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Acta Palaeontol. Pol. 66 (X): xxx–xxx, 2021 https://doi.org/10.4202/app.00879.2021
A new representative of the “orthopteroid” family
Cnemidolestidae from the lower Permian of Germany
ANDRE NEL and MARKUS J. POSCHMANN
Nel, A. and Poschmann, M.J. 2021. A new representative of the “orthopteroid” family Cnemidolestidae from the lower
Permian of Germany. Acta Palaeontologica Polonica 66 (X): xxx–xxx.
Palatinarkema prokopi gen. et sp. nov., the third German representative of the late Carboniferous–early Permian ar-
chaeorthopteran clade Cnemidolestidae is described and figured. It is compared to the other known genera. We discuss
putative aspects of the paleoecology of these stick-insect-like “orthopteroids”, based on their body and leg structures,
plus the presence of a high disparity of color patterns in their forewings, suggesting an important diversity of biology
and habitats for these insects. Their elongate bodies and legs show some similarities with the extant but distantly related
stick insects. Cnemidolestids possibly expressed cryptic behaviours among the vegetation, using disruptive colors but
also ovale eye-like spots, as in the extant insects.
Key words: Insecta, Polyneoptera, Archaeorthoptera, cryptic behavior, insect predation, Palaeozoic, Germany.
André Nel [anel@mnhn.fr, https://orcid.org/0000-0002-4241-7651], Institut de Systématique, Evolution, Biodiversité
(ISYEB), Muséum national d’Histoire naturelle, CNRS, Sorbonne Université, EPHE, Université des Antilles, CP 50, 57
rue Cuvier, 75005 Paris, France.
Markus J. Poschmann [markus.poschmann@gdke.rlp.de, https://orcid.org/0000-0001-9710-1673], Generaldirektion
Kulturelles Erbe RLP, Direktion Landesarchäologie/Erdgeschichte, Niederberger Höhe 1, D-56077 Koblenz, Germany.
Received 11 February 2021, accepted 12 April 2021, available online 8 September 2021.
Copyright © 2021 A. Nel and M.J. Poschmann. This is an open-access article distributed under the terms of the Creative
Commons Attribution License (for details please see http://creativecommons.org/licenses/by/4.0/), which permits unre-
stricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Introduction
The superorder Archaeorthoptera is one of the major poly-
neopteran insect groups with remarkable morphological
and species diversity in ecosystems recorded since the late
Carboniferous. Among their most “basal” subclades, the or-
der Cnemidolestodea Handlirsch, 1937, even in the restricted
sense of Béthoux (2005), is one of the most diverse with
22 genera in the family Cnemidolestidae Handlirsch, 1906,
from the upper Carboniferous and lower Permian of Europe,
North America, Brazil, China, and possibly Madagascar
(Aristov 2014; Gu et al. 2014). The clade Cnemidolestidae is
supported by strong synapomorphies in their forewing ve-
nation, but it also shows an important diversity of venation
and coloration patterns.
Here we describe one further new genus and species from
the early Permian. It is the third representative of this family
from Germany (Dvořák et al. 2021) and the first record from
the Carboniferous–Permian of the Saar-Nahe Basin.
Institutional abbreviations.—PE, Naturhistorisches Museum
Mainz/Landessammlung für Naturkunde Rhein land-Pfalz,
Ger many.
Other abbreviations.—C, costa; CuA, cubitus anterior;
CuP, cubitus posterior; CuPa, most anterior branch of CuP;
CuPb, posterior branch of CuP; MA, median vein anterior;
MP, median vein posterior; ma-mp, specialized crossvein
between MA and MP+CuA+CuPa; R, radius; RA/P, radius
anterior/posterior; ScP, subcostal posterior.
Nomenclatural acts.—This published work and the nomen-
clatural acts it contains, have been registered in urn:lsid:
zoobank.org:pub:96EB973D-87B2-4086-A26F-C35E93
FD12B0
Material and methods
The fossil was observed using a Leica MZ 7.5 stereomicro-
scope both dry and immersed in isopropanol. Photographs
were taken with specimen immersed in isopropanol using a
Canon EOS 600D SLR camera equipped with a Canon EFS
60 mm macro lens. Original photographs were processed us-
ing the image-editing software Adobe Photoshop. Drawings
were made from enlarged photographs using Inkscape. The
wing venation nomenclature generally follows Kukalová-
Peck (1991) and Béthoux and Nel (2002).
2 ACTA PALAEONTOLOGICA POLONICA 66 (X), 2021
Systematic palaeontology
Superorder Archaeorthoptera Béthoux and Nel, 2002
Order Cnemidolestodea Handlirsch, 1937
Family Cnemidolestidae Handlirsch, 1906
Genus Palatinarkema nov.
ZooBank LSID: urn:lsid:zoobank.org:act:66B028D6-D771-47A1-95
DC-873DDDBDD7E3
Etymology: From Latin palatium, origin of Palatinate, the type region,
and the genus name Narkema.
Type species: Palatinarkema prokopi sp. nov., by monotypy; see below.
Diagnosis.—Forewing characters only. Posterior branch
of MP+CuA+CuPa anteriorly pectinate, with small bifur-
cations on only two branches; stem of anterior branch of
MP+CuA+CuPa very short; veinlet ma-mp very strong
and not aligned with anterior branch of MP+CuA+CuPa;
angle between anterior branch and posterior branch of
MP+CuA+CuPa very acute; no well-defined vein in area
between MA, MP+CuA+CuPa, ma-mp crossvein present;
branches of anterior branch of MP+CuA+CuPa and of RP
simple.
Palatinarkema prokopi sp. nov.
Fig. 1.
ZooBank LSID: urn:lsid:zoobank.org:act:D32AD59D-F659-4837-83
C8-0896E4CCFC45
Etymology: Named after our friend Jakub Prokop, specialist in fossil
insects.
Holotype: PE 2020/5004-LS a, b, part and counterpart of a complete
wing with parts of the basal region hidden by remains of an undeter-
mined insect wing and/or a mineral stain.
Type locality: East of the village of Niedermoschel, Saar-Nahe Basin,
Germany.
Type horizon: Niedermoschel black shale, Jeckenbach Subformation,
Meisenheim Formation, Lower Rotliegend, lower Permian (sensu
Schindler 1997), probably Asselian–?Sakmarian (Schneider and Wer-
neburg 2012; Schneider et al. 2020).
Material.—Holotype only.
Diagnosis.—As for the genus; two parallel darkened bands
in distal half of forewing plus some spots in mid part.
Description.—Based on forewing venation: estimated total
wing length about 17.1 mm, maximum width at midwing
4.4 mm; wing membrane probably originally hyaline with
oblique colored bands; concave ScP slightly curved, run-
ning parallel with costal margin, merging with RA at distal
two-thirds of wing; costal and subcostal areas nearly as
broad where preserved; stem of R diverging from M+CuA
near base of wing; division of RA and RP proximal of mid-
wing, 2.3 mm basal to connection of ScP with RA; strongly
convex ScP+RA simple ending on costal margin basal of
wing apex; numerous oblique crossveins present between
RA and costal wing margin; concave RP mostly posteriorly
pectinate ending with six simple branches terminating at the
wing tip; neutral (neither really convex not concave) vein
MA diverging from M+CuA and running parallel to stem of
R/RP; MA deeply forked into two simple branches, anterior
one MA1 shortly connected to RP and posterior one MA2
nearly straight; CuPa not visible at base, hidden by debris;
stem of MP+CuA+CuPa elongate, 1.7 mm long, bifurcating
into two branches, the posterior branch subdivided into four
branches; and an anterior branch with a short stem and bifur-
cating into two elongate branches, defining a long but rather
narrow area between it, MA and crossvein ma-mp, without
well-defined and strong vein inside it; angle between ante-
rior and posterior branches of MP+CuA+CuPa very acute,
ca. 18°; both branches of anterior branch of MP+CuA+CuP
simple; crossvein ma-mp between MA and anterior branch
of MP+CuA+CuPa very strong, apparently convex, and not
aligned with anterior branch of MP+CuA+CuPa; area be-
tween branches of RP, concave CuP basally dividing into
CuPa and CuPb, CuPb running parallel to MP+CuA+CuPa
towards posterior wing margin; anal area incomplete with
partly preserved first and second anal veins running parallel
to CuP and CuPb.
Stratigraphic and geographic range.—Type locality and
horizon only.
Discussion
Palatinarkema gen. nov. can be attributed to the Archae-
orthoptera as it shares the main diagnostic character of this
superorder, namely the basal fusion of CuA with M and sub-
sequent connection with the anterior branch of CuP as CuPa
(Béthoux and Nel 2002). Furthermore, it displays characters
typical of the order Cnemidolestodea (sensu Béthoux 2005),
such as ScP merging with RA, CuPa merging with MP+CuA,
MP+CuA+CuPa emitting an anterior branch weakly conver-
gent to MA, with a specialized crossvein ma-mp between it
and MA, thus defining a large median area between it and
MA (Béthoux 2005; Gu et al. 2014: figs. 1, 2). The archae-
orthopteran family Tococladidae and “lobeattids” show some
similarities in the forewing venation with Palatinarkema gen.
nov. in the subcostal, radial and cubital veins, with a funda-
mental difference concerning the vein MP separated from
CuA+CuP, resulting in the absence of a median area in the
former (Béthoux et al. 2003, 2012; Chen et al. 2020).
Aristov (2014), using a different diagnosis and wing vena-
tion nomenclature for the Cnemidolestodea, proposed a key
to families. On the basis of the character “MP weak, ending
on CuA or MA, or absent”, Palatinarkema gen. nov. falls
in the family Cnemidolestidae sensu Aristov 2014, which
includes also families Ischnoneuridae Handlirsch, 1906,
Aetophlebidae Handlirsch, 1906, Narkemidae Handlirsch,
1911, and Narkeminidae Pinto and Ornellas, 1991). Indeed,
in Palatinarkema gen. nov., MP is clearly basally fused to
CuA. Béthoux (2005) and Gu et al. (2014: fig. 2 A1, C1)
supposed that the vein MP can be reduced to a weak veinlet
between MA and CuA+CuP.
NEL AND POSCHMANN—NEW PERMIAN INSECT FROM GERMANY 3
The family Cnemidolestidae comprises the following
genera (Aristov 2014; Gu et al. 2014; Dvořák et al. 2021):
Aeto phlebia Scudder, 1885; Amphiboliacridites Lan giaux
and Parriat, 1974; Anarkemina A r i stov, 2014; Argen tino-
nar ke mina Martins-Neto, Gallego, and Brauck mann, 2007;
Bouleites Lameere, 1917; Carbonokata Aristov, 2013; Cne-
mi dolestes Handlirsch, 1906; Evenkiophlebia Ari stov,
2013; Irajanar kemina Martins-Neto, Gallego, and Brauck-
mann, 2007; Ischnoneura Brongniart, 1893; Long zhua Gu,
Béthoux, and Ren, 2011; Narkema Handlirsch, 1911; Nar-
ke mina Martynov, 1930; Narkeminopsis Whalley, 1979;
Nar ke minuta Aristov, 2013; Narkemulla Arist ov, 2013;
Para nar kemina Pinto and Ornellas, 1980; Piesbergopterum
Dvořák, Pecharová, Leipner, Nel, and Prokop, 2021; Proto-
Fig. 1. Forewing of the cnemidolestid archaeorthopteran Palatinarkema prokopi gen. et sp. nov. (holotype, PE 2020/5004-LS) from Niedermoschel,
Saar-Nahe Basin, Germany; Niedermoschel black shale, Lower Rotliegend, lower Permian. Photographs: part (A1), counterpart (A2), interpretive drawing
(A3); small lines along wing represent the limits of the different fields. Abbreviations: CuA, cubitus anterior; CuPa, most anterior branch of cubitus pos-
terior; CuPb, posterior branch of cubitus posterior; MA, median vein anterior; MP, median vein posterior; ma-mp, specialized crossvein between MA and
MP+CuA+CuPa; R, radius; RA/P, radius anterior/posterior; ScP, subcostal posterior. Scale bars 5 mm.
4 ACTA PALAEONTOLOGICA POLONICA 66 (X), 2021
diam phipnoa Brongniart, 1885; Tshunoptera Aristov, 2013;
Velizphlebia Martins-Neto, Gallego, and Brauck mann,
2007; and Xixia Gu, Béthoux, and Ren, 2014.
Palatinarkema gen. nov. shares with Narkema and Long-
zhua a narrow area between MA and MP+CuA+CuPa,
but differs from it in the quite longer anterior branch of
MP+CuA+CuPa with a simple anterior subdivision, and a
pectinate (with only a branch having a short fork) posterior
branch of MP+CuA+CuPa (Béthoux 2005: fig. 3; Gu et
al. 2011). Xixia has also a narrow area between MA and
MP+CuA+CuPa but the crossvein ma-mp is weaker than
in Palatinarkema gen. nov. and has a posterior branch of
MP+Cu A+CuPa simple or with two branches (Gu et al. 2014).
Ischnoneura, Cnemidolestes, Aetophlebia, Carbonokata,
and Anarkemina differ from Palatinarkema gen. nov. in
the crossvein ma-mp aligned with the anterior branch of
MP+CuA+CuPa and the presence of a well-defined sig-
moidal vein in the area between MA and MP+CuA+CuPa
(Béthoux and Nel 2005: figs. 18, 19; Aristov 2013: fig. 1g;
2014; Gu et al. 2014). Narkemulla has also a well-defined
Fig. 2. Habitus of the cnemidolestid archaeorthopteran Protodiamphipnoa gaudryi (Brongniart, 1885) (MNHN.F.R51393) from Commentry, France;
Gzhelian, Carboniferous. Photographs: part (A1) and counterpart (A2). Scale bars 10 mm.
NEL AND POSCHMANN—NEW PERMIAN INSECT FROM GERMANY 5
sigmoidal vein in the area between MA and MP+CuA+CuPa
(holotype specimen PIN 3115/119), but PIN 5384/16 has a
very different area between MA and MP+CuA+CuPa from
that of the holotype of this genus, much narrower and with-
out any vein (Aristov 2013: fig. 1c–e).
Argentinonarkemina, Protodiamphipnoa, Bouleites, and
Irajanarkemina share with Palatinarkema gen. nov. a nar-
row area between MA and MP+CuA+CuPa, but this area
is also very long compared to its width and with a very
short crossvein ma-mp and a very long anterior branch of
MP+CuA+CuPa (Béthoux and Nel 2005; Martins-Neto et al.
2007: figs. 2, 7). Paranarkemina differs from Pala tinarkema
gen. nov. in the absence of the crossvein ma-mp (Martins-
Neto et al. 2007: fig. 6). Narkeminuta has also a very weak
crossvein ma-mp (Aristov 2013: fig. 1i).
Palatinarkema gen. nov. differs from Narkemina, Pies-
bergopterum, Amphiboliacridites, Evenkiophlebia, Tshu no-
ptera, and Veliz p hleb i a in the clearly narrower area between
MA and MP+CuA+CuPa, the more acute angle between
the anterior and the posterior branches of MP+CuA+CuPa,
and the specialized crossvein ma-mp not aligned with the
anterior branch of MP+CuA+CuPa (Martynov 1930: fig. 6;
Langiaux and Parriat 1974; Martins-Neto et al. 2007: figs.
3–5; Aristov 2012, 2013: fig. 1a, f, h; Dvořák et al. 2021).
Their patterns of coloration are also different.
Narkeminopsis has a narrow area between MA and
MP+CuA+CuPa and a pectinate posterior branch of MP+
CuA+CuPa, but a more open angle than Palatinarkema gen.
nov., and a longer stem of anterior branch of MP+CuA+CuPa
(Béthoux and Nel 2005: fig. 20; Aristov 2013: fig. 1b). The
second species Narkeminopsis inversa Aristov, 2013, differs
from the type species Narkeminopsis eddi Whalley, 1979,
and Palatinarkema prokopi gen. et sp. nov. in the presence
of several branches of MP+CuA+CuPa and RP with forks.
Conclusions
The present discovery of Palatinarkema prokopi gen. et
sp. nov. increases our knowledge on the diversity of the
Cnemidolestidae. Unfortunately, the body structures of
the Cnemidolestidae remain poorly known, thus few mor-
pho-functional inferences can be drawn from their bi-
ology, and potentially shed light on the causes for their
extinction. Some late Carboniferous representatives (e.g.,
Protodiamphipnoa tertrini Brongniart, 1893, Cnemidolestes
woodwardi [Brongniart, 1893]) have elongate bodies and
very strong and long legs (but apparently not jumping hind
legs), somewhat reminiscent of those of some extant stick
insects (e.g., Eurycantha calcarata Lucas, 1869; Béthoux
and Nel 2005; Béthoux 2005), possibly suggesting similar
cryptic behavior among leaves and plants. The shape of their
hind wings remains poorly know, especially the relative size
of their anal fan (see Aristov 2012: figs. 2, 3). Thus we can-
not accurately establish their flight ability, but their massive
bodies and large legs suggest a rather poor flight ability. The
hypothesis of a cryptic lifestyle could be supported by the
spectacular diversity of color patterns of their forewings.
These are hyaline in some cases (in Narkemina kata A ristov,
2013), while some have a series of more or less parallel
bands of colors (in Narkeminopsis eddi or Cnemidolestes
woodwardi). Others have a large ovale eye-like spot situated
in the middle of the wing, as in Protodiamphipnoa gaudryi
(Brongniart, 1885) (Fig. 2), or a series of numerous spots
distributed in the distal part of the wing (Piesbergopterum
punctatum Dvořák, Pecharová, Leipner, Nel, and Prokop,
2021), or even a pattern of dark patches distributed over
the entire forewing as in Xixia huban Gu, Béthoux, and
Ren, 2014 (Béthoux and Nel 2005; Aristov 2013; Gu et al.
2014; Dvořák et al. 2021). Palatinarkema gen. nov. has a
pattern of colored parallel bands, probably with a disrup-
tive function during flight or at rest, which has been ob-
served rather frequently among Palaeozoic insects, e.g. also
in Palaeodictyoptera (Jarzembowski 2005; Li et al. 2013).
Stevens et al. (2006) noticed that this strategy is efficient
for insects to escape flying predators such as birds. Large
ovale eye-like spots are clearly less frequent during the
same period, but become more common during the Jurassic
and Cretaceous with the kalligrammatid lacewings, and
later with the Lepidoptera Saturniidae and Nymphalidae
Satyrinae. The eyespots can have multiple functions, such
as intimidation of a predator, deflection of the attack to a
non-vital zone of the body, and in sexual selection (Stevens
2005, Collins 2013; Crees et al. 2021). It is nearly impos-
sible to determine the exact function of the eyespots of
Protodiamphipnoa gaudryi, but their large size and con-
centric ellips of different colors would suggest a function of
intimidation as in some Recent Lepidoptera (Blest 1957), or
sexual selection. In the case of Piesbergopterum punctatum,
the small spots could have had a function of deflection of
attacks as they are small and mainly distributed in the non-
vital distal part of the forewing. In conclusion, such a dis-
parity in the wing patterns strongly suggests an outstanding
diversity with regards to the biology of the Cnemidolestidae.
The “giant” griffenflies (Odonatoptera Meganeuridae)
were the major flyi ng predators dur ing the late Carboni ferous
and until the middle Permian (Nel et al. 2018), before the
emergence of gliding or flying vertebrates. However, they
were probably not able to capture insects hidden among
the vegetation. The development of various strategies of
cryptic behavior among the late Carboniferous and early
Permian Cnemidolestidae suggests that the predation pres-
sure of the small terrestrial vertebrates greatly increased
at that time. The balance between the costs/benefit was
favorable for the development of cryptic coloration in some
of their potential prey.
Acknowledgements
MJP thanks Olivier Béthoux (Muséum national d’Histoire naturelle,
Paris, France) for offering generous help with software. We also thank
Andrew Ross (Museum of Scotland, Edinburgh, UK) and an anonymous
referee for their very useful remarks on the first version of the paper.
6 ACTA PALAEONTOLOGICA POLONICA 66 (X), 2021
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