Cretaceous Research 143 (2023) 105424

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Cretaceous Research
journal homepage: www.elsevier.com/locate/CretRes

Short communication

A new tarsophlebiid dragonfly from the Lower Cretaceous of western

Liaoning, NE China (Insecta: Odonatoptera, Panodonata)
Rui Fang a, b, Daran Zheng a, *
a
State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology and Center for Excellence in Life and
Paleoenvironment, Chinese Academy of Sciences, 39 East Beijing Road, Nanjing 210008, China
b
Shangdong Province Key Laboratory of Depositional Mineralization & Sedimentary Minerals, Shandong University of Science and Technology, Qingdao,
Shandong, 266590, China

a r t i c l e i n f o a b s t r a c t

Article history: A well-preserved dragonfly Turanophlebia liaoningensis sp. nov., belonging to the family Tarsophlebiidae,
Received 23 August 2022 is described from the Lower Cretaceous Yixian Formation in the Huangbanjigou Village, western
Received in revised form Liaoning, China. This is the second species of Turanophlebia recovered in China, although it was widely
11 October 2022
distributed in Eurasia during the Late Jurassic and Early Cretaceous. Turanophlebia liaoningensis sp. nov.
Accepted in revised form 17 November 2022
Available online 23 November 2022
differs from all other species of Turanophlebia in the hindwing pterostigma covering 6e8 cells and
cubito-anal area with five rows of cells between CuA and the posterior wing margin. The discovery
provides new insight into the biodiversity and evolution of Tarsophlebiidae.
Keywords:
Insecta
© 2022 Elsevier Ltd. All rights reserved.
Tarsophlebiidae
Turanophlebia
Yixian Formation
Earliest Aptian
China

1. Introduction In China, only one tarsophlebiid dragonfly, Turanophlebia sinica,

The extinct family Tarsophlebiidae comprises two genera: Tar-
sophlebia Hagen, 1866 and Turanophlebia Pritykina, 1968. Tarso-
phlebia contains two species: Tarsophlebia eximia Hagen, 1866 and
T. minor Fleck et al., 2004, both found in the Upper Jurassic Sol-
nhofen Formation of SW Germany (Fleck et al., 2004). Turano-
phlebia comprise seven species: Turanophlebia martynovi Pritykina,
1968, T. neckini Martynov, 1927, T. anglicana Fleck et al., 2004,
T. mongolica Fleck et al., 2004, T. vitimensis Fleck et al., 2004,
T. sibirica Pritykina, 1977, and T. sinica Huang and Nel, 2009 (Fleck
et al., 2004; Huang and Nel, 2009). Except the former two species
recorded in the Upper Jurassic of Karatau, Southern Kazakhstan, the
remaining were from the Lower Cretaceous: T. anglicana from the
lower Weald Clay Formation (upper Hauterivian) of United
Kingdom, T. mongolica from the Bon-Tsagaan series (Lower Creta-
ceous) of Central Mongolia, T. vitimensis and T. sibirica from the Zaza
Formation (Lower Cretaceous) of Russia, and T. sinica from the
Yixian Formation (lowermost Aptian) of NE China (Fleck et al.,
2004; Huang and Nel, 2009).
* Corresponding author.
E-mail address: drzheng@nigpas.ac.cn (D. Zheng).
was described from the Yixian Formation of western Liaoning (Huang
and Nel, 2009). Here we describe a new well-preserved dragonfly
from the Yixian Formation of the Huangbanjigou outcrop, western
Liaoning and attribute it to the genus Turanophlebia.
2. Material and methods
The specimen described herein was collected from the lower
Yixian Formation of the Huangbanjigou outcrop in Beipiao (Fig. 1),
western Liaoning Province, China (47
370 N, 120
500 E). The
Mesozoic strata near Huangbanjigou Village consist of the Yixian
Formation and the underlying Tuchengzi Formation. The layer
yielding the dragonfly belongs to the Jianshangou Bed of the Yixian
Formation and was dated at ca. 125 Ma (earliest Aptian; Chang
et al., 2009; Cohen et al., 2013).
The specimen was examined dry using a Nikon SMZ1000 ste-
reomicroscope. Photographs were taken using a Canon 5D digital
camera and the line drawings were prepared from photographs
using image-editing software (CorelDraw X7.0 and Adobe Photo-
shop CS6). The specimen is housed in the Nanjing Institute of Ge-
ology and Palaeontology, Chinese Academy of Sciences (NIGPAS).
All taxonomic acts established in the present work have been
registered in ZooBank (see below), together with the electronic

https://doi.org/10.1016/j.cretres.2022.105424
0195-6671/© 2022 Elsevier Ltd. All rights reserved.
R. Fang and D. Zheng
Fig. 1. Geological sketch map and photograph showing the location of the Huangbanjigou outcrop.
publication LSID: urn:lsid:zoobank.org:pub: 4D1952E2-8202-
4AF4-9129-CEA76194DFA9.
The nomenclature of the dragonfly wing venation used in this
paper is based on the interpretations of Riek (1976) and Riek and
Kukalov
a-Peck (1984), as modified by Nel et al. (1993) and Bechly
(1996). Vein abbreviations are as follows: AA, anterior anal; Arc,
arculus; Ax0, Ax1, Ax2, primary antenodal crossveins; Cr, nodal
crossvein; CuAa, distal branch of anterior cubitus; CuAb, proximal
branch of anterior cubitus; IR1, IR2, intercalary radial veins; MA,
anterior median; MP, posterior median; Msp1, median supplement;
N, nodus; ‘O’, oblique vein; Pt, pterostigma; RA, anterior radius; RP,
posterior radius; Rsp1, radial supplement; ScP, posterior subcosta; Sn,
subnodal crossvein; T, discoidal triangle.
The higher classification of fossil and extant Odonatoptera, as
well as family and generic characters followed in the present work,
is based on the phylogenetic system proposed by Bechly (1996,
2014) and Fleck et al. (2004) for Panodonata.
3. Systematic palaeontology
Order Odonatoptera Martynov, 1932
Suborder Panodonata Bechly, 1996
Family Tarsophlebiidae Handlirsch, 1906
Genus Turanophlebia Pritykina, 1968
Turanophlebia liaoningensis sp. nov.
(urn:lsid:zoobank.org:act: 4D1952E2-8202-4AF4-9129-CEA761
94DFA9)
(Fig. 2)
Derivation of name. Named after Liaoning Province yielding the
dragonfly.
Type Material. NIGP163539, deposited in the Nanjing Institute of
Geology and Palaeontology, Chinese Academy of Sciences, Nanjing,
China.
2
Cretaceous Research 143 (2023) 105424
Locality and Horizon. Huangbanjigou Village, Beipiao City, Liaoning
Province, China; Jianshangou Bed of the Yixian Formation, Lower
Cretaceous (lowermost Aptian).
Diagnosis. Five rows of cells between CuA and posterior hind wing
margin; pterostigma covering 6e8 cells; subdiscoidal area divided
into two or three cells.
Description. Left hind wing hyaline (Figs. 2e4); preserved wing
47.3 mm long, 14.7 mm wide; distance from base to arculus 8.1 mm;
from arculus to nodus 18.9 mm; median and submedian areas free
of crossveins; CuP straight, midway between Ax1 and Ax2, basally
closing subdiscoidal area; primary antenodal braces Ax1 and Ax2
stronger than secondary antenodal crossveins, 3.1 mm apart, with
no secondary crossvein between them; Ax1 4.4 mm from wing
base; arculus aligned with Ax2; 14 secondary antenodal crossveins
between ScP and costal margin distal of Ax2, not aligned with 13
crossveins of second row between ScP and RA; about 14 crossveins
in area between RA and RP, and between arculus and subnodus;
MP þ CuA curved just basal of its fusion with MAb; a sharp angle
between MP þ CuA and MAb; a long fusion between MAb and
MP þ CuA before CuA separated from MP, 0.7 mm long, charac-
teristic of the Tarsophlebiidae; RP þ MA, MA and MAb,
MP þ CuA þ MAb, and basal free part of CuA well aligned with
arculus, as in other Tarsophlebiidae; discoidal area basally opened;
presence of a two-celled ‘tarsophlebiid pseudodiscoidal area’ just
distal of MAb in postdiscoidal area; subdiscoidal area divided into
two cells by one crossvein; AA without any strong posterior
branches; anal area with one or two rows of cells; posterior wing
margin rounded; petiole short, 3.4 mm long; AA reaching free part
of CuA at sharp angle; no CuAb; CuA without strong posterior
branches; five rows of small cells between CuA and posterior wing
margin; a relatively long not zigzagged secondary vein (‘postero-
CuA vein’) closely parallel to distal part of CuA in cubito-anal area,
and another one in area between MP and CuA (‘antero-CuA vein’);
CuA reaching posterior wing margin just distal to nodus level; area
R. Fang and D. Zheng
Fig. 2. Turanophlebia liaoningensis sp. nov., holotype (NIGP163539), photograph of the specimen. Scale bar ¼ 10 mm.
between MP and CuA with one row of cells in its basal part but
greatly widened in its distal half, with about 24 rows of cells along
posterior wing margin; postdiscoidal area with two rows of cells in
its basal part, narrowed in its mid part and widened distally, with
five rows of cells between MA and MP along posterior wing margin;
bases of RP3/4 and IR2 present between arculus and nodus,
distinctly nearer to arculus; base of RP3/4 10.3 mm basal of nodus;
base of IR2 apparently on RP3/4; nodal Cr and subnodus strongly
oblique; base of RP2 aligned with subnodus; oblique vein ‘O’ three
small cells distal of base of RP2; numerous Bq crossveins, but
apparently no crossvein in basal part of areas between RA and RP, and
between RP3/4 and IR2; numerous Bq crossveins; about 15 postnodal
crossveins between C and RA, not aligned with 16 postsubnodal
crossveins; base of IR1 eight cells distal of that of RP2; IR1 well-
defined, basally zigzagged and slightly curved distally; one row of
cells between RP1 and IR1; one rows of cells (posterior wing margin
not preserved) in area between IR1 and RP2 basally; area between
RP2 and IR2 distinctly widened distally, area between IR2 and RP3/4
distally widened; area between RP3/4 and MA distally widened.
Right forewing hyaline, 59.2 mm long and 15.0 mm wide. Right
hindwing 60.1 mm long, 14.6 mm wide; pterostigmal area well
preserved (Fig. 5). Pterostigma dark brown in color, elongated and
narrow, 7.8 mm long and 1.2 mm wide, covering 6e8 cells, not
basally recessed; pterostigmal brace oblique and strong, well
aligned with pterostigma base.
4. Discussion
The attribution of Turanophlebia liaoningensis sp. nov. to the family
Tarsophlebiidae is supported by the following synapomorphies
3
Cretaceous Research 143 (2023) 105424
provided by Fleck et al. (2004): primary antenodal crossveins Ax1 and
Ax2 stronger than secondary antenodal crossveins; in all fore- and
hind- wings, pairs of secondary longitudinal concave veins present
“above” and “below” convex veins CuA, MA, and IR2, and closely
parallel to them.
Within the family Tarsophlebiidae, Turanophlebia liaoningensis sp.
nov. can be easily excluded from the genus Tarsophlebia Hagen, 1866
by the following characteristics (Fleck et al., 2004): more than 20
postnodal crossveins (instead of 16 or less); 14 secondary antenodal
crossveins in hind wing (instead of less than ten); five rows of cells
between CuA and posterior hind wing margin (against less than five
rows); larger forewing, ca. 58e60 mm (against 35e41 mm long in
Tarsophlebia eximia Hagen, 1866 and about 26 mm long in T. minor
Fleck et al., 2004); and IR1 longer than in T. eximia.
Within Turanophlebia, T. martynovi, T. neckini, T. anglicana and
T. mongolica can be easily excluded from T. liaoningensis sp. nov. by
their small size, and wing lengths around 39e41 mm instead of
58e60 mm in the latter (Fleck et al., 2004). T. liaoningensis sp. nov.
further differs from T. martynovi in its subdiscoidal area divided into
two or three cells instead of three, the hindwing pterostigma covering
6e8 cells instead of six, IR1 zigzagged for 10 cells instead of four, and
five rows of cells present between CuA and the posterior wing margin
instead of six (Huang and Nel, 2009). T. liaoningensis sp. nov. differs
from T. sibirica and T. anglicana in the following characters: pter-
ostigma covering 6e8 cells, arculus opposite to Ax2, and two rows of
cells present between C and RA distal of pterostigma. T. liaoningensis
sp. nov. further differs from T. mongolica as follows: no secondary
crossveins between Ax1 and Ax2 in hindwing, vein ‘O’ oblique, and 24
rows of cells between MP and CuA along posterior wing margin
instead of only 15. Affinities between T. liaoningensis and T. vitimensis
can be excluded (Fleck et al., 2004), since the former has five rows of
R. Fang and D. Zheng Cretaceous Research 143 (2023) 105424

Fig. 3. Turanophlebia liaoningensis sp. nov., holotype (NIGP163539), line drawing of wing. Scale bar ¼ 10 mm.

cells in the cubito-anal area instead of 9e10 rows in the latter, CuA not with T. sinica in having a similar wing length and more than 10 sec-
reaching the posterior wing margin distinctly distal of the nodus ondary antenodal crossveins in the hindwing, but well differs from
level, and the oblique vein ‘O’ three cells distal of base of RP2 instead the latter in having the hindwing pterostigma covering 6e8 cells
of four in the latter. T. liaoningensis sp. nov. shows close relationship instead of 5e6 in the latter, no secondary crossvein between Ax1 and

Fig. 4. Turanophlebia liaoningensis sp. nov., holotype (NIGP163539), line drawing of left hind wing showing wing venation. Scale bar ¼ 10 mm.

4
R. Fang and D. Zheng
Fig. 5. Turanophlebia liaoningensis sp. nov., holotype (NIGP163539), photograph of pterostigmal part. Scale bar ¼ 2 mm.
Ax2 in the hindwing, the subdiscoidal area divided into two or three
cells instead of three in the latter, and five rows of cells between CuA
and the posterior wing margin instead of six in the latter.
5. Conclusions
Turanophlebia liaoningensis sp. nov. recovered from the Lower
Cretaceous Yixian Formation of western Liaoning represents the
second Turanophlebia species in China, although it is widely
distributed in the Upper Jurassic-Lower Cretaceous of Eurasia. The
discovery increases the diversity of Tarsophlebiidae in China, and it
is anticipated that more fossils will be found predating the later
Cretaceous environmental changes in freshwater ecosystems.
Acknowledgments
We offer our sincere gratitude to editors and two anonymous re-
viewers for their very useful comments on the earlier version of this
manuscript. This research was supported by the Strategic Priority
Research Program of the Chinese Academy of Sciences (XDB26000000),
the Chinese Academy of Sciences (E221020002), and the National
Natural Science Foundation of China (42125201, 41688103).
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