ZooKeys 1092: 19-30 (2022) A peer-reviewed open-access journal
doi: 10.3897/zookeys.1092.81701 RESEARCH ARTICLE $Z00Ke y S

https:/ / ZOO keys. pensoft.net Launched to accelerate biodiversity research

Nothotytthonyx, a new genus of Malthininae
(Coleoptera, Cantharidae) from
mid-Cretaceous amber of northern Myanmar

Yan-Da Li'?, Gabriel Biff?, Robin Kundrata*, Di-Ying Huang!', Chen-Yang Cai!”

I State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, and
Centre for Excellence in Life and Paleoenvironment, Chinese Academy of Sciences, Nanjing 210008, China
2 School of Earth Sciences, University of Bristol, Life Sciences Building, Tyndall Avenue, Bristol BS8 1TQ,
UK 3 Museu de Zoologia, Universidade de Séo Paulo, Av. Nazaré, 481— Ipiranga, 04263-000, Sao Paulo,
SP. Brazil 4 Department of Zoology, Faculty of Science, Palacky University, 77900 Olomouc, Czech Republic

Corresponding author: Chen-Yang Cai (cycai@nigpas.ac.cn)

Academic editor: Vinicius S. Ferreira | Received 4 February 2022 | Accepted 12 March 2022 | Published 4 April 2022
Attp://zoobank. org/688 1 6AB2-2FA3-4B8B-A7F8-4987 BAGDF5A8

Citation: Li Y-D, Biff G, Kundrata R, Huang D-Y, Cai C-Y (2022) Nothotytthonyx, a new genus of Malthininae
(Coleoptera, Cantharidae) from mid-Cretaceous amber of northern Myanmar. ZooKeys 1092: 19-30. https://doi.
org/10.3897/zookeys.1092.81701

Abstract

A new fossil genus and species of Cantharidae, Nothotytthonyx serratus Li, Bifh, Kundrata & Cai gen. et
sp. nov., is reported from mid-Cretaceous Burmese amber. The new species is tentatively attributed to
the extant subfamily Malthininae based on a combination of characters, including the symmetrical apical
maxillary palpomeres, shortened elytra, pronotum with arched margins and well-defined borders, tibiae
with apical spurs, and tarsal claws simple, although its well-developed gonostyli are atypical in Malthini-
nae. The discovery of Nothotytthonyx also suggests a possible Gondwanan origin for Malthininae.

Keywords

Burmese amber, Cretaceous, fossil, paleontology, soft-bodied Elateroidea, soldier beetles, systematics

Introduction

Cantharidae is a diverse group among the soft-bodied Elateroidea, with over 5000
species distributed worldwide (Ramsdale 2010). Cantharid adults are highly active,
and may feed on foliage-frequenting invertebrates, nectar or pollen (Crowson 1972;

Copyright Yan-Da Li 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.

20 Yan-Da Li et al. / ZooKeys 1092: 19-30 (2022)

Ramsdale 2010). Both larvae and adults of cantharids have paired lateral glandular
pores for chemical defense against predators, although the pores may sometimes be
inconspicuous and hard to determine. The family appears to be closely related to the
elaterid-lampyroid group. However, its accurate position remains unsettled, as several
recent phylogenomic studies have produced inconsistent results (Zhang et al. 2018;
McKenna et al. 2019; Douglas et al. 2021; Cai et al. 2022). Brancucci (1980) con-
ducted a comprehensive study on the taxonomy of Cantharidae, and divided it into
five subfamilies, namely Cantharinae, Chauliognathinae, Dysmorphocerinae, Silinae
and Malthininae. The relationships among the subfamilies varied in different studies
and were not well understood (e.g., Brancucci 1980; Kundrata et al. 2014; McKenna
et al. 2015, 2019; Zhang et al. 2018; Hsiao et al. 2021; Cai et al. 2022).

Records of fossil cantharids are relatively abundant, especially in amber deposits.
More than 80 species have been reported from Eocene Baltic amber (e.g., Fanti and
Kupryjanowicz 2017; Fanti and Damgaard 2020; Fanti 2021). Additional cantharids
have been described from Early Cretaceous Spanish amber (Peris and Fanti 2018), Late
Cretaceous Agdzhakend amber (Kazantsev and Perkovsky 2019a), Eocene Rovno am-
ber (Kazantsev and Perkovsky 2020), Eocene Sakhalinian amber (Kazantsev and Perk-
ovsky 2019b), and Miocene Dominican amber (Fanti and Damgaard 2019; Fanti and
Pankowski 2021). From the fossiliferous mid-Cretaceous Burmese amber, about 20
species have been described in nine genera (e.g., Hsiao et al. 2021; Yang et al. 2021).
Most cantharids from Burmese amber were originally assigned to Cantharinae diag-
nosed by, for example, securiform apical maxillary palpomeres, unmodified pronotal
margins and well-developed elytra concealing the wings and abdomen. However, the
subfamilial placement of at least some of the genera (Ornatomalthinus Poinar & Fanti
and Sanaungulus Fanti et al.) is controversial due to their conflicting diagnostic charac-
ters (Fanti 2018; Hsiao and Huang 2018; Hsiao et al. 2021). In the morphology-based
phylogenetic analysis by Hsiao et al. (2021), such group of genera was retrieved form-
ing the “Burmite Cantharinae” clade, as sister to Silinae, and their placement within
Cantharinae was rejected. In the same work, Archaeomalthodes Hsiao et al., originally
placed in Malthininae (Hsiao et al, 2017), was suggested as a member of Dysmorpho-
cerinae (Hsiao et al. 2021). Here, we describe a new fossil cantharid from Burmese
amber, and tentatively assign it to subfamily Malthininae.

Materials and methods

The Burmese amber specimen studied herein (Figs 1-5) originated from an amber
mine near Noije Bum (26°20'N, 96°36'E), Hukawng Valley, Kachin State, northern
Myanmar. Jewellery-grade Burmese amber specimens are commonly carried and sold
legally in Ruili, Dehong Prefecture on the border between China and Myanmar. The
specimen in this study was purchased in late 2016, and is permanently deposited in
the Nanjing Institute of Geology and Palaeontology (NIGP), Chinese Academy of Sci-
ences, Nanjing, China. The amber piece was trimmed with a small table saw, ground
with emery paper of different grit sizes, and finally polished with polishing powder.

New cantharid from mid-Cretaceous Burmese amber 21

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 combined with a fluorescence imaging system. Confocal images were
obtained with a Zeiss LSM710 confocal laser scanning microscope, using the 488 nm
(Argon) or 561 nm (DPSS 561-10) laser excitation lines (Fu et al. 2021). Images un-
der incident light and widefield fluorescence were stacked in Helicon Focus 7.0.2 or
Zerene Stacker 1.04. Confocal images were stacked with Helicon Focus 7.0.2 and
Adobe Photoshop CC. Microtomographic data were obtained with a Zeiss Xradia 520
Versa 3D X-ray microscope at the micro-CT laboratory of NIGP and analyzed in VG-
Studio MAX 3.0. Scanning parameters were as follows: isotropic voxel size, 6.1511 um;
power, 4 W; acceleration voltage, 50 kV; exposure time, 1.5 s; projections, 2401. Im-
ages were further processed in Adobe Photoshop CC to adjust brightness and contrast.

Data availability

The original confocal and micro-CT data are available in Zenodo repository (https://

doi.org/10.5281/zenodo.6336149).

Systematic paleontology

Order Coleoptera Linnaeus, 1758
Superfamily Elateroidea Leach, 1815
Family Cantharidae Imhoff, 1856
Subfamily Malthininae Kiesenwetter, 1852

Genus Nothotytthonyx Li, Biff, Kundrata & Cai, gen. nov.
http://zoobank.org/A56DECAD-2C71-4822-B48C-690DD67B4C3E

Type species. Nothotytthonyx serratus sp. nov.

Etymology. ‘The generic name is derived from the Greek “nothos”, false, and the
generic name Tytthonyx LeConte. The name is masculine in gender.

Diagnosis. Antennae strongly serrate (Figs 3F, 4C). Mandibles with a prominent
tooth on incisor edge (Fig. 5D). Apical maxillary palpomere symmetrical, fusiform
(Fig. 4A). Gular sutures confluent (Figs 3A, 5B). Elytra shortened; surface somewhat
punctate. Tibial spurs present (Fig. 4D). Gonostyli well developed (Fig. 4F).

Nothotytthonyx serratus Li, Bifh, Kundrata & Cai, sp. nov.
http://zoobank.org/44E 1AFDB-31BC-4B43-B8FA-D3F566934E3F
Figs 1-5

Material. Holotype, NIGP 179427, female.

22 Yan-Da Li et al. / ZooKeys 1092: 19-30 (2022)

.

Figure |. General habitus of Nothotytthonyx serratus Li, Bifh, Kundrata & Cai sp. nov., holotype,
NIGP179427, under incident light A dorsal view B ventral view. Scale bars: 1.5 mm.

Etymology. ‘The specific name refers to its distinctly serrate antennae.

Locality and horizon. Amber mine located near Noije Bum Village (26°20'N,
96°36'E), Tanai Township, Myitkyina District, Kachin State, Myanmar; unnamed ho-
rizon, mid-Cretaceous, Upper Albian to Lower Cenomanian.

Diagnosis. As for the genus.

Description. Adult female. Body weakly sclerotized, elongate, about 5.3 mm
long, 1.4 mm wide (widest across abdomen).

Head (Fig. 3A,F) fully exposed, prognathous, subquadrate, weakly narrowed pos-
teriorly, including eyes almost as wide as pronotum; dorsal surface flat, without protu-
berance or depression. Compound eyes moderately large and weakly protruding, finely
facetted, without interfacetal setae. Antennal insertions located anteriorly, dorsally ex-
posed, separated by approximately the length of antennomere 1. Subantennal grooves
absent. Antennae (Fig. 4B,C) with 11 antennomeres; antennomere 1 moderately
broad; antennomeres 2 short; antennomere 3—10 distinctly serrate. Mandibles with

New cantharid from mid-Cretaceous Burmese amber 23

one prominent tooth on incisor edge (Fig. 5D). Apical maxillary and labial palpomeres
elongate, symmetrical, fusiform, not unequally expanded, apex acute (Fig. 4A). Gular
sutures confluent (Figs 3A, 5B).

Pronotal disc (Fig. 3G) transverse; anterior and posterior angles broadly rounded;
lateral and posterior margins clearly bordered. Elytra (Fig. 3H) relatively short, cov-
ering only about half of posterior body; surface somewhat punctate. Procoxae (Fig.
5B) conical, well projecting, contiguous. Mesocoxae (Fig. 5B) conical, well project-
ing, narrowly separated. Metaventrite (Fig. 3C) large, with distinct discrimen and me-
takatepisternal suture. Metacoxae (Fig. 3C) transverse, almost contiguous.

Figure 2. General habitus of Nothotytthonyx serratus Li, Biff, Kundrata & Cai sp. nov., holotype,
NIGP179427, under widefield fluorescence A dorsal view. B ventral view. Scale bars: 1.5 mm.

24 Yan-Da Li et al. / ZooKeys 1092: 19-30 (2022)

Figure 3. Details of Nothotytthonyx serratus Li, Bifh, Kundrata & Cai sp. nov., holotype, NIGP179427,
under confocal microscopy A head, ventral view, showing the confluent gular suture (arrowhead) B pro-
thorax, ventral view C metathorax, ventral view D abdominal base, ventral view E abdominal apex,
ventral view F head, dorsal view G prothorax, dorsal view H elytral base, dorsal view H abdominal apex,
dorsal view. Abbreviations: anl—11, antennomeres 1-11; el, elytron; ey, compound eye; md, mandible;
msf, mesofemur; mtc, metacoxa; mttb, metatibia; mtts, metatarsus; mtv, metaventrite; mxp, maxillary

palp; pc, procoxa; pn, pronotum; ps, prosternum; vl-—7, ventrites 1—7. Scale bars: 300 pm.

Legs slender. Trochanters obliquely articulated to femoral bases. ‘Tibiae with weak
spurs (at least as seen on left mesotibia; Fig. 4D). Tarsal formula 5—5—5; tarsomere 4
ventrally bilobed (Figs 4D, 5E). Claws simple.

Abdomen with seven free ventrites. Gonostyli well developed (Fig. 4F).

New cantharid from mid-Cretaceous Burmese amber 25

Figure 4. Details of Nothotytthonyx serratus Li, Bifh, Kundrata & Cai sp. nov., holotype, NIGP179427,

under confocal microscopy A mouthparts, ventral view B antennal base, dorsal view C antennal apex

D mid leg, showing the two weak tibial spurs (arrowhead) E abdominal apex, dorsal view F ovipositor,
ventral view. Abbreviations: anl—11, antennomeres 1—11; gc, gonocoxite; gs, gonostylus; lbp, labial palp;

mstl—5, mesotarsomeres 1—5; mstb, mesotibia; mxp, maxillary palp. Scale bars: 200 pm.

Discussion

Within soft-bodied elateroids, Nothotytthonyx is firmly placed in Cantharidae, primar-
ily based on the fully exposed prognathous head (Fig. 3A,F), 11-segmented antennae
(Fig. 1), and ventrally bilobed tarsomere 4 (Figs 4D, 5E).

The current classification of Cantharidae into five subfamilies is solely based on
extant species (Brancucci 1980). However, there are no incontestable diagnostic char-
acters for most subfamilies (except for Chauliognathinae). Many characters may have
evolved independently in separate subfamilies, and some characters used for diagnosis
may be absent in certain lineages within a subfamily, which hampers the precise sys-
tematic placement of some genera (e.g., Tytthonyx). In the case of fossils, this problem
is aggravated by the impossibility of observation of important characters, especially the
genitalia and wing venation, leading to the conflicting hypotheses of placement (e.g.,
Fanti 2018; Hsiao and Huang 2018; Hsiao et al. 2021). For instance, Malthininae
and a few members of Dysmorphocerinae have radially symmetrical apical maxillary

26 Yan-Da Li et al. / ZooKeys 1092: 19-30 (2022)

Figure 5. X-ray microtomographic reconstruction of Nothotytthonyx serratus Li, Bifh, Kundrata &

Cai sp. nov., holotype, NIGP179427 A dorsal view B ventral view C lateral view D anterodorsal view

E anterolateral view. Scale bar: 2 mm.

palpomeres, while in other subfamilies and most of Dysmorphocerinae the apical max-
illary palpomeres are securiform, except for Tytthonyx, currently classified as incertae
sedis in Silinae. Dysmorphocerinae, however, generally have a wide pronotum and

New cantharid from mid-Cretaceous Burmese amber DF

complete elytra. The elytra are reduced in most of Malthininae genera, although this
feature is also present in species in most other subfamilies (e.g., Chauliognathinae:
Ichthyurus Westwood, Lobetus Kiesenwetter; Cantharinae: some Lycocerus Gorham;
Silinae: some Polemius LeConte, Brachysilidius Pic).

Nothotytthonyx is herein tentatively assigned to the subfamily Malthininae by a
combination of characters, such as the radially symmetrical apical maxillary palpomer-
es, shortened elytra, pronotum with arched margins and well-defined borders, tibiae
with apical spurs, and tarsal claws simple. However, the ovipositor with long gonostyli
of Nothotytthonyx seems to be quite aberrant in Malthininae. No extant species of
Malthininae (and Dysmorphocerinae and Silinae) has long and clearly defined styli.
According to Brancucci (1980), the well-defined coxites and styli are the “primitive
form’, and they are typical of the subfamily Cantharinae. In Malthininae, the styli are
indistinct; according to Brancucci, they are either extremely reduced or, most probably,
solidly fused to the coxites, and correspond to the pubescent area of the coxites.

Within Malthininae, Malthinini have confluent or almost confluent gular sutures,
while Malthodini and Malchinini have separated gular sutures (Brancucci 1980). Be-
sides, there is a prominent tooth on the incisor edge of mandibles in Malthinini, while
in Malthodini and Malchinini the mandibles are armed with a weak tooth, with a
row of small teeth, or simple (without teeth) (Brancucci 1980). Nothotytthonyx differs
from genera in the Malthodini and Malchinini in having confluent gular sutures and
mandibles with a prominent tooth on the incisor edge. Yet, Nothotytthonyx is distinc-
tive among genera in Malthinini for having strongly serrate antennae. Most genera in
Malthinini have filiform or weakly serrate antennae, and Paramalthinus Brancucci has
pectinate antennae (even though the antennae of Paramalthinus are pectinate, its an-
tennomere bodies are rather elongate). Nothotytthonyx is different from other genera in
Malthinini additionally in the combination of the moderately shortened elytra, clearly
confluent gular sutures, presence of tibial spurs, and unelongated metacoxae (Bran-
cucci 1980; Fanti and Castiglione 2017; Fanti and Vitali 2017).

It is notable that Nothotytthonyx is somewhat similar to Tytthonyx. This genus
shares characters with both Malthininae (e.g., mandibles with retinaculum, radially
symmetrical apical maxillary palpomeres, the shape of pronotum, reduced elytra and
wing venation) and Silinae (e.g., the structures of terminal ventrites and tergites and
the aedeagus). Tytthonyx has been kept in its own tribe Tytthonyxini as incertae sedis
in Silinae (Brancucci 1980); however, in a recent morphology-based phylogenetic
analysis, Tytthonyx was revealed as the sister group of Malthininae (Hsiao et al. 2021).
Nothotytthonyx shares with Tytthonyx a similar habitus, symmetrical apical maxillary
palpomeres, and shortened elytra. In some species of Tytthonyx (subgenus Tytthonyx),
the antennae are also distinctly serrate. Nothotytthonyx nevertheless differs from
Tytthonyx in the confluent gular sutures (separated in Zytthonyx) and the structure of
abdomen (gonostyli absent in Tytthonyx; Brancucci 1980).

Malthininae today generally have a Holarctic (Laurasian) distribution, with only
limited fauna known from Gondwanan parts of the World, whereas Dysmorphoceri-
nae have a strictly Gondwanan distribution. Although Archaeomatlthodes from Burmese
amber was once classified in Malthininae (Hsiao et al. 2017), it was later revised as a

28 Yan-Da Li et al. / ZooKeys 1092: 19-30 (2022)

member of Dysmorphocerinae (Hsiao et al. 2021). Thus, as the first fossil of Malth-
ininae from Burmese amber and also from the Mesozoic Era, Nothotytthonyx has im-
portant biogeographical implications. If we accept that the Burmese amber, which
comes from the mines located on the West Burma Block, is of a Gondwanan origin
(Poinar 2019), then our current discovery indicates that Malthininae were present in
Gondwanan lands in the Mesozoic, and taking into consideration that they are not
known from any northern-hemisphere Mesozoic deposits, they may have originated in
the south, and only later dispersed to north where they greatly diversified and survived
until now whereas they became rare in the south. However, this hypothesis will need
to be tested in future.

Acknowledgements

We are grateful to Su-Ping Wu for technical help in micro-CT reconstruction, and
Rong Huang for technical help in confocal imaging. Vinicius Ferreira (Editor), Gareth
Powell and Yun Hsiao provided helpful comments on an early version of this paper.
Financial support was provided by the Second Tibetan Plateau Scientific Expedition
and Research project (2019QZKKO0706), the Strategic Priority Research Program of
the Chinese Academy of Sciences (XDB26000000), the National Natural Science
Foundation of China (41688103), and grants from University of Sao Paulo Support
Foundation (FUSP 3587- ITV/MZ) to GB.

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