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Sinopyrophorinae, a new subfamily of Elateridae (Coleoptera, Elateroidea) with the first record of a luminous click beetle in Asia and evidence for multiple origins of bioluminescence in Elateridae


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The new subfamily Sinopyrophorinae within Elateridae is proposed to accommodate a bioluminescent species, Sinopyrophorus schimmeli Bi & Li, gen. et sp. nov., recently discovered in Yunnan, China. This lineage is morphologically distinguished from other click-beetle subfamilies by the strongly protruding frontoclypeal region, which is longitudinally carinate medially, the pretarsal claws without basal setae, the hind wing venation with a well-defined wedge cell, the abdomen with seven (male) or six (female) ventrites, the large luminous organ on the abdominal sternite II, and the male genitalia with median lobe much shorter than parameres, and parameres arcuate, with the inner margin near its apical third dentate. Molecular phylogeny based on the combined 14 mitochondrial and two nuclear genes supports the placement of this taxon far from other luminescent click-beetle groups, which provides additional evidence for the multiple origin of bioluminescence in Elateridae. Illustrations of habitus and main diagnostic features of S. schimmeli Bi & Li, gen. et sp. nov. are provided, as well as the brief description of its luminescent behavior.
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Sinopyrophorinae, a new subfamily of Elateridae (Coleoptera, Elateroidea) with the rst ... 79
Sinopyrophorinae, a new subfamily of Elateridae
(Coleoptera, Elateroidea) with the first record of a
luminous click beetle in Asia and evidence for multiple
origins of bioluminescence in Elateridae
Wen-Xuan Bi1,2*, Jin-Wu He1*, Chang-Chin Chen3, Robin Kundrata4, Xue-Yan Li1
1 State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of
Sciences, Kunming 650223, China 2 Room 401, No. 2, Lane 155, Lianhua South Road, Shanghai, 201100,China
3NPS oce, Tianjin New Wei San Industrial Company, Ltd., Tianjing, China 4 Department of Zoology, Faculty
of Science, Palacky University, 17. listopadu 50, 77146, Olomouc, Czech Republic
Corresponding author: Xue-Yan Li (, Wen-Xuan Bi (
Academic editor: Hume Douglas| Received 15 May 2018 |Accepted 15 May 2019| Published 17 July 2019
Citation: Bi W-X, He J-W, Chen C-C, Kundrata R, Li X-Y (2019) Sinopyrophorinae, a new subfamily of Elateridae
(Coleoptera, Elateroidea) with the rst record of a luminous click beetle in Asia and evidence for multiple origins of
bioluminescence in Elateridae. ZooKeys 864: 79–97.
e new subfamily Sinopyrophorinae within Elateridae is proposed to accommodate a bioluminescent
species, Sinopyrophorus schimmeli Bi & Li, gen. et sp. nov., recently discovered in Yunnan, China. is
lineage is morphologically distinguished from other click-beetle subfamilies by the strongly protruding
frontoclypeal region, which is longitudinally carinate medially, the pretarsal claws without basal setae, the
hind wing venation with a well-dened wedge cell, the abdomen with seven (male) or six (female) ventrites,
the large luminous organ on the abdominal sternite II, and the male genitalia with median lobe much shorter
than parameres, and parameres arcuate, with the inner margin near its apical third dentate. Molecular
phylogeny based on the combined 14 mitochondrial and two nuclear genes supports the placement of this
taxon far from other luminescent click-beetle groups, which provides additional evidence for the multiple
origin of bioluminescence in Elateridae. Illustrations of habitus and main diagnostic features of S. schimmeli
Bi & Li, gen. et sp. nov. are provided, as well as the brief description of its luminescent behavior.
China, mitochondrial genome, molecular phylogeny, new genus, new species, taxonomy
* ese authors contributed equally to this work.
ZooKeys 864: 79–97 (2019)
doi: 10.3897/zookeys.864.26689
Copyright Wen-Xuan Bi et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC
BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
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Wen-Xuan Bi et al. / ZooKeys 864: 79–97 (2019)
e cosmopolitan family Elateridae currently contains approximately 600 genera and
almost 10,000 species (Costa et al. 2010, Kundrata and Bocak 2011). Based on a
synthesis of the work over the past several decades, Costa et al. (2010; and references
therein) provided the main characteristics of adult and larval elaterid morphology and
recognized 17 subfamilies. ereafter, slight modications of the above classication
were proposed by Kundrata and Bocak (2011), Kundrata et al. (2016, 2018), Bocak
et al. (2018), and Douglas et al. (2018) based on molecular analyses. Currently, 18
subfamilies and 37 tribes of Elateridae are recognized (Kundrata et al. 2018).
Approximately 200 species of Elateridae are able to emit light, and these belong to
Agrypninae: Pyrophorini (most species), ylacosterninae (Balgus schnusei Heller) and
Campyloxeninae (Campyloxenus pyrothorax Fairmaire) (Costa et al. 2010). e vast major-
ity of bioluminescent species is known from the Neotropical region, with several species oc-
curring in small Melanesian islands (Costa 1975, 1984b; Costa et al. 2010). e position of
the luminous organs varies among adults of dierent elaterid lineages; they can be found on
both prothorax and abdomen (Pyrophorini: Hapsodrilina and Pyrophorina), only on the
prothorax (Balgus Fleutiaux, Campyloxenus Fairmaire, Pyrophorini: Nyctophyxina), or only
on the abdomen (Pyrophorini: Hifo Candèze) (Costa 1975; 1984a, b; Costa et al. 2010).
In 2017, during an expedition to the western Yunnan in China, in which the rst
author participated, a remarkable dusk-active bioluminescent click beetle with a single
luminous organ on the abdomen was discovered. Since no bioluminescent representa-
tive of Elateridae has been recorded in Asia to date, morphological study and molecular
phylogenetic analysis were undertaken simultaneously to clarify the identity and phy-
logenetic placement of the new taxon. He et al. (2019) published the mitochondrial
genome of this species and provided a preliminary phylogenetic hypothesis for it based
on the analysis of 13 protein-coding genes. Here, we formally describe this species in
a new genus, which we propose to place into a new subfamily within Elateridae, and
provide more robust phylogenetic hypothesis for this taxon.
Materials and methods
All specimens of the new species were collected from the vicinity of Longchuan Coun-
ty and Yingjiang County in western Yunnan, in subtropical evergreen broadleaf forests
by searching for ashes or by setting light traps during the night. Specimens are de-
posited in the following collections: Kunming Institute of Zoology, Chinese Academy
of Sciences, Kunming, China (KIZ-CAS), e Insect Collection of Shanghai Normal
University, Shanghai, China (SNUC), collection of Wen-Xuan Bi, Shanghai, China
(CBWX), and the collection of Chang-Chin Chen, Tianjin, China (CCCC). Morpho-
logical terminology follows Calder (1996) and Costa et al. (2010), except for the wing
venation, for which we follow Kukalova-Peck and Lawrence (1993, 2004).
Sinopyrophorinae, a new subfamily of Elateridae (Coleoptera, Elateroidea) with the rst ... 81
Phylogenetic analysis
To explore the phylogenetic position of the new Chinese luminescent click beetle, we
newly generated 14 mitochondrial genes (13 protein-coding genes and 16S) and two
nuclear rRNA genes (18S and 28S) for this species and merge them with the available
four-gene Elateridae dataset (18S, 28S, 16S, cox1) by Kundrata et al. (2016, 2018)
and Douglas et al. (2018) plus all publicly available Elateridae mitochondrial genomes
downloaded from the GenBank (altogether 179 terminals representing 13 subfami-
lies). Four species of Phengodidae were used as outgroups following Kundrata et al.
(2016, 2018) (Suppl. material 1, Table S1).
e mitogenome of the new species was obtained from GenBank (accession num-
ber MH065615) (He et al. 2019). Because we were not able to obtain high qual-
ity reference sequences of 18S and 28S from the Illumina reads previously used for
assembling the mitogenome, we assembled 18S and 28S based on another batch of
Illumina reads sequenced using fresh specimens collected from type locality later in
June 13, 2018, and immediately frozen in liquid nitrogen and stored at -80 °C before
use. Total genomic DNA (gDNA) was isolated from two male adultswith Sodium
Dodecyl Sulfonate method. Library (150-bp insert size) was prepared and sequenced
on the Illumina HisSeq4000. Total 38 Gb clean reads were used for assembling 18S
and 28S based on the following method: 1) reads were mapped to reference genes of
rDNA using mrsFAST v3.3.0 with several iterations (Hach et al. 2014) (rst iteration
referred to genes downloaded from GenBank (NCBI Resource Coordinators 2016);
later iterations referred to assembled contigs in the latest iteration); 2) mapped reads
were then assembled using SPAdes v3.11 (Bankevich et al. 2012); 3) tandem contigs
were ltered using mereps v2.6 because a huge number of repetitive reads greatly low-
ers assembly eciency (Kolpakov et al. 2003); 4) steps one to three were automatically
performed for 3–10 iterations until all genes were recovered.
e individual genes were aligned using Mat online version 7 (https://mat.cbrc.
jp/alignment/server/) (Kuraku et al. 2013) with default parameters. e alignment was
displayed and manually curated in Mega7 (Kumar et al. 2016). All 16 aligned genes
were concatenated using SequenceMatrix version 1.7.8 (Vaidya et al. 2011). e con-
catenated matrix was used to calculate the best-t evolutionary model (GTR+I+G) us-
ing PartitionFinder 2.1.1 (Lanfear et al. 2012, 2017). Maximum likelihood (ML) anal-
ysis were carried out using RAxML version 7.0.4. (Stamatakis et al. 2008) with 1000
bootstrap replicates. Phylograms were drawn using Interactive Tree of Life (ITOL)
(Letunic and Bork 2016).
Phylogenetic inference
Our phylogenetic analysis recovered similar topology (Fig. 1) to that of Kundrata et
al. (2018), only Tetralobinae were placed in an unsupported terminal clade with Car-
Wen-Xuan Bi et al. / ZooKeys 864: 79–97 (2019)
diophorinae + Negastriinae, and Dendrometrinae including Diplophoenicus Candèze
(Morostomatinae) formed a separate clade. Similar to previously published phylog-
enies, we obtained low statistical support for the backbone of tree. All subfamilies were
monophyletic, except for Dendrometrinae (due to the inclusion of Diplophoenicus)
and Lissominae (due to the inclusion of ylacosterninae). e newly sequenced lu-
minescent species from China was found in an unsupported clade with Oestodinae
and Hemiopinae as follows: Oestodes tenuicollis (Randall) + (Hemiops sp. + new lu-
minescent taxon), far from other bioluminescent groups like ylacosterninae and
Agrypninae: Pyrophorini.
Figure 1. Inferred phylogenetic position of Sinopyrophorus schimmeli Bi & Li, gen. et sp. nov. within
Elateridae based on the concatenated 14 mitochondrial genes (13 protein-coding genes and 16S) and two
nuclear ribosomal genes (18S, 28S) using the Maximum Likelihood (ML) analysis. Numbers near each
branch indicate ML bootstrap values with 1000 replicates. e same colored shaded areas at the terminals
denote the same subfamily. Green bold lines indicate luminescent taxa. e bold red line indicates the
presence of luminescent species within the same genus.
Sinopyrophorinae, a new subfamily of Elateridae (Coleoptera, Elateroidea) with the rst ... 83
Sinopyrophorus Bi & Li, gen. nov.
Figs 2–23
= Sinopyrophorus He et al., 2019: 565 [nomen nudum; published without description,
unavailable name according to the ICZN (1999, Art. 13)].
Type species. Sinopyrophorus schimmeli Bi & Li, sp. nov., here designated.
Diagnosis. Head with frontoclypeal region (Fig. 4) strongly protruding, longitu-
dinally strongly carinate medially; antennomeres II and III short, subequal in length;
clicking mechanism (i.e., prosternal process tting into mesoventral cavity) fully devel-
oped; prosternal process straight in lateral view, pretarsal claw (Fig. 13) lacking setae at
base; hind wing (Fig. 14) with well-dened wedge cell; abdomen with seven (male) or
six (female) ventrites; large transverse luminous organ present on abdominal sternite
II (Fig. 16); aedeagus (Fig. 20) with parameres arcuate and median lobe much shorter
than parameres.
Description. Male. Body elongate, ~ 4.6 times as long as wide, weakly convex in
lateral view. Vestiture of ne, suberect setae.
Head with frontoclypeal region strongly protruding, inexed at apex, medially
longitudinally carinate; carina setose and apparently not joined to supra-antennal cari-
nae, basally half as wide as frons, then narrowed and subparallel-sided, with cuticle
between edges at (Fig. 4). Antennal insertions concealed from above. Labrum free,
transverse, anterior margin convex in dorsal view. Maxilla (Fig. 7) with galea scoop-
like, anterior part covered with setae, denser on inner edge; lacinia elongate, densely
pilose; palp with apical palpomere slightly expanded anteriorly. Labium (Fig. 6) with
prementum elongate, trilobed anteriorly. Mandibles (Fig. 5) bidentate, apical tooth
narrowly acute, subapical mesal tooth small. Antenna with 11 antennomeres, liform;
antennomeres II and III subequal in length (together 1/3 as long as antennomere IV),
globular; remaining antennomeres at least ve times longer than wide.
Prothorax with chin piece of prosternum short, bisinuate, not concealing la-
bium; prosternal process slightly constricted between coxae in ventral view, almost
straight in lateral view. Pronotosternal suture almost straight. Procoxae narrowly
separated, externally broadly open. Scutellar shield (Fig. 9)trapezoidal, moderately
elevated, narrowed posteriorly, posterior apex slightly emarginate. Mesocoxal cavi-
ties narrowly separated, open laterally to both mesepimeron and mesepisternum;
mesotrochantin visible. Mesoventrite (Figs 10, 11) with posterior area lower than
metaventrite. Meso-metaventral suture distinct. Metacoxae extending laterally to
meet metepimeron, metacoxal plates not covering trochanters when legs withdrawn.
Hind wing (Fig. 14) 2.35 times as long as wide; apical eld ~ 0.25 times as long as
total wing length, apical eld with three sclerites forming an epsilon gure; radial
cell longer than wide, with inner posterobasal angle acute; cross-vein r3 horizontal;
Wen-Xuan Bi et al. / ZooKeys 864: 79–97 (2019)
MP3+4 with basal cross-vein and basal spur; CuA2 meet
ing MP4; wedge cell present,
~ 3.5 times as long as wide, with obliquely truncate apex. Leg with trochanter-femur
joint oblique; tibial spurs double, tarsal formula 5-5-5; pretarsal claws (Fig. 13) simple,
lacking setae at base; empodium weakly developed, bisetose.
Abdomen with seven ventrites (sternites III–IX, Figs 2b, 15). Sternite II with trans-
verse, semicircular luminescent organ occupying more than half of its width (Fig. 16).
Intercoxal process of ventrite I (i.e., sternite III) narrowly rounded. First ve ventrites
subequal in length; ventrite V with posterior margin emarginate. Tergite VIII (Fig. 18);
sternite VIII (Fig. 17) ~ 0.7 times as long as ventrite V (i.e., sternite VII), posteromedi-
ally emarginate, with each lobe slightly emarginate posteriorly; sternite IX (Fig. 19b)
acute at base, more sclerotized at anterior half, connected to tergite IX by membrane;
tergites IX and X partly fused (Fig. 19a), narrowly pointedanteriorly. Aedeagus (Fig.
20) with median lobe very short, only approximately half as long as aedeagus, with
short basal struts; basal half wider, apical half narrow, slightly concave to subparallel-
sided, with rounded apex.
Female. Slightly larger than male. Abdomen with six ventrites. Sternite VIII (Fig.
21) with spiculum ventrale 0.8 times total length of sternite. Ovipositor (Fig. 22) long,
with paraprocts 3.5 times longer than gonocoxites; gonocoxites partially sclerotized;
styli attached subapically. Internal genital tract (Fig. 23) simple; vagina long, mem-
branous, slightly enlarged near entry of common oviduct; bursa copulatrix elongate,
slightly widened anteriorly, sclerotized near base, with single spermathecal gland duct
and ne spinules internally; colleterial gland absent.
Etymology. e generic name is derived from the Latin prex sino-, which means
Chinese, and Pyrophorus, a bioluminescent click-beetle genus from Central and South
America. Gender masculine.
Distribution. China: Western Yunnan.
Sinopyrophorus schimmeli Bi & Li, sp. nov.
Figs 2–23
= Sinopyrophorus schimmeli He et al., 2019: 565 [nomen nudum; published without
description, unavailable name according to the ICZN (1999, Art. 13)].
Type locality. China, Yunnan, Yingjiang, Shangbangzhong, 24°26'N, 97°45'W, 1650 m.
Type material. Holotype: male, “China, Yunnan, Yingjiang, Shangbangzhong,
24°26’N, 97°45’W, 1650 m, 2017.VI.23, leg. Wen-Xuan Bi”; “Holotype Sinopyropho-
russchimmeli sp. nov.” [red handwritten label] (SNUC). Paratypes (9 males, 3 females):
1 female, same data as holotype (KIZ-CAS); 1 male, 1 female, “China, Yunnan,
Longchuan, Husa, 1770 m, 2017.VI.13–14, leg. Wen-Xuan Bi” (CBWX); 1 male,
ditto except 2000 m, 2017.VI.16 (CBWX); 1 male, 1 female, ditto except 1700 m,
2017.VI.24, leg. Yu-Tang Wang (CBWX); 2 males, ditto except 1770 m, 2016.V.31
Sinopyrophorinae, a new subfamily of Elateridae (Coleoptera, Elateroidea) with the rst ... 85
Figures 2–3. Habitus of Sinopyrophorus schimmeli Bi & Li, gen. et sp. nov. paratypes 2 male 3 female. a,
dorsal view; b, ventral view; c, lateral view.
(CCCC); 1 male, ditto except leg. Xiao-Dong Yang (CCCC); 3 males, ditto except
2017.VI.25, leg. Wen-Xuan Bi (KIZ-CAS).
Other material examined. 2 males, China, Yunnan, Longchuan, Husa, 1770 m,
2017.VI.13–16, leg. Wen-Xuan Bi, damaged, partially used for extracting genomic
DNA in a project of mitogenome (accession number MH065615; He et al. 2019); 2
males, China, Yunnan, Longchuan, Husa, 1770 m, 2018.VI.13, leg. Wen-Xuan Bi,
the whole body of both specimens was used for the extraction of genomic DNA in an
ongoing project of de novo genome sequencing and assembly of 18S and 28S.
Diagnostic description. Male (Fig. 2). Body length 9.6–11.3 mm (holotype: 11.3
mm). Body brown to dark brown, with posterior portion of prothorax, fore- and mid-legs
Wen-Xuan Bi et al. / ZooKeys 864: 79–97 (2019)
paler; elytra with broad subapical pale band, zigzagged anteriorly, rounded posteriorly. Body
surface with ne, suberect brown setae, denser on legs; elytral light band with pale setae.
Head transverse, weakly convex, 0.75 times as long as wide, same width as prono-
tal anterior edge; sparsely and nely punctate. Frons rectangular, 1.4 times longer than
width, 0.3 times as wide as head width across the eyes. Frontoclypeal region concave
at sides beneath; with one small median depression. Eyes protuberant, median width
of each eye ~ 0.7 times interocular distance in dorsal view. Mouthparts directed anter-
oventrally. Labrum (Fig. 5) 2.2 times wider than long. Antenna long, reaching second
half of elytral length, ~ 0.7 times as long as body length; scape 2.3 times as long as
Figures 4–16. Sinopyrophorus schimmeli Bi & Li, gen. et sp. nov. Male 4 head (anterior view) 5 labrum
and mandibles (dorsal view) 6 labium 7 maxilla 8 prothorax 9 scutellum (dorsal view) 10 mesoventrite
(ventral view) 11 mesoventrite (lateral view) 12 tarsomeres II–IV (lateral view) 13 tarsal claw (lateral
view) 14 hind wing 15 ventrites IV–VII 16 abdominal luminescent organ (pale area above ventrite I). a,
dorsal view; b, ventral view; c, lateral view. Scale bars: 0.25 mm (4–11); 0.1 mm (12, 13); 1 mm (14);
not to scale (15, 16).
Sinopyrophorinae, a new subfamily of Elateridae (Coleoptera, Elateroidea) with the rst ... 87
combined length of antennomeres II and III, and of approximately same length as
antennomere 4, slightly curved; antennomeres 4–11 successively weakly lengthened,
with ne and very long, distinct setae; setae almost as long as apical antennomere.
Prothorax (Fig. 8) slightly convex in lateral view, tallest anteriorly; ~ 1.2 times as
long as wide in dorsal view, weakly narrowed anteriorly, slightly narrower than elytral
humeral width; sides slightly sinuate; pronotal lateral carina complete; anterior angles
short, subacute; hind angles narrowly acute, moderately produced posterolaterally, each
with short carina; posterior edge straight from dorsal view, medially elevated; pronotal
disk sparsely and nely punctate, with eight shallow depressions: single median and
posteromedian, and three pairs at sides. Prosternum and hypomeron more coarsely
punctate than pronotum. Elytra ~ 3.0 times as long as combined width, ~2.9 times
Figures 17–23. Sinopyrophorus schimmeli Bi & Li, gen. et sp. nov. Male 17 sternite VIII 18 tergite
VIII 19 tergites IX–X with sternite IX 20 aedeagus. Female 21 sternite VIII 22 ovipositor (dorsal view)
23internal genital tract. Abbreviations: eco, the entry of the common ovdiduct; sgd, spermathecal gland
duct. a, dorsal view; b, ventral view; c, lateral view. Scale bars: 1 mm; not to scale (23).
Wen-Xuan Bi et al. / ZooKeys 864: 79–97 (2019)
as long as pronotum, parallel-sided; each elytron with three low and evenly spaced
swellings, longitudinally arranged at basal half near suture; with nine punctate striae;
apices conjointly rounded; epipleura short, abruptly narrowed near metacoxa. Legs
long; tarsomeres I–III elongate, tarsomere I ~ 1.4 times as long as tarsomere II, tar-
somere II as long as combined lengths of tarsomeres III and IV or as long as tarsomere
V; tarsomeres III and IV ventrally lobate (Fig. 12).
Abdomen with each ventrite with paired depressions posterolaterally. Aedeagus
(Fig. 20) robust, ~ 1.6 times as long as wide. Phallobase slightly longer than wide,
narrowed dorsally, emarginate posteriorly. Median lobe approximately half as long as
aedeagus, with basal struts 0.25 times total length of median lobe. Parameres ~ 1.5
times longer than median lobe, shorter than phallobase, with dorsal surface ~ 1.6 times
longer than ventral one, partially fused basally in dorsal view; each paramere with dor-
sal surface angulate on inner margins at basal 2/5, strongly narrowed and curved mesad
with dentate inner margin near apical 1/3 and rounded apex; bearing 20–30 ne setae
near apex, longer on ventral surface.
Female (Fig. 3). Body length 12.1–14.5 mm. Similar to male in its general appear-
ance but with integument paler. Eyes smaller, median width of each eye ~ 0.4 times
interocular distance in dorsal view. Antenna shorter, only reaching elytral humeri, ~
0.3 times as long as body length. Pronotum relatively shorter with lateral margins more
rounded, narrowed anteriorly, with rounded anterior angles. Elytra relatively longer,
~ 3.3 times as long as wide, ~ 3.0 times as long as pronotum. Legs relatively shorter.
Abdominal luminescent organ smaller, occupying approximately one third of basal
abdominal sternite width.
Immature stages. Unknown.
Etymology. is species is named in honor of late Mr. Rainer Schimmel, a special-
ist in Elateridae, who kindly provided valuable comments at the beginning of this study.
Biological notes. All specimens of the new species were collected during the late
May to June (i.e., the middle of the rainy season) from the mountain area in vicinity
of Longchuan County or Yingjiang County, western Yunnan in subtropical evergreen
broadleaf forests by searching for ashes or by light trapping during night. e adults
of both sexes emitted a continuous yellowish green light from the abdominal luminous
organs while in ight, or during a short time when preparing for ight or afterwards.
During this process the luminous organ is exposed ventrally by raising and extending
the abdomen from the metaventrite (Suppl. material 2, Movie S1). e reaction of the
adults when disturbed during a ight is to retract their abdomen, hide their luminous
organ, and show a death-feigning behavior (thanatosis). anatoid adults remained
inactive for a long time, which was obviously dierent from the observation in Pyro-
phorini because the latter begin to emit light and are very active after being disturbed
(Costa 1975). At least three lampyrid species (Luciola sp., Diaphanes sp., and Pyrocoelia
sp.) occurred sympatrically with the new elaterid species and can be found simultane-
ously during night but can be easily distinguished by dierent ash patterns. Although
some specimens of S. schimmeli were previously collected using light traps in 2016, this
species was found to be luminescent only in 2017 when its bioluminescent behavior
Sinopyrophorinae, a new subfamily of Elateridae (Coleoptera, Elateroidea) with the rst ... 89
was observed in the eld. is was caused by the abdominal luminescent organ being
hidden when the beetle is not active.
Nomenclatural notes. He et al. (2019) reported the mitochondrial genome of S.
schimmeli gen. et sp. nov. and used the genus and species name of the here described
taxon in their study. e paper of He et al. (2019) was intended to be published after
the formal description of S. schimmeli Bi & Li, gen. et sp. nov. but unfortunately, it
was published earlier, causing nomenclatural problems by reporting the genus and spe-
cies names without available descriptions and thus unavailable according to the Code
(ICZN 1999).
Sinopyrophorinae Bi & Li, subfam. nov.
Type genus. Sinopyrophorus Bi & Li, gen. nov., here designated.
Diagnosis. e molecular phylogenetic analysis (Fig. 1) and morphology (Figs
2–23) justify the establishment of a new monogeneric subfamily Sinopyrophorinae Bi
& Li, subfam. nov. within Elateridae. Sinopyrophorinae are easily recognizable by the
strongly protruding frontoclypeal region (Fig. 4), which is medially distinctly longi-
tudinally carinate, antennomeres II and III subequal in length and together less than
half as long as antennomeres IV–XI, pronotal hind angles (Fig. 8) acute, produced
posterolaterally, prosternal process (Fig. 8c) straight in lateral view, tarsomeres III and
IV (Fig. 12) with ventral lobes, abdomen with seven (male) or six (female) ventrites,
with a luminous organ (Fig. 16) on sternite II, and aedeagus (Fig. 20) with a median
lobe shorter than phallobase, and arcuate parameres.
Phylogenetic placement and morphology of Sinopyrophorus schimmeli Bi & Li,
gen. et sp. nov.
He et al. (2019) provided the rst preliminary phylogenetic hypothesis for S. schimmeli
based on the analysis of 13 protein-coding mitochondrial genes. ey found this spe-
cies sister to a clade containing Dendrometrinae and Elaterinae; however, their sam-
pling of the click-beetle lineages was limited and the position of S. schimmeli was not
statistically supported.
Here, we used the most comprehensive dataset of Elateridae to date to elucidate
the phylogenetic position of the rst known Asian luminescent species. Our analysis
(Fig. 1) placed S. schimmeli in a clade with Hemiops and Oestodes, which are both the
type genera of the subfamilies Hemiopinae and Oestodinae, respectively, and which
were not included in the study by He et al. (2019). However, the relationships among
these lineages are not statistically supported and the long branches (see Kundrata et al.
Wen-Xuan Bi et al. / ZooKeys 864: 79–97 (2019)
2016) indicate a long-term independent evolution of these groups, which is supported
by their distinct morphology. Sinopyrophorinae dier from both Hemiopinae and
Oestodinae by having the strongly protruding frontoclypeal region with median longi-
tudinal carina, the antennomeres II and III subequal in length and much shorter than
remaining antennomeres (present only in the hemiopine genus Plectrosternus Lacord-
aire), the posterior margin of pronotum simple, without sublateral incisions or carinae,
the prosternal process straight in lateral view, the posterior end of scutellar shield emar-
ginate, the wide mesoventrite with an elongate mesoventral cavity which is only gradu-
ally narrowed posteriorly, the bisetose empodium (present in Oestodes, multisetose in
Hemiopinae), the abdomen with six or seven ventrites and with a luminous organ, the
male genitalia with a median lobe shorter than parameres, and each paramere strongly
curved mesad with a dentate inner margin, the female genitalia with a more sclerotized
ovipositor, longer paraprocts and styli attached subapically to gonocoxites (apically
in Oestodes and Hemiopinae; A. Prosvirov, pers. comm.). ese morphological dier-
ences in combination with the molecular phylogeny support the subfamilial rank of
Sinopyrophorinae, especially when there are no obvious synapomorphies which would
dene the clade of Sinopyrophorinae + Oestodinae + Hemiopinae, or placement of
Sinopyrophorus in any other subfamily. Since the backbone of the Elateridae phyloge-
netic tree is not or only weakly supported (Kundrata et al. 2016, 2018; this study),
future genomic or transcriptomic data might help to resolve the relationships among
the deep splits including Sinopyrophorinae.
Our molecular analysis in combination with the morphological investigation con-
rmed that S. schimmeli does not belong to any of described subfamilies containing
bioluminescent species. e clade of Sinopyrophorinae, Oestodinae and Hemiopinae
formed one of the basal radiations in Elateridae, far from the Agrypninae: Pyroph-
orini, which contains the majority of luminescent click beetles (Fig. 1). e position
of S.schimmeli far from Pyrophorini is additionally supported by the morphological
features such as the pretarsal claws without setae at base (present in Pyrophorini),
hind wings with a well-dened wedge cell (absent in Pyrophorini), abdomen with six
or seven ventrites (ve in Pyrophorini) and the presence of styli on the gonocoxites of
ovipositor (styli absent in Pyrophorini). ylacosterninae, which include the lumines-
cent Balgusschnusei, are regularly recovered inside Lissominae in recent DNA-based
analyses (Kundrata et al. 2014, 2016, 2018; Bocak et al. 2018; this study). ey dif-
fer from Sinopyrophorinae in having the abellate antennae, deep antennal cavities
lying beneath the hypomera, and membranous tarsal lobes (Costa 1984a, Costa et
al. 2010). e members of the subfamily Campyloxeninae, which have not been in-
cluded in any DNA-based study to date, dier from Sinopyrophorinae by much wider
frontoclypeal region with a complete frontal carina (frontoclypeal region strongly pro-
truding, relatively narrow and high, longitudinally carinate medially in S.schimmeli),
narrowed apical portions of lateral lobes of mesoventrite (wide in S. schimmeli) oval-
shaped mesoventral cavity (elongate, gradually narrowed posteriorly in S. schimmeli),
abdomen with ve ventrites and without a luminous organ (six or seven ventrites,
and a large abdominal luminous organ in S. schimmeli), distinctly longer paraprocts,
dierently shaped aedeagus with the median lobe longer than parameres, and the
Sinopyrophorinae, a new subfamily of Elateridae (Coleoptera, Elateroidea) with the rst ... 91
paramere elongate, with a subapical hook, and apex oriented posteriorly (Costa 1975,
Arias-Bohart 2015).
Southern China including Yunnan is a world biodiversity hotspot (Myers et al.
2000) and hosts many endemic beetle species including Elateridae. Intensive entomo-
logical research in southern China and neighboring regions is necessary to nd out
if the newly established Sinopyrophorinae contains more species. Additionally, the
discovery of a larva of S. schimmeli may help to better understand the morphological
distinctiveness and systematic position of this interesting beetle lineage.
Evolution of bioluminescence in Elateridae
Bioluminescence has evolved independently many times in various organisms (Day et
al. 2004). Within Coleoptera, bioluminescence is best known in Lampyridae (reies),
but also present in Phengodidae, Rhagophthalmidae, and some Elateridae, all within
the superfamily Elateroidea (e.g., Oba 2009). Additionally, it was reported also for the
larvae of two species belonging to Staphylinidae (Costa et al. 1986, Rosa 2010), which
are not closely related to Elateroidea (e.g., Zhang et al. 2018). Within the Elateroidea,
bioluminescence independently evolved several times (Bocakova et al. 2007, Sagegami-
Oba et al. 2007, Amaral et al. 2014, Fallon et al. 2018) but despite the recent progress
in elucidating the phylogenetic relationships within this superfamily (Kundrata et al.
2014), the relationships between the luminescent lineages remain unresolved.
Within Elateridae, nearly all luminescent taxa belong to the agrypnine tribe Py-
rophorini, and each of the remaining three smaller groups contain only a single bio-
luminescent species. All other bioluminescent subfamilies, except Sinopyrophorinae,
also include non-luminescent species. Campyloxeninae were placed into Pyrophorini by
Stibick (1979) but this was not accepted by later authors (see e.g., Costa et al. 2010). In
fact, the members of Campyloxeninae dier diagnostically from Pyrophorini in e.g., the
absent setae on tarsal claws, presence of the wedge cell in the hind wing venation and
the ovispositor with styli (Costa 1975, Arias-Bohart 2015). Current molecular phyloge-
ny suggests at least three independent origins of bioluminescence within Elateridae, i.e.,
in Agrypninae: Pyrophorini, Sinopyrophorinae and ylacosterninae. e presence of
a single luminous organ located on abdomen without any prothoracic bioluminescent
organs, which is shared by S. schimmeli and the pyrophorine genus Hifo, is therefore an
apparent homoplasy. Sequences of the fresh DNA-grade material of Campyloxeninae
would help us to better understand the position of this group and if it represents a
fourth origin of bioluminescence in Elateridae. All available phylogenetic analyses indi-
cate that the ancestral state of Elateridae was nonluminescent and the luminescence was
later obtained in several independent lineages (Bocakova et al. 2007, Sagegami-Oba et
al. 2007, Douglas 2011, Kundrata and Bocak 2011, Kundrata et al. 2014).
In Lampyridae, bioluminescence was rst gained by larvae as an aposematic warn-
ing display, and subsequently gained by adults and co-opted as a sexual signal; the over-
all trend of courtship is the use of pheromones in ancestral species, then pheromones
used in conjunction with photic signals, then the sole use of photic signal (Branham and
Wen-Xuan Bi et al. / ZooKeys 864: 79–97 (2019)
Wenzel 2001, 2003; Ohba 2004; Stanger-Hall et al. 2018). Males of S.schimmeli have
a large luminous organ on abdomen plus very long antennae (longer than in females,
hypothesized to be for pheromone detection) and large eyes (larger than in females, and
hypothesized to be for bioluminescence detection), which suggest the use of multiple
communication channels for mate attraction (Stanger-Hall et al. 2018, and references
therein). It is hypothesized that in elaterids, bioluminescence of the abdominal lantern
is an optical signal for the intraspecic sexual communication, while the signals from
the prothoracic lanterns (if present) serve to warn predators and may also provide illu-
mination in ight (Lall et al. 2000, 2010). Additionally, antennae are usually relatively
longer in species which rely greatly on pheromones as sexual signals (Stanger-Hall et al.
2018). erefore, the external morphology of S. schimmeli suggests the combined sexual
communication of both light signals and pheromones. e relatively larger sexual signal
sensors of S. schimmeli males in comparison to conspecic females are in agreement with
situation in many other insect groups, in which males are the more actively searching
sex, with more sensitive sensors (Steinbrecht 1987, Stanger-Hall et al. 2018).
e discovery of S. schimmeli as the rst record of a bioluminescent click beetle in Asia
shed new light on the geographic distribution and evolution of luminescent click beetles.
As a representative of a unique lineage, only distantly related to all other luminescent click
beetles, S. schimmeli may serve as a new model taxon in the research of bioluminescence
within Coleoptera. A project of de novo genome sequencing of S.schimmeli has already
started and should help answer the questions related to the genome characteristics of this
taxon (e.g., genome size, heterozygosity etc.), its genomic dierence from those of lumines-
cent pyrophorine click beetles, and the genomic basis of the origin of its bioluminescence.
We give special thanks to Mr. Yu-Tang Wang (Taiwan, China) for his proposal to collect
the specimens of Lampyridae, which enabled the discovery of the new luminescent click
beetle, and for his assistance in collecting specimens. We also thank Mr. Xiao-Dong
Yang (Sichuan, China) for collecting specimens, Dr. Zi-Wei Yin (SNUC, China) for
capturing the photomicrographs of the new species, Mr. Xing Chen (Yunnan, China)
for assembling nuclear ribosomal DNA, Dr. Alexander Prosvirov (Moscow, Russia) for
the photographs of genitalia of Hemiopinae and Oestodinae, Prof. Cleide Costa (São
Paulo, Brazil) for the discussion on the abdominal luminous organ in Pyrophorini,
Dr. Kathrin Stanger-Hall (Athens, USA) and Dr. Luiz Silveira (Athens, USA) for their
comments on earlier versions of the draft, and Dr. Hume Douglas (Ottawa, Canada)
for valuable comments leading to the improvement of our manuscript. We also thank
Prof. Wen Wang (Yunnan, China) and other members in his lab for their helps and
supports in this study. Special thanks are due to the late Mr. Rainer Schimmel (Vin-
ningen, Germany) for his valuable comments at the beginning of this study. is work
was supported by grants from the National Natural Science Foundation of China (No.
31472035), Yunnan Provincial Science and Technology Department (No. 2014FB179)
and Chinese Academy of Sciences (CAS “Light of West China” Program) to LXY.
Sinopyrophorinae, a new subfamily of Elateridae (Coleoptera, Elateroidea) with the rst ... 93
Amaral DT, Arnoldi FGC, Rosa SP, Viviani VR (2014) Molecular phylogeny of Neotropical
bioluminescent beetles (Coleoptera: Elateroidea) in southern and central Brazil. Lumines-
cence 29: 412–422.
Arias-Bohart ET (2015) Malalcahuello ocaresi gen. & sp. n. (Elateridae, Campyloxeninae).
ZooKeys 508: 1–13.
Bankevich A, Nurk S, Antipov D, Gurevich AA, Dvorkin M, Kulikov AS, Lesin VM, Niko-
lenko SI, Pham S, Prjibelski AD, Pyshkin AV, Sirotkin AV, Vyahhi N, Tesler G, Alekseyev
MA, Pevzner PA (2012) SPAdes: A new genome assembly algorithm and its applications
to single-cell sequencing. Journal of Computational Biology 19: 455–477. https://doi.
Bocak L, Motyka M, Bocek M, Bocakova M (2018) Incomplete sclerotization and phylogeny:
e phylogenetic classication of Plastocerus (Coleoptera: Elateroidea). PLoS ONE 13:
Bocakova M, Bocak L, Hunt T, Teräväinen M, Vogler AP (2007) Molecular phylogenetics
of Elateriformia (Coleoptera): evolution of bioluminescence and neoteny. Cladistics 23:
Branham MA, Wenzel JW (2001) e evolution of bioluminescence in canthar-
oids (Coleoptera: Elateroidea). Florida Entomologist 84: 565–586. https://doi.
Branham MA, Wenzel JW (2003) e origin of photic behavior and the evolution of sexual
communication in reies (Coleoptera: Lampyridae). Cladistics 19: 1–22. https://doi.
Calder AA (1996) Click Beetles: Genera of Australian Elateridae (Coleoptera). Monographs
on Invertebrate Taxonomy. CSIRO Publishing, Collingwood, 401 pp. https://doi.
Costa C (1975) Systematics and evolution of the tribes Pyrophorini and Heligmini with de-
scription of Campyloxeninae, new subfamily (Coleoptera, Elateridae). Arquivos de Zoolo-
gia 26: 49–190.
Costa C (1984a) Note on the bioluminescence of Balgus schnusei (Heller, 1974) (Trixagidae,
Coleoptera). Revista Brasileira de Entomologia 28: 397–398.
Costa C (1984b) On the systematic position of Hifo Candèze, 1881 (Elateridae, Coleoptera).
Revista Brasileira de Entomologia 28: 399–402.
Costa C, Vanin SA, Colepicolo Neto P (1986) Larvae of Neotropical Coleoptera. XIV. First
record of bioluminescence in the family Staphylinidae (Xantholinini). Revista Brasileira de
Entomologia 30: 101–104.
Costa C, Lawrence JF, Rosa SP (2010) Elateridae Leach, 1815. In: Leschen RAB, Beu-
tel RG, Lawrence JF (Eds) Handbook of Zoology, Coleoptera, Beetles, vol 2: Mor-
phology and Systematic (Elateroidea, Bostrichiformia, Cucujifomia partim). Wal-
ter de Gruyter GmbH & Co. KG, Berlin and New York, 75–103. https://doi.
Day JC, Tisi LC, Bailey MJ (2004) Evolution of beetle bioluminescence: the origin of beetle
luciferin. Luminescence 19: 8–20.
Wen-Xuan Bi et al. / ZooKeys 864: 79–97 (2019)
Douglas H (2011) Phylogenetic relationships of Elateridae inferred from adult morphology,
with special reference to the position of Cardiophorinae. Zootaxa 2900: 1–45. https://doi.
Douglas H, Kundrata R, Janosikova D, Bocak L (2018) Molecular and morphological evidence
for new genera in the click-beetle subfamily Cardiophorinae (Coleoptera: Elateridae). En-
tomological Science 21: 292–305.
Fallon TR, Lower SE, Chang C, Bessho-Uehara M, Martin GJ, Bewick AJ, Behringer M,
Debat HJ, Wong I, Day JC, Suvorov A, Silva CJ, Stanger-Hall KF, Hall DW, Schmitz RJ,
Nelson DR, S.M. L, Shigenobu S, Bybee SM, Larracuente AM, Oba Y, Weng JK (2018)
Firey genomes illuminate parallel origins of bioluminescence in beetles. Elife 7: e36495.
Hach F, Sarra I, Hormozdiari F, Alkan C, Eichler EE, Sahinalp SC (2014) MrsFAST-Ultra: a
compact, SNP-aware mapper for high performance sequencing applications. Nucleic Acids
Research 42: W494–W500.
He JW, Bi WX, Dong ZW, Liu GC, Zhao RP, Wang W, Li XY (2019) e mitochondrial genome
of the rst luminous click-beetle (Coleoptera: Elateridae) recorded in Asia. Mitochondrial
DNA Part B-Resources 4: 565–567.
Kolpakov R, Bana G, Kucherov G (2003) Mreps: Ecient and exible detection of tandem re-
peats in DNA. Nucleic Acids Research 31: 3672–3678.
Kumar S, Stecher G, Tamura K (2016) MEGA7: Molecular Evolutionary Genetics Analysis
version 7.0 for bigger datasets. Molecular Biology and Evolution 33: 1870–1874. https://
Kukalova-Peck J, Lawrence JF (1993) Evolution of the hind wing in Coleoptera. Canadian
Entomologist 125: 181–258.
Kukalova-Peck J, Lawrence JF (2004) Relationships among coleopteran suborders and major
endoneopteran lineages: Evidence from hind wing characters. European Journal of Ento-
mology 101: 95–144.
Kundrata R, Bocak L (2011) e phylogeny and limits of Elateridae (Insecta, Coleoptera):
is there a common tendency of click beetles to soft-bodiedness and neoteny? Zoologica
Scripta 40: 364–378.
Kundrata R, Bocakova M, Bocak L (2014) e comprehensive phylogeny of the superfam-
ily Elateroidea (Coleoptera: Elateriformia). Molecular Phylogenetics and Evolution 76:
Kundrata R, Gunter NL, Douglas H, Bocak L (2016) Next step toward a molecular phylogeny
of click-beetles (Coleoptera: Elateridae): redenition of Pityobiinae, with a description
of a new subfamily Parablacinae from the Australasian Region. Austral Entomology 55:
Kundrata R, Gunter NL, Janosikova D, Bocak L (2018) Molecular evidence for the subfamilial
status of Tetralobinae (Coleoptera: Elateridae), with comments on parallel evolution of
some phenotypic characters. Arthropod Systematics & Phylogeny 76: 137–145.
Kuraku SC, Zmasek M, Nishimura O, Katoh K (2013) aLeaves facilitates on-demand explora-
tion of metazoan gene family trees on MAFFT sequence alignment server with enhanced
interactivity. Nucleic Acids Research 41: W22–W28.
Sinopyrophorinae, a new subfamily of Elateridae (Coleoptera, Elateroidea) with the rst ... 95
Lall AB, Cronin TW, Carvalho AA, de Souza JM, Barros MP, Stevani CV, Bechara EJH, Ventu-
ra DF, Viviani VR, Hill AA (2010) Vision in click beetles (Coleoptera: Elateridae): pig-
ments and spectral correspondence between visual sensitivity and species bioluminescence
emission. Journal of Comparative Physiology A - Neuroethology Sensory Neural and Be-
havioral Physiology 196: 629–638.
Lall AB, Ventura DSF, Bechara EJH, de Souza JM, Colepicolo-Nito P, Viviani VR (2000)
Spectral correspondence between visual spectral sensitivity and bioluminescence emission
spectra in the click beetle Pyrophorus punctatissimus (Coleoptera : Elateridae). Journal of
Insect Physiology 46: 1137–1141.
Lanfear R, Calcott B, Ho SYW, Guindon S (2012) PartitionFinder: Combined selection of
partitioning schemes and substitution models for phylogenetic analyses. Molecular Biology
and Evolution 29: 1695–1701.
Lanfear R, Frandsen PB, Wright AM, Senfeld T, Calcott B (2017) PartitionFinder 2: New
methods for selecting partitioned models of evolution for molecular and morphologi-
cal phylogenetic analyses. Molecular Biology and Evolution 34: 772–773. https://doi.
Letunic I, Bork P (2016) Interactive tree of life (iTOL) v3: an online tool for the display and
annotation of phylogenetic and other trees. Nucleic Acids Research 44: W242–W245.
Myers N, Mittermeier RA, Mittermeier CG, da Fonseca GAB, Kent J (2000) Biodiversity hot-
spots for conservation priorities. Nature 403: 853–858.
NCBI Resource Coordinators (2016) Database resources of the National Center for Biotechnolo-
gy Information. Nucleic Acids Research 44: D7–D19.
Rosa SP (2010) Second record of bioluminescence in larvae of Xantholinus Dejean (Staphyli-
nidae, Xantholinini) from Brazil. Revista Brasileira de Entomologia 54: 147–148. https://
Oba Y (2009) On the origin of beetle luminescence. In: Meyer-Rochow VB (Ed) Biolumines-
cence in focus - A collection of illuminating essays. Research Signpost, Kerala, 277–290.
Ohba N (2004) Flash communication systems of Japanese reies. Integrative and Compara-
tive Biology 44: 225–233.
Sagegami-Oba R, Oba Y, Ôhira H (2007) Phylogenetic relationships of click beetles (Coleop-
tera: Elateridae) inferred from 28S ribosomal DNA: Insights into the evolution of biolumi-
nescence in Elateridae. Molecular Phylogenetics and Evolution 42: 410–421. https://doi.
Stamatakis A, Hoover P, Rougemont J (2008) A rapid bootstrap algorithm for the RAxML web
servers. Systematic Biology 57: 758–771.
Stanger-Hall KF, Lower SES, Lindberg L, Hopkins A, Pallansch J, Hall DW (2018) e evolu-
tion of sexual signal modes and associated sensor morphology in reies (Lampyridae, Co-
leoptera). Proceedings of the Royal Society B-Biological Sciences 285: 20172384. https://
Steinbrecht RA (1987) Functional morphology of pheromone-sensitive sensilla. In: Prestwich
GD, Blomquist GJ (Eds) Pheromone biochemistry. Academic Press, Orlando, FL, 353–
Wen-Xuan Bi et al. / ZooKeys 864: 79–97 (2019)
Stibick JNL (1979) Classication of the Elateridae (Coleoptera) - Relationships and classica-
tion of the subfamilies and tribes. Pacic Insects 20: 145–186.
Vaidya G, Lohman DJ, Meier R (2011) SequenceMatrix: concatenation software for the fast
assembly of multi-gene datasets with character set and codon information. Cladistics 27:
Viviani VR (2002) e origin, diversity, and structure function relationships of insect lucif-
erases. Cellular and Molecular Life Sciences 59: 1833–1850.
Zhang SQ, Che LH, Li Y, Liang D, Pang H, Ślipiński A, Zhang P (2018) Evolutionary history
of Coleoptera revealed by extensive sampling of genes and species. Nature Communica-
tions 9: 205.
Supplementary material 1
Table S1.
Authors: Jin-Wu He
Data type: molecular data
Explanation note: e GenBank accession numbers of mitochondrial genes (PCGs
and 16S) and nuclear rRNA genes (18S and 28S) in Elateridae and outgroup used
for the phylogenetic analysis and references in rRNA assembly.
Copyright notice: is dataset is made available under the Open Database License
( e Open Database License
(ODbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.
Supplementary material 2
Movie S1.
Authors: Wen-Xuan Bi
Data type: multimedia
Explanation note: Luminescent behavior of Sinopyrophorus schimmeli.
Copyright notice: is dataset is made available under the Open Database License
( e Open Database License
(ODbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.
Sinopyrophorinae, a new subfamily of Elateridae (Coleoptera, Elateroidea) with the rst ... 97
Supplementary material 3
Sequence/Matrix data.
Copyright notice: is dataset is made available under the Open Database License
( e Open Database License
(ODbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.
... Within Coleoptera, bioluminescence can be found almost exclusively within the so-called "elaterid-lampyroid clade", including Elateridae, Lampyridae, Phengodidae, Rhagophthalmidae, and Sinopyrophoridae, and probably the extinct Cretophengodidae (Oba et al. 2011;Fallon et al. 2018;Bi et al. 2019;Li et al. 2021b;Kusy et al. 2021;Powell et al. 2022). In Phengodidae, all known larvae and females are bioluminescent, as are males of some species (Costa and Zaragoza-Caballero 2010). ...
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 typical representatives have a compact body and a pro-mesothoracic clicking mechanism (Costa et al., 2010); however, this group also includes several soft-bodied lineages (Kundrata and Bocak, 2019). Despite recent progress in understanding the composition, phylogeny, and classification of Elateridae, all these aspects remain 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 Elateridae (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 Elateridae clade comprises more than 13 500 extant species worldwide (Costa et al., 2010;Douglas et al., 2021). ...
Full-text available
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 character among fossil Elateridae. We discuss the diagnostic characters of Cretopachyderes gen. nov. and tentatively place it to subfamily Agrypninae close to extant genus Pachyderes Guérin-Méneville, 1829.
... The elaterid subfamily Sinopyrophorinae, based on Sinopyrophorus schimmeli Bi & Li in Bi et al. (2019) from Yunnan, China, represents the first bioluminescent click beetle from Asia. In a cladogram based on 14 mitochondrial genes and a Maximum Likelihood analysis, the group formed part of a clade including Hemiopinae and Oestodinae and far removed from other bioluminescent elaterids (Balgus within Lissominae and a terminal agrypnine clade). ...
The hind wings of all known families and most subfamilies of Coleoptera are illustrated, annotated and discussed utilising the terminology of Kukalová-Peck and Lawrence (2004), with a few changes in nomenclature suggested by the senior author. The beetle families are discussed in 21 groups, based on recent classifications of Coleoptera. For each of these groups, the most recent works on phylogeny and classification are reviewed, and the wing characters are discussed to determine if some of the wing features might support or refute relationships based on recent molecular and morphological analyses. Part 1 includes a general discussion of wing structure divided into the following sections: hind wing fields, veinal systems (including the history of wing nomenclature), wing folding, wing edge and embayments, hinges and bending zones, cross-veins and braces, cells and other landmarks. It is followed by discussion of the first 14 groups (Archostemata to Elateroidea), 15 figures supporting general discussions, and 426 labelled wing images of the discussed groups, representing 380 genera.
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The soft-bodied click-beetle genus Malacogaster Bassi, 1834 from the Mediterranean region has never been taxonomically revised to date. Information on its morphology, intra- and interspecific variability, systematics and distribution is fragmented and most species have not been properly studied since their description. Therefore, in this study we summarize all available information on the genus Malacogaster. Altogether, we recognize 10 valid species from the area including the Canary Islands, Iberian Peninsula, Balearic Islands, northern coast of Africa, Sardinia, and Sicily. Malacogaster ruficollis Dodero, 1925, stat. nov., which was originally described as a variety of M. bassii Lucas, 1870 and later synonymized with it, is considered a separate species. Malacogaster parallelocollis Reitter, 1894, syn. nov. and M. olcesei var. reductus Pic, 1951, syn. nov. are synonymized with M. maculiventris Reitter, 1894. Malacogaster notativentris Pic, 1951, syn. nov. and M. olcesei Pic, 1951, syn. nov. are synonymized with M. passerinii Bassi, 1834. Lectotypes are designated for M. maculiventris Reitter, 1894, M. nigripes heydeni Reitter, 1894, M. parallelocollis Reitter, 1894, M. thoracica Redtenbacher, 1858, M. olcesei Pic, 1951, and M. rubripes Peyerimhoff, 1949 to fix their identity.
Fireflies are one of the best-known bioluminescent organisms, and the reaction mechanism and ecological utility of bioluminescence have been well-studied. Genome assemblies of six species of bioluminescent beetles have recently been published. These studies have focused on the evolution of novelties; luciferase, and the biosynthesis of luciferin and defensive chemicals. For example, clustering of the luciferase gene with acyl-CoA synthetase genes on a chromosome in luminous beetle genomes suggests the involvement of tandem gene duplications and neofunctionalization during the evolution of beetle bioluminescence. Several candidate genes for critical roles in beetle bioluminescence have been identified, but their functional analyses are still ongoing. The establishment of a long-term mass-rearing system and strain will be the key for the post-genome research on bioluminescent beetles. Lastly, the application of contemporary chromosome-scale genome assembly techniques to luminous beetles will help resolve outstanding evolutionary questions, such as how many times bioluminescence evolved in this clade.
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We here report a new elateroid, Anoeuma lawrencei Li, Kundrata and Cai gen. et sp. nov., from mid-Cretaceous Burmese amber. Though superficially similar to some soft-bodied archostematans, Anoeuma could be firmly placed in the polyphagan superfamily Elateroidea based on the hind wing venation. Detailed morphological comparisons between extant elateroids and the Cretaceous fossils suggest that the unique character combination does not fit with confidence into any existing soft-bodied elateroid group, although some characters indicate possible relationships between Anoeuma and Omalisinae. Our discovery of this new lineage further demonstrates the past diversity and morphological disparity of soft-bodied elateroids.
A new species of the genus Elathous Reitter, 1890 from Lebanon is described and illustrated. Elathous nemeri sp. nov. is similar to other Levantine species, but can be easily recognised from its congeners by its elongated pronotum as well as by the shape of the male genitalia.
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Background Fireflies (Coleoptera: Lampyridae) are commonly recognized by adult traits, such as a soft exoskeleton, lanterns and associated glow and flash patterns, but their larval stage is far less appreciated. However, fireflies spend most of their lives as larvae, and adults of most species rely solely on resources previously obtained. Therefore, studying the immature stages is imperative towards a comprehensive understanding of fireflies. This paper reviews and indicates key gaps in the biology of firefly larvae based on available literature. Methodology We reviewed the literature on firefly larvae to identify key issues and important taxonomic, geographic, and subject biases and gaps. Results We found 376 papers that included information on firefly larvae. Only 139 species in 47 genera across eight of eleven lampyrid subfamilies have been studied during larval stages. These numbers reveal a staggering gap, since 94% of species and over half of the genera of fireflies were never studied in a crucial stage of their life cycle. Most studies on firefly larvae focus on two subfamilies (Luciolinae and Lampyrinae) in four zoogeographic regions (Sino-Japanese, Oriental, Nearctic, and Palearctic), whereas the other subfamilies and regions remain largely unstudied. These studies mainly dealt with morphology and behavior, other subjects remaining greatly understudied by comparison, including habitats, life cycle, physiology and interactions. Conclusions Together, these literature biases and gaps highlight how little is known about firefly larvae, and warmly invite basic and applied research, in the field and in the lab, to overcome these limitations and improve our understanding of firefly biology to better preserve them.
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Bioluminescence is the emission of of perceptible cold light from organisms; therefore, they are recognized as living lights. Bioluminescent organisms are present everywhere on the earth predominately in the ocean. The terrestrial bioluminescent organism includes very well‐known Fireflies; microorganisms like bacteria; insects like flies and springtails. Mushrooms, earthworms, snail, centipedes, millipedes and beetles are some other terrestrial bioluminescent organisms. Attracting prey, intraspecies visual communication and escape from a predator are some major significance of bioluminescence in living organisms. Bioluminescent organisms produce light by the chemical reaction; therefore, it is known as chemiluminescence. Luciferase catalyzes the chemical oxidation of the substrate (luciferin) which leads to the emission of light. About 40 bioluminescence systems have been reported on the earth in which D‐luciferin, coelenterazine and Cypridina luciferin‐based systems are well studied. About 81 known species of bioluminescent fungi are recorded worldwide. These fungi belong to four dissimilar lineages – Omphalotus, Armillaria, mycenoid and Lucentipes. Lightening, monitoring biological process, clinical diagnosis, drug discovery, cancer research, biosensor development and immunoassays are the future scope of bioluminescence technology. Future lighting is the most exciting scope of bioluminescence technology. Recently, eukaryotic yeast cell autonomously glowed after the transfer of a genetically encoded system of bioluminescent fungus Neonothopanus nambi . The bioluminescence system of the same fungus was used to produce an engineered tobacco plant that also has the self glowing property. The fungal bioluminescence system was also successfully used as a biosensor for the environmental monitoring of mercury, copper and zinc. The virulence and growth of some fungi like Candida albicans, Aspergillus fumigatus and Cryptococcus neoformans were real‐time monitored using bioluminescence‐based non‐invasive method in vivo. It is known as bioluminescence imaging (BLI) that allows the temporal and spatial progression of disease in living beings. So, the present chapter aims to give insights about bioluminescence in fungi, its mechanism and future scope.
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We present the latest version of the Molecular Evolutionary Genetics Analysis (MEGA) software, which contains many sophisticated methods and tools for phylogenomics and phylomedicine. In this major upgrade, MEGA has been optimized for use on 64-bit computing systems for analyzing bigger datasets. Researchers can now explore and analyze tens of thousands of sequences in MEGA. The new version also provides an advanced wizard for building timetrees and includes a new functionality to automatically predict gene duplication events in gene family trees. The 64-bit MEGA is made available in two interfaces: graphical and command line. The graphical user interface (GUI) is a native Microsoft Windows application that can also be used on Mac OSX. The command line MEGA is available as native applications for Windows, Linux, and Mac OSX. They are intended for use in high-throughput and scripted analysis. Both versions are available from free of charge.
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The nearly complete mitochondrial genome (mitogenome) of Sinopyrophorus schimmeli Bi et Li, the luminous click beetle recorded in Asia, is described in this study. It totalizes 15,951 bp and contains 13 protein-coding genes (PCGs), two rRNA genes, 22 tRNA genes, and most part of AT-rich region. Thirteen PCGs totalize 11,136 bp, start with ATN, stop with TAA/G, except for cox2 and cox3 stopping with T. The rrnL and rrnS are 1280 and 862 bp, respectively. The AT-rich region contains several structures characteristic of the Coleoptera. The phylogenetic analyses of 13 PCGs confirm the position of S. schimmeli in Elateridae.
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Fireflies and their luminous courtships have inspired centuries of scientific study. Today firefly luciferase is widely used in biotechnology, but the evolutionary origin of bioluminescence within beetles remains unclear. To shed light on this long-standing question, we sequenced the genomes of two firefly species that diverged over 100 million-years-ago: the North American Photinus pyralis and Japanese Aquatica lateralis. To compare bioluminescent origins, we also sequenced the genome of a related click beetle, the Caribbean Ignelater luminosus, with bioluminescent biochemistry near-identical to fireflies, but anatomically unique light organs, suggesting the intriguing hypothesis of parallel gains of bioluminescence. Our analyses support independent gains of bioluminescence in fireflies and click beetles, and provide new insights into the genes, chemical defenses, and symbionts that evolved alongside their luminous lifestyle.
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Tetralobinae is a distinct click-beetle lineage containing 78 species in seven genera. Adults are large-bodied, and larvae live in termite nests and are grub-like unlike typical elaterid wireworms. Their taxonomic position in the Elateridae has been unstable and they were treated either as a separate elaterid subfamily or a tribe within Agrypninae. Here, we provide the first molecular investigation of Tetralobinae to test their phylogenetic position using two nuclear and two mitochondrial molecular markers from three total taxa, one from each of the following genera: Tetralobus Lepeletier & Audinet-Serville, Sinelater Laurent, and Pseudotetralobus Schwarz. Two different datasets were analyzed, Elateridae (181 terminals) and Elateroidea (451 terminals), both composed by the earlier published datasets supplemented with the newly produced tetralobine sequences. The results suggest that Tetralobinae is the sister lineage to the remaining Elateridae and that warrants the subfamilial status instead of an subordinate position in the Agrypninae. Pseudotetralobus (Australia) was sister to the Tetralobus (Africa) + Sinelater (China) consistent with previously published morphological analysis. Additionally, we discuss the homoplastic phenotypic characters which were used for building the earlier click-beetle classification, and which indicated the relationships between Tetralobinae and Agrypninae.
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The Cardiophorinae is consistently at the subfamily rank in recent classifications. Conversely, generic limits and relationships remain unstable. Here, we use two nuclear (18S and 28S) and three mitochondrial (rrnL, cox1–3′ and cox1–5′) markers to test phylogenetic hypotheses for the Cardiophorinae. We investigate the positions of Paracardiophorus buettikeri Chassain from the Arabian Peninsula and a simple‐clawed Argentinian Horistonotus Candèze species, neither of which matches the diagnoses of their assigned genera. Additionally, we test the monophyly of the widely defined Cardiophorus Eschscholtz by including representatives of Cardiophorus (Cardiophorus), Cardiophorus (Coptostethus Wollaston) and the former subgenus Zygocardiophorus Iablokoff‐Khnzorian and Mardjanian. Molecular analyses recover three clades, which are also supported by morphological traits. Using inferred relationships we propose three new genera: Arandelater Douglas and Kundrata gen. nov., Chassainphorus Douglas and Kundrata gen. nov., and Huarpelater Douglas and Kundrata gen. nov. for Horistonotus canescens Steinheil, P. buettikeri and an undescribed species perhaps historically mistaken for H. canescens. Horistonotus tumidicollis Schwarz is synonymized under H. canescens. Coptostethus is raised from a subgenus of Cardiophorus to the genus level with revision of its definition. Eighteen Cardiophorus (Cardiophorus) and Dicronychus Brullé species are newly assigned to Coptostethus. All species currently included in Coptostethus are listed. All Coptostethus species from South Africa are transferred to Cardiophorus (Cardiophorus). Huarpelater cordobae Douglas and Kundrata sp. nov. is described, and Horistonotus quillu Aranda is transferred to Huarpelater gen. nov. The replacement name Coptostethus cobosi nom. nov. and comb. nov. is proposed for Cardiophorus inflatus Cobos. We provide diagnoses and illustrated descriptions for the new genera and their types.
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The relationships of the monogeneric family Plastoceridae Crowson, 1972 (Coleoptera: Elateroidea) have remained contentious due to its modified morphology, incorrect information on incomplete metamorphosis of females and the absence of molecular data. We produced the sequences for P. angulosus (Germar, 1844) (the type-species of Plastocerus Schaum, 1852) and performed molecular phylogenetic analyses to estimate its position. The analyses of Elateroidea (186 spp.) and Elateridae (110 spp.) molecular datasets of two mitochondrial and two nuclear gene fragments repeatedly placed Plastocerus Schaum, 1852 in relationships with the elaterid genera Oxynopterus Hope, 1842 and Pectocera Hope, 1842. Alternative topologies were rejected by likelihood tests. Therefore, Plastoceridae Crowson, 1972 are down-ranked to the subfamily Plastocerinae in Elateridae Leach, 1815. We suggest that the morphology-based placement and high rank for some elateroid lineages were inferred from the presence of homoplasies which evolved due to incomplete sclerotization. Distantly related soft-bodied elateroids share freely movable and transverse coxae, a shortened prosternum, and a weakly sclerotized abdomen with free ventrites. Importantly, the apomorphic structures characteristic for their closest relatives, such as the prosternal process, mesoventral cavity, and intercoxal keel in the first abdominal ventrite are regularly absent. Consequently, morphology-based phylogenetic analyses suggest deeply rooted positions for lineages without expressed apomorphic character states. Molecular data represent an independent character system that is not affected by the convergent morphological evolution, and therefore molecular phylogenies can elucidate the relationships of incompletely sclerotized lineages.
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Beetles (Coleoptera) are the most diverse and species-rich group of insects, and a robust, time-calibrated phylogeny is fundamental to understanding macroevolutionary processes that underlie their diversity. Here we infer the phylogeny and divergence times of all major lineages of Coleoptera by analyzing 95 protein-coding genes in 373 beetle species, including ~67% of the currently recognized families. The subordinal relationships are strongly supported as Polyphaga (Adephaga (Archostemata, Myxophaga)). The series and superfamilies of Polyphaga are mostly monophyletic. The species-poor Nosodendridae is robustly recovered in a novel position sister to Staphyliniformia, Bostrichiformia, and Cucujiformia. Our divergence time analyses suggest that the crown group of extant beetles occurred ~297 million years ago (Mya) and that ~64% of families originated in the Cretaceous. Most of the herbivorous families experienced a significant increase in diversification rate during the Cretaceous, thus suggesting that the rise of angiosperms in the Cretaceous may have been an 'evolutionary impetus' driving the hyperdiversity of herbivorous beetles.
This is the first monograph on the Australian genera of Elateridae - click beetles. The book deals with 74 genera, among which 14 are newly recognised. The volume documents the entire Australian fauna and provides lavish illustrations of representative species, and typical examples of the male and female genitalia for each genus. The phylogeny of the genera is analysed and there is a checklist of all described species and appropriate bibliographic and type locality details are given.
Animals employ different sexual signal modes (e.g. visual, acoustic, chemical) in different environments and behavioural contexts. If sensory structures are costly, then evolutionary shifts in primary signal mode should be associated with changes in sensor morphology. Further, sex differences are expected if male and female signalling behaviours differ. Fireflies are known for their light displays, but many species communicate exclusively with pheromones, including species that recently lost their light signals. We performed phylogenetically controlled analyses of male eye and antenna size in 46 North American taxa, and found that light signals are associated with larger eyes and shorter antennae. In addition, following a transition from nocturnal light displays to diurnal pheromones, eye size reductions occur more rapidly than antenna size increases. In agreement with the North American taxa, across 101 worldwide firefly taxa in 32 genera, we found light displays are associated with larger eye and smaller antenna sizes in both males and females. For those taxa with both male and female data, we found sex differences in eye size and, for diurnal species, in antenna size.
PartitionFinder 2 is a program for automatically selecting best-fit partitioning schemes and models of evolution for phylogenetic analyses. PartitionFinder 2 is substantially faster and more efficient than version 1, and incorporates many new methods and features. These include the ability to analyze morphological datasets, new methods to analyze genome-scale datasets, new output formats to facilitate interoperability with downstream software, and many new models of molecular evolution. PartitionFinder 2 is freely available under an open source license and works on Windows, OSX, and Linux operating systems. It can be downloaded from The source code is available at