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First Member of ‘Higher Endomychidae’ (Coleoptera: Coccinelloidea) from the Mid-Cretaceous Amber of Myanmar and New Insights into the Time of Origin of the Handsome Fungus Beetles

Authors:

Abstract

A new genus and species of the family Endomychidae (Coleoptera: Coccinelloidea): Cretostenotarsus striatus Tomaszewska, Szawaryn and Arriaga-Varela gen. et sp. nov. are described, diagnosed and illustrated from the mid-Cretaceous amber from northern Myanmar. To test the systematic placement of the new extinct genus and species within the family, a phylogenetic analysis was conducted. A dataset of 38 morphological characters scored for 29 species (including the new fossil taxon), members of Endomychidae sensu stricto and representatives of Coccinelloidea as outgroups were analyzed using maximum parsimony. The results of the analysis indicate unequivocally that Cretostenotarsus striatus is a member of the Stenotarsus clade within a monophyletic ‘endomychine complex’ sensu Robertson et al. (2015), which corresponds to ‘Higher Endomychidae’ sensu Tomaszewska (2005). The present discovery confirms at least the Jurassic origin of Coccinelloidea and indicates a much older origin of Endomychidae than previously hypothesized.
Citation: Tomaszewska, W.;
Szawaryn, K.; Arriaga-Varela, E. First
Member of ‘Higher Endomychidae’
(Coleoptera: Coccinelloidea) from the
Mid-Cretaceous Amber of Myanmar
and New Insights into the Time of
Origin of the Handsome Fungus
Beetles. Insects 2022,13, 690. https://
doi.org/10.3390/insects13080690
Academic Editor: Markus Friedrich
Received: 13 July 2022
Accepted: 28 July 2022
Published: 31 July 2022
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insects
Article
First Member of ‘Higher Endomychidae’ (Coleoptera:
Coccinelloidea) from the Mid-Cretaceous Amber of Myanmar
and New Insights into the Time of Origin of the Handsome
Fungus Beetles
Wioletta Tomaszewska *, Karol Szawaryn and Emmanuel Arriaga-Varela
Museum and Institute of Zoology, Polish Academy of Sciences, Wilcza 64, 00-679 Warsaw, Poland;
k.szawaryn@gmail.com (K.S.); arriagavarelae@gmail.com (E.A.-V.)
*Correspondence: wiolkat@miiz.waw.pl
This published work and the nomenclatural acts it contains have been registered in ZooBank, the online
registration system for the ICZN (International Code of Zoological Nomenclature). The LSID (Life Science
Identifier) for this publication are:urn:lsid:zoobank.org:act:37F51B30-D88E-461D-852B-DBBAECF1024D;
urn:lsid:zoobank.org:act: 6DEC503D-BF61-48F9-9202-BA6A5B92FBFF urn:lsid:zoobank.org:act:
99D602C3-01AF-4C63-B355-84A7D9C650C5.
Simple Summary:
We discovered an extinct handsome fungus beetle almost 100 million years old
embedded in amber from Myanmar. Comparing and analyzing the characteristics of its body with
those of other beetles of this family living today, we were able to find out which ones could be their
closest relatives. What we discovered suggests that our beetle, representing a new genus and species,
is a part of a group called “Higher Endomychidae”. This group shares some features with members of
the family Coccinellidae, the well-known ladybugs or ladybirds. Our finding supports the hypothesis
that handsome fungus beetles (family Endomychidae) originated at least at the beginning of the
Cretaceous period, and most probably in the Jurassic, coinciding with the heyday of dinosaurs
on earth.
Abstract:
A new genus and species of the family Endomychidae (Coleoptera: Coccinelloidea): Cre-
tostenotarsus striatus Tomaszewska, Szawaryn and Arriaga-Varela
gen. et sp. nov.
are described,
diagnosed and illustrated from the mid-Cretaceous amber from northern Myanmar. To test the
systematic placement of the new extinct genus and species within the family, a phylogenetic analysis
was conducted. A dataset of 38 morphological characters scored for 29 species (including the new
fossil taxon), members of Endomychidae sensu stricto and representatives of Coccinelloidea as out-
groups were analyzed using maximum parsimony. The results of the analysis indicate unequivocally
that Cretostenotarsus striatus is a member of the Stenotarsus clade within a monophyletic ‘endomy-
chine complex’ sensu Robertson et al. (2015), which corresponds to ‘Higher Endomychidae’ sensu
Tomaszewska (2005). The present discovery confirms at least the Jurassic origin of Coccinelloidea
and indicates a much older origin of Endomychidae than previously hypothesized.
Keywords:
burmite; Cenomanian; fossil; handsome fungus beetles; new genus; new species;
endomychine complex
1. Introduction
The family Endomychidae (handsome fungus beetles) is a moderately diverse family
of mycophagous beetles that has been a subject of a major taxonomic rearrangement
in the last decade. Traditionally, Endomychidae contained 12 subfamilies [
1
] and was
classified in the superfamily Cucujoidea [
2
5
], in the derived group called ‘Cerylonid
Series’ [
6
]. However, the most recent, comprehensive molecular research on Cucujoidea
by Robertson et al. [
7
] has resulted in the formal recognition of the Cerylonid Series as
Insects 2022,13, 690. https://doi.org/10.3390/insects13080690 https://www.mdpi.com/journal/insects
Insects 2022,13, 690 2 of 13
an independent superfamily Coccinelloidea and the redefinition of Endomychidae by
removing Anamorphidae, Mycetaeidae, and Eupsilobiidae as separate families.
Endomychidae currently contains over 1600 described species classified in about
90 genera
distributed in all zoogeographical realms, with the highest diversity in the tropi-
cal and subtropical regions of the world [
1
,
3
,
4
]. The study of Robertson et al. [
7
] recovered
two main clades within the family, the ‘merophysiine complex’ and the ‘endomychine
complex’. The merophysiine complex includes the subfamilies Leiestinae, Merophysiinae,
and Pleganophorinae, the basal lineages of the family according to Tomaszewska [
4
], while
the endomychine complex includes Cyclotominae, Endomychinae, Epipocinae, and Ly-
coperdininae, and corresponds to ‘Higher Endomychidae’ sensu Tomaszewska [
4
]. The
endomychine complex is supported morphologically by the pseudotrimerous tarsi in
adults and the V- or U-shaped frontal arms on the head, and four pairs of stemmata in lar-
vae [
4
]. The study of Robertson et al. [
7
], however, did not include exemplars of subfamilies
Danascelinae and Xenomycetinae, this last one a sister group to ‘Higher Endomychidae’
in the analysis of Tomaszewska [
4
], so their relationships with the rest of the handsome
fungus beetles remain unclear. Moreover, some current subfamilies, e.g., Endomychinae in
the new sense, which now includes the genus Endomychus plus all genera of the former
subfamily Stenotarsinae, are an anatomically heterogeneous group with no potential sup-
porting synapomorphies. All this makes the phylogeny and evolution of the group still
unclear and the classification of the family preliminary, requiring further molecular and
morphological research, including fossil data.
Fossil taxa of handsome fungus beetles have been poorly known to date. Only
14 species
of Endomychidae sensu stricto have been formally described to the present.
Among them, eight species, representatives of subfamilies Leiestinae, Merophysiinae,
Pleganophorinae, and Endomychinae, have been described from Cenozoic ambers (Eocene
Baltic, Bitterfeld, and French amber from Oise) [
8
10
]. Even more intriguing and system-
atically significant fossil Endomychidae, important for elucidating the origin and early
evolution of the family, come from Mesozoic deposits. Recently, Tomaszewska et al. [
11
] de-
scribed the new genera Cretolestes,Burmalestes (Leiestinae), Cretoparamecus (Merophysiinae),
and Palaeomycetes (Xenomycetinae), and Li et al. [
12
] described the genus Rhomeocalpsua (of
incertae sedis placement in Endomychidae), the oldest known endomychids from northern
Myanmar amber, known as the Burmese amber or burmite which contains an outstandingly
diverse biota from the early Upper Cretaceous (approx. 99 Ma) [13].
Noticeable is the diversity of extinct forms in the subfamilies belonging to the basal
lineages of Endomychidae [
4
]. Apart from described taxa, additional undescribed rep-
resentatives of Leiestinae and Merophysiinae are abundant in the Myanmar amber, and
represent new species and new genera (WT and EAV, personal observation). This can
suggest at least an Early Cretaceous origin of Endomychidae ancestor, which agrees with
McKenna et al. [
14
] molecular study and estimates indicating that Coccinelloidea appeared
during the Middle Jurassic.
However, recent observations on additional specimens embedded in Myanmar amber
have suggested the existence of a member of the more derived evolutionary lineage ‘Higher
Endomychidae’, with overall similarity to Stenotarsus and its allies, at that time. This
discovery would suggest older than previously hypothesized, presumably Jurassic origin
of Endomychidae, agreeing with results of, e.g., Cai et al. [
15
], indicating that Coccinelloidea
diverged in the Late Triassic to Early Jurassic.
Most endomychid subfamilies, as defined by the molecular analyses of Robertson
et al. [
7
], are also supported by synapomorphies recovered in the studies of
Tomaszewska [3,4]
.
Nevertheless, some groups, like Endomychinae, are difficult to characterize morpholog-
ically. Moreover, the synapomorphies for some subfamilies, e.g., those belonging to the
endomychine complex (‘Higher Endomychidae’), like Epipocinae, Cyclotominae, and
former Stenotarsinae are difficult or impossible to observe in fossil specimens, as they
refer to the structures of male and female genitalia. Therefore, to avoid ambiguity and a
narrative/subjective assignment of the new fossil taxon within the family, cladistic analysis
Insects 2022,13, 690 3 of 13
of morphological data was conducted, with non-visible morphological characters of the
fossil taxon scored as ‘?’.
In the present contribution, we describe and illustrate a new genus and species from the
Upper Cretaceous Myanmar amber. Based on the results of our analyses, the new taxon is
classified in the subfamily Endomychinae as currently defined [
7
]. Our discovery represents
the oldest fossil taxon of the endomychine complex sensu Robertson et al. [
7
] (=‘Higher
Endomychidae’ sensu Tomaszewska [
4
]), thus filling an important gap in our knowledge
of the early evolution of the Endomychidae. Completing the gaps in our knowledge of
the fossil taxa of handsome fungus beetles will contribute to future, objective fossil-based
calibration in phylogenetic analyses (EAV et al. in preparation) and gradually bring us
closer to getting a complete picture of the origin and evolution of the Endomychidae and
the Coccinelloidea superfamily.
2. Materials and Methods
2.1. Examination and Deposition of Fossil Taxon
The study is based on a specimen of Endomychidae embedded in the Myanmar amber
originated from the Kachin State of northern Myanmar. The age of these amber deposits is
generally considered the earliest Cenomanian [
16
] or possibly the latest Albian [
17
], thus,
near the boundary between Upper and Lower Cretaceous. U-Pb zircon dating conducted
by Shi et al. [
13
] restricted its age at 98.79
±
0.62, which is equivalent to the earliest
Cenomanian, Upper Cretaceous. The type specimen of the new species described here is
deposited in the Museum of Amber Inclusions at the University of Gda´nsk, Poland (MAIG),
and is in accordance with all ethical standards for studying fossils from Myanmar [18].
The amber was cut, ground, and polished prior to the study. The specimen was
examined and documented with a Leica M205A stereomicroscope with a Leica DM6000
digital camera operating under Leica Application SuiteVR LAS 3.7 (MAIG). The following
measurements were made and are used in the description: total length—from the apical
margin of clypeus to apex of elytra; pronotal length—from the middle of anterior margin
to the margin of basal foramen; pronotal width—at widest part; elytral length—across
sutural line including scutellum; elytral width—across both elytra at the widest part. The
morphological terminology and the generic attribution and comparison with extant and
extinct taxa follow Tomaszewska [35].
Taxonomic acts established in the present work have been registered in ZooBank (see
below), together with the electronic publication LSID: urn:lsid:zoobank.org:pub: 37F51B30-
D88E-461D-852B-DBBAECF1024D.
2.2. Morphological Dataset and Phylogenetic Analysis
The main aim of the cladistics analysis of morphological characters conducted during
this study was a recognition of a correct subfamily/genus group placement of the new
fossil genus and species within the family Endomychidae. Despite some contradictions
between the results of analyses based solely on molecular data [
7
] or morphology [
3
,
4
],
having at disposal a somewhat limited set of morphological characters of a new taxon from
the past, the cladistics analysis of morphological characters was the most objective way to
recognize closest relatives of the newly described extinct genus and species.
Taxon sampling and characters used for the analysis are based in a broad sense on
Tomaszewska [
3
,
4
], with some modifications made in an attempt to provide resolution
among genera of Endomychidae sensu stricto. Representatives of Endomychidae, Eupsilobi-
idae, Mycetaeidae, and Coccinellidae were sampled, including the type species for genera
when specimens were available for study. A member of Hobartiidae, Hobartius eucalypti
(Blackburn), was used as a more distant outgroup. In total, our morphological data matrix
comprised 29 taxa (22 ingroup taxa including new fossil and seven outgroups), scored for
38 multistate characters. Unknown character states were treated as missing and coded
with ‘?’. The list of morphological characters and the full character matrix can be found in
Appendices Aand B, respectively.
Insects 2022,13, 690 4 of 13
The maximum parsimony (MP) analysis of our dataset was conducted in TNT 1.5 [
19
]
to find the most parsimonious trees (MPTs), using the New Technology (NT) option, us-
ing Driven Search with Sectorial Search, Ratchet, and Tree fusing options activated with
standard settings; and the Traditional Search (TS) option under the following parameters:
memory set to hold 1,000,000 trees, tree bisection—reconnection (TBR) branch-swapping al-
gorithm with 1000 replications saving 100 trees per replicate; zero-length branches collapsed
after the search. All characters were treated as unordered, and analysis was performed
under equal weights. Bremer support was calculated using the TNT Bremer function,
using suboptimal trees up to 20 steps longer. Character mapping was done in Winclada
v1.00.08 [20] using unambiguous optimization.
3. Results
3.1. Phylogenetic Analysis of Morphological Dataset
The phylogenetic analysis conducted under different search strategies resulted in
sets of at least partly similar resolution. The maximum parsimony analysis under New
Technology Search (MP NT) resulted in a single most parsimonious tree (MPT) with a
length (L) of 87 steps, consistency index (CI) = 52, and retention index (RI) = 75 (Figure 1).
Insects 2022, 13, x FOR PEER REVIEW 5 of 13
Figure 1. Single maximum parsimony tree based on morphological matrix obtained in TNT. Bremer
support values are shown over the nodes. Fossil Cretostenotarsus species indicated with ‘†’.
3.2. Systematic Palaeontology
Order Coleoptera Linnaeus, 1758
Suborder Polyphaga Emery, 1886
Superfamily Coccinelloidea Latreille, 1807
Family Endomychidae Leach, 1815
Subfamily Endomychinae Leach, 1815
Cretostenotarsus gen. nov.
LSID. urn:lsid:zoobank.org:act: 6DEC503D-BF61-48F9-9202-BA6A5B92FBFF.
Derivation of name. The name of the new genus is a combination of the Creto-, re-
ferring to the Cretaceous age of the deposit with the name Stenotarsus, the genus group
where the new taxon belongs. The gender is masculine.
Composition. The new genus is monotypic, represented by the type species only.
Type species. Cretostenotarsus striatus sp. nov.
Diagnosis. Cretostenotarsus can be distinguished from all known extant Endomych-
inae genera by the following combination of characters: body comparatively small, oval,
pubescent (Figure 2C), with irregularly punctate elytra; pronotum weakly narrower than
the base of elytra (Figure 2B), widely margined laterally, with anterior angles rounded not
produced, with distinct basal and triangular lateral sulci; sutural striae complete through-
out the elytra (Figure 2B); scutellar shield transverse, semi-rectangular; and tarsi 4-4-4,
simple with tarsomeres 1 and 2 weakly lobed (Figure 2H).
Figure 1.
Single maximum parsimony tree based on morphological matrix obtained in TNT. Bremer
support values are shown over the nodes. Fossil Cretostenotarsus species indicated with ‘†’.
The MP analysis under Traditional Search (MP TS) resulted in 16 MPTs with some trees
displaying the same or similar resolution as the single tree from the NT option. However, a
strict consensus tree calculated from the 16 MPTs from the TS (L = 97 steps; CI = 47;
RI = 68
)
displays almost all recognized subfamilies used in the analysis, mostly as unresolved
Insects 2022,13, 690 5 of 13
polytomy (Appendix C). The sister relationships of Eupsilobiidae + Coccinellidae and
Mycetaea + (Pleganophorinae + Merophysiinae), as well as fossil taxon recovered as part of
the Stenotarsus clade, are recovered in all trees from TS and the tree from NT analysis.
From the obtained resolutions, we selected as our preferred tree the best-resolved
topology—the single tree from the MP NT search (Figure 1), which mostly agrees with
Tomaszewska [4] and at least partly agrees with Robertson et al. [7] results.
Our results of morphology-based analysis are in contradiction with the current,
molecular-based classification of Endomychidae [
7
] in a few aspects: (1) The family En-
domychidae+ is recovered as a monophyletic group, including Mycetaeidae while Myc-
etaeidae is recovered as sister group to Eupsilobiidae + Coccinellidae in Robertson et al. [
7
];
(2) Leiestinae recovered as sister group to a clade comprising Xenomycetinae, Danascelinae
and “Higher Endomychidae”, while Leiestinae is a sister group to Pleganophorinae +
Merophysiinae in Robertson et al. [
7
]; and (3) Endomychus coccineus recovered as sister
group to Cyclotominae, and not forming a clade with former Stenotarsinae, what makes
the currently recognized subfamily Endomychinae sensu Robertson et al. [7] polyphyletic.
The group of ‘Higher Endomychidae’ sensu Tomaszewska [
4
] (=endomychine complex
sensu Robertson et al. [7] is recovered in our analysis, as shown in Figure 1.
The present results seem to be morphologically well justified. However, we are aware
of the limitation of morphology-only analysis. Therefore we do not intend to change current
classifications, leaving the above problems, particularly the monophyly of Endomychinae,
to further research with an integrative approach. For the present paper, we follow the
current classification and name the clade containing the former Stenotarsinae genera as
subfamily Endomychinae.
The following taxonomical decision is based on the preferred tree.
3.2. Systematic Palaeontology
Order Coleoptera Linnaeus, 1758
Suborder Polyphaga Emery, 1886
Superfamily Coccinelloidea Latreille, 1807
Family Endomychidae Leach, 1815
Subfamily Endomychinae Leach, 1815
Cretostenotarsus gen. nov.
LSID. urn:lsid:zoobank.org:act: 6DEC503D-BF61-48F9-9202-BA6A5B92FBFF.
Derivation of name.
The name of the new genus is a combination of the Creto-,
referring to the Cretaceous age of the deposit with the name Stenotarsus, the genus group
where the new taxon belongs. The gender is masculine.
Composition. The new genus is monotypic, represented by the type species only.
Type species. Cretostenotarsus striatus sp. nov.
Diagnosis.
Cretostenotarsus can be distinguished from all known extant Endomychi-
nae genera by the following combination of characters: body comparatively small, oval,
pubescent (Figure 2C), with irregularly punctate elytra; pronotum weakly narrower than
the base of elytra (Figure 2B), widely margined laterally, with anterior angles rounded not
produced, with distinct basal and triangular lateral sulci; sutural striae complete through-
out the elytra (Figure 2B); scutellar shield transverse, semi-rectangular; and tarsi 4-4-4,
simple with tarsomeres 1 and 2 weakly lobed (Figure 2H).
The new genus most resembles the extant genera Stenotarsus Perty, 1832, and Chondria
Gorham, 1887, in overall body shape (oval with elytra rounded laterally and pronotum
weakly narrower than the base of elytra). However, elytra with sutural striae, scutellar
shield short and transverse, anterior angles of pronotum rounded and not produced, and
terminal labial palpomere at most as long as wide and widening apically separate Cretosteno-
tarsus from both genera. Additionally, simple tarsomeres distinguish it from Stenotarsus.
Insects 2022,13, 690 6 of 13
Insects 2022, 13, x FOR PEER REVIEW 7 of 13
palpomere 2 longer than 1 and 3, about 2 times longer than 3; terminal palpomere longer
than remaining palpomeres combined, narrow at the base, widest toward middle length
then tapering and weakly obliquely truncate apically, about 2.8 times as long as wide in
the widest part. Labium with mentum transverse, widest near the base; prementum trans-
verse with ligula distinctly widened laterally; labial palpomeres moderately widely sepa-
rated; palpomere 1 shortest/smallest; palpomere 2 transverse; terminal palpomere
scarcely widened towards the apex, about as long as wide, nearly 2 times longer than
palpomere 2, truncate at apex.
Figure 2. Cretostenotarsus striatus sp. nov., holotype, MAIG 5999. (A), habitus, ventral. (B), habitus,
dorsal. (C), detail of lateral margin of elyt ra with seta e. (D), head and mouthparts, ventral. (E), meso-
and metaventrite, ventral. F, antennal club, dorsal. (G), antenna, ventral. (H), hind leg. Abbrevia-
tions: a1–a10, antennomeres; bsul, basal sulcus; cl, tarsal claw; fem, femur; lp1–lp3, labial palpo-
meres; lsul, lateral sulci; men, mentum; mes, mesoventrite; met, metaventrite; mo1–mp4, maxillary
palpomeres; pment, prementum; pros, prosternum; smen, submentum; str, sutural striae; t1–t4, tar-
someres; tib, tibia; tro, trochanter; v1–v5, ventrites. Scale bars represent: 500 µm (A); 250 µm (D,E,H).
Figure 2.
Cretostenotarsus striatus sp. nov., holotype, MAIG 5999. (
A
), habitus, ventral. (
B
), habitus,
dorsal. (
C
), detail of lateral margin of elytra with setae. (
D
), head and mouthparts, ventral. (
E
), meso-
and metaventrite, ventral. F, antennal club, dorsal. (
G
), antenna, ventral. (
H
), hind leg. Abbreviations:
a1–a10, antennomeres; bsul, basal sulcus; cl, tarsal claw; fem, femur; lp1–lp3, labial palpomeres; lsul,
lateral sulci; men, mentum; mes, mesoventrite; met, metaventrite; mo1–mp4, maxillary palpomeres;
pment, prementum; pros, prosternum; smen, submentum; str, sutural striae; t1–t4, tarsomeres; tib,
tibia; tro, trochanter; v1–v5, ventrites. Scale bars represent: 500 µm (A); 250 µm (D,E,H).
Moreover, Cretostenotarsus can be distinguished from the genus Danae Reiche, 1847
by having a more oval body, the pronotum without produced anterior angles and with
deeply impressed sulci, and by five abdominal ventrites; from Saula Gerstaecker, 1858
and Africanasaula Pic, 1946 by a more oval body, pronotum weakly narrower than base
of elytra, widely margined laterally with deeply impressed lateral and basal sulci, and
five abdominal ventrites; from Tragoscelis Strohecker, 1953 by having not as stout antenna
and not produced and acute front angles of pronotum, and by having lateral margins of
Insects 2022,13, 690 7 of 13
elytra flattened and well visible from above; from Ectomychus Gorham, 1887 by a more
oval body, symmetrical antennal club, distinct/deep basal sulcus on the pronotum and
not produced anterior pronotal angles; from Perrisina Strand, 1921 by light brown body,
terminal labial palpomere short and widening apically, and pronotal lateral margins not
so strongly elevated; from Paniegena Heller, 1916 by uniformly light brown body and
elytra irregularly and finely punctate; from Tharina Arriaga-Varela et al., 2018 by the
differently shaped mouthparts (maxillary and labial structures), well developed pronotal
sulci, prosternal process narrowing towards acute apex, and posterior angles of pronotum
regular without indentation; and from Endomychus Panzer, 1795 by the distinctly pubescent
body, different shape of scutellar shield, pronotum widely margined laterally, sutural striae
on the elytra, much more elongate terminal maxillary palpomere, ligula widening laterally
and not elongate terminal labial palpomere.
Cretostenotarsus is distinguished from the Cenozoic fossil member of the subfam-
ily Endomychinae from Baltic amber, Zemyna Tomaszewska, 2018 (=Laima Alekseev &
Tomaszewska, 2018) by having a brown, oval body, pronotum only weakly narrower
than the base of elytra with distinct triangular lateral sulci, and sutural striae complete
throughout the elytra.
Cretostenotarsus striatus sp. nov. (Figure 2A–H)
LSID. urn:lsid:zoobank.org:act:99D602C3-01AF-4C63-B355-84A7D9C650C5
Derivation of the name.
The species name refers to sutural striae on the elytra, present
very rarely in the subfamily.
Holotype.
MAIG No. 5999, sex unknown. The holotype is embedded in a small,
flattened, subquadrate piece (8.0
×
8.3
×
3.0 mm) of clear-yellow colored Myanmar amber
(burmite). Complete specimen. Syninclusions are present in the form of specimens of
Diptera and Hymenoptera and several small organic pieces.
Diagnosis. As stated for the new genus.
Description.
Body length 2.25 mm; width (at the widest point in about the middle
part of elytra) 1.14 mm. Body oval (about 1.97 times as long as wide) and moderately
convex, brown (Figure 2B); dorsal surfaces shiny and distinctly covered with moderately
long, pale pubescence (Figure 2C).
Head transverse, retracted into prothorax to the hind margin of eyes, finely punctate
and pubescent. Clypeus transverse and flat. Eyes large, rounded, prominent, and coarsely
faceted (Figure 2D). Antennal insertions visible from above. Antenna 11-segmented
(
Figure 2G
), relatively long, extending well behind bases of elytra, about as long as the
width of pronotum; scape stout, distinctly larger than pedicel; antennomeres 3–8 subequal,
regular, scarcely widening from their bases towards apices, each about 1.6 times as long
as wide, about 0.5 times as long pedicel; antennomeres 9–11 form elongate, narrow, loose
scarcely flattened club (Figure 2F); antennomeres 9 and 10 about the same length, and
antennomere 11 slightly longer, obliquely truncate at apex. Maxillary palpomeres with
membranous insertions on their inner apical edges (Figure 2D); palpomere 1 shortest,
palpomere 2 longer than 1 and 3, about 2 times longer than 3; terminal palpomere longer
than remaining palpomeres combined, narrow at the base, widest toward middle length
then tapering and weakly obliquely truncate apically, about 2.8 times as long as wide
in the widest part. Labium with mentum transverse, widest near the base; prementum
transverse with ligula distinctly widened laterally; labial palpomeres moderately widely
separated; palpomere 1 shortest/smallest; palpomere 2 transverse; terminal palpomere
scarcely widened towards the apex, about as long as wide, nearly 2 times longer than
palpomere 2, truncate at apex.
Prothorax 0.57 times as long as wide (Figure 2B); widest near anterior third, slightly
constricted at basal third; disc rather finely punctate with interspaces about 1.5–2.5 diame-
ters apart. Lateral sides widely margined; anterior angles rounded and not produced, pos-
terior angles nearly right-angled; disc evenly convex; lateral sulci in the form of triangular,
concave impressions, basal sulcus well developed. Posterior margin nearly straight, ante-
rior margin scarcely rounded medially. Prosternal process moderately widely separating
Insects 2022,13, 690 8 of 13
procoxae (Figure 2A), extending posteriorly distinctly beyond them, narrowing continu-
ously towards the apex, which is almost acute. Procoxae circular in outline
(Figure 2A)
.
Prosternum with anterior margin straight; in front of coxae about 0.8 times as long as
longitudinal coxal diameter.
Mesoventrite with two large oval pits on sides near anterior margin (Figure 2E);
with longitudinal ridge medially and weak concavities on its sides; mesocoxal cavi-
ties circular, widely separated with mesoventral intercoxal process about 1.35 times as
wide as coxal diameter; mesocoxal cavities open laterally; exposure of mesotrochantin
is uncertain. Scutellar shield moderately large (Figure 2B), strongly transverse, about
2.9 times
wider than long, and of semi-rectangular shape. Elytra 1.42 times as long as
wide,
1.43 times
as wide as pronotum; moderately convex; lateral margins moderately
widely flattened, visible from above; irregularly punctate, punctures sparser than those on
head and pronotum, about
2.0–4.0 diameters
apart; sutural stria distinct, entire; epipleuron
narrow
(Figure 2A,H)
, incomplete at the apex. Metaventrite transverse, nearly 1.7 times as
broad as long
(Figure 2A,E)
; more than twice longer than mesoventrite; distinctly bordered
anteriorly; intercoxal process straight between mesocoxae; postcoxal pits two pairs, large.
Metacoxae transverse, broadly separated (Figure 2A,H). Hind wings not visible.
Legs comparatively long (Figure 2A,H). Femur subclavate, pubescent. Tibiae weakly
widening towards apices without modifications, densely covered with short pubescence;
tibial spurs absent. Tarsal formula 4-4-4 (Figure 2H). Tarsi long, tarsomeres 1–3 grad-
ually slightly shorter and weakly lobed; tarsomere 4 slightly longer than tarsomeres
2–3 combined.
Abdomen with five ventrites (Figure 2A); ventrite 1 weakly shorter than the remaining
four ventrites combined. Ventrites 2, 3, and 4 subequal in length; terminal ventrite rounded
apically. Postcoxal lines absent. Abdominal punctation similar to those on metathorax.
Locality and horizon.
Myanmar, Kachin State, Hukawng Valley, Myanmar amber;
unnamed horizon; Cretaceous, Upper Albian to Lower Cenomanian.
4. Discussion
Knowledge of fossil faunas is crucial for making conclusions about the origin and
evolution of studied groups. Although the fossil fauna of handsome fungus beetles is still
poorly known, the situation has improved constantly, especially in recent years.
Representatives of Endomychidae sensu lato in the fossil record have appeared in
several publications [
1
,
21
27
], but only very few Cenozoic taxa have been properly named
and formally described until recently [
8
10
]. Among 16 genera of Endomychidae sensu lato
from the Eocene reported in the literature, only 13 species are formally described, 12 from
the Baltic and Bitterfeld ambers (ca. 40–45 Ma), and one species from the French amber
from Oise (ca. 53 Ma). Many of them are recently described members of Anamorphidae
from the Eocene Baltic and Bitterfeld amber, one classified in Symbiotes Redtenbacher, and
four in extinct monotypic genera: Gramboale,Kleinzaches,Palaecoryphus, and Giltine [
9
].
Some species of Endomychidae from the Eocene Baltic amber have been, at least initially,
assigned to extant genera, e.g., Trochoideus,Holoparamecus,Mycetina, or Lycoperdina. Myc-
etaeidae are only known through an unnamed Mycetaea species reported by Klebs [
24
], and
Eupsilobiidae lack any reported fossil species.
Of the properly described taxa of Endomychidae sensu stricto from the Eocene, Palaeoestes
Kirejtshuk & Nel (French Oise amber), Phymaphoroides Motschulsky, and Glesirhanis Shockley
& Alekseev (Baltic amber) represent extinct genera of Leiestinae and Zemyna Tomaszewska
(=Laima Alekseev & Tomaszewska) (Baltic amber) is the only known extinct genus of
the subfamily Endomychinae. Two species of Trochoideus described by Alekseev and
Tomaszewska [
9
] are the only Eocene species classified in the extant genus of the subfam-
ily Pleganophorinae. While two species of Holoparamecus described by Reike et al. [
10
]
represent the only Eocene species of this extant genus of the endomychid subfamily Mero-
physiinae (although treated as family Merophysiidae by these authors, ignoring the results
of numerous research based on morphological and molecular data sets (e.g., [4,5,7])).
Insects 2022,13, 690 9 of 13
Nearly all those discoveries that appeared over 170 years since the first record of
fossil Endomychidae [
21
] originally seemed to confirm Carpenter’s suggestion that the
family Endomychidae sensu lato originated during the Eocene [
27
]. Only the description of
Palaeoestes eocenicus Kirejtshuk & Nel, a member of the subfamily Leiestinae from French
amber (53 Ma) [
28
], changed this situation, suggesting an older origin of the group, probably
in the Paleocene (65–54.8 Ma). First estimates of the Mesozoic origin of the Endomychidae
by Ponomarenko [
29
] indicated that the group originated during the Late Cretaceous. And
subsequent reports of Endomychidae in Myanmar (99 Ma) and Lebanese amber (125 Ma)
by Poinar and Poinar [30] further supported a much older origin for the family.
The recent discovery and the formal description of the Endomychidae sensu stricto
members in Myanmar amber confirmed at least the Cretaceous origin of Endomychidae [
11
].
Tomaszewska et al. [
11
] described the oldest known endomychids from Myanmar amber,
representatives of the subfamilies Leiestinae (Cretolestes,Burmalestes), Merophysiinae (Cre-
toparamecus), and Xenomycetinae (Palaeomycetes), all belonging to the basal evolutionary
lineages of the handsome fungus beetles [4].
The species (described and undescribed) known to date from the Cretaceous Myanmar
amber display much greater taxonomic diversity than the taxa from the Eocene ambers
described so far. Particularly noticeable is the diversity of extinct forms in the subfamily
Leiestinae in the Myanmar amber. The diversity and distribution of this group seem to
have decreased in time, with a few members known from Eocene amber and currently
being a small and exclusively Holarctic group [
3
,
31
]. While the subfamily Xenomycetinae
is not recorded from the Eocene deposits until now. The occurrence of Merophysiinae in
the Myanmar amber may be correlated with the increased availability of termite and ant
colonies inhabited by many species of this subfamily. However, the species of Trochoideus,
a member of Pleganophorinae, likely a sister group to Merophysiinae, being the typical
symphile, is known only from the Eocene so far.
The abundance and diversification of the basal Endomychidae in the mid-Cretaceous
environments suggest at least the Early Cretaceous origin of this group, which agrees with
McKenna et al. [
14
] molecular study indicating the Middle Jurassic (187–157 Ma) origin of
the superfamily Coccinelloidea.
However, some other, more recent estimates suggest that Coccinelloidea diverged
in the Late Triassic to Early Jurassic (217–182 Ma), with most extant families originating
in the Jurassic to Early Cretaceous [
15
], or even this superfamily could occur in mid-
Permian to mid-Triassic (ca. 267–238 Ma) [
32
]. And our current discovery of a member of a
more derived evolutionary lineage of Endomychidae in Upper Cretaceous amber seems to
confirm these estimates and indicate the Jurassic origin of the family.
5. Conclusions
The good preservation and visibility of the fossil specimen from Myanmar amber from
the Upper Cretaceous allowed us to examine, describe and illustrate its morphology and
test its position within the family Endomychidae. The results of our phylogenetic analysis
based on a morphological dataset indicate that Cretostenotarsus striatus new genus and
species described here belongs to the subfamily Endomychinae, therefore representing the
oldest known member of the endomychine complex [
7
] or “Higher Endomychidae” [
4
].
Although Cretostenotarsus striatus shows a second tarsomere only weakly lobed, it repre-
sents the oldest example of the pseudotrimerous tarsal configuration characteristic for the
endomychine complex and family Coccinellidae (ladybugs). The abundance and diversity
of members of the merophysiine complex or “basal Endomychidae” in the Upper Creta-
ceous ambers suggest at least a Lower Cretaceous origin of this group agreeing with the
estimations of McKenna et al. [
14
]. But the discovery here reported of a member of “Higher
Endomychidae” present in the Upper Cretaceous environments pushes back in time the
possible time of origin and subsequent diversification of Endomychidae, which agrees more
with Toussaint et al. [
32
] or Cai et al. [
15
] estimates on Coccinelloidea origin. Our discovery
Insects 2022,13, 690 10 of 13
will be of great value in future testing of the hypotheses of the causes and consequences of
diversification of this superfamily that require a robust time-calibrated phylogeny.
Author Contributions:
Conceptualization, W.T. and E.A.-V.; methodology, W.T and K.S.; formal
analysis, W.T. and K.S.; data curation, K.S and E.A.-V.; writing—original draft preparation, W.T.;
writing—review and editing, W.T., K.S. and E.A.-V.; visualization, K.S.; funding acquisition, W.T. and
E.A.-V. All authors have read and agreed to the published version of the manuscript.
Funding:
This research was funded by the National Science Centre of Poland (Narodowe Centrum
Nauki; grant No. 2018/29/B/NZ8/02745 to WT and grant No. 2020/36/C/NZ8/00584 to EAV).
Institutional Review Board Statement: Not applicable.
Informed Consent Statement: Not applicable.
Data Availability Statement: The data presented in this study are available in the present article.
Acknowledgments:
We would like to thank El ˙
zbieta Sontag (University of Gda´nsk, Poland) for the
loan of material used in this study. Bła˙
zej Bojarski (University of Gda´nsk, Poland) is acknowledged
for help in preparing the amber piece. Three anonymous reviewers are acknowledged for their
comments that helped to improve the manuscript.
Conflicts of Interest:
The authors declare no conflict of interest. The funder of this research had no
role in the design of the study; in the collection, analysis, or interpretation of data; in the writing of
the manuscript, or in the decision to publish the results.
Appendix A. List of Morphological Characters Used in Cladistics Analyses
1. Head with occipital file: absent (0); present (1).
2. Head with antennal grooves: short (0); long (1), absent (2).
3. Antenna: 4–5/7 (0); 8–9 (1); 10 (2); 11 (3).
4. Antennal club: 3-segmented (0); 1 or 2-segmented (1).
5. Fronto-clypeal suture: absent (0); present (1)
6. Mandible with mola: well-developed, large (0); absent or reduced (1).
7. Mandible with prostheca: only setose (0); with setae and apical, sclerotized, elongate
projections (1).
8. Mandible with subapical teeth/serrations (0), with at most one subapical tooth
present (1).
9. Mentum: widest basally or medially (0); widest anteriorly (1).
10. Labium with mentum: flat, smooth (0); with small, triangular, setose hill placed
medially (1); with large, triangular, raised area (2).
11. Labial palp with palpomere 2: subcylindrical or transverse (0); oval, inflated (1).
12. Maxillary palpomere 2 and 3 with membranous insertions on inner apical edges:
absent (0); present (1).
13. Tentorium with anterior arms: separate (0); meeting medially (1).
14. Corpotentorium absent (0); present (1).
15. Anterior margin of pronotum with stridulatory membrane: absent (0); present (1).
16. Pronotum laterally: narrowly bordered (0); with wide lateral margins (1).
17. Pronotum with basal and/or lateral sulci/modifications: absent (0); present (1).
18. Prothoracic ventral antennal grooves: absent (0); present on prosternum (1); present
on hypomeron (2).
19. Mesoventrite with intercoxal process bicarinate (boat-shaped): absent (0);
present (1)
.
20. Mesocoxal cavity: open outwardly (0); narrowly closed outwardly (1).
21. Mesotrochantin: at least partially exposed (0); concealed (1).
22. Metaventrite with postcoxal lines: absent (0); present (1).
23. Tarsal formula: 5-5-5 (or 5-5-4 in male) (0); 4-4-4 (1); 3-3-3 (2).
24. Penultimate mesotarsomere: not or slightly reduced and not enclosed within lobe
of antepenultimate tarsomere (0); highly reduced and partly or entirely enclosed within
ventral lobe of antepenultimate tarsomere (1).
Insects 2022,13, 690 11 of 13
25. Pretarsal claws: simple (0); modified (1).
26. Number of functional abdominal spiracles: 7 pairs (0); 5 pairs (1).
27. Abdominal ventrite 1: without postcoxal lines (0); with postcoxal lines (1).
28. Sternite (IX) of male genital segment with lateral edges deeply, asymmetrically
curved inwardly: absent (0); present (1).
29. Tegmen: without penis guide (0); with penis guide (1); tegmen absent (2).
30. Aedeagus with median lobe: nearly straight or curved (0); coiled (1).
31. Median lobe with T-shaped capsule: absent (0); present (1).
32. Median lobe with curled 1/3 of its basal part: absent (0); present (1).
33. Ovipositor without infundibulum (0); with infundibulum-like structure/sperm
duct modified (1); with infundibulum (2).
34. Ovipositor with sperm duct attached: directly to spermatheca (0); to broad connec-
tion between spermatheca and accessory gland (1).
35. Ovipositor: with deeply divided basal part of coxites (1); not divided basal part of
coxites (0).
36. Mesoventral postcoxal pits: absent (0); present (1).
37. Metaventral postcoxal pits (anterior margin of metaventrite, adjacent to posterior
margin of mesocoxal cavities): absent (0); present (1).
38. Antennal sockets: visible from above (0); not visible (1).
Appendix B. Morphological Matrix Scored for 29 Taxa and 38 Characters Used for
Cladistics Analyses in TNT
Hobartius_eucalypti 02301001000011000000000000000000000000
Coccinella_septempunctata 02300101100100000000111111101010200000
Chilocorus_cacti 021?0101100100000000111111101010000001
Scymnomorphus_fulvus 0021010110000000000011??01101010000000
Mycetaea_subterranea 02301000010011011000101001000000000000
Holoparamecus_sp. 02311000001010001001102001000000000001
Cholovocera_beloni 01111011021010001101112001102000000001
Merophysia_sp. 01111011021010001101102001002000000001
Leiestes_seminiger 02301001000011001010101001000000000111
Phymaphora_pulchella 02301000000011001010101001000000000111
Rhanidea_unicolor 02301000000011001010101001000000000111
Cyclotoma_testudinaria 02301001000011000000001101000001010000
Meilichius_impressicollis 02301001000011001000001101000001010010
Epipocus_tibialis 02301001000011001000001101010000000110
Epopterus_sp. 02301001000011001000001101010000000110
Endomychus_coccineus 02301001000011001000001101000000000110
Stenotarsus_nobilis 02301001000011011000001101000000011110
Tharina_antennalis 02301001000011011000002001000000001110
Aphorista_laeta 12301001000011101000001101000000000110
Eumorphus_quadriguttatus 12301001000011101000001101000000000110
Lycoperdina_bovistae 12311001000011101000001001000000000110
Pleganophorus_bispinosus 0201100000001100100110100100????000000
Trochoideus_desjardinsi 02011000000011001001101001000000000000
Xenomycetes_morrisoni 02301001000011001000001001000000000110
Danascelis_elongata 02301001000011001000001001000000000110
Eidoreus_minutus 00211001020011000000011001100110100000
Chileolobius_notatus 01310101020011000200111001100110100000
Danae_sp. 02301001000011011000001101000000001000
Cretostenotarsus striatus ?2301???0001??011000?0100?0????????110
Insects 2022,13, 690 12 of 13
Appendix C. Strict Consensus Tree from 16 Most Parsimonious Trees Calculated in
TNT under Traditional Search Option
Insects 2022, 13, x FOR PEER REVIEW 12 of 13
Xenomycetes_morrisoni 02301001000011001000001001000000000110
Danascelis_elongata 02301001000011001000001001000000000110
Eidoreus_minutus 00211001020011000000011001100110100000
Chileolobius_notatus 01310101020011000200111001100110100000
Danae_sp. 02301001000011011000001101000000001000
Cretostenotarsus striatus ?2301???0001??011000?0100?0????????110
Appendix C. Strict Consensus Tree from 16 Most Parsimonious Trees Calculated in TNT under
Traditional Search option.
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Article
A new extinct handsome fungus beetle species of the extant Palaearctic genus Leiestes Chevrolat, L. tomaszewskae Alekseev and Bukejs sp. nov., is described from the Baltic amber. The new species is studied and illustrated in detail using X-ray micro-computed tomography (μCT). A list of described Endomychidae known from fossil resins (Baltic, Bitterfeld, Oise and Burmese ambers) is compiled.
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