Rhagophthalmidae Olivier, 1907 (Coleoptera, Elateroidea): described genera and species, current problems, and prospects for the bioluminescent and paedomorphic beetle lineage

Robin Kundrata; Johana Hoffmannova; Kevin R. Hinson; Oliver Keller; Gabriela Packova

doi:10.3897/zookeys.1126.90233

Research Article

Rhagophthalmidae Olivier, 1907 (Coleoptera, Elateroidea): described genera and species, current problems, and prospects for the bioluminescent and paedomorphic beetle lineage

Robin Kundrata^‡, Johana Hoffmannova^‡, Kevin R. Hinson^§, Oliver Keller^|, Gabriela Packova^‡

‡ Palacky University, Olomouc, Czech Republic

§ EpiLogic GmbH Agrarbiologische Forschung und Beratung, Freising, Germany

| 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

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.

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

ZooBank: urn:lsid:zoobank.org:pub:0ABE7C8D-BD9C-44ED-89D3-CACB78D12AB9

Abstract

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.

Keywords

Catalogue, classification, Drilidae, Lampyridae, neoteny, Oriental Region, Phengodidae

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).

Figure 1.

Morphology of Rhagophthalmidae A 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).

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 21^st 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

ESRI Endemic Species Research Institute, JiJi, Nantou, Taiwan, Taiwan

ICM Insect Center, Moscow, Russia

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

MNHN Museum National d’Histoire Naturelle, Paris, France

MSNG Museo Civico di Storia Naturale, Genova, Italy

MZB Bogor Zoology Museum, Bogor, Indonesia

NHMB Naturhistorisches Museum, Basel, Switzerland

NHMUK Natural History Museum, London, The United Kingdom

NKME Naturkundemuseum Erfurt, Germany

NMNS Department 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

RMNH Naturalis Biodiversity Center, Leiden, The Netherlands

SMNH Swedish Museum of Natural History, Stockholm, Sweden

SMNS Staatliches Museum für Naturkunde, Stuttgart, Germany

TARI Taiwan Agricultural Research Institute, Taichung, Taiwan

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

YCM Yokosuka City Museum, Yokosuka, Japan

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

Systematics

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].

Vietnam.

Literature

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

Genus Bicladum Pic, 1921

Bicladum Pic, 1921b: 12. Gender: neuter. Type species. Bicladum multipunctatum Pic, 1921; by monotypy.

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 D. adamsii

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.

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.

Genus 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].

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).

Genus Dodecatoma Westwood, 1849

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.

Genus 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.

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.

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.

Falsophrixothrix humeralis Pic, 1937

Falsophrixothrix humeralis Pic, 1937: 138.

Falsophrixothrix humeralis ab. unicolor Wittmer, 1938: 301 [unavailable name, ICZN 1999].

Haplocladon indicum Gorham, 1903: 330.

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).

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.

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.

Genus Mimoochotyra Pic, 1937

Mimoochotyra Pic, 1937: 137. Gender: feminine. Type species. Mimoochotyra ocularis Pic, 1937; by monotypy.

Vietnam.

Literature

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

Genus 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).

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).

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].

Genus 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].

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.

Rhagophthalmus confusus Olivier, 1912

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).

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.

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.

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).

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.

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.

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.

Rhagophthalmus ohbai Wittmer in Wittmer and Ohba 1994

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!]].

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 Rhagophthalmidae

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 Rhagophthalmidae

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 (Jurasai Rosa et al., 2020 being more paedomorphic than Tujamita Rosa 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 Rhagophthalmidae

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 Rhagophthalmidae

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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﻿Abstract

Keywords

﻿Introduction

﻿Materials and methods

﻿Type depositories

﻿Systematics

﻿ Rhagophthalmidae Olivier, 1907

Literature

Remarks

﻿Genus Bicladodrilus Pic, 1921

Literature

Remarks

﻿ Bicladodrilus bakeri Pic, 1921

Type depository

Type locality

Distribution

Literature

﻿ Bicladodrilus laticollis Pic, 1923

Type depository

Type locality

Distribution

Literature

﻿Genus Bicladum Pic, 1921

Literature

Remarks

﻿ Bicladum mjobergi Pic, 1930

Type depositories

Type locality

Distribution

Literature

Remarks

﻿ Bicladum multipunctatum Pic, 1921

Type depository

Type locality

Distribution

Literature

﻿Genus Dioptoma Pascoe, 1860

Literature

Remarks

﻿ Dioptoma adamsii Pascoe, 1860

Type depository

Type locality of D. adamsii

Distribution

Literature

Remarks

﻿ Dioptoma atripennis Pic, 1916

Type depository

Type locality

Distribution

Literature

﻿Genus Diplocladon Gorham, 1883

Literature

Remarks

﻿ Diplocladon atripenne Yiu, 2017

Type depository

Type locality

Distribution

Literature

﻿ Diplocladon hasseltii hasseltii Gorham, 1883

Type depository

Type locality

Distribution

Literature

Remarks

﻿ Diplocladon hasseltii testaceum Pic, 1921

Type depository

Type locality

Distribution

Literature

Remarks

﻿Genus Dodecatoma Westwood, 1849

Literature

Remarks

﻿ Dodecatoma bicolor Westwood, 1849

Type depository

Type locality

Distribution

Literature

﻿ Dodecatoma fuscicornis fuscicornis Gorham, 1895

Type depository

Abstract

Introduction

Materials and methods

Type depositories

Systematics

Rhagophthalmidae Olivier, 1907

Genus Bicladodrilus Pic, 1921

Bicladodrilus bakeri Pic, 1921

Bicladodrilus laticollis Pic, 1923

Genus Bicladum Pic, 1921

Bicladum mjobergi Pic, 1930

Bicladum multipunctatum Pic, 1921

Genus Dioptoma Pascoe, 1860

Dioptoma adamsii Pascoe, 1860

Dioptoma atripennis Pic, 1916

Genus Diplocladon Gorham, 1883

Diplocladon atripenne Yiu, 2017

Diplocladon hasseltii hasseltii Gorham, 1883

Diplocladon hasseltii testaceum Pic, 1921

Genus Dodecatoma Westwood, 1849

Dodecatoma bicolor Westwood, 1849

Dodecatoma fuscicornis fuscicornis Gorham, 1895

Dodecatoma fuscicornis testaceicornis Pic, 1921

Dodecatoma gracilis Wittmer, 1995

Dodecatoma parvicornis Wittmer, 1979

Dodecatoma riedeli Wittmer, 1995

Dodecatoma saluki Kazantsev, 2012

Dodecatoma schmidti Kazantsev, 2012

Dodecatoma testaceiceps Pic, 1924

Genus Falsophrixothrix Pic, 1937

Falsophrixothrix costata Pic, 1951

Falsophrixothrix flava Wittmer, 1939

Falsophrixothrix humeralis Pic, 1937

Falsophrixothrix javana (Pic, 1914)

Falsophrixothrix punctata (Pic, 1921)

Falsophrixothrix pygmaea (Olivier, 1911)