Cretaceous Research 52 (2015) 501e506

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

Revision of the unique Early Cretaceous Mecoptera from Koonwarra

(Australia) with description of a new genus and family
Wies1aw Krzemin
ski a, Agnieszka Soszyn

ska-Maj b, *, Alexei S. Bashkuev c,
Katarzyna Kope

cd
a
Institute of Systematic and Evolution of Animals, Polish Academy of Sciences, 31-016 Kraków, Poland
b
Department of Invertebrate Zoology and Hydrobiology, University of Łód
z, 90-237 Łód
z, Poland
c
Paleontological Institute, Russian Academy of Sciences, 117997 Moscow, Russia
d
Pedagogical University of Cracow, 30-084 Kraków, Poland

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

Article history: The study provides revision of three species of Mecoptera: Choristopanorpa drinnani Jell & Duncan,
Received 20 January 2014 Cretacochorista parva Jell & Duncan, Prochoristella leongatha Jell & Duncan and one fossil specimen
Accepted in revised form 4 April 2014 considered previously as Diptera (No. NMVP103336), described by Jell and Duncan (1986) from Koon-
Available online 5 June 2014
warra (Australia). Using fluorescence under UV light allowed the detailed studies of wings venations and
led to reconsideration of taxonomical position of some specimens. New drawings, photographs and some
Keywords:
description were made. Based on the unique early forking of M1þ2, the species P. leongatha was trans-
Scorpionflies
ferred to the newly established genus Australochorista gen. nov. and the new family Australochoristidae
Mesozoic
Koonwarra
fam. nov. The wing venation pattern of A. leongatha comb. nov. resembles that of Prochoristella Riek, but
Orthophlebiidae also shows characters not typical of Permochoristidae and possibly synapomorphic with some Pan-
Australochoristidae fam. nov. orpoidea (sensu Novokshonov, 2002), particularly of orthophlebiid and panorpoid grades. The phylo-
Australochorista gen. nov. genetic relationships of Australochoristidae are not quite clear, but observed taxonomical characters do
Australochorista leongatha comb. nov. not support it as an ancestral group to any living group of Mecoptera including the Australian endemic
Choristidae. This newly described family probably emerged in Australia and extinct without any recently
living representatives.
Ó 2014 Elsevier Ltd. All rights reserved.

1. Introduction
The Koonwarra Fossil Bed (south Gippsland, Victoria) in Australia
was discovered in 1961 (Jell and Duncan, 1986). Using plant micro-
and macrofossils the sediments from Konwarra were dated as
Aptian, Early Cretaceous (126e113 Ma) (Dettmann, 1963; Douglas,
1969, 1973) and the outcrop was named as “one of the great fossil
localities of the Mesozoic Era”. Twenty years later, second extensive
exploration of this site resulted in a collection of a considerable
number of invertebrate taxa (Jell and Duncan, 1986). The Koonwarra
Fossil Bed is considered to be a very diverse and one of the only few
sites with Cretaceous insects in Australia (Grimaldi and Engel, 2005).
The Permian and Triassic insect faunas of Australia including
Mecoptera from southern Queensland and New South Wales were
* Corresponding author. Tel.: þ48 42 635 46 64; fax: þ48 42 635 644 40.

ski),
E-mail addresses: krzeminski@muzeum.pan.krakow.pl (W. Krzemin

ska-Maj),
agasosz@biol.uni.lodz.pl, agnieszka.soszynska.maj@gmail.com (A. Soszyn
fossilmec@gmail.com (A.S. Bashkuev), k_slazyk@poczta.onet.pl (K. Kope

c).
extensively studied by Tillyard (1916, 1917) and Riek (1950, 1953,
1970). The Jurassic Australian Mecoptera are reported only as rare
and mostly fragmentary unidentified specimens, based on frag-
ments of wings (Martin, 2008; Baettie and Avery, 2012), while the
fossil Cretaceous records are very poor (Jell, 2004). Before the
Koonwarra Bed was discovered, the only known Cretaceous
Australian insects were dragonflies (Woodward, 1884; Riek, 1954).
The Koonwarra fauna is more related to the extant one with ma-
jority of the described representatives belonging to, or closely
related to, the currently living genera. Thus, Jell and Duncan (1986)
suggested that Australian fauna is at least 120 million years old and
its “modernization” occurred probably in the Jurassic. We revised
the fossils of Cretaceous Mecoptera from Australia on account of
their uniqueness and importance in understanding the phyloge-
netic relationships of fossil and recent Mecoptera.
Jell and Duncan (1986) described five taxa of Mecoptera from
Australia: four species from single fossils of adults and aquatic
larvae from genus Nannochorista Tillyard, 1917. We present a revi-
sion of this unique Australian Cretaceous material supplementing it

http://dx.doi.org/10.1016/j.cretres.2014.04.004
0195-6671/Ó 2014 Elsevier Ltd. All rights reserved.
502
ski et al. / Cretaceous Research 52 (2015) 501e506
W. Krzemin
by the new detailed drawings, photos and species description and
establish a new genus and new family of fossil Mecoptera.
2. Material and methods
The study was based on six fossils dated as Aptian, Early
Cretaceous (w125e113 Ma) collected from The Koonwarra Fossil
Bed, in the Gippsland Basin (Jell, 2004). This sedimentatory basin
laid down in freshwater and fluviatile environment and belonged
to Strzelecki Ranges high of Korumburra Group (Drinnan and
Chambers, 1986). This outcrop is situated 3 km east of Koonwarra
and 142 km south-east of Melbourne on the South Gippsland
Highway near Tarwin, Victoria (Fig. 1). This locality is entered as
NMVPL425 in the Museum of Victoria Paleontological Locality
Register. The investigated specimens are deposited in the Paleon-
tological Collection, Museum Victoria, Melbourne, Australia
(NMVP).
The specimens were studied with use of Leica MZFLII stereo-
microscope, under transmitted and reflected light. The material
was also treated with UV light to find some fluorescing remains of
organic matter invisible in normal light. The photographs were
made with the Leica DFC295 camera attached to a microscope
under reflected light. Drawings were made from the photographs
and digitally processed.
The terminology of wing venation follows the combination of
Issiki (1933) and Willmann (1989). The nomenclature of fossil taxa
is based on Novokshonov (2002). The geological time scale follows
Cohen et al. (2013).
3. Systematic paleontology
Order: Mecoptera Hyatt & Arms, 1891
Superfamily: Panorpoidea Latreille, 1805
Family: Orthophlebiidae Handlirsch, 1906
Genus: Choristopanorpa Riek, 1950
Type species. Choristopanorpa bifasciata Riek, 1950
Choristopanorpa drinnani Jell and Duncan, 1986
Fig. 1. Map of Victoria showing the location of Koonwarra Fossil Bed (redrawn from Tosolini et al., 1999).
Material. Holotype NMVP102506, fore and hind wings.
Figs. 2e3
Remarks. We provide new drawings of the fore and hind wings of
the holotype (Figs. 3AB), corrected and improved, with venation
visible only under UV light. Description of the species follows Jell
and Duncan (1986).
Genus: Cretacochorista Jell and Duncan, 1986
Type species. Cretacochorista parva Jell and Duncan, 1986
Cretacochorista parva Jell and Duncan, 1986
Material. Holotype NMVP103248, wings.
Figs. 4e5
Remarks. We confirm the genus Cretacochorista established by Jell
and Duncan (1986). However, we transfer this taxon from the
family Choristidae to Orthophlebiidae based on the number of
veins, corroborating the previous suggestion by Novokshonov
(1997a). The representatives of Choristidae have similar venation
in both wings, Rs as well as M are 4-branched and fused at the base
with Cu1 (Willmann, 1989). However, venation in Orthophlebiidae
differs between fore and hind wings. Rs sector of Orthophlebiidae is
at least 5-branched in both wings, while the medial sector is at least
4-branched (Willmann, 1989; Qiao, et al. 2012), such as in Ortho-
phlebia liassica Tillyard, 1933. The new corrected drawings of fore
and hind wings are improved with venation visible only under UV
light (Figs. 5AB).
Orthoplebiidae incerta sedis
Material. NMVP103336, fragment of the wing.
Remarks. Jell and Duncan (1986) classified this specimen as
dipteran and named it Limoniidae indet. (page 170, Fig. 49B,C).
However, we identified it as a representative of family Ortho-
phlebiidae. An identification to a lower taxonomical level is
impossible as some crucial characters are missing in the only pre-
served wing fragment.
Family Australochoristidae fam. nov.
Type genus Australochorista gen. nov., by monotypy.
Diagnosis. Distinguishable from the other families by the following
possible apomorphies: oval shape of the wing, only two times longer

ski et al. / Cretaceous Research 52 (2015) 501e506
W. Krzemin 503

Fig 2. Choristopanorpa drinnani Jell and Duncan, 1986; Holotype NMVP102506; Photograph of holotype.

Fig. 3. Choristopanorpa drinnani Jell and Duncan, 1986; Holotype NMVP102506; Line drawings of: (A) fore wing, (B) hind wing.

than wide; numerous crossveins between R1, Rs, and M branches, and without anterior branches, Mb and CuA not fused, connected only by
the undoubtedly main autapomorphy, which is very early forking short crossvein, Cu diverging into two main branches very close to
vein M1þ2 (near mid-length of wing); followed by combination of wing base (possible synapomorphies with Panorpoidea), M five
other characters: short Sc, slightly extending beyond midwing, branched setae on the legs delicate, not arranged in rings.

Fig. 4. Cretacochorista parva Jell and Duncan, 1986; Holotype NMVP103248; Photograph of holotype.
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W. Krzemin

Fig. 5. Cretacochorista parva Jell and Duncan, 1986; Holotype NMVP103248; Line drawings of: (A) fore wing, (B) hind wing.

Genus: Australochorista gen. nov.
Type species. Australochorista leongatha (Jell and Duncan, 1986)
comb. nov.
Etymology. Combination of Australo- for the occurrence of the
fossil, and Chorista, a genus of Choristidae, usually used as a com-
mon suffix for mecopteroid insects. Gender feminine.
Diagnosis. The same as for the family by monotypy.
Australochorista leongatha (Jell and Duncan, 1986) comb. nov.
Prochoristella leongatha Jell and Duncan, 1986
Material. Holotype NMVP102512, the only specimen known.
Figs. 6e7
Description. Small-size scorpionfly, body 5.5 mm long. Head only
partly visible, with extremely large eyes. Antennae with long, slender
flagellum, only 16 segments visible; flagellomeres short, wide,
covered with delicate setae. Legs with scattered soft/delicate setae,
not organized in rings. Fore wing shape oval with a ratio of 2:1 (length
to width), numerous crossveins, the vein Sc reaches midwing, no
branches present. The pterostigmal spot well marked. Rs with 5
branches, R2 forks on R2a and R2b, R2þ3 more than twice longer than
R4þ5.Vein Mb divided into five branches, M1þ2 fork very close to Mb
fork and only 2.5 times longer than M3þ4þ5.Veins Mb and CuA not
fused, connected only by short crossvein. Cu diverges into CuA and
CuP almost at the base of the wing. A1 and A2 possibly connected at
the base of the wing.
Remarks. The diagnosis and description of the wing venation by Jell
and Duncan (1986) (pages 161e162, Fig. 43AeC) is formulated
incorrectly and not completely. We studied the holotype in details,
provided the new description and drawing of fore wing (Fig. 7) and
supplemented the description with important character consid-
ering legs pubescence.
4. Discussion
Jurassic fossil Mecoptera are rare from Australia and the
discovered fossils were not studied in details (Martin, 2008, Baettie
and Avery, 2012). Thus it is impossible to reconstruct the transitions
of mecopteran fauna on this continent in Mesozoic. In early
Cretaceous the environment had started to change according to
expansion of angiosperms. It was followed by huge changes in
Cretaceous fauna. The Koonwarra materials are dated Aptian, Early
Cretaceous (126e113 Ma) (Dettmann, 1963; Douglas, 1969, 1973).
Thus the detailed and correct description of this material is crucial.
Jell and Duncan (1986) described more than 70 species of insects
belonging to 12 orders from the Koonwarra Fossil Bed with
Mecoptera as a small component of the Cretaceous fauna. The au-
thors described four species of scorpionflies from single fossils of
adults and a few specimens of aquatic larvae from genus Nanno-
chorista. Taking into account that the Jurassic stage of Mecoptera
evolution in Australia is not reflected in fossil record, this small
collection of Cretaceous material is extremely precious.
The most valuable effect of this revision is a description of the
new family of Mecoptera e Australochoristidae. The type species
was originally assigned to the genus Prochoristella Riek, 1953
(placed in Mesopanorpodidae, but likely belonging to Permo-
choristidae: Bashkuev, 2011) based on the combination of char-
acters figured by Jell and Duncan (1986). Our re-examination of
the material reveals that its venation differs substantially from
that provided in the original paper, and makes questionable its
relationship to Permochoristidae. Although Australochorista leon-
gatha indeed resembles some unusual representatives of Permo-
choristidae (most of all Prochoristella ignara Novokshonov, 1997 as
well as some other species from the Triassic Madygen Fm:
Novokshonov, 1997b, 2001), it also shows characters not typical of
Permochoristidae, but possibly synapomorphic with some Pan-
orpoidea (sensu Novokshonov, 2002): 1) Sc being shortened and
lacking anterior branches (common in Parachoristidae, Ortho-
phlebiidae, Panorpidae, and Choristidae); and especially 2) Cu
diverging into two main branches very close to wing base, with
base of CuA gently inclined and connected with M by a short
crossvein, as typical for orthophlebiid and panorpoid grades
(Archibald, et al. 2013) in contrast to one-point connection (X-

ski et al. / Cretaceous Research 52 (2015) 501e506
W. Krzemin 505

Fig. 6. Australochorista leongatha comb. nov. (Jell and Duncan, 1986); Holotype NMVP102512; Photograph of holotype.

vein) of most Mesozoic Permochoristidae (Ansorge, 1995) or reg-
ular Y-vein of most Permian representatives of the family
(Novokshonov, 1997a). Vein M generally corresponds to the typical
5-branched M of both Orthophlebiidae and Permochoristidae, but
its branching pattern with very early forking of M1þ2 is unique in
Mecoptera. At the same time, in all Panorpoidea (except Eomer-
opidae and Dinopanorpidae) (Archibald, 2005), setae on legs are
arranged in distinctly encircling pattern, as opposed to both Per-
mochoristidae and Australochorista. That does not support Aus-
tralochoristidae as an ancestral group to any living group of
Mecoptera including Choristidae e another Australian endemic.
However, a secondary disorganization of setae cannot be
completely ruled out, taking into account the evidence from some
Pseudopolycentropodidae (Grimaldi, et al. 2005).
Thus, the taxonomic relationships of the newly described family
with other known taxa are not entirely clear; it represents most
probably a highly specified lateral branch of orthophlebiid stem,
which probably evolved in Australia and extinct without any
recently living representatives.
5. Concluding remarks
The re-examination and revision of Lower Cretaceous Mecop-
tera from Koonwarra Fossil Bed (Australia) using UV light allowed
reconsideration of the taxonomical position of some specimens. As
a result, the new taxonomic position of Australochorista leongatha
comb. nov. in family Australochoristidae fam. nov. is proposed. This
unique fossil represents most probably a branch of scorpionflies,

Fig. 7. Australochorista leongatha comb. nov. (Jell and Duncan, 1986); Holotype NMVP102512; Line drawing of fore wing.
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ski et al. / Cretaceous Research 52 (2015) 501e506
W. Krzemin
which extinct in Australia without any recently living representa-
tives. The newly established family is distinguishable from the
other Panorpoidea families by the main apomorphy e very early
forking vein M1þ2 followed by combination of other possible syn-
apomorphies and characters. We provide also the new corrected
reconstruction of the wings of two holotypes of Orthophlebiidae
from Koonwarra: Choristopanorpa drinnani and Cretacochorista
parva, supplemented with venation visible only under UV light.
This allows us to transfer the second taxon from the family Chori-
stidae to Australochoristidae.
Acknowledgments
We would like to thank Micha1 Grabowski (University of qód
z)
for some linguistic comments to the first version of the manuscript.
The research was partly supported by the Polish National Science
Center, grant no. 2013/09/B/NZ8/03270 as well as The Russian
Foundation for Basic Research no. 12-04-01177.
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