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Bioluminescent beetles of the superfamily Elateroidea (fireflies, fire beetles, glow-worms) are the most speciose group of terrestrial light-producing animals. The evolution of bioluminescence in elateroids is associated with unusual morphological modifications, such as soft-bodiedness and neoteny, but the fragmentary nature of the fossil record discloses little about the origin of these adaptations. We report the discovery of a new bioluminescent elateroid beetle family from the mid-Cretaceous of northern Myanmar (ca 99 Ma), Cretophengodidae fam. nov. Cretophengodes azari gen. et sp. nov. belongs to the bioluminescent lampyroid clade, and would appear to represent a transitional fossil linking the soft-bodied Phengodidae + Rhagophthalmidae clade and hard-bodied elateroids. The fossil male possesses a light organ on the abdomen which presumably served a defensive function, documenting a Cretaceous radiation of bioluminescent beetles coinciding with the diversification of major insectivore groups such as frogs and stem-group birds. The discovery adds a key branch to the elateroid tree of life and sheds light on the evolution of soft-bodiedness and the historical biogeography of elateroid beetles.
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royalsocietypublishing.org/journal/rspb
Research
Cite this article: Li Y-D, Kundrata R, Tihelka
E, Liu Z, Huang D, Cai C. 2021
Cretophengodidae, a new Cretaceous beetle
family, sheds light on the evolution of
bioluminescence. Proc. R. Soc. B 288:
20202730.
https://doi.org/10.1098/rspb.2020.2730
Received: 31 October 2020
Accepted: 17 December 2020
Subject Category:
Palaeobiology
Subject Areas:
evolution, palaeontology
Keywords:
Elateroidea, biogeography, fossil,
bioluminescence, Cretaceous Terrestrial
Revolution
Author for correspondence:
Chenyang Cai
e-mail: cycai@nigpas.ac.cn
Electronic supplementary material is available
online at https://doi.org/10.6084/m9.figshare.
c.5253502.
Cretophengodidae, a new Cretaceous
beetle family, sheds light on the
evolution of bioluminescence
Yan-Da Li1,2, Robin Kundrata3, Erik Tihelka4, Zhenhua Liu5,6, Diying Huang1
and Chenyang Cai1,4
1
State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, and
Centre for Excellence in Life and Palaeoenvironment, Chinese Academy of Sciences, Nanjing 210008,
Peoples Republic of China
2
School of Life Sciences, Peking University, Beijing 100871, Peoples Republic of China
3
Department of Zoology, Faculty of Science, Palacký University, 77900 Olomouc, Czech Republic
4
School of Earth Sciences, University of Bristol, Life Sciences Building, Tyndall Avenue, Bristol BS8 1TQ, UK
5
State Key Laboratory of Biocontrol, School of Life Sciences, Sun Yat-Sen University, Guangzhou 510275,
Peoples Republic of China
6
Australian National Insect Collection, CSIRO National Research Collections Australia, GPO Box 1700, Canberra,
Australian Capital Territory 2601, Australia
Y-DL, 0000-0002-9439-202X; RK, 0000-0001-9397-1030; ET, 0000-0002-5048-5355;
ZL, 0000-0002-2739-3305; DH, 0000-0002-5637-4867; CC, 0000-0002-9283-8323
Bioluminescent beetles of the superfamily Elateroidea (fireflies, fire beetles,
glow-worms) are the most speciose group of terrestrial light-producing animals.
The evolution of bioluminescence in elateroids is associated with unusual
morphological modifications, such as soft-bodiedness and neoteny, but the
fragmentary nature of the fossil record discloses little about the origin of
these adaptations. We report the discovery of a new bioluminescent elateroid
beetle family from the mid-Cretaceous of northern Myanmar (ca 99 Ma),
Cretophengodidae fam. nov. Cretophengodes azari gen.etsp.nov.belongsto
the bioluminescent lampyroid clade, and would appear to represent a transi-
tional fossil linking the soft-bodied Phengodidae + Rhagophthalmidae clade
and hard-bodied elateroids. The fossil male possesses a light organ on the abdo-
men which presumably served a defensive function, documenting a Cretaceous
radiation of bioluminescent beetles coinciding with the diversification of
major insectivore groups such as frogs and stem-group birds. The discovery
adds a key branch to the elateroid tree of life and sheds light on the evolution
of soft-bodiedness and the historical biogeography of elateroid beetles.
1. Introduction
Bioluminescence, the production of light by living organisms, evolved over 30
times independently on diverse branches of the tree of life including unicellular
algae, cnidarians, fishes and marine annelid worms [1]. On land, light-producing
beetles are the most widespread and abundant bioluminescent organisms [2,3];
their elaborate flash displays play a role in mate recognition as well as in
aposematic signalling, communication and luring prey [46]. The majority of
bioluminescent beetles (approx. 2300 species) belong to the megadiverse poly-
phagan superfamily Elateroidea and have been grouped into a single clade by
recent molecular studies: the elaterid-lampyroid clade, i.e. Elateridae (click
beetles, fire beetles), Sinopyrophoridae, Lampyridae (fireflies), Phengodidae
(glow-worm beetles) and Rhagophthalmidae [3,711]. The evolution of biolumi-
nescence in beetles is associated with a number of unusual morphological
modifications such as soft-bodiedness and extremely modified neotenic females
that retain larviform features into adulthood [10,17], but little is known about the
transitions leading to the evolution of these peculiar morphologies and modes of
life since the fossil record of soft-bodied elateroids is exceedingly scarce.
© 2021 The Author(s) Published by the Royal Society. All rights reserved.
Within the bioluminescent lampyroid clade, Phengodidae
and Rhagophthalmidae were traditionally considered to be clo-
sely related based on morphological evidence [18,19], and
indeed, they have been regularly recovered as sister groups
in recent molecular phylogenies [7,9,12,14]. Rhagophthalmidae
is a relatively small group of beetles occurring mainly in the
East Palearctic and Oriental regions, with about 60 species
describedtodate[20].ThemorespeciosePhengodidae
includes nearly 300 species, with the centre of diversity in the
Neotropical region [21], and the isolated subfamily Cydistinae
distributed in West Asia. Although fossil Lampyridae have
been reported from Cretaceous Burmese amber and a few Cen-
ozoic deposits [2225], fossils belonging to the Phengodidae +
Rhagophthalmidae clade remain unknown. Here, we describe
Cretophengodes azari gen. et sp. nov., based on a specimen pre-
served in mid-Cretaceous amber from northern Myanmar. The
peculiar genus is classified in a new family, Cretophengodidae,
belonging to the specialized bioluminescent lampyroid clade.
This fossil possesses an intriguing combination of characters,
shedding light on the early evolution of the lampyroid clade
and the origin of bioluminescence in beetles.
2. Systematic palaeontology
Order Coleoptera Linnaeus, 1758
Suborder Polyphaga Emery, 1886
Superfamily Elateroidea Leach, 1815
Cretophengodidae Li, Kundrata, Tihelka and Cai fam. nov.
Type genus. Cretophengodes gen. nov.
Diagnosis (male). Mandibles slender, sickle-shaped. Fron-
toclypeal region moderately declined anteriorly. Eyes large,
strongly protruding. Antennae 12-segmented; antennomere
1 stout, expanding apically; antennomeres 2 and 3 short;
antennomeres 411 elongate, bipectinate. Prosternum in front
of coxae longer than diameter of procoxal cavity. Prosternal
process narrow and elongate, acute apically, reaching posterior
edge of procoxae. Elytra oblong, sub-parallel sided, nearly com-
pletely covering abdomen, leaving at most only apex of ultimate
tergite exposed. Tarsal formula 5-5-5; tarsomeres 24eachwith
membranous lobe. Abdomen with six apparently immovable
ventrites; photic organ present on median portions of the
basal three abdominal ventrites.
Composition and distribution. Monogeneric, with Creto-
phengodes gen. nov. known only from mid-Cretaceous
Burmese amber.
Cretophengodes Li, Kundrata, Tihelka and Cai gen. nov.
Type species. Cretophengodes azari sp. nov., here designated.
Etymology. The generic name is derived partly from
Cretaceous, in reference to the age of the fossil, and the
genus Phengodes, the type genus of the morphologically simi-
lar and presumably closely related Phengodidae. The gender
is masculine.
Diagnosis. As for the family with additional characters:
body moderate (approx. 7.3 mm long); pronotum sub-pentago-
nal, wider than long; elytra irregularly punctate, with several
raised interstrial intervals forming indistinct carinae; claws
simple.
Cretophengodes azari Li, Kundrata, Tihelka and Cai sp. nov.
Figures 1a,band 2.
Etymology. After Prof. Dany Azar, palaeoentomologist
extraordinaire.
(a)(b)(e)(f)
(c)(d)
Figure 1. General habitus of Cretophengodidae and representatives of the closely related Phengodidae and Rhagophthalmidae, under incident light.
(a,b)Cretophengodes azari gen. et sp. nov., dorsal and ventral views, respectively, with arrowhead showing the photic organ. (c,d)Zarhipis sp. (Phengodidae),
dorsal and ventral views, respectively. (e,f)Rhagophthalmus sp. (Rhagophthalmidae), dorsal and ventral views, respectively. Scale bars: (a,b,e,f) 2 mm;
(c,d) 4 mm. (Online version in colour.)
royalsocietypublishing.org/journal/rspb Proc. R. Soc. B 288: 20202730
2
Type material. Holotype, NIGP173775, male, mid-Cretac-
eous (upper Albian to lower Cenomanian [26,27]), from amber
mine near Noije Bum Village, Hukawng Valley, Tanai Township,
Myitkyina District, Kachin State, northern Myanmar.
Diagnosis. As for the genus.
Description. For a full description of Cretophengodes azari
gen. et sp. nov., refer to the electronic supplementary material.
3. Discussion
(a) Systematic placement of Cretophengodes
Cretophengodes possesses a unique combination of characters
within Elateroidea, including antennae with 12 antenno-
meres, which are bipectinate from antennomere 4, a
relatively long prosternal process, and six abdominal ven-
trites in males (figure 2ce). Within Elateroidea, only a few
groups have 12-segmented bipectinate (or bilamellate) anten-
nae, including Asiopsectra (Brachypsectridae), Elateridae,
Rhagophthalmidae and most Phengodidae. Although some
other lineages like Lampyridae and Eucnemidae also contain
species with bipectinate antennae, these do not consist of 12
antennomeres [28]. The antennae of Asiopsectra and Elateri-
dae are more lamellate, relatively flat and short [29], while
in Cretophengodes the rami are much longer and more slender.
In Rhagophthalmidae, the antennomere 3 is longer than
antennomere 2, and possesses rami if the antennae are bipec-
tinate (electronic supplementary material, figure S2) [30]. On
the contrary, Phengodidae, as well as Cretophengodes,have
antennomere 3 short and simple, and the long and slender
rami begin from antennomere 4 [30,31]. Just like males of
Phengodidae and Rhagophthalmidae, Cretophengodes pos-
sesses a putative photic organ on the abdominal ventrites
which is preserved as a white foamy layer on abdominal ven-
trites 13 (figure 1b). The region of the abdomen bearing the
light organ corresponds well to the appearance of photic
organs in modern Phengodidae and Rhagophthalmidae
males (e.g. fig. 8 in [31]).
(a)(c)
(d)
(b)
(e)(f)
(g)(h)
Figure 2. Details of Cretophengodes azari gen. et sp. nov., under epifluorescence. (a) Head, dorsal view. (b) Head, ventral view. (c) Prothorax, ventral view.
(d) Abdomen, ventral view. (e) Antenna, dorsal view. ( f) Mesothorax, ventral view. (g) Elytra, dorsal view. (h) Prothorax, dorsal view. Abbreviations: a14,12,
antennomeres 14,12; an, antenna; el, elytron; ey, compound eye; lbp, labial palp; md, mandible; msc, mesocoxa; msv, mesoventrite; mtf, metafemur; mttb,
metatibia; mtts, metatarsus; mtv, metaventrite; mxp, maxillary palp; pf, profemur; pn, pronotum; ps, prosternum; ptb, protibia; pts, protarsus; sc, scutellum;
v16, ventrites 16. Scale bars: 500 µm. (Online version in colour.)
royalsocietypublishing.org/journal/rspb Proc. R. Soc. B 288: 20202730
3
Morphologically, Cretophengodes resembles extant Phen-
godidae (figure 1c) in its head structures (including
antennae and sickle-shaped mandibles) and pronotal shape
(figure 2b). However, other characters of Cretophengodes are
distinctly different from extant Phengodidae. The position
of the abdominal photic organ medially at the basal abdomi-
nal ventrites is more similar to the arrangement in the
rhagophthalmid genus Diplocladon than to any species in
Phengodidae. The long prosternal process of Cretophengodes
is typical of well-sclerotized elateroids, in which it usually
reaches at least to the posterior edge of procoxae (figure 2c).
By contrast, most soft-bodied elateroids, including Phengodi-
dae, have an incomplete or reduced prosternal process
[32,33]. Additionally, the prosternum in front of the coxae is
very short in most Phengodidae and Rhagophthalmidae,
often forming a very narrow transverse bar (figure 1d,f),
whereas in Cretophengodes the prosternum in front of coxae
is not shortened, being longer than the shortest diameter of
the procoxae (figure 2c). The non-shortened prosternum is
present only in the Asian Cydistinae, which, however, have
a considerably reduced prosternal process [30]. All extant
lineages of Phengodidae share the frontal partly membranous
and reduced mesoventrite, whereas in Cretophengodes it is
well-developed and sclerotized. Similarly, the elytra are
slightly to extremely shortened in Phengodidae and some
Rhagophthalmidae, exposing several to all abdominal ter-
gites, whereas in Cretophengodes, the tergites are nearly
completely covered by the elytra, with only the apical portion
of last visible tergite exposed, which may, however, be caused
by compression during fossilization (for similar preservation,
see [34]). An additional character distinguishing Cretophen-
godes from Phengodidae is the number of abdominal
ventrites. This character has been widely used for differentiat-
ing various elateroid families, and varies from five connate
ventrites in some hard-bodied groups to seven or eight free
ventrites connected by extensive membranes in the soft-
bodied families [33]. In the male of Cretophengodes, the six
ventrites are not connected by membranes, giving the abdo-
men a rather compact appearance (figures 1band 2d).
Males of Phengodidae and Rhagophthalmidae, however,
have eight very freely connected ventrites, such that the
abdomen is capable of considerable stretching.
The unique combination of characters in Cretophengodes is
unknown in any currently defined beetle lineage. Therefore,
Cretophengodes deserves family status in Elateroidea, and
due to shared morphological features with Phengodidae,
we hypothesize its position in the previously defined lampyr-
oid clade [79]. Taking into account its divergent
morphology, shared characters with Rhagophthalmidae and
the hard-bodied elateroids, and the dramatic change that
would necessitate for the morphological diagnosis of Phen-
godidae, we prefer to accommodate the new genus as a
separate family. We hypothesize that Cretophengodidae
probably represents a stem group of Phengodidae + Rha-
gophthalmidae (figure 3; electronic supplementary material).
(b) Evolution of beetle bioluminescence: timing
and drivers
Beetles of the lampyroid clade (Sinopyrophoridae, Phengodi-
dae, Rhagophthalmidae and Lampyridae) are rare in the
fossil record. The earliest fossil belonging to the group is a lam-
pyrid from Burmese amber that possesses an abdominal photic
organ [25], while no fossils belonging to the Phengodidae +
Rhagophthalmidae clade have been reported previously. Our
discovery of Cretophengodidae confirms that beetles of the bio-
luminescent lampyroid clade begun to diversify by at least the
mid-Cretaceous, during a period of biotic turnover associated
with the radiation of angiosperms known as the Cretaceous
Terrestrial Revolution (KTR) [35]. This helps to narrow the
previous molecular clock estimates of the origin of the Phengo-
didae+ Rhagophthalmidae clade, which range from the Late
Cretaceous to Palaeogene (95% CI from 116 to 59 Ma [8]).
In elateroids, bioluminescence is more widespread in the
immature stages than in adults [19,36]. Larvae of bioluminescent
groups emit light when disturbed or glow spontaneously in
slow pulses or continuously at night [5,37]. Glowing larvae are
avoided by some predators [4], leading some workers to postu-
late that bioluminescencefirst evolved as a defensive mechanism
and was only later co-opted as a mating and communication
150 125 100 75 50 25
0
Ma
Elateridae
Sinopyrophoridae
Lampyridae
Phengodidae
Rhagophthalmidae
Jur. Cretaceous Palaeogene Neo.
lampyroid clade
1
2
3
4
5
†Cretophengodidae
Q.
Figure 3. Hypothesized phylogenetic relationships within the lampyroid clade, with Cretophengodidae as sister to Phengodidae + Rhagophthalmidae. Node 1:
frontoclypeus strongly and abruptly declined, lateral portion of prosternum 0.52.0 times as long as the length of procoxal cavity. Node 2: bioluminescence.
Node 3: clicking mechanism lost, soft-bodied, females neotenic (exceptions in some Lampyridae). Node 4: head never fully concealed from above by pronotum,
bipectinate or bilamellate 12-segmented antennae (exceptions in some Rhagophthalmidae and Phengodidae), labral apex emarginate. Node 5: prosternum in front
of procoxae very short (exceptions in Phengodidae: Cydistinae), prosternal process incomplete or reduced, males with eight abdominal ventrites, abdominal ventrites
separated by membranes. Abbreviations: Jur., Jurassic; Neo., Neogene; Q., Quaternary. Tree topology after Kusý et al. [9]. (Online version in colour.)
royalsocietypublishing.org/journal/rspb Proc. R. Soc. B 288: 20202730
4
signal in adults [5,19,36,38,39]. The function of bioluminescence
in adults of Phengodidae and Rhagophthalmidae, which pro-
duce a range of colours from green to red, is poorly
understood. Light production in southern American Phengodi-
dae adults does not appear to play a role in searching for mates,
since a bioluminescent display only occurs during mating, so a
defensive function seems likely [40]. Likewise, Rhagophthalmus
males (figure 1e) are luminescent only shortly after eclosion
and so light production is not used in courtship [41]. This
indicates that Cretophengodidae adults may have produced
light primarily for defence, although we cannot rule out the
possibility that it also functioned as a mating signal.
Today, major predators of soft-bodied elateroids include
invertebrate groups such as spiders and ants as well as ver-
tebrates such as frogs and birds [4244]. Crucially, several
of these groups may have initially driven the evolution of bio-
luminescence in beetles diversifying during the KTR. The
earliest ants are known from Cretaceous amber deposits and
stem-group ants co-occurred with Cretophengodes in the
Burmese amber forest [45,46]. Major clades of frogs (Anura)
originated in the Early Cretaceous and the group became
morphologically diverse during this period [47,48]. Cretaceous
stem-group birds (Enantiornithes) included insectivores
specialized for feeding on soft-bodied arthropods [49,50],
and are also known from Burmese amber [51,52]. These visu-
ally hunting predators may have provided a strong selection
pressure favouring the evolution of defensive bioluminescence
in elateroids during the Cretaceous.
(c) Biogeography of the Phengodidae +
Rhagophthalmidae clade
Extant species of Rhagophthalmidae occur mainly in eastern
and southeastern Asia [53]. Phengodidae, however, display a
disjunct distribution pattern. The phengodid subfamily Cydis-
tinae is known exclusively from West Asia [30], while the
remaining and more speciose phengodid subfamilies Phengo-
dinae and Mastinocerinae occur throughout the New World
from southern Canada to Chile ( figure 4) [31,54,55].
Our discovery of Cretophengodes, the first fossil putatively
belonging to the Phengodidae + Rhagophthalmidae clade,
supports the hypothesis that the clade originated in the Old
World. Under this scenario, the diversity of Phengodidae in
the New World resulted from intercontinental dispersal and
subsequent local diversification. Both Rhagophthalmidae
and Phengodidae are neotenic, with adult females retaining
a larviform morphology, which contributes to their extremely
limited dispersal ability [56]. Our finding nevertheless
would demonstrate that neotenic lineages are capable of
long-distance dispersal over long geological time scales.
Kusý et al. [9] suggested that the lampyroid clade originated
in eastern Laurasia, citing an unpublished Phengodidae fossil in
Burmese amber as supporting evidence. However, this appears
to be problematic in light of our new discovery as well as the
palaeobiogeography of Burmese amber fossils. Although
historically the Burma Terrane was considered as part of Laur-
asia [57], the biota of Burmese amber indicates a Gondwanan
affinity of the Burma Terrane [58]. A recent palaeomagnetic
study further confirmed that the Burma Terrane was probably
an isolated island at near-equatorial latitude during the mid-
Cretaceous [59]. Hence, the possible presence of Phengodidae
fossils in Burmese amber cannot be regarded as supporting
evidence of a Laurasian origin of the lampyroid clade.
(d) Evolution of soft-bodiedness in Elateroidea
Elateroids were traditionally divided into two major groups,
the strongly sclerotized Elateroidea sensu stricto and the
soft-bodied Cantharoidea [14,15]. Recent molecular studies,
however, firmly reject the monophyly of Cantharoidea [4,5,8],
indicating that the shared morphological traits of soft-bodied
elateroid beetles were acquired several times independently
in their evolutionary history [8,9]. Moreover, the differences
between hard- and soft-bodied elateroids are often not evident.
Instead, there is a continuum from fully sclerotized lineages to
extremely soft-bodied ones with neotenic forms [12].
The newly discovered male specimen of Cretophengodes is a
good example documenting such gradual change in the
Rhagophthalmidae
extant
Cretophengodes
mid-Cretaceous
Cydistinae
extant
Phengodinae + Mastinocerinae
extant
Figure 4. Geographical distribution of Cretophengodidae (genus Cretophengodes), Phengodidae (subfamilies Cydistinae, Mastinocerinae and Phengodinae) and
Rhagophthalmidae. World map adapted from Natural Earth (NaturalEarthData.com). (Online version in colour.)
royalsocietypublishing.org/journal/rspb Proc. R. Soc. B 288: 20202730
5
evolution towards soft-bodiedness in the Phengodidae +
Rhagophthalmidae lineage. Since both Phengodidae and
Rhagophthalmidae have extremely soft body cuticles and
variously modified morphologies tied with neoteny, Cretophen-
godes represents an intermediate form between hard- and soft-
bodied groups. In well-sclerotized elateroids, the prosternal
process is complete and fits into a deep mesoventral cavity
[32], forming a clicking mechanism that enables beetles lying
on their dorsal side to jump and land on their legs or escape pre-
dators. In most soft-bodied groups, such as Phengodidae
and Rhagophthalmidae, the prosternal process is reduced or
completely absent [53,54]. However, the prosternum of male
Cretophengodes is well-developed, with the prosternal process
elongate and narrow, although it probably does not reach the
mesoventral cavity. Besides, most well-sclerotized elateroids
have five abdominal ventrites (sternites IIIVII), with usually at
least the basal four connate. By contrast, most soft-bodied elater-
oids usually have seven or eight free ventrites, so that the
sternites II, VIII and IX are also exposed [33]. Due to the presence
of well-developed membranes between the individual segments,
soft-bodied elateroids have very flexible abdomens, capable of
twistingandstretchingthataresometimesextendbeyondthe
elytra. The six ventrites in male Cretophengodes do not seem to
be flexible, representing an intermediate stage. The presence of
six ventrites in males was previously known only in the elaterid
tribe Cebrionini [60]. Notably, in the recently described South
American family Jurasaidae, another elateroid family that com-
bines characters of hard- and soft-bodied lineages, males have
only five but freely connected ventrites [16], which indicates
that the increase of ventrite number and disconnection of indi-
vidual ventrites occurred in different soft-bodied lineages in
different sequences during the evolution of the group.
The females of Cretophengodidae remain unknown.
However, given that all known females of their presumable
relatives (i.e. Phengodidae and Rhagophthalmidae) are
more or less larviform [53,54], it is plausible that female
Cretophengodidae were neotenic as well (figure 5). The
neotenic adult females may be easily misidentified as larvae
and therefore ignored by researchers unfamiliar with this
particular group. Even in extant neotenic groups, females
are much less represented in collections or are not known
at all [15,30,61]. Therefore, more attention should be paid to
larviform beetle fossils, which could be crucial for improving
our understanding of the evolution of soft-bodiedness
in beetles.
4. Material and methods
The Burmese amber specimen studied here originates from
amber mines near the Noije Bum Hill (26°20N, 96°36E),
Hukawng Valley, Kachin State, northern Myanmar. The speci-
men is deposited in the Nanjing Institute of Geology and
Palaeontology (NIGP), Chinese Academy of Sciences at Nanjing,
China. The amber piece was trimmed with a small table saw,
ground with emery papers of different grit sizes, and finally
polished with polishing powder.
Photographs under incident light were taken with a Zeiss
Discovery V20 stereo microscope. Widefield fluorescence images
were captured with a Zeiss Axio Imager 2 light microscope com-
bined with a fluorescence imaging system. Confocal images
were obtained with a Zeiss LSM710 confocal laser scanning micro-
scope. Images under incident light and widefield fluorescence
were stacked in Helicon Focus 7.0.2 or Zerene Stacker 1.04. Confo-
cal images were manually stacked in Adobe Photoshop CC.
Images were further processed in Adobe Photoshop CC to
enhance contrast. The classification of Elateroidea follows Kun-
drata et al. [12] and Kusý et al. [9], the classification and limits of
Phengodidae follow Kundrata et al. [30] and morphological
terminology is adapted from Costa & Zaragoza-Caballero [54].
Nomenclatural acts. This published work and the nomenclatural acts have
been registered in ZooBank. The LSID for this publication is urn:lsid:
zoobank.org:pub:FB7D9187-835C-4B32-86DE-A6B2E37A8812.
Data accessibility. The supplementary descriptions and figures support-
ing this article have been uploaded as the electronic supplementary
material. The original series of confocal slices are available on
Zenodo repository (doi:10.5281/zenodo.4313770).
Authorscontributions. C.C. and Y.-D.L. conceived the study, Y.-D.L., Z.L.
and R.K. processed the photomicrograph data. Y.-D.L., E.T. and C.C.
drafted the manuscript, to which R.K., D.H. and Z.L. contributed.
Y.-D.L., C.C., R.K. and E.T. interpreted data.
Competing interests. The authors declare no competing interests.
Funding. This work has been supported by the Strategic Priority
Research Program of the Chinese Academy of Sciences (grant nos.
XDB26000000, XDB18000000), the National Natural Science Foun-
dation of China (grant nos. 41672011, 41688103), and the Second
Tibetan Plateau Scientific Expedition and Research project (grant
no. 2019QZKK0706).
Acknowledgements. We are grateful to Yan Fang for technical help with
confocal imaging, Dinghua Yang for the artistic reconstruction, and
Jyrki Muona and Luiz F. L. da Silveirafor discussion on the antennal mor-
phology of Eucnemidae and Lampyridae, respectively. We also thank the
editors and three anonymous reviewers for their helpful comments.
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Figure 5. Artistic reconstruction of Cretophengodes azari gen. et sp. nov. The
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... The number of genera included in Rhagophthalmidae and also their placement within Elateroidea classification vary by source (e.g., McDermott 1966;Crowson 1972;Lawrence and Newton 1995;Kawashima et al. 2010;Kundrata and Bocak 2011a). In the last decade, Elateroidea systematic research has accelerated and the classification of the superfamily has experienced many taxonomic changes (e.g., Kundrata et al. 2014Bocak et al. 2018;Kusy et al. 2018bKusy et al. , 2021, including the discoveries of two new recent families 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). ...
... Kusy et al. (2021) defined the "lampyroid clade", which contains Lampyridae, Phengodidae, Rhagophthalmidae, and Sinopyrophoridae. Fossil Cretophengodidae were probably also a part of that clade (Li et al. 2021b). ...
... Cretophengodidae were described from mid-Cretaceous amber of northern Myanmar (ca. 99 Mya, Shi et al. 2012;Li et al. 2021b), and Kusy et al. (2021) reported unpublished Phengodidae from the same deposit. Kusy et al. (2021) summarized and reviewed the published molecular dating analyses of the elaterid-lampyroid clade, and showed that median estimates suggest the split of the Lampyridae, Phengodidae, and Rhagophthalmidae clade in the mid-Cretaceous. ...
Article
Full-text available
Rhagophthalmidae are a small beetle family known from the eastern Palaearctic and Oriental realms. Rhagophthalmidae are closely related to railroad worms (Phengodidae) and fireflies (Lampyridae) with which they share highly modified paedomorphic females and the ability to emit light. Currently, Rhagophthalmidae include 66 species classified in the following 12 genera: Bicladodrilus Pic, 1921 (two spp.), Bicladum Pic, 1921 (two spp.), Dioptoma Pascoe, 1860 (two spp.), Diplocladon Gorham, 1883 (two spp.), Dodecatoma Westwood, 1849 (eight spp.), Falsophrixothrix Pic, 1937 (six spp.), Haplocladon Gorham, 1883 (two spp.), Menghuoius Kawashima, 2000 (three spp.), Mimoochotyra Pic, 1937 (one sp.), Monodrilus Pic, 1921 (two spp. in two subgenera), Pseudothilmanus Pic, 1918 (two spp.), and Rhagophthalmus Motschulsky, 1854 (34 spp.). The replacement name Haplocladon gorhami Kundrata, nom. nov. is proposed for Diplocladon hasseltii Gorham, 1883b (described in subgenus Haplocladon) which is preoccupied by Diplocladon hasseltii Gorham, 1883a. The genus Reductodrilus Pic, 1943 is tentatively placed in Lampyridae: Ototretinae. Lectotypes are designated for Pseudothilmanus alatus Pic, 1918 and P. marginalis Pic, 1918. Interestingly, in the eastern part of their distribution, Rhagophthalmidae have remained within the boundaries of the Sunda Shelf and the Philippines demarcated by the Wallace Line, which separates the Oriental and Australasian realms. This study is intended to be a first step towards a comprehensive revision of the group on both genus and species levels. Additionally, critical problems and prospects for rhagophthalmid research are briefly discussed.
... The mesmerizing dances of fireflies on warm summer nights have fascinated generations of our ancestors and sparked centuries of scientific enquiry. With approximately 2500 described species [1], the fireflies (Lampyridae) are by far the most widespread and abundant bioluminescent beetles, sharing their ability to produce light with five related elateroid families-some click beetles (Elateridae) from the Neotropical region and small Melanesian islands, Sinopyrophoridae (Sinopyrophorus Bi & Li) endemic to south China, predominantly New World glow-worms (Phengodidae), the Old World star-worms (Rhagophthalmidae) and the recently described fossil family Cretophengodidae [2][3][4][5][6]. Bioluminescence in fireflies is known to fulfil a dual functionin communication [7] and as an aposematic antipredator mechanism [8]. ...
... Although it had no apparent light organ, we can hypothesize that the species was bioluminescent like all known extant phengodid species, at least as immature stages and larviform females. Li et al. [4] described a transitional bioluminescent elateroid beetle family (Cretophengodidae, represented by Cretophengodes azari Li, Kundrata, Tihelka & Cai) from the same amber deposit. It belongs to the bioluminescent lampyroid clade, representing a transitional fossil linking the soft-bodied Phengodidae+Rhagophthalmidae clade and the primitively hard-bodied elateroids. ...
Article
Full-text available
The beetle superfamily Elateroidea comprises the most biodiverse bioluminescent insects among terrestrial light-producing animals. Recent exceptional fossils from the Mesozoic era and phylogenomic studies have provided valuable insights into the origin and evolution of bioluminescence in elateroids. However, due to the fragmentary nature of the fossil record, the early evolution of bioluminescence in fireflies (Lampyridae), one of the most charismatic lineages of insects, remains elusive. Here, we report the discovery of the second Mesozoic bioluminescent firefly, Flammarionella hehaikuni Cai, Ballantyne & Kundrata gen. et sp. nov., from the Albian/Cenomanian of northern Myanmar (ca 99 Ma). Based on the available set of diagnostic characters, we interpret the specimen as a female of stem-group Luciolinae. The fossil possesses deeply impressed oval pits on the apices of antennomeres 3–11, representing specialized sensory organs likely involved in olfaction. The light organ near the abdominal apex of Flammarionella resembles that found in extant light-producing lucioline fireflies. The growing fossil record of lampyrids provides direct evidence that the stunning light displays of fireflies were already established by the late Mesozoic.
... clade (Li, Kundrata, Tihelka, et al., 2021). Fossils of soft-bodied 'lampyroids' are rare in the Cretaceous fossil record. ...
... Almost 60 species classified in 13 genera have been described from burmite, and single species from Spanish amber, Azerbaijan Agdzhakend amber, and the Koonwarra Fossil Bed in Australia (e.g., Fanti & Müller, 2022;Hsiao et al., 2021;Qu et al., 2023). Li, Kundrata, Packova, et al. (2021) Deng et al. (2019), and several more species followed (e.g., Li, Kundrata, Tihelka, et al., 2021;. ...
Article
Full-text available
Recent progress in beetle palaeontology has incited us to re-address the evolutionary history of the group. The Permian †Tshekardocoleidae had elytra that covered the posterior body in a loose tent-like manner. The formation of elytral epipleura and a tight fit of elytra and abdomen were important evolutionary transformations in the Middle Permian, resulting in a tightly enclosed subelytral space. Permian families were likely associated with dead wood of gymnospermous trees. The end-Permian extinction event resulted in a turnover in the composition of beetle faunas, especially a decline of large-bodied wood-associated forms. Adephaga and Myxophaga underwent a first wave of diversification in the Triassic. Polyphaga are very rare in this period. The first wave of diversification of this suborder occurs in the Jurassic, with fossils of Elateriformia, Staphyliniformia and Cucujiformia. The Cretaceous fossil record has been tremendously enriched by the discovery of amber inclusions. Numerous fossils represent all major polyphagan lineages and also the remaining suborders. Improved analytical methods for documenting and placing extinct taxa are discussed. Different factors have played a role in the diversification of beetles. The enormous number of species associated with flowering plants, and timing and patterns of diversification in phytophagous lineages indicate that the angio-sperm radiation played a major role in beetle macroevolution. Moreover, the evolution of intimate partnerships with symbionts and the acquisition of novel genes-obtained from fungi and bacteria via horizontal gene transfers-facilitated the use of plant material as a food source and were key innovations in the diversification of plant-feeding beetles.
... Several soft-bodied families, however, lack any fossil record at all. It is interesting, though, that from three described extinct families (with unresolved affinities within the superfamily), one is soft-bodied (Berendtimiridae; Winkler, 1987) and remaining two represent some intermediate stages (Mysteriomorphidae and Cretophengodidae;Alekseev &Ellenberger, 2019 andPeris et al., 2020 for the first, Li et al., 2021a for the latter). The genus Anoeuma Li et al., 2021 from Burmese amber, placed as incertae sedis within the superfamily Elateroidea, is also soft-bodied (Li et al., 2021b). ...
... The fossil family Cretophengodidae, which is supposedly closely related to Phengodidae and Rhagophthalmidae, is known exclusively from Burmese amber (Li et al., 2021a). The fact that Cretocydistus gen. ...
Article
Elateroidea superfamily holds a huge diversity, morphological variation and a myriad of habitat specializations. The presence of bioluminescence and paedomorphosis renders the group as an interesting model for several studies. The “lampyroid” clade manifests both features, in a variety of light displays and body-forms, but the small fossil records hampers any advance in understanding the origin of these characteristics, as well as the biogeographic history of the group. We present here the description of a new fossil species, Cretocydistus wittmeri gen. et sp. nov. from the mid-Cretaceous of northern Myanmar, the first fossil of the family Phengodidae. We putatively place the genus in the subfamily Cydistinae, which extant species are distributed in Asia Minor, the Levant, and Iran. We also discuss how the discovery of this fossil taxa influences the study of the family and the “lampyroid” clade evolutionary history and biogeography.
... Since the Burma Terrane was likely an isolated island for large periods of time, numerous taxa were probably also island endemics. This hypothesis is confirmed by numerous taxa being known exclusively from Burmese amber, including the insect families from various orders [90][91][92][93][94][95] . ...
Article
Full-text available
The click beetles (Elateridae) represent the major and well-known group of the polyphagan superfamily Elateroidea. Despite a relatively rich fossil record of Mesozoic Elateridae, only a few species are described from the Upper Cretaceous Burmese amber. Although Elateridae spend most of their lives as larvae, our knowledge on immature stages of this family is limited, which is especially valid for the fossils. So far, only a single larval click beetle has been reported from Burmese amber. Here, we describe two larval specimens from the same deposit which based on their morphology unambiguously belong to the predominantly Southern Hemisphere subfamily Pityobiinae, being the most similar to the representatives of tribe Tibionemini. However, since the larvae of the closely related bioluminescent Campyloxenini have not yet been described, we place our specimens to Tibionemini only tentatively. One species of Pityobiinae was recently described from Burmese amber based on adults, and we discuss if it can be congeneric with the here-reported larvae. Recent representatives of the Tibionemini + Campyloxenini clade are known from South America and New Zealand, and this group is hypothesized to have a Gondwanan origin. Hence, the newly discovered Burmese amber larvae may further contribute to a recently highly debated hypothesis that biota of the resin-producing forest on the Burma Terrane, which was probably an island drifting northward at the time of amber deposition, had at least partly Gondwanan affinities. The discovery of enigmatic click beetle larvae in the Upper Cretaceous Burmese amber sheds further light on the palaeodiversity and distribution of the relatively species-poor Gondwanan clade of click beetles, which contain a recent bioluminescent lineage, as well as on the taxonomic composition of the extinct Mesozoic ecosystem.
... This clade was originally proposed to accommodate Lampyridae, Phengodidae, Rhagophthalmidae, and a recently described enigmatic bioluminescent genus, Sinopyrophorus Bi & Li in Bi et al. (2019), which was originally placed in Elateridae (Bi et al. 2019) but was treated as a separate family by Kusy et al. (2021). Later, Li et al. (2021) suggested that the newly described Mesozoic Cretophengodidae also belonged to the 'lampyroid clade' , hypothesizing its close relationships with Phengodidae and Rhagophthalmidae. Given that the interrelationships between the Elateridae and 'lampyroid' group of families are not yet fully understood (Douglas et al. 2021, Kusy et al. 2021, further phylogenomic studies are needed with an expanded taxon sampling of all above-mentioned families. ...
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Phengodidae (Coleoptera: Elateroidea), commonly known as glowworm beetles, are a small family of bioluminescent and paedomorphic beetles. There are few phylogenetic studies of Phengodidae, and these are mostly discordant, especially when comparing morphology-based and molecular-based phylogenetic hypotheses. Here, we used the anchored hybrid enrichment approach to undertake the first phylogenomic analysis of Phengodidae (≤358 loci and 39 taxa) and evaluate the higher-level classification of the group. In agreement with previous molecular studies, we recovered Phengodidae as sister to Rhagophthalmidae, and the Old World Cydistinae as sister to all New World Phengodidae. In contrast to previous hypotheses, both Phengodinae and Mastinocerinae were each recovered as monophyletic. Cenophengus was found to be sister to Mastinocerinae, in contrast to some previous hypotheses that placed it as sister to all New World Phengodidae. Considering its morphological divergence, we here establish Cenophenginae subfam. nov. Despite the largest and most comprehensive sampling of Phengodidae in any molecular-based study to date, we had only limited success in revealing the relationships among genera within the most species-rich subfamily, Mastinocerinae. Further studies should focus on the phylogeny and classification of this taxonomically neglected subfamily, on the phylogenetic placement of enigmatic Penicillophorinae, and on seeking morphological support for the main clades of Phengodidae.
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More than 4700 nominal family-group names (including names for fossils and ichnotaxa) are nomenclaturally available in the order Coleoptera. Since each family-group name is based on the concept of its type genus, we argue that the stability of names used for the classification of beetles depends on accurate nomenclatural data for each type genus. Following a review of taxonomic literature, with a focus on works that potentially contain type species designations, we provide a synthesis of nomenclatural data associated with the type genus of each nomenclaturally available family-group name in Coleoptera. For each type genus the author(s), year of publication, and page number are given as well as its current status (i.e., whether treated as valid or not) and current classification. Information about the type species of each type genus and the type species fixation (i.e., fixed originally or subsequently, and if subsequently, by whom) is also given. The original spelling of the family-group name that is based on each type genus is included, with its author(s), year, and stem. We append a list of nomenclaturally available family-group names presented in a classification scheme. Because of the importance of the Principle of Priority in zoological nomenclature, we provide information on the date of publication of the references cited in this work, when known. Several nomenclatural issues emerged during the course of this work. We therefore appeal to the community of coleopterists to submit applications to the International Commission on Zoological Nomenclature (henceforth “Commission”) in order to permanently resolve some of the problems outlined here. The following changes of authorship for type genera are implemented here (these changes do not affect the concept of each type genus): CHRYSOMELIDAE: Fulcidax Crotch, 1870 (previously credited to “Clavareau, 1913”); CICINDELIDAE: Euprosopus W.S. MacLeay, 1825 (previously credited to “Dejean, 1825”); COCCINELLIDAE: Alesia Reiche, 1848 (previously credited to “Mulsant, 1850”); CURCULIONIDAE: Arachnopus Boisduval, 1835 (previously credited to “Guérin-Méneville, 1838”); ELATERIDAE: Thylacosternus Gemminger, 1869 (previously credited to “Bonvouloir, 1871”); EUCNEMIDAE: Arrhipis Gemminger, 1869 (previously credited to “Bonvouloir, 1871”), Mesogenus Gemminger, 1869 (previously credited to “Bonvouloir, 1871”); LUCANIDAE: Sinodendron Hellwig, 1791 (previously credited to “Hellwig, 1792”); PASSALIDAE: Neleides Harold, 1868 (previously credited to “Kaup, 1869”), Neleus Harold, 1868 (previously credited to “Kaup, 1869”), Pertinax Harold, 1868 (previously credited to “Kaup, 1869”), Petrejus Harold, 1868 (previously credited to “Kaup, 1869”), Undulifer Harold, 1868 (previously credited to “Kaup, 1869”), Vatinius Harold, 1868 (previously credited to “Kaup, 1869”); PTINIDAE: Mezium Leach, 1819 (previously credited to “Curtis, 1828”); PYROCHROIDAE: Agnathus Germar, 1818 (previously credited to “Germar, 1825”); SCARABAEIDAE: Eucranium Dejean, 1833 (previously “Brullé, 1838”). The following changes of type species were implemented following the discovery of older type species fixations (these changes do not pose a threat to nomenclatural stability): BOLBOCERATIDAE: Bolbocerus bocchus Erichson, 1841 for Bolbelasmus Boucomont, 1911 (previously Bolboceras gallicum Mulsant, 1842); BUPRESTIDAE: Stigmodera guerinii Hope, 1843 for Neocuris Saunders, 1868 (previously Anthaxia fortnumi Hope, 1846), Stigmodera peroni Laporte & Gory, 1837 for Curis Laporte & Gory, 1837 (previously Buprestis caloptera Boisduval, 1835); CARABIDAE: Carabus elatus Fabricius, 1801 for Molops Bonelli, 1810 (previously Carabus terricola Herbst, 1784 sensu Fabricius, 1792); CERAMBYCIDAE: Prionus palmatus Fabricius, 1792 for Macrotoma Audinet-Serville, 1832 (previously Prionus serripes Fabricius, 1781); CHRYSOMELIDAE: Donacia equiseti Fabricius, 1798 for Haemonia Dejean, 1821 (previously Donacia zosterae Fabricius, 1801), Eumolpus ruber Latreille, 1807 for Euryope Dalman, 1824 (previously Cryptocephalus rubrifrons Fabricius, 1787), Galeruca affinis Paykull, 1799 for Psylliodes Latreille, 1829 (previously Chrysomela chrysocephala Linnaeus, 1758); COCCINELLIDAE: Dermestes rufus Herbst, 1783 for Coccidula Kugelann, 1798 (previously Chrysomela scutellata Herbst, 1783); CRYPTOPHAGIDAE: Ips caricis G.-A. Olivier, 1790 for Telmatophilus Heer, 1841 (previously Cryptophagus typhae Fallén, 1802), Silpha evanescens Marsham, 1802 for Atomaria Stephens, 1829 (previously Dermestes nigripennis Paykull, 1798); CURCULIONIDAE: Bostrichus cinereus Herbst, 1794 for Crypturgus Erichson, 1836 (previously Bostrichus pusillus Gyllenhal, 1813); DERMESTIDAE: Dermestes trifasciatus Fabricius, 1787 for Attagenus Latreille, 1802 (previously Dermestes pellio Linnaeus, 1758); ELATERIDAE: Elater sulcatus Fabricius, 1777 for Chalcolepidius Eschscholtz, 1829 (previously Chalcolepidius zonatus Eschscholtz, 1829); ENDOMYCHIDAE: Endomychus rufitarsis Chevrolat, 1835 for Epipocus Chevrolat, 1836 (previously Endomychus tibialis Guérin-Méneville, 1834); EROTYLIDAE: Ips humeralis Fabricius, 1787 for Dacne Latreille, 1797 (previously Dermestes bipustulatus Thunberg, 1781); EUCNEMIDAE: Fornax austrocaledonicus Perroud & Montrouzier, 1865 for Mesogenus Gemminger, 1869 (previously Mesogenus mellyi Bonvouloir, 1871); GLAPHYRIDAE: Melolontha serratulae Fabricius, 1792 for Glaphyrus Latreille, 1802 (previously Scarabaeus maurus Linnaeus, 1758); HISTERIDAE: Hister striatus Forster, 1771 for Onthophilus Leach, 1817 (previously Hister sulcatus Moll, 1784); LAMPYRIDAE: Ototreta fornicata E. Olivier, 1900 for Ototreta E. Olivier, 1900 (previously Ototreta weyersi E. Olivier, 1900); LUCANIDAE: Lucanus cancroides Fabricius, 1787 for Lissotes Westwood, 1855 (previously Lissotes menalcas Westwood, 1855); MELANDRYIDAE: Nothus clavipes G.-A. Olivier, 1812 for Nothus G.-A. Olivier, 1812 (previously Nothus praeustus G.-A. Olivier, 1812); MELYRIDAE: Lagria ater Fabricius, 1787 for Enicopus Stephens, 1830 (previously Dermestes hirtus Linnaeus, 1767); NITIDULIDAE: Sphaeridium luteum Fabricius, 1787 for Cychramus Kugelann, 1794 (previously Strongylus quadripunctatus Herbst, 1792); OEDEMERIDAE: Helops laevis Fabricius, 1787 for Ditylus Fischer, 1817 (previously Ditylus helopioides Fischer, 1817 [sic]); PHALACRIDAE: Sphaeridium aeneum Fabricius, 1792 for Olibrus Erichson, 1845 (previously Sphaeridium bicolor Fabricius, 1792); RHIPICERIDAE: Sandalus niger Knoch, 1801 for Sandalus Knoch, 1801 (previously Sandalus petrophya Knoch, 1801); SCARABAEIDAE: Cetonia clathrata G.-A. Olivier, 1792 for Inca Lepeletier & Audinet-Serville, 1828 (previously Cetonia ynca Weber, 1801); Gnathocera vitticollis W. Kirby, 1825 for Gnathocera W. Kirby, 1825 (previously Gnathocera immaculata W. Kirby, 1825); Melolontha villosula Illiger, 1803 for Chasmatopterus Dejean, 1821 (previously Melolontha hirtula Illiger, 1803); STAPHYLINIDAE: Staphylinus politus Linnaeus, 1758 for Philonthus Stephens, 1829 (previously Staphylinus splendens Fabricius, 1792); ZOPHERIDAE: Hispa mutica Linnaeus, 1767 for Orthocerus Latreille, 1797 (previously Tenebrio hirticornis DeGeer, 1775). The discovery of type species fixations that are older than those currently accepted pose a threat to nomenclatural stability (an application to the Commission is necessary to address each problem): CANTHARIDAE: Malthinus Latreille, 1805, Malthodes Kiesenwetter, 1852; CARABIDAE: Bradycellus Erichson, 1837, Chlaenius Bonelli, 1810, Harpalus Latreille, 1802, Lebia Latreille, 1802, Pheropsophus Solier, 1834, Trechus Clairville, 1806; CERAMBYCIDAE: Callichroma Latreille, 1816, Callidium Fabricius, 1775, Cerasphorus Audinet-Serville, 1834, Dorcadion Dalman, 1817, Leptura Linnaeus, 1758, Mesosa Latreille, 1829, Plectromerus Haldeman, 1847; CHRYSOMELIDAE: Amblycerus Thunberg, 1815, Chaetocnema Stephens, 1831, Chlamys Knoch, 1801, Monomacra Chevrolat, 1836, Phratora Chevrolat, 1836, Stylosomus Suffrian, 1847; COLONIDAE: Colon Herbst, 1797; CURCULIONIDAE: Cryphalus Erichson, 1836, Lepyrus Germar, 1817; ELATERIDAE: Adelocera Latreille, 1829, Beliophorus Eschscholtz, 1829; ENDOMYCHIDAE: Amphisternus Germar, 1843, Dapsa Latreille, 1829; GLAPHYRIDAE: Anthypna Eschscholtz, 1818; HISTERIDAE: Hololepta Paykull, 1811, Trypanaeus Eschscholtz, 1829; LEIODIDAE: Anisotoma Panzer, 1796, Camiarus Sharp, 1878, Choleva Latreille, 1797; LYCIDAE: Calopteron Laporte, 1838, Dictyoptera Latreille, 1829; MELOIDAE: Epicauta Dejean, 1834; NITIDULIDAE: Strongylus Herbst, 1792; SCARABAEIDAE: Anisoplia Schönherr, 1817, Anticheira Eschscholtz, 1818, Cyclocephala Dejean, 1821, Glycyphana Burmeister, 1842, Omaloplia Schönherr, 1817, Oniticellus Dejean, 1821, Parachilia Burmeister, 1842, Xylotrupes Hope, 1837; STAPHYLINIDAE: Batrisus Aubé, 1833, Phloeonomus Heer, 1840, Silpha Linnaeus, 1758; TENEBRIONIDAE: Bolitophagus Illiger, 1798, Mycetochara Guérin-Méneville, 1827. Type species are fixed for the following nominal genera: ANTHRIBIDAE: Decataphanes gracilis Labram & Imhoff, 1840 for Decataphanes Labram & Imhoff, 1840; CARABIDAE: Feronia erratica Dejean, 1828 for Loxandrus J.L. LeConte, 1853; CERAMBYCIDAE: Tmesisternus oblongus Boisduval, 1835 for Icthyosoma Boisduval, 1835; CHRYSOMELIDAE: Brachydactyla annulipes Pic, 1913 for Pseudocrioceris Pic, 1916, Cassida viridis Linnaeus, 1758 for Evaspistes Gistel, 1856, Ocnoscelis cyanoptera Erichson, 1847 for Ocnoscelis Erichson, 1847, Promecotheca petelii Guérin-Méneville, 1840 for Promecotheca Guérin- Méneville, 1840; CLERIDAE: Attelabus mollis Linnaeus, 1758 for Dendroplanetes Gistel, 1856; CORYLOPHIDAE: Corylophus marginicollis J.L. LeConte, 1852 for Corylophodes A. Matthews, 1885; CURCULIONIDAE: Hoplorhinus melanocephalus Chevrolat, 1878 for Hoplorhinus Chevrolat, 1878; Sonnetius binarius Casey, 1922 for Sonnetius Casey, 1922; ELATERIDAE: Pyrophorus melanoxanthus Candèze, 1865 for Alampes Champion, 1896; PHYCOSECIDAE: Phycosecis litoralis Pascoe, 1875 for Phycosecis Pascoe, 1875; PTILODACTYLIDAE: Aploglossa sallei Guérin-Méneville, 1849 for Aploglossa Guérin-Méneville, 1849, Colobodera ovata Klug, 1837 for Colobodera Klug, 1837; PTINIDAE: Dryophilus anobioides Chevrolat, 1832 for Dryobia Gistel, 1856; SCARABAEIDAE: Achloa helvola Erichson, 1840 for Achloa Erichson, 1840, Camenta obesa Burmeister, 1855 for Camenta Erichson, 1847, Pinotus talaus Erichson, 1847 for Pinotus Erichson, 1847, Psilonychus ecklonii Burmeister, 1855 for Psilonychus Burmeister, 1855. New replacement name: CERAMBYCIDAE: Basorus Bouchard & Bousquet, nom. nov. for Sobarus Harold, 1879. New status: CARABIDAE: KRYZHANOVSKIANINI Deuve, 2020, stat. nov. is given the rank of tribe instead of subfamily since our classification uses the rank of subfamily for PAUSSINAE rather than family rank; CERAMBYCIDAE: Amymoma Pascoe, 1866, stat. nov. is used as valid over Neoamymoma Marinoni, 1977, Holopterus Blanchard, 1851, stat. nov. is used as valid over Proholopterus Monné, 2012; CURCULIONIDAE: Phytophilus Schönherr, 1835, stat. nov. is used as valid over the unnecessary new replacement name Synophthalmus Lacordaire, 1863; EUCNEMIDAE: Nematodinus Lea, 1919, stat. nov. is used as valid instead of Arrhipis Gemminger, 1869, which is a junior homonym. Details regarding additional nomenclatural issues that still need to be resolved are included in the entry for each of these type genera: BOSTRICHIDAE: Lyctus Fabricius, 1792; BRENTIDAE: Trachelizus Dejean, 1834; BUPRESTIDAE: Pristiptera Dejean, 1833; CANTHARIDAE: Chauliognathus Hentz, 1830, Telephorus Schäffer, 1766; CARABIDAE: Calathus Bonelli, 1810, Cosnania Dejean, 1821, Dicrochile Guérin-Méneville, 1847, Epactius D.H. Schneider, 1791, Merismoderus Westwood, 1847, Polyhirma Chaudoir, 1850, Solenogenys Westwood, 1860, Zabrus Clairville, 1806; CERAMBYCIDAE: Ancita J. Thomson, 1864, Compsocerus Audinet-Serville, 1834, Dorcadodium Gistel, 1856, Glenea Newman, 1842; Hesperophanes Dejean, 1835, Neoclytus J. Thomson, 1860, Phymasterna Laporte, 1840, Tetrops Stephens, 1829, Zygocera Erichson, 1842; CHRYSOMELIDAE: Acanthoscelides Schilsky, 1905, Corynodes Hope, 1841, Edusella Chapuis, 1874; Hemisphaerota Chevrolat, 1836; Physonota Boheman, 1854, Porphyraspis Hope, 1841; CLERIDAE: Dermestoides Schäffer, 1777; COCCINELLIDAE: Hippodamia Chevrolat, 1836, Myzia Mulsant, 1846, Platynaspis L. Redtenbacher, 1843; CURCULIONIDAE: Coeliodes Schönherr, 1837, Cryptoderma Ritsema, 1885, Deporaus Leach, 1819, Epistrophus Kirsch, 1869, Geonemus Schönherr, 1833, Hylastes Erichson, 1836; DYTISCIDAE: Deronectes Sharp, 1882, Platynectes Régimbart, 1879; EUCNEMIDAE: Dirhagus Latreille, 1834; HYBOSORIDAE: Ceratocanthus A. White, 1842; HYDROPHILIDAE: Cyclonotum Erichson, 1837; LAMPYRIDAE: Luciola Laporte, 1833; LEIODIDAE: Ptomaphagus Hellwig, 1795; LUCANIDAE: Leptinopterus Hope, 1838; LYCIDAE: Cladophorus Guérin-Méneville, 1830, Mimolibnetis Kazantsev, 2000; MELOIDAE: Mylabris Fabricius, 1775; NITIDULIDAE: Meligethes Stephens, 1829; PTILODACTYLIDAE: Daemon Laporte, 1838; SCARABAEIDAE: Allidiostoma Arrow, 1940, Heterochelus Burmeister, 1844, Liatongus Reitter, 1892, Lomaptera Gory & Percheron, 1833, Megaceras Hope, 1837, Stenotarsia Burmeister, 1842; STAPHYLINIDAE: Actocharis Fauvel, 1871, Aleochara Gravenhorst, 1802; STENOTRACHELIDAE: Stenotrachelus Berthold, 1827; TENEBRIONIDAE: Cryptochile Latreille, 1828, Heliopates Dejean, 1834, Helops Fabricius, 1775. First Reviser actions deciding the correct original spelling: CARABIDAE: Aristochroodes Marcilhac, 1993 (not Aritochroodes ); CERAMBYCIDAE: Dorcadodium Gistel, 1856 (not Dorcadodion ), EVODININI Zamoroka, 2022 (not EVODINIINI); CHRYSOMELIDAE: Caryopemon Jekel, 1855 (not Carpopemon ), Decarthrocera Laboissière, 1937 (not Decarthrocerina ); CICINDELIDAE: Odontocheila Laporte, 1834 (not Odontacheila ); CLERIDAE: CORMODINA Bartlett, 2021 (not CORMODIINA), Orthopleura Spinola, 1845 (not Orthoplevra , not Orthopleuva ); CURCULIONIDAE: Arachnobas Boisduval, 1835 (not Arachnopus ), Palaeocryptorhynchus Poinar, 2009 (not Palaeocryptorhynus ); DYTISCIDAE: Ambarticus Yang et al., 2019 and AMBARTICINI Yang et al., 2019 (not Ambraticus , not AMBRATICINI); LAMPYRIDAE: Megalophthalmus G.R. Gray, 1831 (not Megolophthalmus , not Megalopthalmus ); SCARABAEIDAE: Mentophilus Laporte, 1840 (not Mintophilus , not Minthophilus ), Pseudadoretus dilutellus Semenov, 1889 (not P. ditutellus ). While the correct identification of the type species is assumed, in some cases evidence suggests that species were misidentified when they were fixed as the type of a particular nominal genus. Following the requirements of Article 70.3.2 of the International Code of Zoological Nomenclature we hereby fix the following type species (which in each case is the taxonomic species actually involved in the misidentification): ATTELABIDAE: Rhynchites cavifrons Gyllenhal, 1833 for Lasiorhynchites Jekel, 1860; BOSTRICHIDAE: Ligniperda terebrans Pallas, 1772 for Apate Fabricius, 1775; BRENTIDAE: Ceocephalus appendiculatus Boheman, 1833 for Uroptera Berthold, 1827; BUPRESTIDAE: Buprestis undecimmaculata Herbst, 1784 for Ptosima Dejean, 1833; CARABIDAE: Amara lunicollis Schiødte, 1837 for Amara Bonelli, 1810, Buprestis connexus Geoffroy, 1785 for Polistichus Bonelli, 1810, Carabus atrorufus Strøm, 1768 for Patrobus Dejean, 1821, Carabus gigas Creutzer, 1799 for Procerus Dejean, 1821, Carabus teutonus Schrank, 1781 for Stenolophus Dejean, 1821, Carenum bonellii Westwood, 1842 for Carenum Bonelli, 1813, Scarites picipes G.-A. Olivier, 1795 for Acinopus Dejean, 1821, Trigonotoma indica Brullé, 1834 for Trigonotoma Dejean, 1828; CERAMBYCIDAE: Cerambyx lusitanus Linnaeus, 1767 for Exocentrus Dejean, 1835, Clytus supernotatus Say, 1824 for Psenocerus J.L. LeConte, 1852; CICINDELIDAE: Ctenostoma jekelii Chevrolat, 1858 for Ctenostoma Klug, 1821; CURCULIONIDAE: Cnemogonus lecontei Dietz, 1896 for Cnemogonus J.L. LeConte, 1876; Phloeophagus turbatus Schönherr, 1845 for Phloeophagus Schönherr, 1838; GEOTRUPIDAE: Lucanus apterus Laxmann, 1770 for Lethrus Scopoli, 1777; HISTERIDAE: Hister rugiceps Duftschmid, 1805 for Hypocaccus C.G. Thomson, 1867; HYBOSORIDAE: Hybosorus illigeri Reiche, 1853 for Hybosorus W.S. MacLeay, 1819; HYDROPHILIDAE: Hydrophilus melanocephalus G.-A. Olivier, 1793 for Enochrus C.G. Thomson, 1859; MYCETAEIDAE: Dermestes subterraneus Fabricius, 1801 for Mycetaea Stephens, 1829; SCARABAEIDAE: Aulacium carinatum Reiche, 1841 for Mentophilus Laporte, 1840, Phanaeus vindex W.S. MacLeay, 1819 for Phanaeus W.S. MacLeay, 1819, Ptinus germanus Linnaeus, 1767 for Rhyssemus Mulsant, 1842, Scarabaeus latipes Guérin-Méneville, 1838 for Cheiroplatys Hope, 1837; STAPHYLINIDAE: Scydmaenus tarsatus P.W.J. Müller & Kunze, 1822 for Scydmaenus Latreille, 1802. New synonyms: CERAMBYCIDAE: CARILIINI Zamoroka, 2022, syn. nov. of ACMAEOPINI Della Beffa, 1915, DOLOCERINI Özdikmen, 2016, syn. nov. of BRACHYPTEROMINI Sama, 2008, PELOSSINI Tavakilian, 2013, syn. nov. of LYGRINI Sama, 2008, PROHOLOPTERINI Monné, 2012, syn. nov. of HOLOPTERINI Lacordaire, 1868.
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The order Coleoptera (beetles) is arguably the most speciose group of animals, but the evolutionary history of beetles, including the impacts of plant feeding (herbivory) on beetle diversification, remain poorly understood. We inferred the phylogeny of beetles using 4,818 genes for 146 species, estimated timing and rates of beetle diversification using 89 genes for 521 species representing all major lineages and traced the evolution of beetle genes enabling symbiont-independent digestion of lignocellulose using 154 genomes or transcriptomes. Phylogenomic analyses of these uniquely comprehensive datasets resolved previously controversial beetle relationships, dated the origin of Coleoptera to the Carboniferous, and supported the codiversification of beetles and angiosperms. Moreover, plant cell wall-degrading enzymes (PCWDEs) obtained from bacteria and fungi via horizontal gene transfers may have been key to the Mesozoic diversification of herbivorous beetles—remarkably, both major independent origins of specialized herbivory in beetles coincide with the first appearances of an arsenal of PCWDEs encoded in their genomes. Furthermore, corresponding (Jurassic) diversification rate increases suggest that these novel genes triggered adaptive radiations that resulted in nearly half of all living beetle species. We propose that PCWDEs enabled efficient digestion of plant tissues, including lignocellulose in cell walls, facilitating the evolution of uniquely specialized plant-feeding habits, such as leaf mining and stem and wood boring. Beetle diversity thus appears to have resulted from multiple factors, including low extinction rates over a long evolutionary history, codiversification with angiosperms, and adaptive radiations of specialized herbivorous beetles following convergent horizontal transfers of microbial genes encoding PCWDEs.
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Since the description of the genus Cheguevaria as incertae sedis (Lampyridae), the placement of these beetles has been uncertain. This study is the first to address the placement of this genus based on a molecular phylogenetic analysis. We used three genes (18S rRNA, rrnl mitochondrial DNA and cox1 mitochondrial DNA) and a maximum likelihood approach with W-IQ-TREE to support Cheguevaria as a member of the Lampyridae and recognize it as the sole genus in the new subfamily Cheguevariinae stat. nov.
Article
Extinct haidomyrmecine ‘‘hell ants’’ are among the earliest ants known. These eusocial Cretaceous taxa diverged from extant lineages prior to the most recent common ancestor of all living ants and possessed bizarre scythe-like mouthparts along with a striking array of horn-like cephalic projections. Despite the morphological breadth of the fifteen thousand known extant ant species, phenotypic syndromes found in the Cretaceous are without parallel and the evolutionary drivers of extinct diversity are unknown. Here, we provide a mechanistic explanation for aberrant hell ant morphology through phylogenetic reconstruction and comparative methods, as well as a newly reported specimen. We report a remarkable instance of fossilized predation that provides direct evidence for the function of dorsoventrally expanded mandibles and elaborate horns. Our findings confirm the hypothesis that hell ants captured other arthropods between mandible and horn in a manner that could only be achieved by articulating their mouthparts in an axial plane perpendicular to that of modern ants. We demonstrate that the head capsule and mandibles of haidomyrmecines are uniquely integrated as a consequence of this predatory mode and covary across species while finding no evidence of such modular integration in extant ant groups. We suggest that hell ant cephalic integration— analogous to the vertebrate skull—triggered a pathway for an ancient adaptive radiation and expansion into morphospace unoccupied by any living taxon.
Article
Recent discoveries of enantiornithine remains in Burmese amber have provided a wealth of paleobiological data on this extinct clade of Mesozoic birds. Amber, as a unique medium of fossilization, preserves in three dimensions structures with details unmatched elsewhere in the fossil record. This provides the opportunity to combine osteological information with more detailed information on integumentary structures, including soft tissues and the plumage of the specimens. Herein, we describe an isolated bird foot, DIP-V-19354, consisting of complete metatarsals and digits, including the claws. Placement among the Enantiornithes is supported by the presence of a metatarsal IV with a trochlea formed by a single condyle and the large size and curvature of the claws. The bones of the right foot are preserved fully encased in soft tissue. Scutellae scale filaments (SSFs) are present along the metatarsals and digits, and are similar to those previously described in other enantiornithines. The distribution and relative size of the SSFs on the longest digit support the hypothesis of a mechanosensory tactile role of these structures: this may implicate the digit in the feeding strategy of the animal, as has been suggested for Elektorornis. Unlike many of the previously described enantiornithine remains from this deposit, the taphonomic history of DIP-V-19354 suggests that the foot was trapped in resin flows above the forest floor, likely on the trunk of a tree, after the bird had died.
Article
The plesiomorphic antenna with eleven articles is stunningly stable across the over 400.000 beetle species, despite the astonishing diversity of morphologies and ecological settings in which these organisms live. However, a few beetle lineages evolved different antennomere numbers, and these offer an interesting opportunity to understand how an otherwise fixed phenotype may eventually change, and to study what factors shape their patterns of variation. Here, we review for the first time, based on original and literature data, the unique patterns of variation in antennomere numbers in the firefly family (Coleoptera: Lampyridae), which has the greatest range (7 to 62) in antennomere numbers among beetles despite a relatively low species diversity (∼2000 species). We also tested the hypothesis that antennomere numbers are positively related to body size and explored patterns of antennal asymmetry across Amydetes spp, which have the greatest range in antennomere numbers (24 to 62). Out of the ∼100 firefly genera, nine show antennomere numbers lower or higher than 11. Among those nine genera with unusual antennomere numbers, five show intraspecific variation as well. In taxa whose antennae have less than 11 antennomeres, males seem to follow mainly light cues to find mates. In Amydetes, we found that antennomere numbers are positively related and negatively allometric to body size, suggesting that in this group the mechanism shaping antennal segmentation is sensitive to body size. Moreover, fluctuating asymmetry was found to be widespread in Amydetes spp., with a difference to up to six antennomeres between left and right antennae. The combined occurrence of unusual antennomere numbers and within-species variation suggests that in several firefly taxa evolved a destabilization of the mechanism driving antennal segmentation has occurred.
Article
The avian digestive system, like other aspects of avian biology, is highly modified relative to other reptiles. Together these modifications have imparted the great success of Neornithes, the most diverse clade of amniotes alive today. It is important to understand when and how aspects of the modern avian digestive system evolved among neornithine ancestors in order to elucidate the evolutionary success of this important clade and to understand the biology of stem birds and their closest dinosaurian relatives: Mesozoic Paraves. Although direct preservation of the soft tissue of the digestive system has not yet been reported, ingested remains and their anatomical location preserved in articulated fossils hint at the structure of the digestive system and its abilities. Almost all data concerning direct evidence of diet in Paraves comes from either the Upper Jurassic Yanliao Biota or the Lower Cretaceous Jehol Biota, both of which are known from deposits in north‐eastern China. Here, the sum of the data gleaned from the thousands of exceptionally well‐preserved fossils of paravians is interpreted with regards to the structure and evolution of the highly modified avian digestive system and feeding apparatus. This information suggests intrinsic differences between closely related stem lineages implying either strong homoplasy or that diet in each lineage of non‐ornithuromorph birds was highly specialized. Regardless, modern digestive capabilities appear to be limited to the Ornithuromorpha, although the complete set of derived feeding related characters is restricted to the Neornithes.
Article
Cydistinae are a rare monogeneric beetle lineage from Asia with a convoluted history of classification, historically placed in various groups within the series Elateriformia. However, their position has never been rigorously tested. To resolve this long-standing puzzle, we are the first to present sequences of two nuclear and two mitochondrial markers for four species of Cydistinae to determine their phylogenetic position. We included these sequences in two rounds of analyses: one including a broad Elateriformia dataset to test placement at the superfamily/family level, and a second, including a richer, targeted sampling of presumed close relatives. Our results strongly support Cydistinae as sister to Phengodidae in a clade with Rhagophthalmidae. Based on our molecular phylogenetic results and examination of morphological characters, we hereby transfer the formerly unplaced Cydistinae into Phengodidae and provide diagnoses for the newly circumscribed Phengodidae, Cydistinae and Cydistus. Since both Phengodidae and Rhagophthalmidae have bioluminescent larvae and strongly neotenic females, similar features can be hypothesized for Cydistinae. Additionally, Cydistus minor is transferred to the new genus Microcydistus.