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Abstract

The Cerambycids described from Burmese Amber are revised. Qitianniu zhihaoi Lin & Bai, 2017 (Cerambycidae incertae sedis) is recognised as the first known member of the tribe Meroscelisini Thomson, 1861 (Cerambycidae Prioninae). Apophisandra ammytae Molino-Olmedo 2017 and the tribe Apophisandrini Molino-Olmedo 2017 are recognised as no member of the superfamily Cerambycoidea and transferred to the superfamily Cucujiodea, near the family Parandrexidae Kirejtshuk, 1994.
Palaeoentomology 002 (3): 215–218
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Copyright © 2019 Magnolia Press
Short PALAEOENTOMOLOGY
ISSN 2624-2826 (print edition)
ISSN 2624-2834 (online edition)
Accepted by D.-Y. Huang: 20 May 2019; published: 24 Jun. 2019 215
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PE
Communication
Introduction
The description of a new fossil taxon presupposes the global
knowledge of the examined group and of the existence of
possible sibling, mimicking or simply superficially similar
taxa. The older the fossils are, the greater the possibility of
misidentification. Moreover, the knowledge of the assumed
phylogeny and of the evolution centres of the extant taxa
allows understanding the real taxonomy of new fossil
entities, giving consistency and support to the descriptions.
Because of its unusual morphological characters,
the recent description of Apophisandra ammytae Molino-
Olmedo, 2017 (new genus, species and tribe) was
disconcerting to most specialists in cerambycids. This taxon
evidently belongs to another family.
This paper also revises the status of Qitianniu
zhihaoi Lin & Bai, 2017 (whose systematic position inside
Cerambycoidea remained inconclusive in the original
description) and clarifies the systematic position of all
assumed cerambycids in Burmese amber.
Material and methods
Burmite originates from mines located in Myanmar, Kachin
State, Myitkyina District, Hukawng Valley. Based on
radiometric data, Burmese amber is currently dated back
to the earliest Cenomanian, mid-Cretaceous, with an age of
98.79 ± 0.62 Ma (Shi et al., 2012).
The characters and pictures provided in the original
descriptions (Lin & Bai, 2017; Molino-Olmedo, 2017) are
sufficient to discriminate the systematic position of the
assumed cerambycids with the help of several general and
specific works on the systematics of this group (Lamarck,
1817; Thomson, 1860; Mulsant, 1862; Lacordaire, 1869;
Gahan, 1906; Lameere, 1912; Saalas, 1936; Müller, 1949–
1953; Linsley, 1962; Villiers, 1968; Švácha et al., 1997;
Vitali, 2006; Sýkorová, 2008; Nearns, 2013; Švácha &
Lawrence, 2014; Bouyer, 2015).
Systematic palaeontology
Order Coleoptera Linnaeus, 1758
Superfamily Cerambycoidea Latreille, 1802
Family Cerambycidae Latreille, 1802
Subfamily Prioninae Latreille, 1802
Tribe Meroscelisini Thomson, 1861 stat. nov.
Genus Qitianniu Lin & Bai, 2017
Qitianniu zhihaoi Lin & Bai, 2017
According to its authors, Qitianniu zhihaoi is characterised
by minute body size (4.6 mm), body slightly flattened
dorsoventrally, tarsi cryptopentamerous, eyes very large and
coarsely facetted, last segment of palpi not tapered apically,
pronotum with complete lateral margin and antennae longer
than body (Lin & Bai, 2017).
Instead of using these diagnostic characters, the authors,
without explanation, inserted them into the data matrix
proposed by Napp (1994) in order “to clarify the phylogenetic
position” of this taxon. Although they noted and accordingly
corrected some significant errors of this matrix concerning
missing groups (Dorcasominae, Chrysomelidae) or keyed
characters (Saphanus, Atimia, Philus), they overlooked the
fact that the conclusions were still inconsistent even after the
corrections. In fact, the resulting phylogenetic tree appears
incoherent since “the four genera of Spondylidinae [are] very
widely separated and the eight subfamilies of Cerambycidae
are not in a monophyletic clade”. This contradicts the
generally accepted taxonomy of these subfamilies, which
is supported by recent phylogenetic analyses (Sýkorová,
2008). Consequently, the authors admitted that “the trees did
not provide strong evidence to differentiate the cerambycoid
complex or elucidate the position of Qitianniu clearly”.
Afterwards, they briefly examined all known subfamilies
without being able to classify the new taxon. But, this analysis
also revealed uncertainty and several misconceptions of the
peculiar characters of each subfamily. For example, the
reason that Qitianniu does not belong to Lamiinae is not “its
https://doi.org/10.11646/palaeoentomology.2.3.3
http://zoobank.org/urn:lsid:zoobank.org:pub:0688DAE1-3498-46AB-8DF8-274DD5A7A677
Systematic notes on the Cerambycidae (Insecta: Coleoptera) described from
Burmese amber
FRANCESCO VITALI
Nationalmusée fir Naturgeschicht, rue Münster 25, L-2160 Luxembourg, Luxembourg. E-mail address: fvitali@mnhn.lu
VITALI
216 Palaeoentomology 002 (3) © 2019 Magnolia Press
prognathous head” but its truncate palps (Lacordaire, 1869).
Actually, the definition of “prognathous head” is erroneous
as well since the provided photos show that the mandibles
are obliquely directed and the frons is vertical; thus, the
head can not be defined as truly “prognathous”. In addition,
this condition can be also observed in several Lamiinae, as
e.g. in the genus Sternotomis Percheron, 1836, or in most
Tmesisternini.
The authors had evidence of the presence of a complete
lateral margin of the pronotum, which properly excluded
some subfamilies, but they did not draw the logical
conclusion due to misconceptions concerning the phylogeny
of the Prioninae.
The claims that a “prionine ancestor was probably a
larger beetle” and that “the small forms (some Neotropical
Anacolini) are strongly derived” are considered as wrong for
nearly a century (Saalas, 1936; Müller, 1949–1953; Vitali,
2006; Švácha & Lawrence, 2014).
On the contrary, primordial Prioninae were in all
likelihood small forms, similar to other basal cerambycoid
tribes (Cyrtonopini, Disteniini, Dynamostini, Heteropalpini,
Distenini, Vesperini, Philini), which include relatively small
taxa. For a long time, close relationships were noticed among
primitive Prioninae and Philini (Gahan, 1906; Linsley, 1962)
and among Anacolini / Meroscelisini and Philini, Cyrtonopini
and Xystrocerini (Lameere, 1912). Thus, Anacolini are
actually among the most basal tribes of Prioninae and small
representatives of Meroscelisini are even more archaic. In
this last tribe, members of the genus Anoeme Gahan, 1890
can be only 8.5 mm long (Villiers, 1968; Bouyer, 2015).
The complete lateral margin of the pronotum is the
well-known peculiar character of the Prioninae since the
origins of the classification of the cerambycids (Lamarck,
1817; Thomson, 1860; Mulsant, 1862; Lacordaire, 1869).
Moreover, Qitianniu shows other evident diagnostic
characters. The simple pronotum and the shape of the
head suggest some tribes of the transition Prioninae-
Cerambycinae, i.e. Meroscelisini and Xystrocerini, but the
margined pronotum points unequivocally to the former
tribe.
The authors also described a slightly groove at the apex
of the protibiae that might make think to some Cerambycinae
Xystrocerini (Auxesis Thomson, 1858). Nonetheless,
the real existence of this character is rather doubtful. In
the discussion, the protibiae are defined as “sinuate”, a
completely different character. No detailed photo supports
it, but, on the contrary, the published photos seem to suggest
a misinterpreted air bubble. However, the fact that tibial
grooves are present in several unrelated Cerambycoidea
(Disteniidae, Cerambycinae, Lamiinae) implies that this
character evolved many times in the Cerambycoidea without
having a particular phylogenetic importance.
Accordingly, Qitianniu looks similar to Anoeme,
especially to A. murphyi Bouyer, 2015, with which it shares
most characters, while its short elytra are reminiscent of
other congeners. The pronotum looks still similar to more
primitive lepturoid Cerambycidae, e.g. Caraphia Gahan,
1906, but this is perfectly coherent with the phylogenetic
position of this taxon.
Anoeme is currently widespread in the whole Tropical
Africa (Bouyer, 2015) with an endemic species in India
(Gahan, 1906). This split distribution and its systematic
position inside Cerambycidae support the primitiveness
of this taxon, whose origin should be dated before the
separation of India from Africa (Early Cretaceous). The
assumed existence of Anoeme already in the Cretaceous
is harmonious with the classification of Qitianniu inside
Meroscelisini, even suggesting a larger distribution of this
tribe during this period.
Superfamily Cucujoidea Latreille, 1802
Family ?Parandrexidae Kirejtshuk, 1994
Tribe Apophisandrini Molino-Olmedo, 2017
Genus Apophisandra Molino-Olmedo, 2017
Apophisandra ammytae Molino-Olmedo, 2017
According to its author, Apophisandra ammytae is
characterised by minute body size (4.9 mm), flattened body,
pentamerous tarsi and falciform mandibles without teeth or
dorsal ridges (Molino-Olmedo, 2017). The beadlike antennae
are inserted close to the mandibular condyles, without
antennal tubercles and they surpass the elytral base.
Some characters were not considered as belonging
to Parandrini, implying the definition of the new tribe
Apophisandrini: scape elongated and tubular; antennomeres
II–III equal in length and width; eyes projected and
horizontally enlarged, pronotum with rounded basal angles.
The author supported the assignment of this taxon to
Parandrinae (sic!) claiming that this group—having
pentamerous tarsi and short antennae—is considered as
the oldest among cerambycids. He mentioned McKenna
& Farrell (2009), who hypothesised its existence in the
Cretaceous.
Nonetheless, Apophisandra ammytae appears as a
very odd cerambycid. First, the mandibles are exceptionally
narrow, too projected and, especially, having their base too
far from the antennal supports, which are even described as
absent. In all Cerambycidae and in the Oxypeltidae, the base
of the mouthparts is as wide as the frons. The mandibles of
some Disteniidae and Vesperidae might resemble those of
Apophisandra, but these families are completely unrelated
to this fossil. In contrast, these characters can be consistently
observed in many Cucujoidea, such as Cucujidae,
Laemophloeidae, Passandridae and Parandrexidae.
Secondly, the antennae are unusually longer than
in typical Parandrini but they do not show, however, any
cerambycoid aspect. They are “moniliform”, another
NOTES ON THE CERAMBYCIDAE Palaeoentomology 002 (3) © 2019 Magnolia Press 217
bizarre character for cerambycids, which show threadlike,
toothed or eventually comb-like antennae. In addition, the
usual differentiated pedicel is absent, while the last three
antennomeres are not progressively smaller or thinner but
flattened and enlarged, even suggesting the presence of an
apical club.
Thirdly, the abnormally projected eyes do not
correspond to any known cerambycid but are reminiscent of
those of some Cucujoidea, such as Cucujidae sensu lato and
Smicripidae.
Finally, from a phylogenetic point of view,
Apophisandra is doubtfully placeable. As many authors
have indicated (Saalas, 1936; Müller, 1949–1953; Vitali,
2006; Švácha & Lawrence, 2014) and some genetic
analyses (Nearns, 2013) have confirmed, Parandrini are not
a primitive tribe but modified Prioninae, whereas the most
primitive Cerambycoidea are lepturoid families close to the
Chysomeloidea (Švácha et al., 1997). Though living during
the Cretaceous, this taxon shows specialised characters
unknown in any other known cerambycids (beadlike, nearly
club-shaped antennae, falciform mandibles, projected eyes
and rounded basal angles of the pronotum) but no primitive
characters present in other taxa.
The minute size and the pentamerous tarsi confirm the
suspicion that Apophisandra has nothing to do with Parandrini
or other Cerambycoidea. Thus, it is about a cerambycid-like
Cucujoidea, such as Laemophloeus, Uleiota and other ones,
which also show an analogous pronotum.
In reality, all characters of Apophisandra ammytae
fit the superfamily Cucujiodea. The absence of furrows on
the pronotal disc seems to exclude Laemophloeidae, while
the lateral ridge boarding the pronotum resembles some
Passandridae. The characters separating true Cucujidae
from Silvanidae are still unclear (Thomas, 2002) and some
of them, i.e. tarsal proportions and genitalia, are neither
mentioned nor visible in the description of Apophisandra.
However, no extant taxon shows similar large mandibles.
Apophisandra ammytae might belong to the Mesozoic
family Parandrexidae Kirejtshuk, 1994, already confused
with cerambycids in the past (Rohdendorf, 1962), which is
characterised by flattened body, transverse flattened procoxae
with exposed trochantin, large elongated prognathous head,
long palpi and a movable articulation between pronotum
and head (Kirejtshuk, 1994). Many species show analogous
mandibles but also antennae as long as body; nonetheless, a
species with shorter antennae was described from the Lower
Cretaceous of Spain (Soriano et al., 2006).
Though I am unable to perform further analyses, the
tribe Apophisandrini and Apophisandra ammytae should
be removed from Cerambycidae and transferred into
the superfamily Cucujiodea, tentatively near the family
Parandrexidae.
Conclusion
After this systematic review, mid-Cretaceous Burmese
amber now contains only one species of Cerambycidae, i.e.
Qitianniu zhihaoi Lin & Bai, 2017, belonging to Prioninae,
tribe Meroscelisini. The oldest fossil belonging to the
Prioninae is Cretoprionus liutiaogouensis Wang et al., 2014
from the Lower Cretaceous of China (Wang et al., 2014).
Although this fossil shows rather clear diagnostic characters,
the authors remained somewhat uncertain about its tribal
position.
Considering that this taxon already shows some
specialised characters, i.e. saw-toothed antennae, toothed
pronotum, lobed tarsi, the origin of Prioninae should be
dated to even more ancient times.
In fact, the earliest known longhorn beetle is not
Cretoprionus liutiaogouensis, as Wang et al. (2014) claimed,
but Cerambycinus dubius Germar, 1839 from Late Jurassic of
Solnhofen (Germany). Giebel (1856) described this species
again as Mesosa germari, underlining its close resemblance
to this extant genus. Though the attribution to Mesosa seems
almost uncertain, the fossil shows unequivocal long antennae
supporting its belonging to the Cerambycoidea.
In conclusion, it seems evident that the origin of
Cerambycoidea should be dated back at least to the Jurassic,
or, as supposed on the basis of the chorology of some extant
tribes as Macrotomini, maybe even to the Triassic (Vitali,
2008), before the splitting of Laurasia-Gondwana.
Acknowledgements
The author thanks Daniel Heffern, Houston (USA) for
assistance with the English language used in this work.
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https://doi.org/10.1080/14772019.2013.806602
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I describe Apophisandra ammytae gen. and n. sp. (Coleoptera, Cerambyidae) in Cretaceous amber from Myanmar. It can be separated from the rest of Parandrinae, subfamily to which it belongs, by the structure of its antennae with segment I tubular, elongated, somewhat longer than the other two united, and of width similar to theirs, segments II and III of length and width similar to each other and that of the segments IV-X; eyes elongated horizontally ; very flat protorax with very rounded posterior angles ; elytra striated and very flat body, almost glabrous. Given these díferent characteristics, I propose the creation of Apophisandrini, tribus nov. NOTE: Initially included in Cerambycidae Passandrinae, by the antennal insertion and other characters really belongs to Cucujoidea and not to Cerambycidae
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A new cerambycid beetle (Qitianniu zhihaoi gen. et. sp. nov.) is described on the basis of a single specimen embedded in Cretaceous Burmese amber (ca. 99 Ma). Unusual characteristics are hairy antennae and large lateral eyes, a pronotum with lateral margin, and sinuate protibiae. Based on a phylogenetic analysis, the systematic position of Qitianniu is still uncertain and we provisionally place it as Cerambycidae incertae sedis.
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The systematic position of the genus Parandrexis for which a new family is proposed, is discussed. Two new species, P. rotundicollis and P. subtilis, are described, the type species P. parvula Martynov is redescribed, and an identificaiton table of the species of this genus is presented. -Journal summary
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The fossil cerambycid species from Geiseltal (Sachsen-Anhalt, Germany, Middle Eocene) are analysed. Xyleoconites proavus Haupt, 1950, is transferred from Prioninae incertae sedis to Prioninae Macrotomini, while Eocallidium rugulosus Haupt, 1950, corrected in Eocallidium rugulosum Haupt, 1950, is transferred to Tenebrionidae.
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The taxonomic position of Protospondylis florissantensis (Wickham, 1920), Florissant, Oligocene, is analysed. The genus Protospondylis Linsley, 1942 has resulted to be a younger synonym of Parandra Latreille, 1804, whose synonymy is here proposed. Accordingly, Protospondylis florissantensis Wickham, 1920 becomes Parandra florissantensis (Wickham, 1920) nov. comb.