Content uploaded by Robin Kundrata
Author content
All content in this area was uploaded by Robin Kundrata on Nov 03, 2021
Content may be subject to copyright.
Foss. Rec., 24, 347–355, 2021
https://doi.org/10.5194/fr-24-347-2021
© Author(s) 2021. This work is distributed under
the Creative Commons Attribution 4.0 License.
Cretopachyderes gen. nov., a new remarkable click beetle
(Coleoptera: Elateridae: Agrypninae) from the
mid-Cretaceous Burmese amber
Robin Kundrata1, Alexander S. Prosvirov2, Roger Long3,4, and Gabriela Packova1
1Department of Zoology, Faculty of Science, Palacký University Olomouc, 17. listopadu 50,
77146, Olomouc, Czech Republic
2Department of Entomology, Faculty of Biology, Moscow State University, Leninskie gory 1/12, 119234, Moscow, Russia
3Myanmar Amber Museum, 136 37th Street, 1143, Yangon, Myanmar
4private address: 7305 Crape Myrtle Way, Sarasota, Florida 34241, USA
Correspondence: Robin Kundrata (robin.kundrata@upol.cz)
Received: 4 August 2021 – Revised: 25 September 2021 – Accepted: 8 October 2021 – Published: 3 November 2021
Abstract. Although the Mesozoic Era played an important
role in the evolution and diversification of Elateridae, the
Cretaceous click-beetle fauna remains very poorly known.
Here we describe Cretopachyderes burmitinus gen. et sp.
nov. based on a single specimen from the mid-Cretaceous
Burmese amber. This species is remarkable for its extremely
long posterior angles of pronotum, which is a unique charac-
ter among fossil Elateridae. We discuss the diagnostic char-
acters of Cretopachyderes gen. nov. and tentatively place it
to subfamily Agrypninae close to extant genus Pachyderes
Guérin-Méneville, 1829.
1 Introduction
Elateridae, commonly known as click beetles, is the species-
rich cosmopolitan family belonging to the polyphagan su-
perfamily Elateroidea. The typical representatives have a
compact body and a pro-mesothoracic clicking mechanism
(Costa et al., 2010); however, this group also includes sev-
eral soft-bodied lineages (Kundrata and Bocak, 2019). De-
spite recent progress in understanding the composition, phy-
logeny, and classification of Elateridae, all these aspects re-
main open to further study (Kundrata et al., 2018a; Bi et al.,
2019; Kundrata et al., 2019b; Kusy et al., 2021; Douglas et
al., 2021). The most recent study on the phylogeny of Ela-
teridae (Douglas et al., 2021) showed that the Lampyridae
and related bioluminescent families may be in fact derived
click beetles, which means that the widely delimited Elateri-
dae clade comprises more than 13 500 extant species world-
wide (Costa et al., 2010; Douglas et al., 2021). The fossil
record of Elateridae includes 261 species classified in 99 gen-
era and nine subfamilies (Kundrata et al., 2020, 2021). The
Mesozoic click-beetle diversity includes 143 species, with
most of them described from the Jurassic Karatau (Dolin,
1975, 1976, 1980); however, the family placement of many
of them needs revision (Muona et al., 2020; Kundrata et al.,
2021). Only three click-beetle species have been recorded
from the mid-Cretaceous Burmese amber to date (Cockerell,
1917; Otto, 2019), although this deposit is rich in the Elateri-
dae inclusions (Kundrata et al., 2021). In this study, we report
a discovery of a new enigmatic Burmese-amber click-beetle
genus with extremely pronounced posterior angles of prono-
tum, which is a character known only in several present-day
Elateridae lineages.
2 Material and methods
The amber piece was photographed submerged in a Petri
dish of cedar wood oil to help correct for the distortion
caused by the curved faces of the amber. Images were taken
using the following photographic equipment: Kaiser Copy
Stand (stand), Cognisys StackShot Macro Rail Package (im-
age capturing system), Canon 5D Mark II (camera), Canon
MP-E 65 mm f/2.8 1–5×(lens), Canon Macro Twin Lite MT-
24EX TTL (macro main flash), and Lume Cube 2.0 LED
Published by Copernicus Publications on behalf of the Museum für Naturkunde Berlin.
348 R. Kundrata et al.: Cretopachyderes gen. nov., a new remarkable click beetle
fixed light (additional lighting). Adobe Photoshop Lightroom
CC (version 2019-20.0.5) was used for the lens correction
and basic editing, and Adobe Photoshop CC (version 2019-
20.0.5) was used for the noise reduction and image refine-
ment. Stacks of photographs were combined with the soft-
ware Zerene Stacker Professional (version 1.04, Zerene Sys-
tems, Richland, WA, USA), applying the DMap method. Fi-
nal image plates were assembled using Adobe Photoshop CC
(version 2019-20.0.5).
Morphological terminology follows Costa et al. (2010)
and Kundrata et al. (2019b), and the Elateridae classifica-
tion follows Kundrata et al. (2019a). The holotype of the
newly described species was originally deposited in the col-
lection of the Myanmar Amber Museum, Yangon, Myanmar
(MAM), which is currently closed. Therefore, the holotype is
currently in the personal collection of the third author (Roger
Long, Sarasota, FL, USA) and will be eventually sent back to
the MAM when it opens again. The ZooBank LSID number
for this publication is urn:lsid:zoobank.org:pub:1894924B-
A1FE-43AF-A97F-50DAE387E9C0.
3 Systematic palaeontology
Order Coleoptera Linnaeus, 1758
Suborder Polyphaga Emery, 1886
Series Elateriformia Crowson, 1960
Superfamily Elateroidea Leach, 1815
Family Elateridae Leach, 1815
Subfamily Agrypninae Candèze, 1857
Tribe Incertae sedis
Genus Cretopachyderes gen. nov.
urn:lsid:zoobank.org:act:6F4C5567-C295-4E9C-8713-
FEB5E61381EF; Figs. 1–4.
Type species
Cretopachyderes burmitinus sp. nov., here designated.
Etymology
Derived from the words “Creto” (referring to the Creta-
ceous Period) and “Pachyderes” (a genus name in Elateridae)
(Fig. 5). Gender: masculine.
Diagnosis
Cretopachyderes gen. nov. can be readily recognized among
other Mesozoic click-beetle genera by the distinctly pro-
nounced posterior angles of pronotum (Figs. 1–3). Addi-
tionally, it can be recognized by the following combina-
tion of characters: body oval-elongate, rather flattened, an-
tenna serrate, relatively short, almost reaching posterior an-
gles of pronotum, pronotal sublateral carina distinct, com-
ing from subapical inner part of posterior angle and run-
ning medially across whole length of posterior angle, hy-
pomeron with posterior margin subapically subrectangularly
emarginate, pronotosternal sutures anteriorly open, proster-
nal process very slightly gradually narrowed toward apex,
subapically abruptly narrowed, tibia shorter than femur and
also shorter than tarsus, without paired spurs, tarsomeres
simple (Figs. 1–4).
Description
Adult. Body (Fig. 1) oval-elongate, rather flattened. Head
(Figs. 1–3) retracted to prothorax, less than half as wide
as pronotum. Eyes moderately large, with their major part
hidden in prothorax. Antenna (Figs. 2b; 3c–e) serrate from
antennomere 4, relatively short, almost reaching posterior
angles of pronotum; antennomere 3 short, slightly longer
than wide, distinctly shorter than antennomere 4, anten-
nomeres 4–10 elongate, with short serrations; terminal an-
tennomere simple, elongate, apically narrowed and narrowly
rounded. Terminal maxillary palpomere securiform. Prono-
tum (Figs. 1–3) widest at one-third (without posterior an-
gles), otherwise widest at posterior angles when posterior
angles included, distinctly wider than long if measured along
midline (without posterior angles), otherwise slightly longer
than wide when posterior angles included. Anterior an-
gles very short; lateral sides rounded anteriorly, then sub-
parallel; posterior angles very long, surpassing humeral part
of elytra, sharp, oriented posteriorly, with inner margin with
distinct tooth and outer margin slightly concave. Lateral
carina distinct, complete. Sublateral carina distinct, com-
ing from subapical inner part of posterior angle and run-
ning medially across whole length of posterior angle. Hy-
pomeron with posterior margin subapically subrectangularly
emarginate. Pronotosternal sutures almost straight, anteriorly
open. Prosternum (Figs. 2b; 3c) elongate; prosternal lobe
short, rounded; prosternal process robust, very slightly grad-
ually narrowed toward apex, subapically abruptly narrowed.
Scutellar shield (Figs. 1b; 2a; 3a; 4a) longer than wide, nar-
rowly rounded posteriorly. Elytra (Figs. 1a, c; 2a, b; 3a, c;
4a, b) together about twice as long as wide, widest at basal
third, with striae formed by lines of fine punctures; inters-
triae smooth, slightly convex. Metaventrite wider than long.
Metacoxal plate (Fig. 2b) with basal portion wide, poste-
riorly widely roundly emarginate, then obliquely abruptly
narrowed towards body edge. Leg (Figs. 1a–c; 2a–b; 4d–
h) slender, femur robust, elongate; tibia shorter than femur,
densely covered with thick long sharp setae, without distinct
spurs; tarsus slender, elongate, distinctly longer than tibia;
tarsomeres simple; claws simple. Abdomen (Figs. 2b; 4c)
with ventrites 1–4 subequal in length; ventrite 5 longer, nar-
rowly rounded to subacute apically.
Foss. Rec., 24, 347–355, 2021 https://doi.org/10.5194/fr-24-347-2021
R. Kundrata et al.: Cretopachyderes gen. nov., a new remarkable click beetle 349
Figure 1. Cretopachyderes burmitinus gen. et sp. nov., holotype: (a) general view of the inclusion in amber stone; (b) anterior part of body,
dorsal view; (c) habitus, dorsal view. Scale bar equals 2.0 mm (a, c) and 1.0 mm (b).
Composition and distribution
Cretopachyderes gen. nov. is a monotypic genus and is
known exclusively from the Cretaceous Burmese amber.
Cretopachyderes burmitinus sp. nov.
urn:lsid:zoobank.org:act:428E20BE-1DC2-482F-
B81C-A8BC1439A027; Figs. 1–4.
Type material
Holotype, adult specimen, sex unknown, MAM0012
(MAM). A complete beetle is included in a transparent, yel-
low amber piece with dimensions of 13.5 ×15.9 ×4 mm,
without any syninclusions, only with air bubbles.
Type stratum and age
Mid-Cretaceous (upper Albian to lower Cenomanian) (Shi et
al., 2012; Mao et al., 2018; Yu et al., 2019).
Type locality
Hukawng Valley, Tanai Township, Myitkyina District,
Kachin State, northern Myanmar.
Etymology
The specific epithet “burmitinus” is derived from “burmite”
(mid-Cretaceous Burmese amber).
Diagnosis
As for the genus (vide supra).
Description
Adult. Body (Fig. 1) about 7.5 mm long (measured from frons
to apex of elytra) and 2.6 mm wide (measured across widest
part of pronotum), oval-elongate, rather flattened; dorsally
sparsely setose.
Head (Figs. 1–3) subquadrate, retracted to prothorax, 0.45
times as wide as pronotum width. Eyes moderately large,
strongly protuberant, with their major part hidden in protho-
rax. Labrum wider than long, anteriorly slightly concave. An-
tenna (Figs. 2b; 3c–e) serrate from antennomere 4, relatively
short, almost reaching posterior angles of pronotum; scape
robust; pedicel not well visible; antennomere 3 short, slightly
longer than wide, antennomere 4 1.6 times as long as an-
tennomere 3, antennomeres 4–10 elongate, with short serra-
tions; terminal antennomere simple, elongate, about 3 times
as long as wide, about 1.3 times as long as antennomere
10, widest at two-thirds, apically narrowed and narrowly
https://doi.org/10.5194/fr-24-347-2021 Foss. Rec., 24, 347–355, 2021
350 R. Kundrata et al.: Cretopachyderes gen. nov., a new remarkable click beetle
Figure 2. Cretopachyderes burmitinus gen. et sp. nov., holotype: (a) habitus, dorsal view; (b) habitus, ventral view. Scale bar equals 2.0 mm.
rounded. Mandible robust, relatively short, not well visible.
Maxillary palpus moderately long; terminal palpomere secu-
riform, apically flattened.
Pronotum (Figs. 1–3) weakly convex, without posterior
angles widest at one-third, 1.45 times wider than long if
measured along midline, and when posterior angles included
then widest between their apices, and about 1.1 times longer
than wide if length measured between anterior and poste-
rior angles and width between the apices of posterior an-
gles. Anterior margin widely concave; anterior angles very
short; lateral sides rounded anteriorly, then sub-parallel; pos-
terior angles very long, about 0.7 times as long as prono-
tal length when measured along midline, surpassing humeral
part of elytra, gradually narrowed toward apex, relatively
sharp, oriented posteriorly, with inner margin with distinct
tooth and outer margin slightly concave; posterior margin
medially emarginate. Lateral carina distinct, complete. Disc
rather flattened, distinctly punctate; punctures sparser medi-
ally and denser and larger on sides, interstices smooth; sub-
lateral carina distinct, coming from subapical inner part of
posterior angle and running medially across whole length of
posterior angle. Hypomeron somewhat shiny, with posterior
margin subapically subrectangularly emarginate, with punc-
tures distinct, large, separated 1.0–2.0 times of their diam-
eter. Pronotosternal sutures almost straight, anteriorly open.
Prosternum (Figs. 2b; 3c) elongate, about 2.7 times as long as
wide including prosternal process, about 1.8 times as long as
wide without prosternal process; prosternal lobe well devel-
oped, short, rounded; prosternal process robust, very slightly
gradually narrowed toward apex, subapically abruptly nar-
rowed. Scutellar shield (Figs. 1b; 2a; 3a; 4a) about 1.5 times
as long as wide, narrowed subanteriorly, widest medially,
then gradually narrowed toward apex, widely rounded an-
teriorly, narrowly rounded posteriorly, sides sinuate. Ely-
tra (Figs. 1a, c; 2a, b; 3a, c; 4a, b) together about twice
as long as wide (measured at widest place) and 2.5 times
as long as pronotum; weakly convex, widest at basal third,
then gradually slightly narrowed towards apex; each elytron
basally distinctly but weakly emarginate, with striae formed
by lines of fine punctures; interstriae smooth, slightly con-
vex; pubescence sparse, semi-erect, moderately long, ori-
ented apically; elytral epipleuron moderately narrowed be-
hind metacoxae. Mesoventrite not well visible. Metaventrite
rather short and wide, 1.25 times as wide as long, moder-
ately convex, sparsely covered by punctures. Metacoxal plate
(Fig. 2b) with basal portion well developed, wide, poste-
riorly widely roundly emarginate, then obliquely abruptly
narrowed towards body edge. Hind wing not visible. Leg
(Figs. 1a–c; 2a–b; 4d–h) slender, femur robust, elongate,
ventrally with distinct groove for reception of tibia; tibia
shorter than femur, densely covered with thick long sharp
setae, without distinct spurs; tarsus slender, elongate, dis-
Foss. Rec., 24, 347–355, 2021 https://doi.org/10.5194/fr-24-347-2021
R. Kundrata et al.: Cretopachyderes gen. nov., a new remarkable click beetle 351
Figure 3. Cretopachyderes burmitinus gen. et sp. nov., holotype: (a) anterior part of body, dorsal view; (b) head and anterior part of pronotum,
dorsal view; (c) anterior part of body, ventral view; (d) antennomeres 10 and 11; (e) antenna and anterior part of prothorax, ventral view.
Scale bar equals 1.0 mm (a, c), 0.5 mm (b, e), and 0.1 mm (d).
tinctly longer than tibia. Tarsomere I longest, slightly shorter
than tarsomere II and III combined; length ratio of tar-
someres I–V 4.2 : 2.7 : 2.0 : 1.0 : 3.0; tarsomeres I–IV grad-
ually shorter, simple, almost subparallel sided, only api-
cally slightly widened, moderately densely covered by setae,
which are denser and longer apically; apical tarsomere elon-
gate, slender; claws (Fig. 4h) simple, slender, long, moder-
ately curved.
Abdomen (Figs. 2b; 4c) with ventrites moderately densely
covered by distinct punctures; ventrites 1–4 subequal in
length; ventrite 5 about 1.35 times as long as ventrite 4, sub-
triangular, narrowly rounded to subacute apically. Pregenital
segments and genitalia not visible.
4 Discussion
Fossils play an important role not only in our understanding
of the past diversity and evolutionary processes but also in the
more accurate dating of phylogenetic events (Donoghue and
Yang, 2016; Toussaint et al., 2017). Therefore, the reliable
identification and systematic placement of the fossils are cru-
cial tasks for taxonomists. The here described Cretopachy-
deres burmitinus gen. et sp. nov. belongs without any doubt
to Elateridae based on its habitus, exposed labrum, project-
ing posterior angles of pronotum, concealed protrochantins,
well-developed metacoxal plates, and presumably connate
first four abdominal ventrites (Calder, 1996; Costa et al.,
2010). Cretopachyderes gen. nov. is very distinctive due to
https://doi.org/10.5194/fr-24-347-2021 Foss. Rec., 24, 347–355, 2021
352 R. Kundrata et al.: Cretopachyderes gen. nov., a new remarkable click beetle
Figure 4. Cretopachyderes burmitinus gen. et sp. nov., holotype: (a) posterior part of pronotum, scutellar shield and anterior part of elytra,
dorsal view; (b) apex of elytra, dorsal view; (c) apex of abdomen, ventral view; (d) metaleg, ventral view; (e) pro- and mesoleg, dorsal
view; (f) metatibia and metatarsus, dorsal view; (g) metatibia and metatarsus, ventral view; (h) apical part of metatarsus with pretarsal claws,
ventral view. Scale bar equals 0.5 mm (a–e), 0.3 mm (f, g), and 0.1 mm (h).
Figure 5. Live specimen of extant Pachyderes cf. apicalis Candèze, 1865, Singapore: (a, b) habitus, dorsal view; (c) detail of head, antennae
and pronotum, fronto-dorsal view. Credit for photographs: Adeline Goh, Singapore.
its extremely elongate posterior angles of pronotum, which
have never been reported in any fossil Elateridae. Addition-
ally, this character is only rarely present in extant Elateridae,
with the best examples being Dima spicata Schimmel, 1999
(Dendrometrinae: Dimini), and the representatives of genus
Pachyderes Guérin-Méneville, 1829 (Fig. 5). Cretopachy-
deres gen. nov. clearly differs from Dima spicata and other
Dimini in having short antennae (vs. long, usually surpassing
posterior angles of pronotum or even reaching half of elytra),
the pronotal disc flattened (vs. distinctly convex; somewhat
flattened in some Penia spp.), with sides more parallel-sided
(vs. clearly rounded or sinuate) and with a short sublateral ca-
Foss. Rec., 24, 347–355, 2021 https://doi.org/10.5194/fr-24-347-2021
R. Kundrata et al.: Cretopachyderes gen. nov., a new remarkable click beetle 353
rina which reach only the base of posterior angles (vs. long
sublateral carina running along almost whole lateral carina),
the head retracted to prothorax, the elytra more elongate (vs.
oval to elongate-oval in most Dimini), the metacoxal plates
well developed (vs. distinctly reduced towards elytra), and
the tarsomeres simple (vs. at least some tarsomeres ventrally
lobate) (Kundrata et al., 2018b; Qiu et al., 2020).
The new fossil genus is morphologically much more sim-
ilar to Pachyderes – not only in the habitus and shape of
pronotum with extremely pronounced posterior angles, but
also in having the hypomeron with posterior margin subapi-
cally subrectangularly emarginate, body dorsally flattened,
pronotosternal sutures almost straight, open anteriorly, head
more or less depressed above, with straight anterior margin of
frons, antennomeres III–X with elevated area at midline, ab-
sence of basal furrows on pronotum, prosternal lobe well de-
veloped, short and rounded, and prosternal process elongate,
without subapical tooth. However, Pachyderes species have
head not so retracted into prothorax (although this character
in Cretopachyderes gen. nov. might be just an artifact caused
by a defensive behavior possibly connected with the speci-
men preservation in amber), and especially they have tarsi
about as long or only slightly longer than tibiae (vs. tarsi dis-
tinctly longer than tibiae), tarsomeres II–IV relatively much
shorter and wider (vs. slender, distinctly elongate), and all
of them or at least tarsomere IV apically distinctly widened
and often ventrally lobate (vs. all tarsomeres simple). Pachy-
deres contains approximately 20 species known mainly from
the South East Asia, with a few species reaching also India,
Nepal, and China (e.g., Schenkling, 1925; Arimoto, 1993).
This genus was placed either in a separate subfamily Pachy-
derinae (Fleutiaux, 1919; Schenkling, 1925), Conoderinae
(current Oophorini) (Fleutiaux, 1928), tribe Pachyderini in
Denticollinae (current Dendrometrinae) (Stibick, 1979), or
tribe Oophorini in Agrypninae (Cate, 2007; Kundrata et al.,
2019a). Most recent phylogenomic analysis of Elateridae
(Douglas et al., 2021) showed that Pachyderes indeed be-
longs to Agrypninae but not to Oophorini and hence that
the possible redefinition of Pachyderini and its resurrection
from synonymy under Oophorini should be considered in
future studies. Taking into consideration that based on the
available evidence we consider Cretopachyderes gen. nov.
related to Pachyderes and that Pachyderes was found out-
side Oophorini in recent phylogenetic analyses (Douglas et
al., 2021), we tentatively place Cretopachyderes gen. nov. in
Agrypninae incertae sedis until the classification of Pachy-
deres is firmly settled or until further, better preserved, ma-
terial of Cretopachyderes gen. nov. shows that this genus
should belong to another place in the Elateridae tree of life.
One of the characters traditionally used for the definition of
Agrypninae is a presence of setae basally on the pretarsal
claws (Calder, 1996; Costa et al., 2010). There are no long
setae on claws visible on the specimen of Cretopachyderes
gen. nov.; however, they are often not well visible also on
some recent Elateridae species (Robin Kundrata and Alexan-
der S. Prosvirov, personal observation, 2021). Moreover, this
character is not as stable as originally thought, since some
Agrypninae obviously lack such setae and some members of
other subfamilies possess them (Kundrata et al., 2018a).
The peculiar morphology of the newly discovered elaterid
genus Cretopachyderes gen. nov. from the mid-Cretaceous
Burmese amber significantly contributes to a better under-
standing of the palaeodiversity and evolution of the group.
Further material of this interesting lineage may help us not
only with its proper systematic placement but also with the
recognition of the potential sexual dimorphism in this group.
One would expect that the extremely pronounced posterior
angles of pronotum are domain of the males only, but in
extant Dima spicata and Pachyderes spp. there is no strong
sexual dimorphism in this character (Qiu et al., 2020). Some
species of Pachyderes have pectinate antennae, while females
have serrate antennae, such as the examined specimen of
Cretopachyderes gen. nov.; however, we do not know if the
latter genus contained species with pectinate antennae.
Sample availability. The holotype of Cretopachyderes burmitinus
gen. et sp. nov. is deposited in the collection of the Myanmar Amber
Museum (MAM) (temporarily kept in the collection of Roger Long,
USA).
Author contributions. RK conceived and designed the study. RK,
ASP, RL, and GP carried out the morphological investigation. RL,
GP, and RK prepared figure plates. RK wrote the initial manuscript
with help of ASP, RL, and GP. All authors discussed the results and
edited, reviewed, and approved the manuscript.
Competing interests. The contact author has declared that neither
they nor their co-authors have any competing interests.
Disclaimer. Publisher’s note: Copernicus Publications remains
neutral with regard to jurisdictional claims in published maps and
institutional affiliations.
Acknowledgements. We are grateful to Alan Simpson (Ohio, USA)
for help with the photographs and to Adeline Goh (Singapore) for
her kind permission to use her photographs of live Pachyderes cf.
apicalis Candèze, 1865, from Singapore.
Financial support. This study was funded by the internal grant
of the Faculty of Science, UP Olomouc (IGA_PrF_2021_019), to
Robin Kundrata and Gabriela Packova and by the Russian Sci-
ence Foundation (RSF) research project no. 21-74-10024, https:
//rscf.ru/project/21-74-10024/ (last access: 30 October 2021; anal-
ysis of Pachyderes spp.), to Alexander S. Prosvirov.
https://doi.org/10.5194/fr-24-347-2021 Foss. Rec., 24, 347–355, 2021
354 R. Kundrata et al.: Cretopachyderes gen. nov., a new remarkable click beetle
Review statement. This paper was edited by Carolin Haug and re-
viewed by Enrico Ruzzier and one anonymous referee.
References
Arimoto, H.: Some Pachyderes-species (Coleoptera, Elateridae)
from Asia with the description of a new species, Gekkan-Mushi,
270, 9–13, 1993.
Bi, W.-X., He, J.-W., Chen, C.-C., Kundrata, R., and Li,
X.-Y.: Sinopyrophorinae, a new subfamily of Elateridae
(Coleoptera, Elateroidea), with the first record of a lumi-
nous click beetle in Asia and evidence for multiple ori-
gins of bioluminescence in Elateridae, ZooKeys, 864, 79–97,
https://doi.org/10.3897/zookeys.864.26689, 2019.
Calder, A. A.: Click Beetles: Genera of the Australian Elateri-
dae (Coleoptera), Monographs on invertebrate taxonomy, Vol. 2,
CSIRO Publishing, Victoria, Australia, 401 pp., 1996.
Candèze, E. C. A.: Monographie des Élatérides, Tome premier,
Mem. Soc. R. Sci. Liége, 12, 1–400, 1857.
Candèze, E. C. A.: Élatérides nouveaux, Mém. Acad. Belg., 17, 1–
63, 1865.
Cate, P.: Elateridae, in: Catalogue of Palaearctic Coleoptera, edited
by: Löbl, I. and Smetana, A., Volume 4, Apollo Books, Stenstrup,
Denmark, 89–209, 2007.
Cockerell, T. D. A.: Insects in Burmese amber, Ann. Entomol. Soc.
Am., 10, 323–329, 1917.
Costa, C., Lawrence, J. F., and Rosa, S. P.: Elateridae Leach, 1815,
in: Handbook of Zoology, Arthropoda: Insecta; Coleoptera,
Beetles, Volume 2: Morphology and Systematics (Elateroidea,
Bostrichiformia, Cucujiformia partim), edited by: Leschen, R. A.
B., Beutel, R. G., and Lawrence, J. F., Walter de Gruyter GmbH
& Co. KG, Berlin, Germany, 75–103, 2010.
Crowson, R. A.: The phylogeny of Coleoptera, Annu. Rev. Ento-
mol., 5, 111–134, 1960.
Dolin, V. G.: K sistematike mezozoiskikh zhukov-schchelkunov
(Coleoptera, Elateridae), Paleontol. Zhurnal., 9, 51–62, 1975.
Dolin, V. G.: Fossil click beetles (Coleoptera, Elateridae) of the sub-
families Negastriinae and Cardophorinae from Upper Jurassic of
Karatau, Vestn. Zool., 1976, 68–75, 1976.
Dolin, V. G.: Click beetles (Coleoptera, Elateridae) from the Upper
Jurassic of Karatau, in: Fossil Insects of the Mesozoic, edited by:
Dolin, V. G., Panfilov, D. V., Ponomarenko, A. G., and Pritykina,
D. N., Naukova Dumka, Kiev, Ukraine, 17–81, 1980.
Donoghue, P. C. J. and Yang, Z.: The evolution of methods for es-
tablishing evolutionary timescales, Philos. T. Roy. Soc. Lond. B,
371, 20160020, https://doi.org/10.1098/rstb.2016.0020, 2016.
Douglas, H. B., Kundrata, R., Brunke, A. J., Escalona, H. E., Cha-
pados, J. T., Eyres, J., Richter, R., Savard, K., ´
Slipi´
nski, A.,
McKenna, D., and Dettman, J. R.: Anchored Phylogenomics,
Evolution and Systematics of Elateridae: Are All Biolumi-
nescent Elateroidea Derived Click Beetles?, Biology, 10, 451,
https://doi.org/10.3390/biology10060451, 2021.
Emery, C.: Ueber Phylogenie und Systematik der Insekten, Biol.
Zentralbl., 5, 648–656, 1886.
Fleutiaux, E.: Insectes Coléoptères 13, Elateridae, Trixagidae et
Melasidae. in: Voyage de Ch. Alluaud et R. Jeannel en Afrique
Orientale (1911–1912), Résultats Scientifiques, Lhomme, Paris,
119, 1919.
Fleutiaux, E.: Les Élatérides de l’Indo-Chine Française (Cata-
logue raisonné), Deuxième Partie, Encyclopédie Entomologique,
Coléoptères, 3, 120–121, 1928.
Guérin-Méneville, F. E.: Iconographie du règne animal de G. Cu-
vier, ou représentation d’aprés nature de l’une des espèces les
plus remarquables, et souvent non encore figurées, de chaque
genre d’animaux. Avec un texte descriptif mis au courant de la
science. Ouvrage pouvant servir d’atlas a tous les traités de zo-
ologie, Livraison 4, JB Baillière, Paris, pls. 11–12, 1829.
Kundrata, R. and Bocak, L.: Molecular phylogeny reveals the grad-
ual evolutionary transition to soft-bodiedness in click-beetles
and identifies Sub-Saharan Africa as a cradle of diversity for
Drilini (Coleoptera: Elateridae), Zool. J. Linn. Soc., 187, 413–
452, https://doi.org/10.1093/zoolinnean/zlz033, 2019.
Kundrata, R., Gunter, N. L., Janosikova, D., and Bocak, L.:
Molecular evidence for the subfamilial status of Tetralobinae
(Coleoptera: Elateridae), with comments on parallel evolution of
some phenotypic characters, Arthropod. Syst. Phylo., 76, 137–
145, 2018a.
Kundrata, R., Musalkova, M., and Kubaczkova, M.: Anno-
tated catalogue of the click-beetle tribe Dimini (Coleoptera:
Elateridae: Dendrometrinae), Zootaxa, 4412, 1–75,
https://doi.org/10.11646/zootaxa.4412.1.1, 2018b.
Kundrata, R., Kubaczkova, M., Prosvirov, A. S., Douglas, H.
B., Fojtikova, A., Costa, C., Bousquet, Y., Alonso-Zarazaga,
M. A., and Bouchard, P.: World catalogue of the genus-
group names in Elateridae (Insecta, Coleoptera). Part I:
Agrypninae, Campyloxeninae, Hemiopinae, Lissominae,
Oestodinae, Parablacinae, Physodactylinae, Pityobiinae,
Subprotelaterinae, Tetralobinae, ZooKeys, 839, 83–154,
https://doi.org/10.3897/zookeys.839.33279, 2019a.
Kundrata, R., Prosvirov, A. S., Vondracek, D., and Sormova, E.:
Congruence Between Molecular Data and Morphology: Phylo-
genetic Position of Senodoniini (Coleoptera: Elateridae), Insects,
10, 231, https://doi.org/10.3390/insects10080231, 2019b.
Kundrata, R., Packova, G., and Hoffmannova, J.: Fos-
sil genera in Elateridae (Insecta, Coleoptera): A Trias-
sic origin and Jurassic diversification, Insects, 11, 394,
https://doi.org/10.3390/insects11060394, 2020.
Kundrata, R., Packova, G., Prosvirov, A. S., and Hoffmannova, J.:
The Fossil Record of Elateridae (Coleoptera: Elateroidea): De-
scribed Species, Current Problems and Future Prospects, Insects,
12, 286, https://doi.org/10.3390/insects12040286, 2021.
Kusy, D., He, J.-W., Bybee, S. M., Motyka, M., Bi, W.-X., Podsi-
adlowski, L., Li, X.-Y., and Bocak, L.: Phylogenomic relation-
ships of bioluminescent elateroids define the “lampyroid” clade
with clicking Sinopyrophoridae as its earliest member, Syst. En-
tomol., 46, 111–123, https://doi.org/10.1111/syen.12451, 2021.
Leach, W. E.: Entomology, in: Edinburgh Encyclopaedia; Part I,
edited by: Brewster, D., Volume 9, Blackwood, W., Waugh, J.
etc., Edinburgh, UK, 57–172, 1815.
Linnaeus, C.: Systema Naturae per Regna Tria Naturae: Secundum
Classes, Ordines, Genera, Species, Cum Characteribus, Differ-
entiis, Synonymis, Locis, Tomus I, Editio Decima, Reformata,
Laurentius Salvius, Stockholm, Sweden, 1–823, 1758.
Mao, Y., Liang, K., Su, Y., Li, J., Rao, X., Zhang, H., Xia, F., Fu, Y.,
Cai, C., and Huang, D.: Various amberground marine animals on
Burmese amber with discussions on its age, Palaeoentomology,
Foss. Rec., 24, 347–355, 2021 https://doi.org/10.5194/fr-24-347-2021
R. Kundrata et al.: Cretopachyderes gen. nov., a new remarkable click beetle 355
1, 91–103, https://doi.org/10.11646/palaeoentomology.1.1.11,
2018.
Muona, J., Chang, H., and Ren, D.: The clicking Elateroidea from
Chinese Mesozoic deposits (Insecta, Coleoptera), Insects, 11,
875, https://doi.org/10.3390/insects11120875, 2020.
Otto, R. L.: Descriptions of two new elateroid beetles (Coleoptera:
Eucnemidae, Elateridae) from Burmese amber, Insecta Mundi,
702, 1–6, https://doi.org/10.5281/zenodo.3673247, 2019.
Qiu, L., Németh, T., Prosvirov, A., and Kundrata, R.: Dima spi-
cata Schimmel, 1999 (Elateridae: Dimini), a morphologically
remarkable species endemic to China, Zootaxa, 4768, 415–424,
https://doi.org/10.11646/zootaxa.4768.3.8, 2020.
Schenkling, S.: Elateridae I., in: Coleopterorum Catalogus, Pars
80, edited by: Schenkling, S., W. Junk, Berlin, Germany, 1–263,
1925.
Schimmel, R.: Neue und wenig bekannte Elateriden sowie eine
neue Gattung, Acumenator n. gen., aus Südostasien (Insecta:
Coleoptera, Elateridae), Mitt. Pollichia., 85 [1998], 231–259,
1999.
Shi, G., Grimaldi, D. A., Harlow, G. E., Wang, J., Wang, J., Yang,
M., Lei, W., Li, Q., and Li, X.: Age constraint on Burmese am-
ber based on U–Pb dating of zircons, Cret. Res., 37, 155–163,
https://doi.org/10.1016/j.cretres.2012.03.014, 2012.
Stibick, J. N. L.: Classification of the Elateridae (Coleoptera), Rela-
tionships and classification of the subfamilies and tribes, Pacific
Insects, 20, 145–186, 1979.
Toussaint, E. F. A., Seidel, M., Arriaga-Varela, E., Hájek, J., Král,
D., Sekerka, L., and Fikáˇ
cek, M.: The peril of dating beetles,
Syst. Entomol., 42, 1–10, https://doi.org/10.1111/syen.12198,
2017.
Yu, T., Kelly, R., Mu, L., Ross, A., Kennedy, J., Broly, P., Xia, F.,
Zhang, H., Wang, B., and Dilcher, D.: An ammonite trapped in
Burmese amber, P. Natl. Acad. Sci. USA, 116, 11345–11350,
https://doi.org/10.1073/pnas.1821292116, 2019.
https://doi.org/10.5194/fr-24-347-2021 Foss. Rec., 24, 347–355, 2021
