma riedeli Wittmer, 1995 Dodecatoma riedeli Wittmer, 1995: 112. Type depository.

Holotype, male (SMNS). Three paratypes, males (NHMB).

 Type locality.

India: Uttarakhand [“Uttar Pradesh”], Rishikesh.

 Distribution.

India (Uttarakhand).

 Literature.

Wittmer (1995: 112): original description, figures of male antenna and genitalia; Bocak (2007: 225): catalogue; Johnson et al. (2012: 179): ICZN case; Kazantsev (2012: 349): comparison with D. saluki and D. schmidti, identification key.

 Animalia Coleoptera Rhagophthalmidae CCA99148-96F4-5779-BE25-25814566734A Dodecatoma saluki Kazantsev, 2012 Dodecatoma saluki Kazantsev, 2012: 349. Type depository.

Holotype, male (ICM). One paratype, male (NKME).

 Type locality.

India: Uttarakhand [Uttaranchal], Nainital Distr., 5 km SE Mukteshwar, Satkhol.

 Distribution.

India (Uttarakhand), Nepal.

 Literature.

Kazantsev (2012: 349): original description, figures of male habitus, pregenital segments, and genitalia.

 Animalia Coleoptera Rhagophthalmidae F99B3A22-23E1-58EA-804B-087B98DA69F4 Dodecatoma schmidti Kazantsev, 2012 Dodecatoma schmidti Kazantsev, 2012: 349. Type depository.

Holotype, male (NKME).

 Type locality.

Nepal: Kali Gandaki valley, Upper Lete.

 Distribution.

Nepal.

 Literature.

Kazantsev (2012: 349): original description, drawings of male basal antennomeres and genitalia.

 Animalia Coleoptera Rhagophthalmidae 76678D3E-3F2D-5EF5-960E-7436140C45C8 Dodecatoma testaceiceps Pic, 1924 Dodecatoma testaceiceps Pic, 1924: 713. Dodecatoma testaceipes : Wittmer 1944: 212 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Type depository.

Described based on an unknown number of specimens (but probably only one). One syntype, male (MNHN).

 Type locality.

Philippines: Luzon, Mt. Maquiling.

 Distribution.

Philippines.

 Literature.

Pic (1924: 713): original description; Wittmer (1941: 197): catalogue; Wittmer (1944: 212): catalogue [as D. testaceipes [sic!]].

 Remarks.

This species clearly does not represent a member of Rhagophthalmidae and needs to be transferred into a proper family in a future revision.

 Animalia Coleoptera Rhagophthalmidae 7D56A634-B479-5222-9EA0-68DFEE97FAEE Falsophrixothrix Pic, 1937 Fig. 1D Falsophrixothrix Pic, 1937: 138. Gender: feminine. Type species. Phrixothrix javanus [sic!] Pic, 1914; by original designation (Pic 1937: 138). Literature.

Olivier (1911: 20): species description [as Phrixothrix]; Pic (1914: 13): species description [as Phrixothrix]; Pic (1921a: 16): species description [as Phrixothrix]; Pic (1937: 138): original generic description; Wittmer (1938: 301): description of an aberration [term used to denote a class of individuals within a species; unavailable name; see Glossary in ICZN (1999)]; Wittmer (1939: 23): species description; Wittmer (1944: 217): catalogue; Pic (1951: 5): species description; Crowson (1972: 52): remark; Paulus (1975: 78): remark; Herring (1978: 471): checklist; Herring (1987: 157): checklist; Viviani and Bechara (1993: 615): remark; Lawrence and Newton (1995: 857): catalogue, remark; Viviani and Bechara (1997: 389): remark; O‘Keefe (2002: 182): remark; Li et al. (2008b: 495): review; Kawashima et al. (2010: 139): book chapter; Lawrence et al. (2010b: 175): remark; Kundrata and Bocak (2011a: 57): remark; Oba et al. (2011: 777): remark; Janisova and Bocakova (2013: 3): remark; Kovalev and Kirejtshuk (2016: 205): remark; Kundrata et al. (2019: 1263): molecular phylogeny; Douglas et al. (2021: 2): molecular phylogeny. In addition to the aforementioned literature, this genus was included in PhD theses by Jeng (2008) and Roza (2022).

 Remarks.

Falsophrixothrix currently contains six described and several undescribed species from Southeast Asia. It can be recognized by its small body size, strongly bipectinate antennae (Fig. 1D), and usually shortened elytra which do not cover the entire abdomen. It should be noted that all previous authors treated the gender of Falsophrixothrix as masculine; however, -thrix (hair in Greek) is feminine.

 Animalia Coleoptera Rhagophthalmidae BA0BB5FF-34D5-5AA2-9C55-4849D168BF3E Falsophrixothrix costata Pic, 1951 Falsophrixothrix costatus [sic!] Pic, 1951: 5. Type depository.

Described based on an unknown number of specimens. Syntype, male (MNHN).

 Type locality.

Vietnam: Ho Chi Minh City [Saigon].

 Distribution.

Vietnam.

 Literature.

Pic (1951: 5): original description.

 Animalia Coleoptera Rhagophthalmidae BB72D281-72BD-5DA6-8A28-163F3AB82B7A Falsophrixothrix flava Wittmer, 1939 Falsophrixothrix flavus [sic!] Wittmer, 1939: 23. Type depository.

Described based on two specimens. Holotype, male (NHMB); paratype, male (?MZB; in Drescher coll. according to the original description).

 Type locality.

Indonesia: Java, Parahyangan (= Priangan, Preanger), Tangkuban Perahu [G. Tangkoeban Prahoe].

 Distribution.

Indonesia (Java).

 Literature.

Wittmer (1939: 23): original description; Wittmer (1944: 217): catalogue.

 Animalia Coleoptera Rhagophthalmidae C75568F4-C11D-547A-A137-0243FBD502C6 Falsophrixothrix humeralis Pic, 1937 Falsophrixothrix humeralis Pic, 1937: 138. Falsophrixothrix humeralis ab. unicolor Wittmer, 1938: 301 [unavailable name, ICZN 1999]. Type depository.

Described based on an unknown number of specimens. Syntype (labelled as “Holotypus”), male (NHMB).

 Type locality.

Indonesia: Java, Parahyangan (= Priangan, Preanger), Tangkuban Perahu [G. Tangkoeban Prahoe] [only “Java” in the original description, remaining information taken from the locality label under the syntype].

 Distribution.

Indonesia (Java).

 Literature.

Pic (1937: 138): original description; Wittmer (1938: 301): description of F. humeralis ab. unicolor; Wittmer (1939: 24): comparison with F. flavus [sic!]; Wittmer (1944: 217): catalogue; Pic (1951: 5): comparison with F. costatus [sic!]. In addition to the aforementioned literature, this species was included in a PhD thesis by Jeng (2008).

 Remarks.

Wittmer (1938: 301) described the aberration of F. humeralis (ab. unicolor) from Tangkuban Perahu [“G. Tangkoeban Prahoe”] based on material from the collection of F. C. Drescher (possibly in MZB); however, this name is deemed to be infrasubspecific according to the Code (ICZN 1999, Article 45.6.2.).

 Animalia Coleoptera Rhagophthalmidae B3DC8E75-AE47-57D9-86F2-DC1F022D5A39 Falsophrixothrix javana (Pic, 1914) Phrixothrix javanus [sic!] Pic, 1914: 13. Falsophrixothrix javanus [sic!]: Pic 1937: 138. Type depository.

Described based on an unknown number of specimens. Syntype, male (MNHN).

 Type locality.

Indonesia: Java.

 Distribution.

Indonesia (Java).

 Literature.

Pic (1914: 13): original description [as Phrixothrix]; Pic (1921a: 16): comparison with F. punctatus [sic!] (Pic, 1921) [as Phrixothrix]; Pic (1937: 138): comparison with F. humeralis; Wittmer (1939: 24): comparison with F. flavus [sic!]; Wittmer (1944: 217): catalogue.

 Animalia Coleoptera Rhagophthalmidae F5F99FE0-E573-55B0-B6C8-C9CE52DA4F6A Falsophrixothrix punctata (Pic, 1921) Phrixothrix punctatus [sic!] Pic, 1921a: 16. Falsophrixothrix punctatus [sic!]: Wittmer 1944: 217. Type depository.

Described based on an unknown number of specimens. Syntype, male (MNHN).

 Type locality.

Singapore.

 Distribution.

Singapore.

 Literature.

Pic (1921a: 16): original description [as Phrixothrix]; Wittmer (1944: 217): catalogue.

 Animalia Coleoptera Rhagophthalmidae 59B93D8D-12EB-5D5A-885B-DADB41FE9500 Falsophrixothrix pygmaea (Olivier, 1911) Phrixothrix pygmaeus [sic!] E. Olivier, 1911: 19. Falsophrixothrix pygmaeus [sic!]: Wittmer 1939: 24. Type depository.

Described based on an unknown number of specimens. At least one syntype, male (RMNH). Five additional male specimens in RMNH (originally from the Zoological Museum, Amsterdam, ZMAN) may also be syntypes (RK pers. obs.).

 Type locality.

Indonesia, Java: Banyuwangi [Banjoewangi].

 Distribution.

Indonesia (Java).

 Literature.

Olivier (1911: 19): original description [as Phrixothrix]; Wittmer (1939: 24): comparison with F. flavus [sic!]; Wittmer (1944: 217): catalogue.

 Animalia Coleoptera Rhagophthalmidae EAF9466D-67F3-565B-8A06-24E3B422EDC4 Haplocladon Gorham, 1883 Haplocladon Gorham, 1883b: 249 [as a subgenus of Diplocladon Gorham, 1883]. Gender: neuter. Type species. Haplocladon gorhami Kundrata, 2022, nom. nov. [replacement name for Diplocladon hasseltii Gorham, 1883b]; by monotypy. Literature.

Gorham (1883b: 249): original description; Gorham (1895: 310): remark; Gorham (1903: 330): species description; Olivier (1910: 8): catalogue; Wittmer (1944: 211): catalogue; Crowson (1955: 68): remark; Paulus (1972: 49): remark; Li and Liang (2008: 109): remark [as D. haplocladon [sic!]]. In addition to the aforementioned literature, this genus was included in a PhD thesis by Jeng (2008).

 Remarks.

Gorham (1883b) originally described Haplocladon as a subgenus of Diplocladon but later treated it as a separate genus (Gorham 1895, 1903). Unfortunately, he named type species of both Diplocladon and Haplocladon as “hasseltii” (Gorham 1883a, b), which probably confused some subsequent authors who treated Haplocladon as a synonym of Diplocladon (Olivier 1910; Wittmer 1944; Li and Liang 2008). Crowson (1955) and Paulus (1972) again considered Haplocladon a separate genus. Since Haplocladon differs at first sight from Diplocladon by the unipectinate antennae (versus bipectinate in Diplocladon), we prefer to keep Haplocladon at a generic level. Because Diplocladon hasseltii Gorham, 1883a and Diplocladon hasseltii Gorham, 1883b (described in subgenus Haplocladon) are primary homonyms, the latter junior name is permanently invalid (Art. 57.2 of the Code; ICZN 1999) and should be replaced by a new name (see below). Currently, Haplocladon contains two species, one from Indonesia and one from southern India. Based on the available figures, specimens reported by Haneda (1950) from Singapore and identified as Diplocladon hasseltii, which were later mentioned by other authors (e.g., Harvey 1952; Haneda 1955; Lloyd 1971; Lloyd 1978; Crowson 1981; Haneda 1985; Kawashima et al. 2010), are probably members of Haplocladon.

 Animalia Coleoptera Rhagophthalmidae 2DB5AC3A-A0E0-56B0-BB74-4998C7E54BC8 Haplocladon gorhami Kundrata nom. nov. Diplocladon hasseltii Gorham, 1883b: 250 (described in subgenus Haplocladon). Preoccupied by Diplocladon hasseltii Gorham, 1883a: 6. Haplocladon haselti : Gorham 1903: 330 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Type depository.

Described based on an unknown number of specimens. Two syntypes, males (one from Sumatra, one from Java) (RMNH).

 Type locality.

Indonesia: Sumatra, Lampung, Soekadana; Java, Batavia.

 Distribution.

Indonesia (Sumatra, Java).

 Literature.

Gorham (1883b: 250): original description; Gorham (1903: 330): remark [as H. haselti [sic!]]; Olivier (1910: 8): catalogue [as D. hasselti]; Wittmer (1944: 211): catalogue [as D. hasselti].

 Remarks.

Gorham (1883b: 250) also reported an unnamed variety of H. hasseltii as “var. totum testaceum” (i.e., colour description but not the official name of the variety) from Ardjoeno and Batavia in Java. At least one specimen from Batavia labelled as “var.” is present in MNHN. Two specimens from Ardjoeno and one specimen from Batavia deposited in RMNH bear the label “Type”; however, based on Article 72.4.1. of the Code (ICZN 1999), they should not be considered a part of the type series for Haplocladon gorhami.

 Animalia Coleoptera Rhagophthalmidae 684F1691-F8CD-5669-A2A3-38C48B6B9D97 Haplocladon indicum Gorham, 1903 Haplocladon indicum Gorham, 1903: 330. Diplocladon indicum : Olivier 1910: 8. Type depository.

Holotype, male (MNHN).

 Type locality.

India: Nilgiri Hills.

 Distribution.

India (Nilgiri Hills).

 Literature.

Gorham (1903: 330): original description; Olivier (1910: 8): catalogue [as D. indicum]; Wittmer (1944: 211): remark [as D. indicum]; Li and Liang (2008: 109): remark [as D. haplocladon indicum [sic!]]; Yiu (2017: 64): comparison with Diplocladon atripennis [sic!] [as D. indicum].

 Animalia Coleoptera Rhagophthalmidae 3074AEFA-C43E-53EA-9545-A062C81DFCED Menghuoius Kawashima, 2000 Menghuoius Kawashima, 2000: 132. Gender: masculine. Type species. Rhagophthalmus ingens Fairmaire, 1896, by original designation. Menghouius : Bocak 2007: 225 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Menhuoius : Li et al. 2008a: 264 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Distribution.

China (Anhui, Guangxi, ?Hong Kong, Yunnan, Zhejiang), Myanmar, Vietnam.

 Literature.

Kawashima (2000: 132): original description; Kawashima (2002: 487): species description, figures of habitus, body parts, and male genitalia; Bocak (2007: 225): catalogue [as Menghouius [sic!]]; Li et al. (2008a: 264): distribution, morphology, biology, figures of male antenna and genitalia, and larval and female habitus [as Menghouius [sic!]]; Li et al. (2008b: 495): review; Kawashima et al. (2010: 136): book chapter, drawings of head, tarsi, and antenna; Chen et al. (2019: 3): molecular phylogeny; Liu et al. (2020: 46a): luciferase, phylogeny, figures of male and female habitus, and female bioluminescence. In addition to the aforementioned literature, this genus was included in a PhD thesis by Jeng (2008).

 Remarks.

Menghuoius currently contains three described species from China, Myanmar, and Vietnam. It is similar to Rhagophthalmus in habitus, short, serrate antennae, and deeply emarginate eyes but differs in the large size and robust mandibles (Kawashima 2000). Menghuoius was implicitly considered a junior synonym of Rhagophthalmus by Li et al. (2008a) based on the similar morphology of females of both genera. However, since the morphology of highly paedomorphic, larva-like females of Rhagophthalmidae is much less informative than the morphology of adult males, we consider Menghuoius a separate genus until a detailed revision of Rhagophthalmus and related genera is carried out.

 Animalia Coleoptera Rhagophthalmidae B7212A60-A337-5C85-96F3-09FF095A9050 Menghuoius giganteus (Fairmaire, 1888) Rhagophthalmus giganteus Fairmaire, 1888: 25. Menghuoius giganteus : Kawashima 2000: 139. Rhagophthalmus gigantus : Moreira et al. 2022: 7 [unavailable name, incorrect subsequent spelling not in prevailing usage; page number may be changed when the publication is printed]. Type depository.

Described based on an unknown number of specimens. Syntype, male (MNHN).

 Type locality.

China: Yunnan.

 Distribution.

China (Anhui, Guangxi, Yunnan, Zhejiang).

 Literature.

Fairmaire (1888: 25): original description [as R. giganteus]; Fairmaire (1896: 227): comparison with R. ingens [as R. giganteus]; Olivier (1902: 88): catalogue [as R. giganteus]; Olivier (1910: 1): catalogue [as R. giganteus]; Jakobson (1911: 687): catalogue [as R. giganteus]; Olivier (1912: 469): revision [as R. giganteus]; Winkler (1925: 522): catalogue [as R. giganteus]; Wu (1937: 385): catalogue [as R. giganteus]; McDermott (1966: 122): catalogue [as R. giganteus]; Kawashima (2000: 139): comparison with R. ingens [as R. giganteus]; Hua (2002: 71): catalogue [as R. giganteus]; Bocak (2007: 225): catalogue; Li et al. (2008a: 264): distribution, morphology, biology, figures of male antenna and genitalia, and larval and female habitus [also as R. giganteus]; Li et al. (2008b: 496): review [as R. giganteus]; Chen et al. (2019: 3): molecular phylogeny; Liu et al. (2020: 46a): luciferase, phylogeny, figures of male and female habitus, and female bioluminescence; Moreira et al. (2022: 7): luciferase, molecular phylogeny [as R. gigantus [sic!]]. In addition to the aforementioned literature, this species was included in a PhD thesis by Jeng (2008).

 Animalia Coleoptera Rhagophthalmidae 709374AC-B1BB-5FEC-B833-F11691B0E094 Menghuoius ingens (Fairmaire, 1896) Rhagophthalmus ingens Fairmaire, 1896: 227. Menghuoius ingens : Kawashima 2000: 134. Type depository.

Described based on an unknown number of specimens. Syntype, male (MNHN).

 Type locality.

China: probably Hong Kong (Fairmaire 1896).

 Distribution.

China (?Hong Kong), Vietnam.

 Literature.

Fairmaire (1896: 227): original description [as R. ingens]; Olivier (1902: 88): catalogue [as R. ingens]; Olivier (1910: 1): catalogue [as R. ingens]; Jakobson (1911: 687): catalogue [as R. ingens]; Olivier (1912: 469): revision [as R. ingens]; Winkler (1925: 522): catalogue [as R. ingens]; Wu (1937: 385): catalogue [as R. ingens]; McDermott (1966: 122): catalogue [as R. ingens]; Kawashima (2000: 134): redescription; Hua (2002: 71): catalogue [as R. ingens]; Kawashima (2002: 491): comparison with M. kusakabei; Bocak (2007: 225): catalogue [also as R. ingens]; Li et al. (2008a: 264): remark, distribution [also as R. ingens]; Li et al. (2008b: 496): review [as R. ingens]; Kawashima et al. (2010: 136): book chapter, drawings of head, tarsi and antenna; Yiu (2017: 60): comparison with R. motschulskyi [as R. ingens]; Chen et al. (2019: 11): molecular phylogeny; Liu et al. (2020: 47): remark. In addition to the aforementioned literature, this species was included in a PhD thesis by Jeng (2008).

 Remarks.

Olivier (1912) mentioned that R. ingens could be conspecific with R. giganteus.

 Animalia Coleoptera Rhagophthalmidae 58188A76-AC6C-5932-9346-8AF65C685E1F Menghuoius kusakabei Kawashima, 2002 Menghuoius kusakabei Kawashima, 2002: 487. Type depository.

Holotype, male (NWU). Four paratypes, males (PCIK).

 Type locality.

Myanmar: Chin state, Natma Taung National Park near Kanpetlet, Mt. Victoria, ca. 2000 m.

 Distribution.

Myanmar.

 Literature.

Kawashima (2002: 487): original description, figures of habitus, body parts, male genitalia.

 Remarks.

Jeng (2008) reported a possible female of M. kusakabei from Myanmar.

 Animalia Coleoptera Rhagophthalmidae CB04E685-D7A4-5142-834F-477A85A3AA11 Mimoochotyra Pic, 1937 Mimoochotyra Pic, 1937: 137. Gender: feminine. Type species. Mimoochotyra ocularis Pic, 1937; by monotypy. Mimochotyra : McDermott 1964: 11 [unavailable name, incorrect subsequent spelling]. Mimotyra : Herring 1987: 158 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Mimochotrya : Lawrence and Newton 1995: 857 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Literature.

Pic (1937: 137): original description; McDermott (1964: 11, 51): revision [as Mimochotyra [sic!]]; McDermott (1966: 122): catalogue [as Mimochotyra [sic!]]; Mikšić and Mikšić (1966: 32): remark [as Mimochotyra [sic!]]; Herring (1987: 158): checklist [as Mimotyra [sic!]]; Lawrence and Newton (1995: 857): catalogue [as Mimochotrya [sic!]]; Kawashima (2000: 131): remark [as Mimochotyra [sic!]]; Bocakova et al. (2007: 484): molecular phylogeny [as Mimochotyra [sic!]]; Hunt et al. (2007: suppl.): molecular phylogeny [as Mimochotyra [sic!]]; Li et al. (2008a: 259): remark [as Mimochotyra [sic!]]; Li et al. (2008b: 495): review [as Mimochotyra [sic!]]; Li and Liang (2008: 111): remark [as Mimochotyra [sic!]]; Chen et al. (2010: 196): remark [as Mimochotyra [sic!]]; Costa and Zaragoza-Caballero (2010: 134): remark [as Mimochotyra [sic!]]; Kawashima et al. (2010: 135): book chapter [as Mimochotyra [sic!]]; Kundrata and Bocak (2011a: 59): remark [as Mimochotyra [sic!]]; Kundrata and Bocak (2011b: 370): molecular phylogeny [as Mimochotyra [sic!]]; Oba et al. (2011: 777): remark [as Mimochotyra [sic!]]; Kundrata et al. (2013: 202): molecular phylogeny; Kundrata et al. (2014: 167): molecular phylogeny; Bocak et al. (2016: suppl.): molecular phylogeny; Kundrata et al. (2016: 296): molecular phylogeny; Bocak et al. (2018: 4): molecular phylogeny; Kundrata et al. (2019: 1263): molecular phylogeny; Liu et al. (2020: 46): remark [as Mimochotyra [sic!]]. In addition to the aforementioned literature, this genus was included in a PhD thesis by Jeng (2008).

 Remarks.

This genus currently contains a single described species from Java, Indonesia. According to Pic (1937), it is characterized by having serrate antennae with thickened median antennomeres, and relatively long elytra. The specimen identified as Mimochotyra [sic!] and used in the molecular phylogenetic analyses by Bocakova et al. (2007) and more recent studies needs serious re-examination, as it was collected in Malaysia.

 Animalia Coleoptera Rhagophthalmidae F72C2B9F-3A3E-5678-AA57-7F1E1E6F2956 Mimoochotyra ocularis Pic, 1937 Mimoochotyra ocularis Pic, 1937: 137. Type depository.

Described based on an unknown number of specimens (probably only one). Syntype, male (NHMB).

 Type locality.

Indonesia: Java, Gunung Raung [Raoeng], “Bajoekidoel” [detailed data taken from the syntype label; only “Java: Bajoekidoe” [sic!] in original description].

 Distribution.

Indonesia (Java).

 Literature.

Pic (1937: 137): original description; McDermott (1964: 51): revision [as Mimochotyra [sic!]]; McDermott (1966: 122): catalogue [as Mimochotyra [sic!]]; Li et al. (2008b: 496): review [as Mimochotyra [sic!]].

 Animalia Coleoptera Rhagophthalmidae 1D2FB02D-5D5E-56A0-B6AB-0A3C3F91D8A5 Monodrilus Pic, 1921 Monodrilus Pic, 1921b: 12. Gender: masculine. Type species. Monodrilus marginatus Pic, 1921; by monotypy. Remarks.

Monodrilus has more or less serrate antennae and relatively long elytra, and currently contains two species from Indonesia (Java) and Vietnam, respectively, each in a monotypic subgenus. Following Wittmer (1944), we retain the concept of Monodrilus with two subgenera; however, Pic (1930b) already suggested Dodecatomorpha could be a separate genus. A proper taxonomic revision should be conducted to resolve the status of Dodecatomorpha.

 Animalia Coleoptera Rhagophthalmidae 8F01BF07-D4C6-5884-9B1E-70C7EA9CBA07 Monodrilus Pic, 1921 Monodrilus Pic, 1921b: 12. Gender: masculine. Type species. Monodrilus marginatus Pic, 1921; by monotypy. Literature.

Pic (1921b: 12): original description; Pic (1928: 86): comparison with Dodecatomorpha Pic, 1928; Pic (1930b: 321): remark; Wittmer (1944: 212): catalogue; Lawrence et al. (2010b: 175): remark; Kundrata and Bocak (2011a: 57): remark; Janisova and Bocakova (2013: 3): remark. In addition to the aforementioned literature, Monodrilus was included in a PhD thesis by Jeng (2008).

 Remarks.

This subgenus currently contains a single described species from Java, Indonesia.

 Animalia Coleoptera Rhagophthalmidae F26B13F4-5C8E-5512-859D-E3D3A411915F Monodrilus marginatus Pic, 1921 Monodrilus marginatus Pic, 1921b: 12. Type depository.

Described based on an unknown number of specimens. Syntype, male (MNHN).

 Type locality.

Indonesia: Java.

 Distribution.

Indonesia (Java).

 Literature.

Pic (1921b: 12): original description; Pic (1928: 87): comparison with Dodecatomorpha roberti Pic, 1928; Wittmer (1944: 212): catalogue. In addition to the aforementioned literature, this species was included in a PhD thesis by Jeng (2008).

 Animalia Coleoptera Rhagophthalmidae 239836B9-2B1C-5852-852A-BA6FA2ACEA24 Dodecatomorpha Pic, 1928 Dodecatomorpha Pic, 1928: 86 [as a subgenus of Monodrilus Pic, 1921]. Gender: feminine. Type species. Monodrilus roberti Pic, 1928 [in subgenus Dodecatomorpha]; by monotypy. Literature.

Pic (1928: 86): original description; Pic (1930b: 321): remark; Wittmer (1944: 212): catalogue.

 Remarks.

Dodecatomorpha currently contains a single described species from Vietnam.

 Animalia Coleoptera Rhagophthalmidae 90CD2D3D-A7BE-5F3E-BB66-1075D0783AEF Monodrilus (Dodecatomorpha) roberti Pic, 1928 Monodrilus roberti Pic, 1928: 86 [in subgenus Dodecatomorpha]. Type depository.

Described based on an unknown number of specimens. Three syntypes, males (MNHN).

 Type locality.

Vietnam [“Darsa, en Cochinchine”].

 Distribution.

Vietnam.

 Literature.

Pic (1928: 86): original description; Pic (1930b: 321): remark; Wittmer (1944: 212): catalogue.

 Animalia Coleoptera Rhagophthalmidae 60995E64-A2AF-528A-BE13-BF96AB15E6B6 Pseudothilmanus Pic, 1918 Pseudothilmanus Pic, 1918: 2. Gender: masculine. Type species: Pseudothilmanus alatus Pic, 1918; by monotypy. Drilothilmanus Pic, 1918: 3. Type species: Drilothilmanus marginatus, 1918; by monotypy. Synonymized by Kundrata and Bocak (2011a: 58). Literature.

Pic (1918: 2, 3): original descriptions of Pseudothilmanus and Drilothilmanus, respectively; Wittmer (1944: 215): catalogue [also as Drilothilmanus]; Kundrata and Bocak (2011a: 58): revision, synonymization of Drilothilmanus; Liu et al. (2020: 46): remark; Roza (2020: 421): morphology, distribution, figures of male habitus, pronotum, and hind wing [2018 erroneously used as the date of the original description of this genus in figure caption]. In addition to the aforementioned literature, this genus was included in a PhD thesis by Roza (2022).

 Remarks.

This genus has relatively long, serrate antennae and long elytra. It contains two species distributed in the Himalayas (India, Nepal).

 Animalia Coleoptera Rhagophthalmidae BF7F1F35-F3D7-5DDC-AC1B-6DAE0FEC59C0 Pseudothilmanus alatus Pic, 1918 Pseudothilmanus alatus Pic, 1918: 2. Type depository.

Described based on an unknown number of specimens. Lectotype by present designation, with the following label data: “Type [red printed label] / Type [handwritten] / Nov. genus India [handwritten] / Pseudothilmanus alatus Pic [handwritten]” (treated as the holotype and figured by Kundrata and Bocak 2011a), male (MNHN).

 Type locality.

India (without any further details).

 Distribution.

India (Uttarakhand), Nepal.

 Literature.

Pic (1918: 2): original description; Wittmer (1944: 215): catalogue; Kundrata and Bocak (2011a: 59): revision, figures of male habitus, antenna, pronotum, leg, elytral apex, pregenital segments, and genitalia; Roza (2020: 421): morphology, distribution, figures of male habitus, pronotum, and hind wing [2018 erroneously used as the date of the original description of this species in figure caption]. In addition to the aforementioned literature, this species was included in a PhD thesis by Roza (2022).

 Remarks.

Roza (2020) listed Uttar Pradesh for the distribution of this species; however, it was based on the specimen from NHMUK mentioned by Kundrata and Bocak (2011a), which was collected in western Almora, Kumaon which lies in Uttarakhand (considered to be part of Uttar Pradesh prior to 2000).

 Animalia Coleoptera Rhagophthalmidae 998E819E-5DD3-5F78-904D-DF6AEB9C23A1 Pseudothilmanus marginatus Pic, 1918 Drilothilmanus [as a subgenus of Pseudothilmanus] marginatus Pic, 1918: 3. Pseudothilmanus marginatus : Kundrata and Bocak (2011a: 58). Type depository.

Described based on an unknown number of specimens. Lectotype by present designation, with the following label data: “Type [red printed label] / Type [handwritten] / Darjeeling Juni Fruhstorfer leg. [printed] / Drilothilmanus marginatus Pic [handwritten]” (treated as the holotype and figured by Kundrata and Bocak 2011a), male (MNHN).

 Type locality.

India: West Bengal, Darjeeling.

 Distribution.

India (West Bengal).

 Literature.

Pic (1918: 3): original description; Wittmer (1944: 215): catalogue [as Drilothilmanus]; Kundrata and Bocak (2011a: 60): revision, figures of male habitus, antenna, pronotum, leg, and genitalia; Roza (2020: 422): morphology, distribution, figures of male habitus, pronotum, and hind wing [2018 erroneously used as the date of the original description of this species in figure caption].

 Animalia Coleoptera Rhagophthalmidae 798AC722-75B9-542E-96DF-5B8ED0E4448F Rhagophthalmus Motschulsky, 1854 Fig. 1E, F Rhagophthalmus Motschulsky, 1854: 45. Gender: masculine. Type species: Rhagophthalmus scutellatus Motschulsky, 1854, by monotypy. Ochotyra Pascoe, 1862: 323. Gender: feminine. Type species: Ochotyra semiusta Pascoe, 1862: 323, by monotypy. Synonymized with Rhagophthalmus Motschulsky, 1854 by Wittmer in Wittmer and Ohba (1994: 342). Ochotiza : Bourgeois 1903: 479 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Ochrotyra : Lefroy 1909: 327 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Rhagophthalma : Crowson 1981: 274 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Ochotrya : Lawrence 1988: 15 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Rhagophthalums : Suzuki 1997: 38 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Ragophthalmus : Viviani et al. 1999: 8274 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Rhagophtha : Chen 2003: 52 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Rhagophtalmus : Stanger-Hall et al. 2007: 38 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Rhagopthalmus : McKenna et al. 2015: 849 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Literature.

Motschulsky (1854: 45): original description; Motschulsky (1859: 59): remark; Motschulsky (1861: 134): comparison with Pachytarsus Motschulsky, 1861; Pascoe (1862: 323): original description of Ochotyra; Gerstaecker (1863: 409): remark [as Ochotyra]; Gemminger (1869: 1647, 1655): catalogue [also as Ochotyra]; Marschall (1873: 223, 239): remark [also as Ochotyra]; Gorham (1881: 63): remark [as Ochotyra]; Olivier (1885: 372): species description; Heyden (1886: 286): remark; Fairmaire (1888: 25): species description [currently in Menghuoius]; Fairmaire (1889: 352): species description; Gorham (1890: 550): catalogue [as Ochotyra]; Bourgeois (1892: 236): distributional note; Cardon (1892: 238): checklist; Gorham (1895: 310): distributional note [also as Ochotyra]; Fairmaire (1896: 227): species descriptions [one currently in Menghuoius]; Fairmaire (1899: 624): species description; Olivier (1902: 87): catalogue; Bourgeois (1903: 479): distributional note [as Ochotiza [sic!]]; Gorham (1903: 330): distributional note [as Ochotyra]; Bourgeois (1905: 130): distributional record [as Ochotyra]; Olivier (1907: 63): catalogue [also as Ochotyra]; Olivier (1908: 17): remark; Lefroy (1909: 327): catalogue [also as Ochrotyra [sic!]]; Olivier (1910: 1): catalogue [also as Ochotyra]; Jakobson (1911: 687): catalogue; Olivier (1912: 467): revision, key [also as Ochotyra]; Pic (1916: 9): species description; Pic (1917: 3): species description; Lucas (1920: 567): catalogue; Pic (1921b: 18): species description [as Ochotyra]; Pic (1923: 25): catalogue; Handlirsch (1925: 589): catalogue; Pic (1925a: 17): species description; Pic (1925b: 72): species description; Winkler (1925: 522): catalogue; Pic (1937: 137): comparison of Ochotyra with Mimoochotyra; Wu (1937: 385): catalogue; Pic (1938: 15): checklist; Harvey (1952: 392): remark [also as Ochotyra]; Brues et al. (1954: 565): classification; Raj (1957: 788): larval biology, photograph of larvae; McDermott (1964: 11, 50): revision [also as Ochotyra]; McDermott (1966: 121): catalogue [also as Ochotyra]; Mikšić and Mikšić (1966: 32): remark [also as Ochotyra]; Nakane (1968: 3): remark; Crowson (1972: 52): remark; Herring (1978: 471): checklist; Lloyd (1978: 254): remark; Ohba (1980: 14): remark; Crowson (1981: 274): remark [as Rhagophthalma [sic!]]; Sivinski (1981: 168): bioluminescence; Herring (1987: 157): checklist [also as Ochotyra]; Lawrence (1988: 15): remark [also as Ochotrya [sic!]]; Wittmer and Ohba (1994: 341): review, synonymy of Ochotyra with Rhagophthalmus, figures of habitus and body parts; Lawrence and Newton (1995: 857): catalogue, remark [also as Ochotrya [sic!]]; Ohba (1995: 13): remark, bioluminescence; Branham (1996: 18): remark; Chen and Ho (1996: 46): distribution, figure of habitus; Ohba et al. (1996a: 1): morphology, biology; Ohba et al. (1996b: 30): remark; Nakane (1997: 36): remark; Ohba (1997a: 5): checklist; Ohba (1997b: 19): remark; Ohba (1997c: 51): breeding; Ohba et al. (1997: 25): remark; Suzuki (1997: 4): phylogeny, biology; Wittmer (1997: 257): species descriptions; Chen and Ho (1998: 34): bioluminescence; Kawashima (1998: 16): female morphology; Ohba (1998: 3): checklist, biology; Costa et al. (1999: 22): remark; Kawashima (1999: 141): remark; Viviani et al. (1999: 8274): remark [as Ragophthalmus [sic!]]; Goto and Kawashima (2000: 143): distributional remark; Jeng et al. (2000: 316): remark; Kawashima (2000: 131): taxonomy; Kim et al. (2000: 214): molecular phylogeny; Ohmiya et al. (2000: 32): luciferase; Viviani and Ohmiya (2000: 267): remark [as Ragophthalmus [sic!]]; Branham and Wenzel (2001: 565): phylogeny; Kawashima and Satô (2001: 423): species descriptions [also as Ochotyra]; Kobayashi et al. (2001: 1): embryogenesis; Viviani et al. (2001: 1287): bioluminescence [as Ragophthalmus [sic!]]; Hua (2002: 71): catalogue; Kawashima (2002: 492): remark; Kobayashi et al. (2002: 1): embryogenesis; O‘Keefe (2002: 182): remark; Ugarova and Brovko (2002: 322): bioluminescence; Viviani (2002: 1836): remark [as Ragophthalmus [sic!]]; Viviani et al. (2002: 538): remark [as Ragophthalmus [sic!]]; Branham and Wenzel (2003: 3): phylogeny, remark; Chen (2003: 52): morphology, bioluminescence [also as Rhagophtha [sic!]]; Hayashi and Suzuki (2003: 4): morphology, biology, phylogeny; Kawashima and Sugaya (2003: 353): species description, identification key; Kawashima et al. (2003: 255): catalogue [also Ochotyra]; Kobayashi et al. (2003: 19): embryogenesis, development; Satô and Kawashima (2003: 9): remark; DeCock (2004: 341): bioluminescence; Ohba (2004a: 226): bioluminescence, biology; Ohba (2004b: 6): bioluminescence, biology; Lau and Meyer-Rochow (2006: 20): morphology; Li et al. (2006: 818): molecular phylogeny; Arnoldi et al. (2007: 2): molecular phylogeny; Bocak (2007: 225): catalogue [also as Ochotyra]; Bocakova et al. (2007: 484): molecular phylogeny [as Ochotyra]; Geisthardt and Satô (2007: 234): catalogue [species incertae sedis in Lampyridae]; Hunt et al. (2007: suppl.): molecular phylogeny [as Ochotyra]; Lau et al. (2007: 27): eye morphology; Li et al. (2007: 197): mitochondrial genome, phylogeny; Sagegami-Oba et al. (2007: 105): molecular phylogeny; Stanger-Hall et al. (2007: 38): molecular phylogeny [also as Rhagophtalmus [sic!]]; Bocak et al. (2008: 2019): molecular phylogeny [as Ochotyra]; Dong et al. (2008: 479): phylogeny; Li and Liang (2008: 109): female morphology; Li et al. (2008a: 259): species descriptions, taxonomy, distribution [also as Ochotyra]; Li et al. (2008b: 494): review [also as Ochotyra]; Noguchi et al. (2008: 2): luciferase; Sheffield et al. (2008: 2500): mitochondrial genomes; Bogahawatta et al. (2009: 10): remark; Day et al. (2009: 93): remark; Levkanicova and Bocak (2009: 212): molecular phylogeny; Suzuki and Kobayashi (2009: 30): embryogenesis [also as Ochotyra]; Chen et al. (2010: 196): biology, bioluminescence; Costa and Zaragoza-Caballero (2010: 134): remark [also as Ochotyra]; Kawashima et al. (2010: 135): book chapter [also as Ochotyra]; Lawrence et al. (2010b: 173): remark; Bouchard et al. (2011: 326): family-group names catalogue; Kundrata and Bocak (2011a: 57): remark [also as Ochotyra]; Kundrata and Bocak (2011b: 370): molecular phylogeny [as Ochotyra]; Lawrence et al. (2011: 7): phylogeny; Oba et al. (2011: 777): biology, bioluminescence; Yiu (2011a: 14): remark; Yiu (2011b: 20): biology, bioluminescence; Amaral et al. (2012: 1262): luciferase, phylogeny; Ho et al. (2012: 1): species descriptions; Kazantsev (2012: 352): remark; Timmermans and Vogler (2012: 300): molecular phylogeny; Yiu (2012: 30): catalogue; Kundrata et al. (2013: 202): molecular phylogeny [as Ochotyra]; Yiu (2013: 101): biology, bioluminescence; Amaral et al. (2014: 415): molecular phylogeny [also as Rhagophtalmus [sic!]]; Hosoe et al. (2014: 331): chemical defence; Kundrata et al. (2014: 167): molecular phylogeny; Li et al. (2015: 269): catalogue; Martin et al. (2015: 519): molecular phylogeny; McKenna et al. (2015: 843): molecular phylogeny [also as Rhagopthalmus [sic!]]; Oba (2015: 99): bioluminescence; Amaral et al. (2016: 254): molecular phylogeny; Bocak et al. (2016: 3): molecular phylogeny; Kovalev and Kirejtshuk (2016: 205): remark; Kundrata et al. (2016: 296): molecular phylogeny; Wijekoon et al. (2016: 71): checklist [also as Ochotyra]; Amaral et al. (2017a: 674): mitogenome, phylogeny; Amaral et al. (2017b: 84): phylogeny; Amaral et al. (2017c: 157): phylogeny; Martin et al. (2017: 568): molecular phylogeny; Wang et al. (2017: 2): molecular phylogeny, transcriptome; Yiu (2017: 59): species description; Bocak et al. (2018: suppl): molecular phylogeny; Fallon et al. (2018: 8, 96): genomes, bioluminescence; Kusy et al. (2018a: 5): molecular phylogeny; Kusy et al. (2018b: 4): molecular phylogeny; Stanger-Hall et al. (2018: 8): remark; Yiu and Jeng (2018: 72): remark; Zhang et al. (2018: 3): molecular phylogeny; Amaral et al. (2019: 284): molecular phylogeny; Chen et al. (2019: 8): molecular phylogeny; He et al. (2019: 566): molecular phylogeny; Jeng (2019: 13): biofluorescence, biology; Kundrata et al. (2019: 1263): molecular phylogeny; Liu et al. (2019: 3183): mitogenomic phylogeny; Martin et al. (2019: 3): molecular phylogeny; Liu et al. (2020: 46): luciferase, phylogeny [also as Ochotyra]; Zhang et al. (2020: 5): molecular phylogeny, bioluminescence; Ge et al. (2021: 3): mitogenomic phylogeny; Lawrence et al. (2021: 456): wing morphology; Li et al. (2021b: 2): remark; Cai et al. (2022: 6): molecular phylogeny; Ge et al. (2022: 3): mitogenomic phylogeny; He et al. (2022: 4): mitogenomic phylogeny; Moreira et al. (2022: 7): luciferase, molecular phylogeny. In addition to the aforementioned literature, this genus was included in PhD theses by Ho (2002), Jeng (2008), and Roza (2022).

 Remarks.

Rhagophthalmus is the most speciose genus in the family. It contains 34 species from South, East, and Southeast Asia. This genus is characterized by having deeply emarginate eyes and relatively short antennae (Fig. 1E, F). Wittmer in Wittmer and Ohba (1994) synonymized Ochotyra with Rhagophthalmus, and we follow this concept until a proper revision of the genus is carried out. On the other hand, Menghuoius, which was synonymized with Rhagophthalmus by Li et al. (2008a), is considered here a separate genus (see Remarks under Menghuoius).

 Animalia Coleoptera Rhagophthalmidae 60B37031-B17A-5538-A4B3-F08388D01BAB Rhagophthalmus angulatus Wittmer, 1997 Rhagophthalmus angulatus Wittmer, 1997: 258. Type depository.

Holotype, male (NHMB). One paratype, male (NHMB).

 Type locality.

China: East Hubei, 30 km NE Macheng, 500 m.

 Distribution.

China (Hubei).

 Literature.

Wittmer (1997: 258): original description, figures of male antenna and genitalia; Bocak (2007: 225): catalogue; Li et al. (2008a: 265): distribution; Li et al. (2008b: 496): review.

 Animalia Coleoptera Rhagophthalmidae 48719CAC-C1A6-5F9C-A0BA-4301F6563C52 Rhagophthalmus beigansis Ho in Ho et al. 2012 Rhagophthalmus beigansis Ho in Ho et al. 2012: 4. Type depository.

Holotype, male (TARI). Eight paratypes: four males, four females (ESRI, NMNS).

 Type locality.

China/Taiwan, Lienchiang County, Beigan.

 Distribution.

China/Taiwan.

 Literature.

Ho et al. (2012: 4): original description, figures of male habitus, head, antenna and genitalia, and female habitus, head, and bioluminescence; Yiu (2017: 60): remark.

 Animalia Coleoptera Rhagophthalmidae 6CA53368-0FB8-5FBD-91A9-ACE2A1345C43 Rhagophthalmus brevipennis Fairmaire, 1896 Rhagophthalmus brevipennis Fairmaire, 1896: 227. Type depository.

Described based on an unknown number of specimens. Syntype, male (RMNH). Three additional male specimens (on one pin) from MNHN with labels different from the RMNH syntype are also labelled as “Type” but they probably represent a different species.

 Type locality.

India: Maharashtra, Nagpur.

 Distribution.

India (Maharashtra).

 Literature.

Fairmaire (1896: 227): original description; Olivier (1902: 87): catalogue; Lefroy (1909: 327): catalogue; Olivier (1910: 1); catalogue; Olivier (1912: 470): revision; McDermott (1966: 121): catalogue; Li et al. (2008a: 265): distribution; Li et al. (2008b: 496): review.

 Animalia Coleoptera Rhagophthalmidae 3B4B5D4F-BA3F-5A25-8E70-09E711316B86 Rhagophthalmus burmensis Wittmer in Wittmer and Ohba 1994 Rhagophthalmus burmensis Wittmer in Wittmer and Ohba 1994: 349. Type depository.

Holotype, male (NHMB). Seven paratypes, males (NHMB).

 Type locality.

Myanmar: Kambaiti.

 Distribution.

Myanmar.

 Literature.

Wittmer and Ohba (1994: 349): original description, drawings of male genitalia; Li et al. (2008a: 265): distribution; Li et al. (2008b: 496): review.

 Animalia Coleoptera Rhagophthalmidae EE81D048-065A-5CC8-B54E-EFD34D5A9279 Rhagophthalmus confusus Olivier, 1912 Rhagophthalmus confusus E. Olivier, 1912: 469, 471. Rhagophthalmus confuses : Wijekoon et al. 2016: 71 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Type depository.

Described based on an unknown number of specimens. One syntype, male (NHMUK). One probable syntype, male (MNHN).

 Type locality.

Sri Lanka.

 Distribution.

Sri Lanka.

 Literature.

Olivier (1912: 469, 471): original description; Pic (1916: 9): comparison with R. notaticollis; McDermott (1966: 121): catalogue; Li et al. (2008a: 265): distribution; Li et al. (2008b: 496): review; Ho et al. (2012: 1): remark; Wijekoon et al. (2016: 71): checklist [as R. confuses [sic!]]. In addition to the aforementioned literature, this species was included in a PhD thesis by Roza (2022).

 Animalia Coleoptera Rhagophthalmidae B6E42CE2-749C-5017-A5F4-C1A07C853EFB Rhagophthalmus elongatus Wittmer in Wittmer and Ohba 1994 Rhagophthalmus elongatus Wittmer in Wittmer and Ohba 1994: 348. Type depository.

Holotype, male (NHMB).

 Type locality.

China: Guangxi prov., Duyang Shan [“Mts. Toyen-chan”].

 Distribution.

China (Guangxi).

 Literature.

Wittmer and Ohba (1994: 348): original description, drawings of male genitalia; Kawashima and Satô (2001: 428, 430): comparison with R. flavus and R. minutus, respectively; Bocak (2007: 225): catalogue; Li et al. (2008a: 265): distribution; Li et al. (2008b: 496): review.

 Animalia Coleoptera Rhagophthalmidae 64E102B6-489E-5616-BCBB-33A18EB52D95 Rhagophthalmus filiformis Olivier, 1912 Rhagophthalmus filiformis E. Olivier, 1912: 469, 470. Type depository.

Described based on an unknown number of specimens. One syntype, male (NHMUK).

 Type locality.

Sri Lanka.

 Distribution.

Sri Lanka.

 Literature.

Olivier (1912: 469, 470): original description; Pic (1925a: 17): comparison with R. longipennis; McDermott (1966: 121): catalogue; Kawashima and Satô (2001: 429): comparison with R. minutus; Li et al. (2008a: 265): distribution; Li et al. (2008b: 496): review; Ho et al. (2012: 1): remark; Wijekoon et al. (2016: 74): checklist.

 Animalia Coleoptera Rhagophthalmidae 57D2C221-D994-58FA-8530-47DC3D5A0040 Rhagophthalmus flavus Kawashima & Satô, 2001 Rhagophthalmus flavus Kawashima & Satô, 2001: 424. Type depository.

Holotype, male (NWU). One paratype, male (PCIK).

 Type locality.

Myanmar: Dawna.

 Distribution.

Myanmar, Thailand.

 Literature.

Kawashima and Satô (2001: 424): original description, figures of male habitus, antenna, and genitalia; Li et al. (2008a: 265): distribution; Li et al. (2008b: 496): review; Ho et al. (2012: 9): comparison with R. giallolateralus.

 Animalia Coleoptera Rhagophthalmidae E1DAF0C3-B8A9-545D-9734-CCC1DF400A63 Rhagophthalmus formosanus Kawashima & Sugaya, 2003 Rhagophthalmus formosanus Kawashima & Sugaya, 2003: 354. Type depository.

Holotype, male (NMNS). Two paratypes, males (PCIK).

 Type locality.

China/Taiwan: Nantou Hsien, Meimu.

 Distribution.

China/Taiwan.

 Literature.

Kawashima and Sugaya (2003: 354): original description, identification key, figures of male habitus, antenna, and genitalia; Bocak (2007: 225): catalogue; Li et al. (2008a: 265): distribution; Li et al. (2008b: 496): review; Ho et al. (2012: 1): remark.

 Animalia Coleoptera Rhagophthalmidae F830C2A1-CE32-5E3C-A544-9BF41AF40AC0 Rhagophthalmus fugongensis Li & Liang in Li et al. 2008 Rhagophthalmus fugongensis Li & Liang in Li et al. 2008a: 260. Type depository.

Holotype, male, No. 0058739 (KNHMZ). 22 paratypes: eight males and 13 females (KNHMZ), one paratype, male (YCM). Although Li et al. (2008a) stated in the original description that the “holotype and most paratypes are deposited in KIZ [now KNHMZ]; one paratype (male) is deposited in YCM” (Li et al. 2008a: 260), Li et al. (2015) listed only six paratypes (sex not mentioned) from KNHMZ, under the collection numbers 0058740–0058745.

 Type locality.

China: Yunnan Province, Fugong County, Pihe, Wawa Village, 26.59398°N, 98.90819°E, 1263 m.

 Distribution.

China (Yunnan).

 Literature.

Li et al. (2008a: 260): original description; Li et al. (2008b: 496): review; Li et al. (2015: 269): catalogue.

 Animalia Coleoptera Rhagophthalmidae 739B19AD-4386-5174-B463-75C1BBC9433E Rhagophthalmus giallolateralus Ho in Ho et al. 2012 Rhagophthalmus giallolateralus Ho in Ho et al. 2012: 9. Type depository.

Holotype, male (TARI). Four paratypes: two males and two females (ESRI, NMNS).

 Type locality.

China/Taiwan, Lienchiang County, Dongjyu.

 Distribution.

China/Taiwan.

 Literature.

Ho et al. (2012: 9): original description, figures of male habitus, head, antenna, and genitalia, and female habitus, head, and luminous organ; Yiu (2017: 60): comparison with R. motschulskyi.

 Animalia Coleoptera Rhagophthalmidae 3D8462F6-A51A-563A-AA08-73BFF4A423B3 Rhagophthalmus gibbosulus Fairmaire, 1899 Rhagophthalmus gibbosulus Fairmaire, 1899: 624. Type depository.

Described based on an unknown number of specimens. No type specimen found in MNHN by RK.

 Type locality.

Probably China, “Koua-Toun” (Fujian).

 Distribution.

China (Fujian, ?Guangzhou, Shaanxi, Sichuan).

 Literature.

Fairmaire (1899: 624): original description; Olivier (1902: 88): catalogue; Olivier (1910: 1): catalogue; Jakobson (1911: 687): catalogue; Olivier (1912: 470): revision; Winkler (1925: 522): catalogue; Wu (1937: 385): catalogue; McDermott (1966: 121): catalogue; Hua (2002: 71): catalogue; Bocak (2007: 225): catalogue; Li et al. (2008a: 263): distribution, description and figures of male genitalia; Li et al. (2008b: 496): review.

 Animalia Coleoptera Rhagophthalmidae C9D29873-DD3F-5779-B4FE-36B996FB78DA Rhagophthalmus hiemalis Yiu, 2017 Rhagophthalmus hiemalis Yiu, 2017: 62. Type depository.

Holotype, male (TLES). 15 paratypes: 10 males, five females (TLES).

 Type locality.

China: Hong Kong, Tsuen Kam Au, 22.40728°N, 114.10357°E.

 Distribution.

China (Hong Kong).

 Literature.

Yiu (2017: 62): original description, figures of male and female habitus and male genitalia.

 Animalia Coleoptera Rhagophthalmidae E0787DC4-72C6-5E6B-A31E-490670EEC148 Rhagophthalmus jenniferae Kawashima & Satô, 2001 Rhagophthalmus jenniferae Kawashima & Satô, 2001: 430. Rhagophthalmus jeniferae : Bocak 2007: 225 [unavailable name, incorrect subsequent spelling, not in prevailing usage]. Type depository.

Holotype, male (NWU). Three paratypes, males (NTU, PCIK).

 Type locality.

China/Taiwan, Fenchihu, Chiai Hsien.

 Distribution.

China/Taiwan.

 Literature.

Kawashima and Satô (2001: 430): original description, figures of male habitus, antenna, and genitalia; Kawashima and Sugaya (2003: 353): remark, key; Bocak (2007: 225): catalogue [as R. jeniferae [sic!]]; Li et al. (2008a: 265): distribution; Li et al. (2008b: 496): review; Chen et al. (2010: 197): biology and bioluminescence, figures of male and female habitus, body parts, bioluminescence; Ho et al. (2012: 1): remark; Jeng (2019: 13): biofluorescence, biology, figures of larval and female habitus. In addition to the aforementioned literature, this species was mentioned in a PhD thesis by Ho (2002).

 Animalia Coleoptera Rhagophthalmidae 75465716-710E-56F9-A31A-F95732394388 Rhagophthalmus kiangsuensis Wittmer in Wittmer and Ohba 1994 Rhagophthalmus kiangsuensis Wittmer in Wittmer and Ohba 1994: 346. Rhagophthalmus kinagsuensis : Wittmer in Wittmer and Ohba 1994: 347 [unavailable name, incorrect original spelling (ICZN 1999, Art. 19.3); First Reviser (ICZN 1999, Art. 24.2): Bocak (2007: 225)]. Type depository.

Holotype, male (MNHN). One paratype, male (NHMB).

 Type locality.

China: Jiangsu province (without further data).

 Distribution.

China (Jiangsu).

 Literature.

Wittmer and Ohba (1994: 346): original description, drawings of male genitalia; Bocak (2007: 225): catalogue; Li et al. (2008a: 265): distribution; Li et al. (2008b: 496): review.

 Animalia Coleoptera Rhagophthalmidae B6A57FC6-DED1-50DC-971C-5CD1335BA355 Rhagophthalmus laosensis Pic, 1917 Rhagophthalmus laosensis Pic, 1917: 3. Type depository.

Described based on an unknown number of specimens. One syntype, male (MNHN).

 Type locality.

Laos: Thakhek [“Taket”].

 Distribution.

Laos.

 Literature.

Pic (1917: 3): original description; Pic (1923: 29): catalogue; McDermott (1966: 122): catalogue; Li et al. (2008a: 265): distribution; Li et al. (2008b: 496): review.

 Animalia Coleoptera Rhagophthalmidae 7D0F9A94-F2F7-5642-B205-D6A9FB9F3F66 Rhagophthalmus longipennis Pic, 1925 Rhagophthalmus longipennis Pic, 1925a: 17. Ochotyra longipennis : Bocak 2007: 225. Type depository.

Described based on an unknown number of specimens. Syntype, male (MNHN); syntype, male (NHMUK); four syntypes, males (NHMB).

Type localities. Only “Chine” in the original description (Pic 1925a). More detailed locality data available on the syntype labels: Sichuan, Kangding [“Tatsienlu”] (MNHN), Shaanxi, Qinling [“Kinling” or “Kinlung”] (NHMUK and NHMB, respectively).

 Distribution.

China (Shaanxi, Sichuan).

 Literature.

Pic (1925a: 17): original description; Wu (1937: 385): catalogue; McDermott (1966: 122): catalogue; Hua (2002: 71): catalogue; Bocak (2007: 225): catalogue [as Ochotyra longipennis]; Li et al. (2008a: 265): distribution; Li et al. (2008b: 496): review; Yiu (2017: 62): comparison with R. hiemalis.

 Remarks.

Some previous authors erroneously considered 1923 as the year of original description of this species (as “Pic 1923: 29”); however, it was described as a new species in 1925 (Pic 1925a).

 Animalia Coleoptera Rhagophthalmidae CD1D0C3C-D4BD-51CD-8413-A6BB35C2034C Rhagophthalmus lufengensis Li & Ohba in Li et al. 2008 Rhagophthalmus lufengensis Li, Ogoh, Ohba, Liang & Ohmiya, 2007: 196 [nomen nudum; published without description, unavailable name according to the ICZN (1999, Art. 13)]. Rhagophthalmus lufengensis Li & Ohba in Li et al. 2008a: 262. Rhagophthalmus lufegensis : Amaral et al. 2014: 415 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Type depository.

Holotype, male, No. 0058746 (KNHMZ). 11 paratypes: eight males and three females (KNHMZ, YCM). Li et al. (2015) listed three paratypes (two males, one female) from KNHMZ, under the collection numbers 0058747–0058749.

 Type locality.

China: Yunnan Province, Lufeng County, Dajiuzhuang, 25.09774°N, 101.80204°E, 1827 m.

 Distribution.

China (Yunnan).

 Literature.

Li et al. (2007: 196): nomen nudum, mitochondrial genome, phylogeny; Arnoldi et al. (2007: 2): remark; Li et al. (2008a: 262): original description, figures of male and female habitus, male antenna, and male genitalia; Li et al. (2008b: 496): review; Sheffield et al. (2008: 2500): mitochondrial genome; Timmermans and Vogler (2012: 300): molecular phylogeny; Amaral et al. (2014: 415): molecular phylogeny [as R. lufegensis [sic!]]; Li et al. (2015: 269): catalogue; Amaral et al. (2016: 254): molecular phylogeny; Amaral et al. (2017a: 673): mitogenome, phylogeny [as R. lufegensis [sic!]]; Wang et al. (2017: 6): molecular phylogeny; Chen et al. (2019: 8): molecular phylogeny; He et al. (2019: 566): molecular phylogeny; Zhang et al. (2020: 5): molecular phylogeny; He et al. (2022: 4): mitogenomic phylogeny.

 Animalia Coleoptera Rhagophthalmidae 16367371-C338-5095-BDF5-0622342B215D Rhagophthalmus minutus Kawashima & Satô, 2001 Rhagophthalmus minutus Kawashima & Satô, 2001: 428. Type depository.

Holotype, male (NWU). Three paratypes, males (two in NWU, one in PCIK).

 Type locality.

Thailand: Kohn Kaen Province, “near Ban Lon, Lam Chee Yai”.

 Distribution.

Thailand.

 Literature.

Kawashima and Satô (2001: 428): original description, figures of male habitus, antenna, and genitalia; Kawashima and Sugaya (2003: 358): comparison with R. formosanus; Li et al. (2008a: 265): distribution; Li et al. (2008b: 496): review.

 Animalia Coleoptera Rhagophthalmidae 3C041E30-B128-512F-ABAC-B12B5019F816 Rhagophthalmus motschulskyi Olivier, 1912 Rhagophthalmus motschulskyi E. Olivier, 1912: 469, 472. Type depository.

Described based on an unknown number of specimens. Syntype, male (NHMUK).

 Type locality.

China: Hong Kong.

 Distribution.

China (Hong Kong).

 Literature.

Olivier (1912: 469, 472): original description; Winkler (1925: 522): catalogue; McDermott (1966: 122): catalogue; Hua (2002: 71): catalogue; Bocak (2007: 225): catalogue; Li et al. (2008a: 265): distribution; Li et al. (2008b: 496): review; Yiu (2011a: 14): remark; Yiu (2011b: 20): biology and bioluminescence, figures of female bioluminescence; Ho et al. (2012: 1): remark; Yiu (2012: 30): catalogue, figures of habitus; Yiu (2013: 101): remark, bioluminescence; Yiu (2017: 60): redescription, figures of larva, pupa, and adults, and male genitalia.

 Animalia Coleoptera Rhagophthalmidae 3650E34A-00AD-5904-AF24-42C32B05E5F0 Rhagophthalmus neoobscurus Wittmer in Wittmer and Ohba 1994 Ochotyra obscura Pic, 1921b: 18. Rhagophthalmus neoobscurus Wittmer in Wittmer and Ohba 1994: 342 (replacement name for O. obscura Pic, 1921 (in Rhagophthalmus), not R. obscurus (Pic, 1917)). Type depository.

Described based on an unknown number of specimens. One syntype, male (MNHN).

 Type locality.

India (no further data). “Dekan India” written on the syntype label from MNHN.

 Distribution.

India (no further data).

 Literature.

Pic (1921b: 18): original description [as Ochotyra obscura]; McDermott (1966: 122): remark, catalogue [as Ochotyra obscura]; Wittmer and Ohba (1994: 342): taxonomy; Li et al. (2008a: 265): distribution; Li et al. (2008b: 496): review; Wijekoon et al. (2016: 74): checklist [as Ochotyra obscura].

 Remarks.

Wijekoon et al. (2016) erroneously cited “P. Melong” instead of “Pic” as the author of O. obscura.

 Animalia Coleoptera Rhagophthalmidae 3EC69DE1-B43E-5BC3-BA0E-2226726A0CDF Rhagophthalmus notaticollis Pic, 1916 Rhagophthalmus notaticollis Pic, 1916: 9. Rhagophthalmus notaticolis : Wijekoon et al. 2016: 74 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Type depository.

Described based on an unknown number of specimens. One syntype, male (MNHN).

 Type locality.

Sri Lanka.

 Distribution.

Sri Lanka.

 Literature.

Pic (1916: 9): original description; McDermott (1966: 122): catalogue; Li et al. (2008a: 265): distribution; Li et al. (2008b: 496): review; Ho et al. (2012: 1): remark; Wijekoon et al. (2016: 74): checklist [as R. notaticolis [sic!]].

 Remarks.

Wijekoon et al. (2016) erroneously cited “P. Melong” instead of “Pic” as the author of R. notaticollis.

 Animalia Coleoptera Rhagophthalmidae F1D5A4EC-167C-52AC-B73F-5FC4E7E50E0F Rhagophthalmus obscurus (Pic, 1917) Rhagophthalmus tonkineus var. obscurus Pic, 1917: 4. Rhagophthalmus tonkineus var. obscurus : Winkler 1925: 522 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Rhagophthalmus tonkinensis var. obscurus : McDermott 1966: 122 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Rhagophthalmus tokineus var. obscurus : Wittmer in Wittmer and Ohba 1994: 342 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Rhagophthalmus obscurus : Wittmer in Wittmer and Ohba 1994: 342. Type depository.

Described based on an unknown number of specimens. One syntype, male (MNHN).

 Type locality.

Vietnam: Lào Cai [Tonkin: Lao Kay].

 Distribution.

Vietnam.

 Literature.

Pic (1917: 4): original description [as R. tonkineus var. obscurus]; Pic (1923: 29): catalogue [as R. tonkineus var. obscurus]; Winkler (1925: 522): catalogue [as R. tonkineus var. obscursus [sic!]]; McDermott (1966: 122): catalogue [R. tonkinensis var. obscurus [sic!]]; Wittmer and Ohba (1994: 342): taxonomy, drawings of male genitalia [also as R. tokineus var. obscurus [sic!]]; Wittmer (1997: 258): remark; Li et al. (2008a: 265): distribution; Li et al. (2008b: 496): review.

 Animalia Coleoptera Rhagophthalmidae F05EF9B5-CBEB-55E4-BF0D-E56D05B9C269 Rhagophthalmus ohbai Wittmer in Wittmer and Ohba 1994 Rhagophthalmus ohbai Wittmer in Wittmer and Ohba 1994: 344. Rhagophthalmus ohba : Branham and Wenzel 2001: 567 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Type depository.

Holotype, male (YCM). Three paratypes, sex unknown (YCM), three paratypes, two males and one female (NHMB), two paratypes, sex unknown (NWU).

 Type locality.

Japan: Okinawa Prefecture, Yaeyama Islands, Iriomote Island, Sonai.

 Distribution.

Japan (Yaeyama Islands), Taiwan (Chen and Ho 1996, 1998; Ho et al. 2012).

 Literature.

Wittmer and Ohba (1994: 344): original description, drawings of male and female habitus, male antenna and genitalia; Ohba (1995: 13): remark, bioluminescence; Chen and Ho (1996: 46): distribution, figure of habitus; Ohba et al. (1996a: 1): morphology, biology, figures of habitus, body details, and bioluminescence; Ohba et al. (1996b: 30): remark; Nakane (1997: 36): remark; Ohba (1997a: 5): checklist, biology, figures of larval, male and female habitus, male head, and female bioluminescence; Ohba (1997b: 19): remark; Ohba (1997c: 51): breeding, development, immature stages, figures of habitus; Ohba et al. (1997: 25): remark; Suzuki (1997: 4): phylogeny, biology; Wittmer (1997: 259): comparison with R. angulatus; Chen and Ho (1998: 34): bioluminescence; Kawashima (1998: 16): female morphology, drawing of female habitus; Ohba (1998: 3): checklist, biology; Costa et al. (1999: 22): remark; Viviani et al. (1999: 8274): remark [as Ragophthalmus [sic!]]; Goto and Kawashima (2000: 143): distribution; Kawashima (2000: 131): remarks; Kim et al. (2000: 214): molecular phylogeny; Ohmiya et al. (2000: 32): luciferase; Viviani and Ohmiya (2000: 267): remark [as Ragophthalmus [sic!]]; Branham and Wenzel (2001: 567): phylogeny [also as R. ohba [sic!]]; Kawashima and Satô (2001: 432): comparison with R. jenniferae; Kobayashi et al. (2001: 1): development, eggs; Viviani et al. (2001: 1287): bioluminescence [as Ragophthalmus [sic!]]; Kobayashi et al. (2002: 1): embryogenesis, figures of female habitus and bioluminescence; Ugarova and Brovko (2002: 322): bioluminescence; Viviani (2002: 1836): remark [as Ragophthalmus [sic!]]; Viviani et al. (2002: 538): remark [as Ragophthalmus [sic!]]; Branham and Wenzel (2003: 5): phylogeny [also as R. ohba [sic!]]; Chen (2003: 52): morphology, bioluminescence, figures of adult males and females, larva, and bioluminescence; Hayashi and Suzuki (2003: 4): morphology, biology, figure of mating; Kawashima and Sugaya (2003: 353): remark, identification key; Kawashima et al. (2003: 255): catalogue; Kobayashi et al. (2003: 19): embryogenesis, development; Satô and Kawashima (2003: 9): remark; DeCock (2004: 341): remark; Ohba (2004a: 226): bioluminescence, biology, figures of male and female habitus; Ohba (2004b: 6): bioluminescence, biology, figures of male and female habitus; Lau and Meyer-Rochow (2006: 20): eye morphology, figures of male and female head and eye; Li et al. (2006: 818): molecular phylogeny; Arnoldi et al. (2007: 2): molecular phylogeny; Bocak (2007: 225): catalogue; Geisthardt and Satô (2007: 234): catalogue [in Lampyridaeincertae sedis]; Lau et al. (2007: 27): eye morphology of male; Li et al. (2007: 197): mitochondrial genome, phylogeny; Sagegami-Oba et al. (2007: 110): molecular phylogeny; Stanger-Hall et al. (2007: 38): molecular phylogeny [also as Rhagophtalmus [sic!]]; Li et al. (2008a: 259): comparison with R. lufengensis and M. giganteus, distribution; Li et al. (2008b: 496): review; Noguchi et al. (2008: 2): luciferase; Sheffield et al. (2008: 2500): mitochondrial genome; Suzuki and Kobayashi (2009: 30): embryogenesis, figure of egg; Chen et al. (2010: 203): habitus figure showing bioluminescence; Kawashima et al. (2010: 137): book chapter, figures of male and female habitus, and female ovipositor; Lawrence et al. (2011: 7): phylogeny, figure of female abdomen; Oba et al. (2011: 773): biology, bioluminescence, figures of male and female habitus, and female bioluminescence; Amaral et al. (2012: 1262): luciferase, phylogeny; Ho et al. (2012: 1): remarks, comparison with R. beigansis; Timmermans and Vogler (2012: 300): molecular phylogeny; Amaral et al. (2014: 415): molecular phylogeny; Hosoe et al. (2014: 331): chemical defence, figures of male and female habitus; Kundrata et al. (2014: 167): molecular phylogeny; Martin et al. (2015: 519): molecular phylogeny; Amaral et al. (2016: 254): molecular phylogeny; Bocak et al. (2016: suppl.): molecular phylogeny; Kundrata et al. (2016: 296): molecular phylogeny; Amaral et al. (2017a: 674): remark; Amaral et al. (2017b: 84): phylogeny; Amaral et al. (2017c: 157): phylogeny; Martin et al. (2017: 568): molecular phylogeny; Wang et al. (2017: 6): molecular phylogeny; Bocak et al. (2018: suppl.): molecular phylogeny; Fallon et al. (2018: 8): genomes, bioluminescence; Stanger-Hall et al. (2018: 8): remark; Chen et al. (2019: 8): molecular phylogeny; He et al. (2019: 566): molecular phylogeny; Kundrata et al. (2019: 1263): molecular phylogeny; Liu et al. (2019: 3183): mitogenomic phylogeny; Martin et al. (2019: 3): molecular phylogeny; Liu et al. (2020: 47): luciferase, phylogeny; Zhang et al. (2020: 5): molecular phylogeny, bioluminescence; Ge et al. (2021: 3): mitogenomic phylogeny; Ge et al. (2022: 3): mitogenomic phylogeny; He et al. (2022: 4): mitogenomic phylogeny; Moreira et al. (2022: 7): luciferase, molecular phylogeny. In addition to the aforementioned literature, this species was included in PhD theses by Ho (2002) and Jeng (2008).

 Animalia Coleoptera Rhagophthalmidae C4F415C2-EC04-5569-91FE-F82B3C55EBE0 Rhagophthalmus sausai Wittmer, 1997 Rhagophthalmus sausai Wittmer, 1997: 257. Type depository.

Holotype, male (NHMB). Two paratypes, males (NHMB).

 Type locality.

China: Guizhou, 60 km N Kaili, Shibing, Yuntai Shan.

 Distribution.

China (Guizhou).

 Literature.

Wittmer (1997: 257): original description, drawings of male antenna and genitalia; Bocak (2007: 225): catalogue; Li et al. (2008a: 265): distribution; Li et al. (2008b: 496): review.

 Animalia Coleoptera Rhagophthalmidae BE3064F2-27B5-5EC4-B5C0-DE4690921853 Rhagophthalmus scutellatus Motschulsky, 1854 Rhagophthalmus scutellatus Motschulsky, 1854: 45. Type depository.

Holotype, male (ZMM).

 Type locality.

China: Beijing.

 Distribution.

China (Beijing, Fujian, Jiangsu/Shanghai).

 Literature.

Motschulsky (1854: 45): original description; Motschulsky (1859: 59): remark, drawings of male habitus, lateral head, and leg; Gemminger (1869: 1655): catalogue; Olivier (1885: 372): comparison with R. sumatrensis; Heyden (1886: 286): remark; Fairmaire (1888: 25): comparison with R. giganteus; Fairmaire (1889: 353): comparison with R. tonkineus; Bourgeois (1892: 236): distributional note; Cardon (1892: 238): checklist; Fairmaire (1899: 624): comparison with R. gibbosulus; Olivier (1902: 88): catalogue; Olivier (1910: 1): catalogue; Jakobson (1911: 687): catalogue; Olivier (1912: 470): revision; Pic (1916: 9): comparison with R. notaticollis; Lucas (1920: 567): catalogue; Winkler (1925: 522): catalogue; Pic (1938: 15): checklist; McDermott (1964: 49): remark; McDermott (1966: 122): catalogue; Wittmer and Ohba (1994: 343): taxonomy, morphology, drawings of male genitalia; Wittmer (1997: 261): taxonomy, morphology, distribution, drawings of male genitalia; Kawashima and Satô (2001: 423): remark, comparison with R. jenniferae; Hua (2002: 71): catalogue; Kawashima et al. (2003: 255): remark, catalogue; Kawashima and Sugaya (2003: 358): remark, identification key; Bocak (2007: 225): catalogue; Li et al. (2008a: 264): comparison with R. gibbosulus, distribution; Li et al. (2008b: 494): review; Suzuki and Kobayashi (2009: 30): remark; Chen et al. (2010: 196): remark; Lawrence et al. (2011: 7): phylogeny; Yiu (2017: 60): remark, comparison with R. hiemalis; Lawrence et al. (2021: 456): wing morphology, figure of hind wing. In addition to the aforementioned literature, this species was included in a PhD thesis by Roza (2022).

 Animalia Coleoptera Rhagophthalmidae 701A1362-2971-5FAD-A325-598927A48F4A Rhagophthalmus semisulcatus Wittmer, 1997 Rhagophthalmus semisulcatus Wittmer, 1997: 259. Type depository.

Holotype, male (NHMB). Five paratypes, males (NHMB). According to the original description (Wittmer 1997), there are only five paratypes; however, there are five additional specimens with different labels designated as paratypes in NHMB.

 Type locality.

China: Yunnan: Yulong Shan, 27°10’N, 100°13’E, 3900 m.

 Distribution.

China (Yunnan).

 Literature.

Wittmer (1997: 259): original description, drawings of male antenna and genitalia; Bocak (2007: 225): catalogue; Li et al. (2008a: 264): distribution, biology, figure of female habitus; Li et al. (2008b: 496): review.

 Animalia Coleoptera Rhagophthalmidae 34B3ED28-B807-5E6D-A707-69560ADFB6AD Rhagophthalmus semiustus (Pascoe, 1862) Ochotyra semiusta Pascoe, 1862: 323. Rhagophthalmus (Ochrotyra [sic!]) semiusta [sic!]: Lefroy 1909: 327. Rhagophthalmus semiustus : Wittmer in Wittmer and Ohba 1994: 342. Type depository.

Holotype, male (NHMUK).

 Type locality.

India: “Malabar”.

 Distribution.

India (Karnataka, Kerala, Tamil Nadu) [“Malabar, Coromandel”], Sri Lanka.

 Literature.

Pascoe (1862: 323): original description, drawing of male habitus [as Ochotyra]; Gerstaecker (1863: 409): remark [as Ochotyra]; Gemminger (1869: 1647): catalogue [as Ochotyra]; Gorham (1890: 550): catalogue; Gorham (1895: 310): distributional note, morphology [as Ochotyra]; Bourgeois (1903: 479): distributional note [as Ochotiza [sic!]]; Gorham (1903: 330): distributional note [as Ochotyra]; Bourgeois (1905: 130): distributional note [as Ochotyra]; Lefroy (1909: 327): catalogue [as Rhagophthalmus (Ochrotyra [sic!]) semiusta [sic!]]; Olivier (1910: 1): catalogue [as Ochotyra]; Pic (1921b: 18): comparison with R. neoobscurus [as Ochotyra]; McDermott (1964: 50): revision [as Ochotyra]; McDermott (1966: 122): catalogue [as Ochotyra]; Wittmer and Ohba (1994: 342): taxonomic remark [as Ochotyra]; Bocak (2007: 225): catalogue [as Ochotyra]; Li et al. (2008a: 265): distribution [as R. semiusta [sic!]]; Li et al. (2008b: 496): review [as R. semiusta [sic!]]; Wijekoon et al. (2016: 71): checklist [as Ochotyra]. In addition to the aforementioned literature, this species was included in a PhD thesis by Jeng (2008).

 Animalia Coleoptera Rhagophthalmidae 34E64C06-789B-5251-8936-28D0C9C23DF5 Rhagophthalmus sulcatus Pic, 1925 Rhagophthalmus sulcatus Pic, 1925b: 72. Type depository.

Described based on an unknown number of specimens. No type material was found in MNHN by RK.

 Type locality.

India: West Bengal, Darjeeling.

 Distribution.

India (West Bengal).

 Literature.

Pic (1925b: 72): original description; McDermott (1966: 122): catalogue; Wittmer (1997: 261): comparison with other species; Bocak (2007: 225): catalogue; Li et al. (2008a: 265): distribution; Li et al. (2008b: 496): review.

 Remarks.

This species could be a synonym of R. sulcicollis Olivier, 1912 (see Wittmer 1997 for more information).

 Animalia Coleoptera Rhagophthalmidae 1DEE83C5-F591-5149-84A0-7E28506B61A7 Rhagophthalmus sulcicollis sulcicollis Olivier, 1912 Rhagophthalmus sulcicollis E. Olivier, 1912: 471. Type depository.

Lectotype, male (NHMUK). Five paralectotypes, males (NHMUK) (although only four paralectotypes were reported by Wittmer 1997: 261). There are also two additional specimens in MNHN bearing the labels “lectotype” and “paralectotype”; however, they have locality label data that differ slightly from the original description.

 Type locality.

China: Tibet/Xizang, Yalong, over 3000 m.

 Distribution.

China (Tibet/Xizang).

 Literature.

Olivier (1912: 471): original description; Winkler (1925: 522): catalogue; Pic (1925b: 72): comparison with R. sulcatus; McDermott (1966: 122): catalogue; Wittmer (1997: 261): taxonomy, morphology, drawings of male pronotum and genitalia; Hua (2002: 71): catalogue; Bocak (2007: 225): catalogue; Li et al. (2008a: 265): distribution; Li et al. (2008b: 496): review.

 Animalia Coleoptera Rhagophthalmidae 1599ADFD-D53F-5AB3-9D75-ECBEA9B53E0B Rhagophthalmus sulcicollis bhutanensis Wittmer, 1997 Rhagophthalmus sulcicollis subsp. bhutanensis Wittmer, 1997: 261. Type depository.

Holotype, male (NHMB).

 Type locality.

Bhutan: Karrumphe, 2700 m.

 Distribution.

Bhutan.

 Literature.

Wittmer (1997: 261): original description, drawings of male antenna and pronotum; Bocak (2007: 225): catalogue.

 Animalia Coleoptera Rhagophthalmidae 810E5C7A-7E2A-5BB9-9AFD-DA592473158C Rhagophthalmus sumatrensis Olivier, 1885 Rhagophthalmus sumatrensis Olivier, 1885: 372. Type depository.

Described based on an unknown number of specimens. Three syntypes, males (MSNG).

 Type locality.

Indonesia: Sumatra, Mt. Singalang.

 Distribution.

Indonesia (Sumatra).

 Literature.

Olivier (1885: 372): original description; Fairmaire (1889: 353): comparison with R. tonkineus; Olivier (1902: 88): catalogue; Olivier (1910: 1): catalogue; Olivier (1912: 470): revision, drawings of head, antenna, and tarsus; McDermott (1966: 122): catalogue; Wittmer (1997: 259): comparison with R. angulatus; Li et al. (2008a: 265): distribution; Li et al. (2008b: 496): review; Ho et al. (2012: 1): remark.

 Animalia Coleoptera Rhagophthalmidae 1F8E5DC5-719B-5300-8795-8276BC0C8D33 Rhagophthalmus tienmushanensis Wittmer in Wittmer and Ohba 1994 Rhagophthalmus tienmushanensis Wittmer in Wittmer and Ohba 1994: 346. Type depository.

Holotype, male (NHMB).

 Type locality.

China: Zhejiang, Tianmushan.

 Distribution.

China (Zhejiang, Shanghai).

 Literature.

Wittmer and Ohba (1994: 346): original description, drawings of male genitalia; Bocak (2007: 225): catalogue; Li et al. (2008a: 265): distribution; Li et al. (2008b: 496): review.

 Animalia Coleoptera Rhagophthalmidae B676B7AE-85C4-5DD8-A4FD-CD12B3CB5A45 Rhagophthalmus tonkineus Fairmaire, 1889 Rhagophthalmus tonkineus Fairmaire, 1889: 352. Rhagophthalmus tonkinensis : McDermott 1966: 122 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Rhagophthalmus tokineus : Wittmer and Ohba 1994: 342 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Type depository.

Described based on an unknown number of specimens. No type material was found in MNHN (Wittmer and Ohba 1994; RK, pers. obs.).

 Type locality.

Vietnam [“Tonkin”].

 Distribution.

Vietnam, China (Guangxi) (Li et al. 2008a); Laos (Pic 1923).

 Literature.

Fairmaire (1889: 352): original description; Fairmaire (1896: 228): comparison with R. brevipennis; Olivier (1902: 88): catalogue; Olivier (1910: 1): catalogue; Olivier (1912: 470): revision; Pic (1917: 4): comparison with R. obscurus; Pic (1923: 29): catalogue, distributional note; Winkler (1925: 522): catalogue; McDermott (1966: 122): catalogue [as R. tonkinensis [sic!]]; Wittmer and Ohba (1994: 342): remark, taxonomy [as R. tokineus [sic!]]; Li et al. (2008a: 265): distribution [also as R. tonkinensis [sic!]]; Li et al. (2008b: 496): review [as R. tonkinensis [sic!]].

 Animalia Coleoptera Rhagophthalmidae 272966E5-7C16-5F39-B200-67379585F633 Rhagophthalmus xanthogonus Olivier, 1912 Rhagophthalmus xanthogonus Olivier, 1912: 469, 471. Rhagophthalmus xanthogenus : McDermott 1966: 122 [unavailable name, incorrect subsequent spelling not in prevailing usage]. Type depository.

Described based on an unknown number of male specimens. No type material was found in MNHN by RK.

 Type locality.

China (no further data).

 Distribution.

China (no further data).

 Literature.

Olivier (1912: 469, 471): original description; Pic (1917: 4): comparison with R. laosensis; Winkler (1925: 522): catalogue; McDermott (1966: 122): catalogue [as R. xanthogenus [sic!]]; Hua (2002: 71): catalogue [as R. xanthogenus [sic!]]; Bocak (2007: 225): catalogue [as R. xanthogenus [sic!]]; Li et al. (2008a: 265): distribution [as R. xanthogenus [sic!]]; Li et al. (2008b: 496): review [as R. xanthogenus [sic!]].

Taxa removed from <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Rhagophthalmidae</tp:taxon-name-part></tp:taxon-name> Animalia Coleoptera Rhagophthalmidae B63B8803-6966-5B25-A249-C4110F4B245C Cydistus Bourgeois, 1885 [Phengodidae: Cydistinae] Cydistus Bourgeois, 1885: 272. Type species. Cydistus reitteri Bourgeois, 1885; by monotypy. Composition and distribution.

Six described species from Asia Minor, the Levant, Iraq, and Iran: Cydistus chindaaricus Bolívar y Pieltain, 1913, C. escalerai Bolívar y Pieltain, 1913, C. nigripennis Wittmer, 1979, C. persicus Bolívar y Pieltain, 1913, C. reitteri Bourgeois, 1885, and C. zurcheri Bourgeois, 1908 (Kundrata et al. 2019).

 Remarks.

Cydistus was originally placed in Drilidae (Olivier 1910; Wittmer 1944). Later, Crowson (1955) hypothesized Cydistus might be an intermediate form between Karumiidae (currently a subfamily in Dascillidae) and Phengodidae. Although Paulus (1972) erected Cydistinae within Karumiidae for Cydistus, Crowson (1972) transferred this genus into the widely delimited Phengodidae, which also included Rhagophthalmidae. Lawrence and Newton (1995) and Bocak (2007) classified Cydistus in Phengodidae: Rhagophthalminae. Lawrence et al. (2010a, b) and Lawrence (2016) considered Cydistinae in Elateriformiaincertae sedis. Finally, Kundrata et al. (2019) were the first to include Cydistinae in a molecular phylogenetic analysis, and found them sister to the New World Phengodidae, which are only distantly related to Rhagophthalmidae. This placement was confirmed by a morphology-based analysis by Roza (2022).

 Animalia Coleoptera Rhagophthalmidae 11B2DB26-6DFF-502A-9ACE-3A0E5129FF50 Luciola antipodum Bourgeois, 1884 [Lampyridae: Luciolinae] Luciola antipodum Bourgeois, 1884: 285. Rhagophthalmus antipodum : Olivier 1902: 87; Fauvel, 1904: 140. Bourgeoisia antipodum : Olivier 1908: 17. Distribution.

New Caledonia, Solomon Islands.

 Remarks.

This firefly species was originally described in Luciola Laporte, 1833 (Bourgeois 1884) and later transferred to Rhagophthalmus by Olivier (1902). The same author later placed this species in his new genus Bourgeoisia Olivier, 1908, and McDermott (1966) subsequently designated it the type species of this genus. Bourgeoisia is currently considered a synonym of Luciola (e.g., Ballantyne and Lambkin 2013; Ballantyne et al. 2019). For more information on L. antipodum see e.g., McDermott (1966); Ballantyne (1968); Ballantyne and Lambkin (2013); and Ballantyne et al. (2019).

 Animalia Coleoptera Rhagophthalmidae 7AACD42B-C4E0-5E18-A955-6F482FE6F6C5 Reductodrilus Pic, 1943 [Lampyridae: Ototretinae] Reductodrilus Pic, 1943: 9. Type species. Reductodrilus nigroapicalis Pic, 1943; by monotypy. Composition and distribution.

Only a single species, R. nigroapicalis Pic, 1943 from northern Borneo (Malaysia: Sabah). Reductodrilus nigroapicalis var. latetestaceus Pic, 1943 should have a subspecific status according to Article 45.6.4. of the Code (ICZN 1999).

 Remarks.

Reductodrilus was initially placed in Drilidae (Pic 1943; Wittmer 1948). After most Drilidae genera were transferred to different families (e.g., Lampyridae, Lycidae, Omethidae, and Rhagophthalmidae) by Crowson (1972), Reductodrilus remained in an uncertain position. Kundrata and Bocak (2011a) listed it in Rhagophthalmidae in their revision of Pseudothilmanus. Probable syntypes of both subspecies of R. nigroapicalis are deposited in MNHN. Here, we tentatively transfer Reductodrilus to Lampyridae: Ototretinae based on its suboval and somewhat flattened body, antennae with 11 antennomeres which clearly extend beyond the posterior pronotal margin, head partially covered by pronotum, eyes clearly separated by frons, pronotum transverse, medially elevated, with anterior angles inconspicuous, rounded, and posterior angles projected posteriad (for more details see Janisova and Bocakova 2013). A detailed revision of this genus should improve our understanding of its systematic position.

 Discussion

Although Rhagophthalmidae have been known to entomologists for more than a century, their taxonomy and classification are still poorly known. The number of genera included in Rhagophthalmidae and also their placement within Elateroidea classification vary by source (e.g., McDermott 1966; Crowson 1972; Lawrence and Newton 1995; Kawashima et al. 2010; Kundrata and Bocak 2011a). In the last decade, Elateroidea systematic research has accelerated and the classification of the superfamily has experienced many taxonomic changes (e.g., Kundrata et al. 2014, 2019; Bocak et al. 2018; Kusy et al. 2018b, 2021), including the discoveries of two new recent families (Bocak et al. 2016; Rosa et al. 2020) and one new extinct family (Li et al. 2021b). However, only six new species of Rhagophthalmidae were described in three taxonomic papers in the same period (Ho et al. 2012; Kazantsev 2012; Yiu 2017). This is especially striking when compared to the most closely related family of Rhagophthalmidae (i.e., Phengodidae), where numerous taxonomic studies were published (e.g., Constantin 2014, 2016; Zaragoza-Caballero and Hernández 2014; Roza et al. 2017, 2018; Roza and Mermudes 2019, 2020; Vega-Badillo and Zaragoza-Caballero 2019; Vega-Badillo et al. 2020, 2021a, b), including not only descriptions of several new genera and species but also phylogenetic analyses of the group (Zaragoza-Caballero and Zurita-García 2015; Quintino 2017; Kundrata et al. 2019; Roza 2022). In Rhagophthalmidae, the most important research topics include taxonomic limits, phylogenetic relationships, accurate dating of the origin of the group, the evolution of bioluminescence and paedomorphosis, systematics of all genera (including revisions of already known species as well as descriptions of new taxa), descriptions of paedomorphic females and immature stages for all genera and species, and evaluating the distribution of the group at both generic and family levels.

Phylogenetic relationships, origin, and monophyly of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Rhagophthalmidae</tp:taxon-name-part></tp:taxon-name>

The phylogenetic placement of Rhagophthalmidae within Elateroidea has been controversial based on morphology only (Crowson 1972; Lawrence 1988; Branham and Wenzel 2001; Lawrence et al. 2011), and Rhagophthalmidae were often placed either in or close to Lampyridae or Phengodidae. Molecular phylogenetic analyses using various datasets and analytical approaches repeatedly confirmed that Rhagophthalmidae are sister to Phengodidae (Bocakova et al. 2007; Kundrata and Bocak 2011b; Kundrata et al. 2014; Zhang et al. 2018; Douglas et al. 2021; Kusy et al. 2021; Cai et al. 2022). Both families share soft-bodied males with large eyes, often bipectinate antennae with 12 antennomeres, leathery elytra which are usually shortened and narrowed, larviform females, and larvae that possess bioluminescent organs and feed on millipedes (Kawashima et al. 2010; Zaragoza-Caballero and Hernández 2014; Kundrata et al. 2019). Kusy et al. (2021) defined the “lampyroid clade”, which contains Lampyridae, Phengodidae, Rhagophthalmidae, and Sinopyrophoridae. Fossil Cretophengodidae were probably also a part of that clade (Li et al. 2021b).

The date of the origin of Rhagophthalmidae is unclear, as there are no known fossils of the group. Generally, soft-bodied elateroids are rarely found as fossils, and to date, the most informative fossils are inclusions in various ambers. Cretophengodidae were described from mid-Cretaceous amber of northern Myanmar (ca. 99 Mya, Shi et al. 2012; Li et al. 2021b), and Kusy et al. (2021) reported unpublished Phengodidae from the same deposit. Kusy et al. (2021) summarized and reviewed the published molecular dating analyses of the elaterid-lampyroid clade, and showed that median estimates suggest the split of the Lampyridae, Phengodidae, and Rhagophthalmidae clade in the mid-Cretaceous. However, an earlier date is also possible (Kusy et al. 2021).

Another important issue is the monophyly of Rhagophthalmidae. The group was originally proposed only for Dioptoma, Ochotyra, and Rhagophthalmus (Olivier 1907, 1910), and later Pic (1937) added Mimoochotyra. This concept was adopted by McDermott (1964, 1966). Crowson (1972) transferred some Asian genera (Cydistus, Diplocladon, Falsophrixothrix) from Drilidae to Phengodidae, and these were later added to Rhagophthalmidae together with Dodecatoma (Lawrence and Newton 1995). Cydistus was later transferred to Phengodidae (Kundrata et al. 2019). The current concept of Rhagophthalmidae consists of males which have exactly 12 antennomeres, with antennomere III being longer than antennomere II, a telescopic abdomen that is usually narrowed apically, and females which are more or less larva-like. However, the monophyly of this group as currently defined has never been rigorously tested.

Several genera were included in molecular phylogenetic analyses, including Bicladodrilus, Falsophrixothrix, Mimoochotyra, and Rhagophthalmus (incl. Ochotyra) (Bocakova et al. 2007; Kundrata et al. 2014, 2019). These genera always formed a monophylum. However, it should be noted that at least the generic placements of specimens identified as Bicladodrilus sp. from China and Mimoochotyra sp. from Malaysia are dubious. As Bicladodrilus, Bicladum, and Diplocladon are similar in general appearance and possess biflabellate antennae, this generic complex is in need of revision. While there are no described Bicladodrilus species in China, a species of Diplocladon was recently described from Hong Kong (Yiu 2017). The single described species of Mimoochotyra is known from Java (Pic 1937; Wittmer 1944).

In his unpublished PhD thesis, Jeng (2008) focused on systematics and paedomorphosis (neoteny) in Lampyridae. He also included representatives of the rhagophthalmid genera Dioptoma, Diplocladon, Dodecatoma, Falsophrixothrix, Menghuoius, Monodrilus, and Rhagophthalmus (incl. Ochotyra) in his morphology-based analyses. These genera were monophyletic and sister to Phengodidae. In another unpublished PhD thesis, Roza (2022) focused on phylogenetic relationships of Phengodidae, and included Bicladodrilus, Dioptoma, Diplocladon, Dodecatoma, Falsophrixothrix, Pseudothilmanus, and Rhagophthalmus in his morphology-based analyses. These genera formed a monophylum in all analyses performed. A phylogenomic analysis including representatives (ideally type species) of all rhagophthalmid genera would be a valuable assessment of the monophyly of the group.

 Bioluminescence and paedomorphosis in <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Rhagophthalmidae</tp:taxon-name-part></tp:taxon-name>

Within Coleoptera, bioluminescence can be found almost exclusively within the so-called “elaterid-lampyroid clade”, including Elateridae, Lampyridae, Phengodidae, Rhagophthalmidae, and Sinopyrophoridae, and probably the extinct Cretophengodidae (Oba et al. 2011; Fallon et al. 2018; Bi et al. 2019; Li et al. 2021b; Kusy et al. 2021; Powell et al. 2022). In Phengodidae, all known larvae and females are bioluminescent, as are males of some species (Costa and Zaragoza-Caballero 2010). Bioluminescence is hypothesized for lineages in which larvae and females are unknown (e.g., Cydistinae; Kundrata et al. 2019). All known larvae and females of Rhagophthalmidae are bioluminescent. Both larvae and females were reported for Diplocladon (in fact, it is probably Haplocladon; see Remarks under both genera), Menghuoius, and Rhagophthalmus, whereas only females are known for Dioptoma (Green 1913; Gahan 1925; Coblentz and Hughes 1926; Ridley 1934; Haneda 1950; Raj 1957; Lloyd 1971, 1978; Wittmer and Ohba 1994; Ohba et al. 1996a; Kawashima 2000; Jeng 2008; Li and Liang 2008; Li et al. 2008a; Kawashima et al. 2010). Males of at least some genera (e.g., Dioptoma and Rhagophthalmus) also emit light (Kawashima et al. 2010). At least in some cases, however, there are doubts about the correct genus identification of larvae or females. For example, Jeng (2008) suggested that the giant larviform female from Yunnan, China, identified as Diplocladon by Li and Liang (2008) is “likely of Menghuoius giganteus or the other related species described from there” (Jeng 2008: 135). The correct identification of larviform females is, however, very important for conclusions on the evolution of morphological modifications caused by paedomorphosis (e.g., Jeng 2008; Kawashima et al. 2010). This should be possible by e.g., rearing both sexes of the same species from larvae, finding a mating couple, or by the use of DNA barcoding. Information on the life-history and biology of most genera of Rhagophthalmidae is minimal or entirely absent. Further research should be conducted to confirm the presence of bioluminescence in the remaining rhagophthalmid genera.

Elateroid beetles are well-known not only for bioluminescence but also for morphological modifications caused by paedomorphosis (Crowson 1972; Cicero 1988; Bocak et al. 2008; Ferreira and Ivie 2022). In Elateroidea, mainly adult females are more or less modified, with a gradual series of morphological modifications across many families (Elateridae, Jurasaidae, Lampyridae, Lycidae, etc.), ranging from taxa that possess only a slightly softer body cuticle and a more relaxed abdomen through a number of intermediate stages, with variously reduced mouthparts, antennae, elytra, hind wings, and/or parts of the thorax, and a higher number of free abdominal ventrites, to taxa which are highly larviform (Bocakova et al. 2007; Bocak et al. 2008; Ferreira et al. 2019, 2020, 2022; Kundrata and Bocak 2019; Rosa et al. 2020; Ferreira and Ivie 2022). In both Phengodidae and Rhagophthalmidae, all known females are highly paedomorphic, being wingless and larva-like (Costa and Zaragoza-Caballero 2010; Kawashima et al. 2010). In Rhagophthalmidae, females are known only for Dioptoma, Diplocladon (Haplocladon?), Menghuoius, and Rhagophthalmus (Haneda 1950; Harvey 1952; Ohba 1997c; Kawashima 1998; Chen 2003; Jeng 2008; Li and Liang 2008; Li et al. 2008a); however, only those of Diplocladon (Haplocladon?) and Rhagophthalmus are described in detail (Kawashima et al. 2010). Interestingly, females of both genera exhibit different degrees of paedomorphic modifications, with Diplocladon (Haplocladon?) being completely larviform (including having stemmata, antennae with three antennomeres, tibiotarsus with a single pretarsal claw, and missing ovipositor) and Rhagophthalmus being incompletely larviform (having compound eyes, antennae with six or seven antennomeres, tarsi with five tarsomeres and two pretarsal claws, and ovipositor; Kawashima et al. 2010). Similar cases of different levels of morphological modifications in females of different genera were also reported for e.g., Elateridae (Drilus Olivier, 1790 being more paedomorphic than Omalisus Geoffroy, 1762 or Cebrio Olivier, 1790; Kundrata and Bocak 2019), Jurasaidae (JurasaiRosa et al., 2020 being more paedomorphic than TujamitaRosa et al., 2020; Rosa et al. 2020), and Lampyridae (Lamprigera Motschulsky, 1853 or Stenocladius Fairmaire, 1878 being more paedomorphic than Lampyris Geoffroy, 1762 or Lamprohiza Motschulsky, 1853; Ohba et al. 1997; Dong et al. 2021). It would be, therefore, very interesting to discover and describe in detail the females of all remaining rhagophthalmid genera.

 Generic classification and systematics of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Rhagophthalmidae</tp:taxon-name-part></tp:taxon-name>

It is clear from the above text that the classification and systematics of Rhagophthalmidae is in a very poor state of knowledge. Species of Bicladodrilus, Bicladum, Falsophrixothrix, Mimoochotyra, and Monodrilus have not been taxonomically treated since their descriptions, and their names have usually appeared only in catalogues, if at all. Taxonomic revisions are urgently needed for all genera currently included in Rhagophthalmidae with the exception of Pseudothilmanus, which was revised recently (Kundrata and Bocak 2011a). Although the most species-rich genus Rhagophthalmus received some taxonomic attention in the last decades (e.g., Wittmer and Ohba 1994; Wittmer 1997; Kawashima and Satô 2001; Kawashima and Sugaya 2003; Li et al. 2008a; Ho et al. 2012; Yiu 2017), a comprehensive revision is still needed.

Due to a scarcity of information on the morphology of most rhagophthalmid taxa, an identification key which would help taxonomists to recognize genera and species in collections and subsequently enhance knowledge on their diversity, variability, and distributions, is also missing. Most importantly, it is necessary to delimit generic boundaries in some problematic generic complexes. For example, Bicladodrilus, Bicladum, and Diplocladon share biflabellate antennae and relatively long elytra, and are not clearly distinguished from one other. Detailed taxonomic studies should also be conducted to revise the status of Ochotyra (currently a synonym of Rhagophthalmus) and Menghuoius (currently a separate genus but treated by some authors as a synonym of Rhagophthalmus). Some genera contain species which are probably not congeneric with their type species (e.g., some Dodecatoma spp. resemble Pseudothilmanus more than D. bicolor), and e.g., Dodecatoma testaceiceps should be removed from Rhagophthalmidae after a detailed revision. Taxonomic attention should be given not only to currently described taxa, but also to numerous undescribed Rhagophthalmidae mainly from Southeast Asia, which are housed in various institutional and personal collections (RK pers. obs.).

Taxonomic revisions are usually hampered by missing, lost, or otherwise unavailable type specimens, especially in long-neglected groups, such as Rhagophthalmidae. However, the vast majority of name-bearing rhagophthalmid type specimens are available in European and Asian museum collections. To date, we have been unable to locate name-bearing type specimens of only five species described by either Pic, Fairmaire, or Olivier, four of which belong to Rhagophthalmus. Name-bearing type specimens of species in 10 smaller genera are each deposited in one to three museum collections only; however, those of Dodecatoma and Rhagophthalmus are in seven and 12 institutions, respectively.

 Distribution of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Rhagophthalmidae</tp:taxon-name-part></tp:taxon-name>

Rhagophthalmidae are distributed in the Oriental realm and the Palaearctic bioregion of East Asia, in the area from Afghanistan and Pakistan, through the Himalayas, Indian Peninsula, Sri Lanka, China, and mainland Southeast Asia, to Sumatra, Java, Bali, Borneo, and the Philippines. The center of genus-level diversity of Rhagophthalmidae lies in Southeast Asia. Nine out of 12 genera have at least one species distributed in Southeast Asia, with five genera (i.e., Bicladodrilus, Bicladum, Falsophrixothrix, Mimoochotyra, and Monodrilus) being endemic to the region. However, this only accounts for approximately one third of described species. The genera Dodecatoma and Pseudothilmanus are known only from the Himalayas and surrounding regions (one species and one subspecies of Dodecatoma from Southeast Asia should be removed from that genus), and Dioptoma is endemic to the Indian Peninsula and Sri Lanka. Regarding the most species-rich genus Rhagophthalmus, only seven out of 34 species are known from Southeast Asia, including only a single species from the Greater Sunda Islands. Another seven species are known from the Indian Peninsula and Sri Lanka, and the remaining majority of species are distributed in mainland China and among the islands of East Asia.

Interestingly, in the eastern part of their distribution, Rhagophthalmidae have remained within the boundaries of the Sunda Shelf and the Philippines, i.e, west of the originally proposed Wallace Line, which was demarcated to separate Indo-Malayan (Oriental) and Austro-Malayan (Australasian) realms (Wallace 1863; Voris 2000; Lohman et al. 2011). The Sunda Shelf is a southward expansion of the continental shelf of Southeast Asia that was intermittently exposed by lowered sea levels during the Pleistocene (Hall 1998; Voris 2000; Lomolino et al. 2017). The Wallace line separates Bali and Borneo on the west from Lombok and Sulawesi on the east. It is a strong dispersal barrier to many terrestrial animals because of deep oceanic trenches (Lomolino et al. 2017). Rhagophthalmidae have a limited dispersal propensity due to their highly modified larviform females and, therefore, it is not surprising that they remained within the boundaries of the continental shelf of Southeast Asia, with a single described species from the Philippines. Additionally, it should be noted that there are no Rhagophthalmidae from east of the Wallace Line among the extensive material of non-type specimens (including numerous new species) that reside in major European museums and several personal collections, which the first author examined for a planned generic revision of the group. A single, unreported rhagophthalmid species from Bali is the closest that the family has been observed to the Wallace Line. A taxonomic revision of Rhagophthalmidae will further improve our knowledge of the distributions of individual genera and their species, some of which are currently known only from a single specimen.

 Conclusions

Here we provide the first comprehensive catalogue of the currently defined Rhagophthalmidae. The only catalogues of the group were those of Olivier (1910) and McDermott (1966; Rhagophthalminae as a subfamily of Lampyridae) but they contained only three genera and nine species, and four genera and 21 species, respectively. Here, we recognize 12 genera and 66 species. However, all genera but Pseudothilmanus urgently need taxonomic revisions, and numerous new species await formal descriptions. The phylogenetic position of Rhagophthalmidae as a sister group to Phengodidae is now generally accepted; however, interrelationships within the group and generic classification remain poorly known. Although morphology-based analyses in two PhD studies that were focused on related families (Jeng 2008; Roza 2022) confirm the monophyly of currently circumscribed Rhagophthalmidae, molecular analysis including representatives of all genera would be desirable. Additionally, little information is known of the biology of the group. Although various studies have been published on the ontogeny, biology, and behaviour of the most common genus Rhagophthalmus, there is virtually nothing known about the majority of other genera. Because known females in Rhagophthalmidae are highly morphologically modified and remain larviform as adults, they are interesting subjects for researching the evolution of paedomorphosis in Elateroidea. However, only a few have been studied in detail (Kawashima et al. 2010). Additionally, all known larvae and females (and some males) are bioluminescent, and therefore are an important source of information for research on the evolution of bioluminescence in beetles. However, this phenomenon is also understudied in Rhagophthalmidae, as larvae and females are unknown for most genera. Discoveries, field observations, morphological studies, and correct genus and species identifications of larvae and larviform females of Rhagophthalmidae are therefore crucial not only for our improved knowledge of the diversity, systematics, and morphology of the group, but also for a better understanding of the evolution of paedomorphosis and bioluminescence in Elateroidea and beetles in general.

 Acknowledgements

We are very grateful to the following colleagues for the loan of material and/or photographs of specimens and/or information on type specimens deposited in their collections: I. Zürcher-Pfander (NHMB), M. Geiser, M. V. L. Barclay (both NHMUK), A. Mantilleri, O. Montreuil, A. Taghavian (all MNHN), M. Hartmann (NKME), I. Plonski, M. Jäch, H. Schillhammer (all NHMW, Vienna, Austria), R. Poggi (MSNG), J. Hájek (NMPC, Prague, Czech Republic), H. Huijbregts (RMNH), B. Brugge (ZMAN, now in RMNH), J. Bergsten (SMNH), and A. Spooner (OUMNH). We thank P. Bouchard (CNCI, Ottawa, Canada) and N. Evenhuis (BM, Honolulu, USA) for help with ICZN issues and for dating problematic literature. Furthermore, we thank M. A. Alonso-Zarazaga (Spain) for his invaluable help identifying the correct gender of some genus-group names. Special thanks from RK go to M. Geiser (NHMUK) for numerous helpful comments (not only) on Rhagophthalmidae, and to A. Roza (Brazil) for discussions on Phengodidae and Rhagophthalmidae. OK would like to thank the Interlibrary loan section at the University of Florida for their help with literature, and the Florida Department of Agriculture and Consumer Services, Division of Plant Industries (FDACS-DPI), for support of this work. RK would like to thank M. París, M. A. Alonso Zarazaga, P. Alvárez Fidalgo, J. L. Zapata de la Vega, J. Cabarga, M. García París, M. Sánchez Ruiz, and I. Rey Fraile for their hospitality, kindness and help during his stay in MNCN, Madrid, and M. Geiser, M. Barclay, K. Matsumoto, D. Telnov, J. Wilkinson, and L. Stevens for their hospitality, kindness and help during his stay in NHMUK, London. We thank E. C. Powell and P. E. Skelley (both FDACS-DPI) for pre-submission reviews, M. A. Alonso-Zarazaga and A. Roza for reviews which led to improvement of this study, and V. S. Ferreira (Editor) for handling the manuscript. This study was funded by an internal grant of the Faculty of Science, UP Olomouc (IGA_PrF_2022_024; to RK, JH, and GP). RK received support from the SYNTHESYS+ project which is financed by European Community Research Infrastructure Action under the H2020 Integrating Activities Programme, Project number 823827 (projects ES-TAF TA1-2536 and GB-TAF-8656 for stays in MNCN, Madrid and NHMUK, London, respectively).

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