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https://doi.org/10.11646/zootaxa.4748.1.12
http://zoobank.org/urn:lsid:zoobank.org:pub:F45EA07B-C354-437B-A60D-078C95328010
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
195
Zootaxa 4748 (1): 195–200
https://www.mapress.com/j/zt/
Copyright © 2020 Magnolia Press Correspondence
Accepted by A.-P. Liang: 5 Feb. 2020; published: 5 Mar. 2020
On the taxonomic position of the genera Krundia, Breukoscelis, and Uphodato
(Hemiptera, Auchenorrhyncha, Fulgoroidea) described from the Insect
Limestone of the Isle of Wight
VLADIMIR M. GNEZDILOV & ALEXANDR F. EMELJANOV
Zoological Institute, Russian Academy of Sciences, 1 Universitetskaya Emb., St Petersburg 199034, Russia.
E-mails: vmgnezdilov@mail.ru, vgnezdilov@zin.ru
Three fossil genera, Krundia Szwedo, 2019, Breukoscelis Szwedo, 2019, and Uphodato Szwedo, 2019, described in the
family Issidae are transferred to other families according to the shape and venation of forewings. Krundia Szwedo is
transferred to the family Tropiduchidae. Breukoscelis Szwedo is placed in synonymy under Uphodato Szwedo and trans-
ferred to the family Cixiidae. Forewing venation of the members of Issidae, Tropiduchidae, and Cixiidae are discussed
and illustrated.
Key words: systematics, morphology, new synonym, new combinations, Eocene, Tropiduchidae, Cixiidae, Bucini,
Eucarpiini
In a recently published comprehensive study on Hemipteran insects from the Late Eocene of Isle of Wight (Szwedo et al.
2019) three new genera with four new species were described by J. Szwedo in the family Issidae Spinola, 1839. However,
our study of imprint photos (Szwedo et al. 2019, Pl. 8, figs 5–8) and drawings (Szwedo et al. 2019, figs 45–48) provided
for these new taxa suggests that they are better placed in different families. Krundia korba Szwedo, 2019 belongs to the
family Tropiduchidae Stål, and may be tentatively placed in the tribe Bucini Gnezdilov, Bartlett et Bourgoin, 2016 cur-
rently comprised of the genus Buca Walker with three extant species from South America (Gnezdilov et al., 2016). Breu-
koscelis vadimgratshevi Szwedo, 2019, B. phrikkosus Szwedo, 2019, and Uphodato garwoterus Szwedo, 2019 belong
to the family Cixiidae Spinola, 1839 and may be placed in the tribe Eucarpiini Emeljanov, 2002. Moreover, Breukoscelis
Szwedo, 2019 and Uphodato Szwedo, 2019 are synonyms.
Material and methods
Morphological terminology generally follows Anufriev & Emeljanov (1988) and Gnezdilov & Bartlett (2018) for fore-
wing venation.
Taxonomy
Family Cixiidae Spinola, 1839
Subfamily Cixiinae Spinola, 1839
Tribe Eucarpiini Emeljanov, 2002
Genus Uphodato Szwedo, 2019
Note. We suggest placement of Breukoscelis Szwedo, 2019 in synonymy under Uphodato Szwedo, 2019 as indicated by
the basally concaved costal margin of forewing and peculiar venation of the wing discussed below.
A basal concavity in the costal margin of the forewing is a characteristic feature of the tribe Eucarpiini Emeljanov of
Cixiidae and some Achilidae (the genera Tropiphlepsia Muir, 1924, Apateson Fowler, 1900, and Achilixius Muir, 1923),
with their wings held steeply tectiform in repose. This concavity serves for optimal position of the hind legs before jump-
ing. Issidae have no such concavity, as their legs are covered by forewings before jumping and are angled differently at
GNEZDILOV & EMELJANOV
196 · Zootaxa 4748 (1) © 2020 Magnolia Press
jumping (Anufriev & Emeljanov, 1988, fig. 11: 7). The forewing in Issidae has the bulla in basal half under R1 close to first
furcation of the radius. Most issid species have convex forewings except, for example, Oronoqua Fennah, 1947 and Eupi-
lis Walker, 1857 with flat wings; however, these genera have forewings with convex costal margins, without a concavity.
The forewing imprint of Uphodato garwoterus Szwedo, 2019 exhibits a distinct “cixiid clavus”, with Pcu + A1 joining the
wing margin before the claval apex, and with tubercles on setal bases on the veins (Fig. 1) which are characteristic features
of Cixiidae, but not known for Achilidae.
Uphodato Szwedo, 2019 and Breukoscelis Szwedo, 2019 are distinguished by the stem R + M on the forewing (Figs
1, 2). Such a condition is known for extant Dilacreon orpheus (Fennah, 1956) (Fig. 4), described from Caroline Islands
(Fennah, 1956), within the Eucarpiini. Uphodato and Breukoscelis are also characterized by a unique vein which may
be treated as a transverse m-cua crossvein based on its position on the wing (Figs 1–3), but very likely it is the “root” of
CuA, becoming a medio-cubital vein as a result of “interception” of the CuA by the M. In favor of this interpretation is
the wing venation in members of the genus Dilacreon Fennah, 1980 illustrated by Fennah (1956, 1980), in particular, D.
orpheus, which has a short stem R + M. However, on the drawings for Uphodato and Breukoscelis (Szwedo et al., 2019,
figs 46–48) CuA is not furcating, which is not typical for Cixiidae. If our interpretation of vein “interception” is true, then
CuA acquires the usual character of furcating. Thus, the species of the genus Dilacreon have the basal part of CuA1 and
prenodal vein m-cua situated symmetrically as though ready for “interception”. The extravenal pterostigma is not visible
on the imprints of fossil taxa; however, within extant Eucarpiini it is small or completely obsolete.
Uphodato Szwedo, 2019
Uphodato Szwedo, 2019: 367 (in Szwedo et al. 2019). Type species: Uphodato garwoterus Szwedo, 2019.
Breukoscelis Szwedo, 2019: 365 (in Szwedo et al. 2019), syn. n. Type species: Breukoscelis vadimgratshevi Szwedo, 2019.
Uphodato garwoterus Szwedo, 2019
Uphodato garwoterus Szwedo, 2019: 367, pl. 8: 8, fig. 48 (in Szwedo et al. 2019).
Uphodato vadimgratshevi (Szwedo, 2019), comb. n.
Breukoscelis vadimgratshevi Szwedo, 2019: 365, pl. 8: 6, fig. 46 (in Szwedo et al. 2019).
Uphodato phrikkosus (Szwedo, 2019), comb. n.
Breukoscelis phrikkosus Szwedo, 2019: 367, pl. 8: 7, fig. 47 (in Szwedo et al. 2019).
Family Tropiduchidae Stål, 1866
Subfamily Elicinae Melichar, 1915
Tribe Bucini Gnezdilov, Bartlett et Bourgoin, 2016
Genus Krundia Szwedo, 2019
Krundia Szwedo, 2019: 365 (in Szwedo et al. 2019). Type species: Krundia korba Szwedo, 2019.
Krundia korba Szwedo, 2019
Krundia korba Szwedo, 2019: 365, pl. 8: 5, fig. 45 (in Szwedo et al. 2019).
Note on forewing veneation. Forewing enlarged apically. First furcation of R and M at the same level in basal third of the
wing. R and M multibranched in the distal part of the wing. CuA single. Transverse veins ir, r-m, im, and m-cua form ap-
pearance of nodal line (Fig. 5). According to these features K. korba is close to species of the genus Buca Walker, 1858, in
particular, the venation is similar with R1 furcating apically known for Buca truncoptera Gnezdilov, Bartlett et Bourgoin,
2016 (Fig. 6).
The apically enlarged forewing is not a typical feature for Issidae. In Issidae R furcates before M, main veins poorly
furcate apically or not furcate, nodal line never present (Figs 7, 8).
TAXONOMIC POSITION OF KRUNDIA, BREUKOSCELIS, AND UPHODATO Zootaxa 4748 (1) © 2020 Magnolia Press · 197
FIGURES 1–4. Eucarpiini, forewing. 1. Uphodato garwoterus Szwedo; 2. U. vadimgratshevi (Szwedo); 3. U. phrikkosus (Szwedo);
4. Dilacreon orpheus (Fennah). 1–3 after Szwedo et al. (2019), modified, 4 after Fennah (1956).
GNEZDILOV & EMELJANOV
198 · Zootaxa 4748 (1) © 2020 Magnolia Press
FIGURES 5–6. Bucini, forewing. 5. Krundia korba Szwedo (after Szwedo et al. (2019), modified); 6. Buca truncoptera Gnezdilov,
Bartlett et Bourgoin (after Gnezdilov et al. (2018), modified).
TAXONOMIC POSITION OF KRUNDIA, BREUKOSCELIS, AND UPHODATO Zootaxa 4748 (1) © 2020 Magnolia Press · 199
FIGURES 7–8. Issidae, forewing. 7. Kovacsiana abyssinica Synave (after Gnezdilov, 2017, modified); 8. Waorania jaguarina Gnez-
dilov et Bartlett (after Gnezdilov & Bartlett, 2018).
Acknowledgements
We wish to thank Dr. Charles R. Bartlett (Newark, Delaware, USA) and Dr. Michael R. Wilson (Cardiff, UK) for their
comments on English and Dr. Jacek Szwedo (Gdańsk, Poland) for his review of the manuscript.
The study was performed in the framework of the Russian State Research project (No. АААА-А19-119020690101-6).
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