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Rhagophthalmidae Olivier, 1907 (Coleoptera,
Elateroidea): described genera and species,
current problems, and prospects for the
bioluminescent and paedomorphic beetle lineage
Robin Kundrata1, Johana Homannova1,
Kevin R. Hinson2, Oliver Keller3, Gabriela Packova1
1Department of Zoology, Faculty of Science, Palacky University, 17. listopadu 50, 77900, Olomouc, Czech
Republic 2 EpiLogic GmbH Agrarbiologische Forschung und Beratung, Hohenbachernstr. 19–21, 85354,
Freising, Germany 3Florida State Collection of Arthropods, Florida Department of Agriculture and Consumer
Services, P.O. Box 147100, Gainesville, FL, 32614-7100, USA
Corresponding author: Robin Kundrata (robin.kundrata@upol.cz)
Academic editor: Vinicius S. Ferreira|Received 12 July 2022|Accepted 12 September 2022|Published 1 November 2022
https://zoobank.org/0ABE7C8D-BD9C-44ED-89D3-CACB78D12AB9
Citation: Kundrata R, Homannova 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
Abstract
Rhagophthalmidae are a small beetle family known from the eastern Palaearctic and Oriental realms. Rhago-
phthalmidae are closely related to railroad worms (Phengodidae) and reies (Lampyridae) with which they
share highly modied paedomorphic females and the ability to emit light. Currently, Rhagophthalmidae
include 66 species classied 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.). e replacement name Haplocladon gorhami Kundrata, nom. nov. is proposed for Diplocladon hasseltii
Gorham, 1883b (described in subgenus Haplocladon) which is preoccupied by Diplocladon hasseltii Gor-
ham, 1883a. e 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. is
study is intended to be a rst step towards a comprehensive revision of the group on both genus and species
levels. Additionally, critical problems and prospects for rhagophthalmid research are briey discussed.
ZooKeys 1126: 55–130 (2022)
doi: 10.3897/zookeys.1126.90233
https://zookeys.pensoft.net
Copyright Robin Kundrata et al. 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.
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Robin Kundrata et al. / ZooKeys 1126: 55–130 (2022)
56
Keywords
Catalogue, classication, Drilidae, Lampyridae, neoteny, Oriental Region, Phengodidae
Introduction
Rhagophthalmidae are a small elateroid family distributed in South, East, and South-
east Asia (Wittmer 1979; Kawashima et al. 2010; Kundrata and Bocak 2011a; Ka-
zantsev 2012). Soft-bodied males are capable of ight, whereas all known females are
strongly paedomorphic and remain larva-like as adults (Fig. 1). Predaceous larvae oc-
cur in soil and leaf litter where they feed on millipedes. Both larvae and adults are bio-
luminescent, although the biology and ecology of most species are unknown (Li and
Liang 2008; Kawashima et al. 2010). Rhagophthalmidae have a convoluted history of
classication. Most genera were originally placed either in Lampyridae or the widely
dened Drilidae (currently Drilini in Elateridae: Agrypninae; Kundrata and Bocak
2011b). e 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 classication of Rhagoph-
thalmidae have varied greatly, and various authors have recognized 3–11 genera in the
group. At various times, the majority of Rhagophthalmidae have been considered ei-
ther a subgroup of Lampyridae (e.g., McDermott 1964, 1966) or Phengodidae (Crow-
son 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).
e 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 classied 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 classied 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 rst catalogues for Rhagophthalmidae and
Drilidae (Olivier 1910), with the latter including Diplocladon (with Haplocladon as a
Annotated catalogue of Rhagophthalmidae Olivier, 1907 (Coleoptera, Elateroidea) 57
synonym) and Dodecatoma. Jakobson (1911) included many soft-bodied groups, including
“Rhagophthalmini”, in his “Cantharididae”. Olivier (1912) revised Rhagophthalmus and
recognized 12 species, ve 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 Sin-
gapore 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) redened 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 redened
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).
Robin Kundrata et al. / ZooKeys 1126: 55–130 (2022)
58
Kawashima (1998) described the morphology of a larviform adult female of
Rhagophthalmus. He also erected Menghuoius Kawashima, 2000 for two Chinese
species originally classied 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 ailand,
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 conrmed 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 Rhagophthalmi-
dae, 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 scien-
tists 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 com-
prehensive study which would summarize all relevant information of all genera and
species in the group. erefore, 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 ro-
bust framework for subsequent taxonomic revisions of all genera in addition to studies
devoted to diversity, evolution, nature conservation, and ecology of the group.
Annotated catalogue of Rhagophthalmidae Olivier, 1907 (Coleoptera, Elateroidea) 59
Materials and methods
Names of family-, genus-, and species-group taxa are given with the name of the au-
thor, and the year and page of publication. e page given is the page where the taxon
name and description are printed. e year and page given for the incorrect subsequent
spellings are the rst year and page in which they are used. Incorrect subsequent spell-
ings 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 pro-
vide lectotype designations to x the species identity for two species of Pseudothilmanus
Pic, 1918. ese 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 rst be revised in detail. Under
each name, the currently valid name is listed rst, followed by synonyms in chrono-
logical 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, classication, phylogeny,
biology, and ecology. Since PhD or any other student theses are not ocially 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 spe-
cies. Dates of publications and exact bibliographic references (especially problematic
ones, often not cited uniformly by researchers) are taken from the following compre-
hensive 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
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, e United Kingdom
NKME Naturkundemuseum Erfurt, Germany
Robin Kundrata et al. / ZooKeys 1126: 55–130 (2022)
60
NMNS National Museum of Natural Science, Taichung, 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, e 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 Motschul-
sky,1854.
Rhagophtalmidae: Junk 1912: 24 [unavailable name, incorrect subsequent spelling not
in prevailing usage].
Phagophthalmidae: 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; Ga-
han (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): classication; Crowson (1955: 68):
remark, morphology [as Rhagophthalminae]; Harvey (1955: 19): checklist, biolumi-
nescence; Raj (1957: 788): larval biology; McDermott (1964: 49): revision [as Rhago-
phthalminae]; McDermott (1966: preface (unnumbered),121): catalogue, distribution
[as Rhagophthalminae]; Mikšić and Mikšić (1966: 31): remark [as Rhagophthalminae];
Annotated catalogue of Rhagophthalmidae Olivier, 1907 (Coleoptera, Elateroidea) 61
Nakane (1968: 3): remark [as Rhagophthalminae]; Crowson (1972: 50): classication,
morphology [as Rhagophthalminae]; McElroy et al. (1974: 415): remark [as Rhago-
pthalmidae [sic!]]; Lawrence (1982: 512): remark; Haneda (1985: 167): biolumines-
cence [as Rhagopthalmidae [sic!]]; Herring (1987: 158): checklist [as Rhagophthal-
minae]; 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): biol-
ogy [also as Rhagophthalidae [sic!]]; Costa et al. (1999: 22): remark [as Rhagophthalmi-
nae]; Goto and Kawashima (2000: 141): distribution; Jeng et al. (2000: 316): remark;
Kawashima (2000: 131): genus description; Kim et al. (2000: 214): molecular phylog-
eny; 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 Rhagoph-
thalminae]; Branham and Wenzel (2003: 3): phylogeny; Chen (2003: 52): morphol-
ogy, bioluminescence; Hayashi and Suzuki (2003: 4): biology, morphology, phylogeny,
gure 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): molecu-
lar phylogeny; Arnoldi et al. (2007: 2): molecular phylogeny, remark; Bocak (2007:
224): catalogue [as Rhagophthalminae]; Bocakova et al. (2007: 477): molecular phy-
logeny [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): mo-
lecular 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 phy-
logeny; 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): classication; 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
Robin Kundrata et al. / ZooKeys 1126: 55–130 (2022)
62
[as Rhagophthalminae]; Ho et al. (2012: 1): species descriptions; Johnson et al. (2012:
178): ICZN case; Kazantsev (2012: 349): species descriptions; Timmermans and Vo-
gler (2012: 299): remark, molecular phylogeny; Kundrata et al. (2013: 201): molecular
phylogeny; Yiu (2013: 101): remark, bioluminescence; Amaral et al. (2014: 415): mo-
lecular 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 phy-
logeny; McKenna et al. (2015: 843): molecular phylogeny [also as Rhagopthalmidae
[sic!]]; Oba (2015: 99): bioluminescence; Amaral et al. (2016: 255): molecular phylog-
eny; Bocak et al. (2016: 2): molecular phylogeny; Kundrata et al. (2016: 293): molecu-
lar phylogeny; Lawrence (2016: 17): classication; Wijekoon et al. (2016: 69): checklist
[also as Rhagophthalminae]; Amaral et al. (2017a: 674): mitogenome, phylogeny; Kun-
drata 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 phyloge-
ny; 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): biouorescence,
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, distribu-
tion; 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 dened 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., Afghani-
stan). 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 re-
ferred, similar to Lawrence and Newton (1995: 858).
Annotated catalogue of Rhagophthalmidae Olivier, 1907 (Coleoptera, Elateroidea) 63
Genus 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): molecu-
lar 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. is genus currently contains two described species from the Philip-
pines and Vietnam, respectively. e 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. is generic
complex is in need of revision.
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.
Robin Kundrata et al. / ZooKeys 1126: 55–130 (2022)
64
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.
Genus 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. is 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. is generic complex is in need of revision.
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. is 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.
Annotated catalogue of Rhagophthalmidae Olivier, 1907 (Coleoptera, Elateroidea) 65
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!]].
Genus 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; Homann (1984: 230): remark; Herring (1987: 158): checklist;
Cicero (1988: 148): remark; Viviani and Bechara (1993: 615): remark; Wittmer and
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66
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. is 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.
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 publica-
tion, and we believe that this name rst 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
Annotated catalogue of Rhagophthalmidae Olivier, 1907 (Coleoptera, Elateroidea) 67
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!]]; Homann (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:
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, rst author’s collection, lateral view F habitus of Rhagophthalmus sp. from China, male,
rst author’s collection, dorsal view. Scale bars: 5.0 mm (A, F ); 1.0 mm (B , E ); 2.0 mm (C , D ).
Robin Kundrata et al. / ZooKeys 1126: 55–130 (2022)
68
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. is species was referred to as “adamsi” in the majority of publications.
e 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 syn-
types, males (MNHN).
Type locality. India: Tamil Nadu, Madurai [Madura].
Distribution. India (Tamil Nadu).
Literature. Pic (1916: 8): original description; McDermott (1966: 122): cata-
logue; 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].
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, gure of male habitus;
Annotated catalogue of Rhagophthalmidae Olivier, 1907 (Coleoptera, Elateroidea) 69
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; Homann (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, gures
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,
gures of habitus; Yiu (2013: 113): remark, biology; Kovalev and Kirejtshuk (2016:
205): remark; Yiu (2017: 64): description of a new species, identication key; Tan
(2018: 135): possible larva, distribution, gures 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 bipec-
tinate antennae (Fig. 1C) and long elytra. is generic complex is in need of revision.
Robin Kundrata et al. / ZooKeys 1126: 55–130 (2022)
70
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, gures of male habitus, pregenital
segments and genitalia.
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, gure 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!]]; Homann (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, gures 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, gure
Annotated catalogue of Rhagophthalmidae Olivier, 1907 (Coleoptera, Elateroidea) 71
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 gures, adults of both sexes which were reported
by Haneda (1950) from Singapore, and repeatedly mentioned in subsequent stud-
ies (e.g., Harvey 1952; Haneda 1955, 1985; Lloyd 1971, 1978; Crowson 1981; Ka-
washima 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.
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. e name “testaceum” is deemed to be subspecic according to Article
45.6.4. of the Code (ICZN 1999).
Genus 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:
Robin Kundrata et al. / ZooKeys 1126: 55–130 (2022)
72
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, identication 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 Afghani-
stan, India, Nepal, Indonesia, and the Philippines. is genus is in need of revision;
taxa from Southeast Asia should be removed from Dodecatoma, and the generic as-
signment 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).
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; Lu-
cas (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, gure of hind wing. In addition to the aforementioned literature, this
species was included in PhD theses by Jeng (2008) and Roza (2022).
Dodecatoma fuscicornis fuscicornis Gorham, 1895
Dodecatoma fuscicornis Gorham, 1895: 309.
Type depository. Described based on “several examples” (Gorham 1895: 309). ree
syntypes, males (NHMUK). Several specimens from Belgaum deposited in MNHN
are potentially syntypes (RK pers. obs.).
Annotated catalogue of Rhagophthalmidae Olivier, 1907 (Coleoptera, Elateroidea) 73
Type locality. India: Karnataka, Belgaum.
Distribution. India (Karnataka).
Literature. Gorham (1895: 309): original description; Olivier (1910: 8): cata-
logue; Wittmer (1944: 211): catalogue; Wittmer (1979: 90): comparison with other
species; Johnson et al. (2012: 179): ICZN case.
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. e name “testaceicornis” is deemed to be subspecic according to
Art. 45.6.4. of the Code (ICZN 1999). is taxon is not morphologically similar to
D.fuscicornis Gorham, 1895 nor to any other species of Dodecatoma.
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, gures 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, identication key.
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.
Robin Kundrata et al. / ZooKeys 1126: 55–130 (2022)
74
Dodecatoma riedeli Wittmer, 1995
Dodecatoma riedeli Wittmer, 1995: 112.
Type depository. Holotype, male (SMNS). ree paratypes, males (NHMB).
Type locality. India: Uttarakhand [“Uttar Pradesh”], Rishikesh.
Distribution. India (Uttarakhand).
Literature. Wittmer (1995: 112): original description, gures 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, identication key.
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 Muk-
teshwar, Satkhol.
Distribution. India (Uttarakhand), Nepal.
Literature. Kazantsev (2012: 349): original description, gures of male habitus,
pregenital segments, and genitalia.
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.
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 prob-
ably only one). One syntype, male (MNHN).
Type locality. Philippines: Luzon, Mt. Maquiling.
Annotated catalogue of Rhagophthalmidae Olivier, 1907 (Coleoptera, Elateroidea) 75
Distribution. Philippines.
Literature. Pic (1924: 713): original description; Wittmer (1941: 197): catalogue;
Wittmer (1944: 212): catalogue [as D. testaceipes [sic!]].
Remarks. is 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): spe-
cies 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; Witt-
mer (1944: 217): catalogue; Pic (1951: 5): species description; Crowson (1972: 52): re-
mark; 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 chap-
ter; 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 lit-
erature, 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 bi-
pectinate 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.
Robin Kundrata et al. / ZooKeys 1126: 55–130 (2022)
76
Falsophrixothrix ava Wittmer, 1939
Falsophrixothrix avus [sic!] Wittmer, 1939: 23.
Type depository. Described based on two specimens. Holotype, male (NHMB); para-
type, 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,
ICZN1999].
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): descrip-
tion of F. humeralis ab. unicolor; Wittmer (1939: 24): comparison with F. avus [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 infrasubspecic according to the Code (ICZN 1999, Article 45.6.2.).
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.
Annotated catalogue of Rhagophthalmidae Olivier, 1907 (Coleoptera, Elateroidea) 77
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): com-
parison with F. humeralis; Wittmer (1939: 24): comparison with F. avus [sic!]; Witt-
mer (1944: 217): catalogue.
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.
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. avus [sic!]; Wittmer (1944: 217): catalogue.
Genus 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): re-
mark; Gorham (1903: 330): species description; Olivier (1910: 8): catalogue; Wittmer
(1944: 211): catalogue; Crowson (1955: 68): remark; Paulus (1972: 49): remark; Li
Robin Kundrata et al. / ZooKeys 1126: 55–130 (2022)
78
and Liang (2008: 109): remark [as D. haplocladon [sic!]]. In addition to the aforemen-
tioned 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). Unfor-
tunately, 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 diers at rst sight from Diplocladon by the unipectinate an-
tennae (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 re-
placed by a new name (see below). Currently, Haplocladon contains two species, one
from Indonesia and one from southern India. Based on the available gures, specimens
reported by Haneda (1950) from Singapore and identied 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 prob-
ably members of Haplocladon.
Haplocladon gorhami Kundrata, nom. nov.
Replacement name for Diplocladon hasseltii Gorham, 1883b.
Diplocladon hasseltii Gorham, 1883b: 250 (described in subgenus Haplocladon). Preoc-
cupied by Diplocladon hasseltii Gorham, 1883a: 6.
Haplocladon haselti: Gorham 1903: 330 [unavailable name, incorrect subsequent spell-
ing not in prevailing usage].
Type depository. Described based on an unknown number of specimens. Two syn-
types, 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): re-
mark [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 ocial name of the va-
riety) 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.
Annotated catalogue of Rhagophthalmidae Olivier, 1907 (Coleoptera, Elateroidea) 79
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): cata-
logue [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].
Genus 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), Myan-
mar,Vietnam.
Literature. Kawashima (2000: 132): original description; Kawashima (2002:
487): species description, gures of habitus, body parts, and male genitalia; Bocak
(2007: 225): catalogue [as Menghouius [sic!]]; Li et al. (2008a: 264): distribution, mor-
phology, biology, gures 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, gures 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 diers 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.
Robin Kundrata et al. / ZooKeys 1126: 55–130 (2022)
80
Menghuoius giganteus (Fairmaire, 1888)
Rhagophthalmus giganteus Fairmaire, 1888: 25.
Menghuoius giganteus: Kawashima 2000: 139.
Rhagophthalmus gigantus: Moreira et al. 2022: 7 [unavailable name, incorrect subse-
quent 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): com-
parison with R. ingens [as R. giganteus]; Hua (2002: 71): catalogue [as R. giganteus];
Bocak (2007: 225): catalogue; Li et al. (2008a: 264): distribution, morphology, bi-
ology, gures 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, gures of male and
female habitus, and female bioluminescence; Moreira et al. (2022: 7): luciferase, mo-
lecular 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):
Annotated catalogue of Rhagophthalmidae Olivier, 1907 (Coleoptera, Elateroidea) 81
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 conspecic 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, gures 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.
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 subse-
quent 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
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82
[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. is 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. e specimen identied
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
inMalaysia.
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
originaldescription].
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!]].
Genus 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.
Annotated catalogue of Rhagophthalmidae Olivier, 1907 (Coleoptera, Elateroidea) 83
Subgenus 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. is subgenus currently contains a single described species from
Java,Indonesia.
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).
Subgenus 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.
Monodrilus (Dodecatomorpha) roberti Pic, 1928
Monodrilus roberti Pic, 1928: 86 [in subgenus Dodecatomorpha].
Type depository. Described based on an unknown number of specimens. ree syn-
types, males (MNHN).
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84
Type locality. Vietnam [“Darsa, en Cochinchine”].
Distribution. 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 mono-
typy. 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, gures of
male habitus, pronotum, and hind wing [2018 erroneously used as the date of the
original description of this genus in gure caption]. In addition to the aforementioned
literature, this genus was included in a PhD thesis by Roza (2022).
Remarks. is genus has relatively long, serrate antennae and long elytra. It con-
tains 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 [hand-
written] / Nov. genus India [handwritten] / Pseudothilmanus alatus Pic [handwritten]”
(treated as the holotype and gured 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, gures of male habitus, antenna, pronotum,
leg, elytral apex, pregenital segments, and genitalia; Roza (2020: 421): morphology, dis-
tribution, gures of male habitus, pronotum, and hind wing [2018 erroneously used as
the date of the original description of this species in gure 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 Uttara-
khand (considered to be part of Uttar Pradesh prior to 2000).
Annotated catalogue of Rhagophthalmidae Olivier, 1907 (Coleoptera, Elateroidea) 85
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 pre-
sent designation, with the following label data: “Type [red printed label] / Type [handwrit-
ten] / Darjeeling Juni Fruhstorfer leg. [printed] / Drilothilmanus marginatus Pic [handwrit-
ten]” (treated as the holotype and gured 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, gures of male habitus,
antenna, pronotum, leg, and genitalia; Roza (2020: 422): morphology, distribution,
gures of male habitus, pronotum, and hind wing [2018 erroneously used as the date
of the original description of this species in gure 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].
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].
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86
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): classication; 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, gures 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, gure 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:
Annotated catalogue of Rhagophthalmidae Olivier, 1907 (Coleoptera, Elateroidea) 87
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, identication
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; Sheeld
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;
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88
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): biouorescence, 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. is 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).
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, gures of male antenna
and genitalia; Bocak (2007: 225): catalogue; Li et al. (2008a: 265): distribution; Li et
al. (2008b: 496): review.
Annotated catalogue of Rhagophthalmidae Olivier, 1907 (Coleoptera, Elateroidea) 89
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, gures of male habitus,
head, antenna and genitalia, and female habitus, head, and bioluminescence; Yiu
(2017:60):remark.
Rhagophthalmus brevipennis Fairmaire, 1896
Rhagophthalmus brevipennis Fairmaire, 1896: 227.
Type depository. Described based on an unknown number of specimens. Syntype,
male (RMNH). ree additional male specimens (on one pin) from MNHN with
labels dierent from the RMNH syntype are also labelled as “Type” but they probably
represent a dierent species.
Type locality. India: Maharashtra, Nagpur.
Distribution. India (Maharashtra).
Literature. Fairmaire (1896: 227): original description; Olivier (1902: 87): cat-
alogue; 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.
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.
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Rhagophthalmus confusus Olivier, 1912
Rhagophthalmus confusus E. Olivier, 1912: 469, 471.
Rhagophthalmus confuses: Wijekoon et al. 2016: 71 [unavailable name, incorrect subse-
quent spelling not in prevailing usage].
Type depository. Described based on an unknown number of specimens. One syn-
type, male (NHMUK). One probable syntype, male (MNHN).
Type locality. Sri Lanka.
Distribution. Sri Lanka.
Literature. Olivier (1912: 469, 471): original description; Pic (1916: 9): com-
parison 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 litera-
ture, this species was included in a PhD thesis by Roza (2022).
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. avus and
R. minutus, respectively; Bocak (2007: 225): catalogue; Li et al. (2008a: 265): distribu-
tion; Li et al. (2008b: 496): review.
Rhagophthalmus liformis Olivier, 1912
Rhagophthalmus liformis E. Olivier, 1912: 469, 470.
Type depository. Described based on an unknown number of specimens. One syn-
type, male (NHMUK).
Type locality. Sri Lanka.
Distribution. Sri Lanka.
Literature. Olivier (1912: 469, 470): original description; Pic (1925a: 17): com-
parison 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.
Annotated catalogue of Rhagophthalmidae Olivier, 1907 (Coleoptera, Elateroidea) 91
Rhagophthalmus avus Kawashima & Satô, 2001
Rhagophthalmus avus Kawashima & Satô, 2001: 424.
Type depository. Holotype, male (NWU). One paratype, male (PCIK).
Type locality. Myanmar: Dawna.
Distribution. Myanmar, ailand.
Literature. Kawashima and Satô (2001: 424): original description, gures 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.
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, identica-
tion key, gures of male habitus, antenna, and genitalia; Bocak (2007: 225): cata-
logue; Li et al. (2008a: 265): distribution; Li et al. (2008b: 496): review; Ho et al.
(2012: 1): remark.
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.
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92
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, gures of male habitus, head,
antenna, and genitalia, and female habitus, head, and luminous organ; Yiu (2017: 60):
comparison with R. motschulskyi.
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): revi-
sion; 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 gures of male genitalia; Li et al. (2008b: 496): review.
Rhagophthalmus hiemalis Yiu, 2017
Rhagophthalmus hiemalis Yiu, 2017: 62.
Type depository. Holotype, male (TLES). 15 paratypes: 10 males, ve 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, gures of male and female habitus
and male genitalia.
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].
Annotated catalogue of Rhagophthalmidae Olivier, 1907 (Coleoptera, Elateroidea) 93
Type depository. Holotype, male (NWU). ree paratypes, males (NTU, PCIK).
Type locality. China/Taiwan, Fenchihu, Chiai Hsien.
Distribution. China/Taiwan.
Literature. Kawashima and Satô (2001: 430): original description, gures 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, gures of
male and female habitus, body parts, bioluminescence; Ho et al. (2012: 1): remark; Jeng
(2019: 13): biouorescence, biology, gures 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 syn-
type, male (MNHN).
Type locality. Laos: akhek [“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.
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94
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; Mc-
Dermott (1966: 122): catalogue; Hua (2002: 71): catalogue; Bocak (2007: 225): cata-
logue [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 origi-
nal description of this species (as “Pic 1923: 29”); however, it was described as a new
species in 1925 (Pic 1925a).
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 subse-
quent 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, gures of male and female habitus, male antenna, and male genitalia;
Li et al. (2008b: 496): review; Sheeld 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): mitogenomicphylogeny.
Annotated catalogue of Rhagophthalmidae Olivier, 1907 (Coleoptera, Elateroidea) 95
Rhagophthalmus minutus Kawashima & Satô, 2001
Rhagophthalmus minutus Kawashima & Satô, 2001: 428.
Type depository. Holotype, male (NWU). ree paratypes, males (two in NWU, one
in PCIK).
Type locality. ailand: Kohn Kaen Province, “near Ban Lon, Lam Chee Yai”.
Distribution. ailand.
Literature. Kawashima and Satô (2001: 428): original description, gures 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.
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): re-
view; Yiu (2011a: 14): remark; Yiu (2011b: 20): biology and bioluminescence, gures
of female bioluminescence; Ho et al. (2012: 1): remark; Yiu (2012: 30): catalogue,
gures of habitus; Yiu (2013: 101): remark, bioluminescence; Yiu (2017: 60): rede-
scription, gures of larva, pupa, and adults, and male genitalia.
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 syn-
type, 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]; McDer-
mott (1966: 122): remark, catalogue [as Ochotyra obscura]; Wittmer and Ohba (1994:
Robin Kundrata et al. / ZooKeys 1126: 55–130 (2022)
96
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 sub-
sequent spelling not in prevailing usage].
Type depository. Described based on an unknown number of specimens. One syn-
type, male (MNHN).
Type locality. Sri Lanka.
Distribution. Sri Lanka.
Literature. Pic (1916: 9): original description; McDermott (1966: 122): cata-
logue; 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. obscursus: Winkler 1925: 522 [unavailable name, incor-
rect 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 [una-
vailable 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 syn-
type, 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.
Annotated catalogue of Rhagophthalmidae Olivier, 1907 (Coleoptera, Elateroidea) 97
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). ree 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, gure of habitus;
Ohba et al. (1996a: 1): morphology, biology, gures of habitus, body details, and
bioluminescence; Ohba et al. (1996b: 30): remark; Nakane (1997: 36): remark;
Ohba (1997a: 5): checklist, biology, gures of larval, male and female habitus, male
head, and female bioluminescence; Ohba (1997b: 19): remark; Ohba (1997c: 51):
breeding, development, immature stages, gures 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, gures 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, gures of adult males and females,
larva, and bioluminescence; Hayashi and Suzuki (2003: 4): morphology, biology,
gure of mating; Kawashima and Sugaya (2003: 353): remark, identication 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, gures of male and female habitus;
Ohba (2004b: 6): bioluminescence, biology, gures of male and female habitus; Lau
and Meyer-Rochow (2006: 20): eye morphology, gures 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
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98
[in Lampyridae incertae 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; Sheeld et al. (2008: 2500): mitochondrial genome; Suzuki and Kobayashi
(2009: 30): embryogenesis, gure of egg; Chen et al. (2010: 203): habitus gure
showing bioluminescence; Kawashima et al. (2010: 137): book chapter, gures of
male and female habitus, and female ovipositor; Lawrence et al. (2011: 7): phylogeny,
gure of female abdomen; Oba et al. (2011: 773): biology, bioluminescence, gures
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, gures 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).
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.
Annotated catalogue of Rhagophthalmidae Olivier, 1907 (Coleoptera, Elateroidea) 99
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; Mc-
Dermott (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; Ka-
washima et al. (2003: 255): remark, catalogue; Kawashima and Sugaya (2003: 358):
remark, identication key; Bocak (2007: 225): catalogue; Li et al. (2008a: 264): com-
parison 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, gure of hind wing. In addition to the aforementioned
literature, this species was included in a PhD thesis by Roza (2022).
Rhagophthalmus semisulcatus Wittmer, 1997
Rhagophthalmus semisulcatus Wittmer, 1997: 259.
Type depository. Holotype, male (NHMB). Five paratypes, males (NHMB). Accord-
ing to the original description (Wittmer 1997), there are only ve paratypes; how-
ever, there are ve additional specimens with dierent 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, biol-
ogy, gure of female habitus; Li et al. (2008b: 496): review.
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100
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”],
SriLanka.
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): distri-
butional note, morphology [as Ochotyra]; Bourgeois (1903: 479): distributional note
[as Ochotiza [sic!]]; Gorham (1903: 330): distributional note [as Ochotyra]; Bour-
geois (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).
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): cata-
logue; Wittmer (1997: 261): comparison with other species; Bocak (2007: 225): cata-
logue; Li et al. (2008a: 265): distribution; Li et al. (2008b: 496): review.
Remarks. is species could be a synonym of R. sulcicollis Olivier, 1912 (see
Wittmer 1997 for more information).
Annotated catalogue of Rhagophthalmidae Olivier, 1907 (Coleoptera, Elateroidea) 101
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). ere are also
two additional specimens in MNHN bearing the labels “lectotype” and “paralectotype”;
however, they have locality label data that dier slightly from the originaldescription.
Type locality. China: Tibet/Xizang, Yalong, over 3000 m.
Distribution. China (Tibet/Xizang).
Literature. Olivier (1912: 471): original description; Winkler (1925: 522): cata-
logue; Pic (1925b: 72): comparison with R. sulcatus; McDermott (1966: 122): cata-
logue; 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.
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.
Rhagophthalmus sumatrensis Olivier, 1885
Rhagophthalmus sumatrensis Olivier, 1885: 372.
Type depository. Described based on an unknown number of specimens. ree syn-
types, males (MSNG).
Type locality. Indonesia: Sumatra, Mt. Singalang.
Distribution. Indonesia (Sumatra).
Literature. Olivier (1885: 372): original description; Fairmaire (1889: 353): com-
parison 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.
Robin Kundrata et al. / ZooKeys 1126: 55–130 (2022)
102
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.
Rhagophthalmus tonkineus Fairmaire, 1889
Rhagophthalmus tonkineus Fairmaire, 1889: 352.
Rhagophthalmus tonkinensis: McDermott 1966: 122 [unavailable name, incorrect sub-
sequent 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): cata-
logue; Olivier (1912: 470): revision; Pic (1917: 4): comparison with R. obscurus; Pic
(1923: 29): catalogue, distributional note; Winkler (1925: 522): catalogue; McDer-
mott (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).
Annotated catalogue of Rhagophthalmidae Olivier, 1907 (Coleoptera, Elateroidea) 103
Literature. Olivier (1912: 469, 471): original description; Pic (1917: 4): compari-
son with R. laosensis; Winkler (1925: 522): catalogue; McDermott (1966: 122): cata-
logue [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): distribu-
tion [as R. xanthogenus [sic!]]; Li et al. (2008b: 496): review [as R. xanthogenus [sic!]].
Taxa removed from Rhagophthalmidae
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 Rhagophthal-
midae. Lawrence and Newton (1995) and Bocak (2007) classied Cydistus in Phengo-
didae: Rhagophthalminae. Lawrence et al. (2010a, b) and Lawrence (2016) considered
Cydistinae in Elateriformia incertae sedis. Finally, Kundrata et al. (2019) were the rst
to include Cydistinae in a molecular phylogenetic analysis, and found them sister to
the New World Phengodidae, which are only distantly related to Rhagophthalmidae.
is placement was conrmed by a morphology-based analysis by Roza (2022).
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. is rey species was originally described in Luciola Laporte, 1833
(Bourgeois 1884) and later transferred to Rhagophthalmus by Olivier (1902). e same
author later placed this species in his new genus Bourgeoisia Olivier, 1908, and McDer-
mott (1966) subsequently designated it the type species of this genus. Bourgeoisia is
currently considered a synonym of Luciola (e.g., Ballantyne and Lambkin 2013; Ballan-
tyne 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).
Robin Kundrata et al. / ZooKeys 1126: 55–130 (2022)
104
Reductodrilus Pic, 1943 [Lampyridae: Ototretinae]
Reductodrilus Pic, 1943: 9. Type species. Reductodrilus nigroapicalis Pic, 1943; bymonotypy.
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 subspecic 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 dierent families (e.g., Lampyridae,
Lycidae, Omethidae, and Rhagophthalmidae) by Crowson (1972), Reductodrilus re-
mained in an uncertain position. Kundrata and Bocak (2011a) listed it in Rhagoph-
thalmidae 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 attened 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 revi-
sion of this genus should improve our understanding of its systematic position.
Discussion
Although Rhagophthalmidae have been known to entomologists for more than a cen-
tury, their taxonomy and classication are still poorly known. e number of genera in-
cluded in Rhagophthalmidae and also their placement within Elateroidea classication
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 sys-
tematic research has accelerated and the classication 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). is is especially striking when com-
pared 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
Annotated catalogue of Rhagophthalmidae Olivier, 1907 (Coleoptera, Elateroidea) 105
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 Rhagophthalmidae
e phylogenetic placement of Rhagophthalmidae within Elateroidea has been con-
troversial 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 vari-
ous datasets and analytical approaches repeatedly conrmed 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, larvi-
form 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) dened the “lampyroid clade”, which contains Lampyridae,
Phengodidae, Rhagophthalmidae, and Sinopyrophoridae. Fossil Cretophengodidae
were probably also a part of that clade (Li et al. 2021b).
e date of the origin of Rhagophthalmidae is unclear, as there are no known fos-
sils 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 mo-
lecular dating analyses of the elaterid-lampyroid clade, and showed that median esti-
mates 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. e group
was originally proposed only for Dioptoma, Ochotyra, and Rhagophthalmus (Olivier
1907, 1910), and later Pic (1937) added Mimoochotyra. is 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). e 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 dened has never been rigorously tested.
Several genera were included in molecular phylogenetic analyses, including
Bicladodrilus, Falsophrixothrix, Mimoochotyra, and Rhagophthalmus (incl. Ochotyra)
Robin Kundrata et al. / ZooKeys 1126: 55–130 (2022)
106
(Bocakova et al. 2007; Kundrata et al. 2014, 2019). ese genera always formed a
monophylum. However, it should be noted that at least the generic placements of
specimens identied as Bicladodrilus sp. from China and Mimoochotyra sp. from
Malaysia are dubious. As Bicladodrilus, Bicladum, and Diplocladon are similar in
general appearance and possess biabellate 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). e 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. ese 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. ese 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
identication of larvae or females. For example, Jeng (2008) suggested that the giant
larviform female from Yunnan, China, identied as Diplocladon by Li and Liang
(2008) is “likely of Menghuoius giganteus or the other related species described from
there” (Jeng 2008: 135). e correct identication of larviform females is, however,
very important for conclusions on the evolution of morphological modications
Annotated catalogue of Rhagophthalmidae Olivier, 1907 (Coleoptera, Elateroidea) 107
caused by paedomorphosis (e.g., Jeng 2008; Kawashima et al. 2010). is should be
possible by e.g., rearing both sexes of the same species from larvae, nding 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 conrm the presence of bioluminescence in the remaining
rhagophthalmid genera.
Elateroid beetles are well-known not only for bioluminescence but also for
morphological modications caused by paedomorphosis (Crowson 1972; Cicero
1988; Bocak et al. 2008; Ferreira and Ivie 2022). In Elateroidea, mainly adult females
are more or less modied, with a gradual series of morphological modications 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 dierent degrees of paedomorphic modications, 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 ve tarsomeres and two pretarsal
claws, and ovipositor; Kawashima et al. 2010). Similar cases of dierent levels of
morphological modications in females of dierent genera were also reported for e.g.,
Elateridae (Drilus Olivier, 1790 being more paedomorphic than Omalisus Georoy,
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 Georoy, 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 classication 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
Robin Kundrata et al. / ZooKeys 1126: 55–130 (2022)
108
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 identication 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 biabellate 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 unavail-
able type specimens, especially in long-neglected groups, such as Rhagophthalmi-
dae. 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 ve 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. e 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 ve 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. e genera Dodecatoma and
Pseudothilmanus are known only from the Himalayas and surrounding regions (one
Annotated catalogue of Rhagophthalmidae Olivier, 1907 (Coleoptera, Elateroidea) 109
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 re-
mained 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). e Sunda Shelf is a southward expansion of the continen-
tal shelf of Southeast Asia that was intermittently exposed by lowered sea levels dur-
ing the Pleistocene (Hall 1998; Voris 2000; Lomolino et al. 2017). e 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 modied 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 rst author examined
for a planned generic revision of the group. A single, unreported rhagophthalmid spe-
cies 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 rst comprehensive catalogue of the currently dened Rhagoph-
thalmidae. e only catalogues of the group were those of Olivier (1910) and McDer-
mott (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. e
phylogenetic position of Rhagophthalmidae as a sister group to Phengodidae is now gen-
erally accepted; however, interrelationships within the group and generic classication
remain poorly known. Although morphology-based analyses in two PhD studies that
were focused on related families (Jeng 2008; Roza 2022) conrm the monophyly of cur-
rently circumscribed Rhagophthalmidae, molecular analysis including representatives of
Robin Kundrata et al. / ZooKeys 1126: 55–130 (2022)
110
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 modied 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, eld observations, morphological studies, and
correct genus and species identications of larvae and larviform females of Rhagoph-
thalmidae 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 pho-
tographs of specimens and/or information on type specimens deposited in their col-
lections: 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 cor-
rect 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. is study was funded by an internal grant of the Faculty
Annotated catalogue of Rhagophthalmidae Olivier, 1907 (Coleoptera, Elateroidea) 111
of Science, UP Olomouc (IGA_PrF_2022_024; to RK, JH, and GP). RK received
support from the SYNTHESYS+ project which is nanced 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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