ArticlePDF Available

A Supposed Eupelycosaur Body Impression from the Early Permian of the Intra-Sudetic Basin, Poland


Abstract and Figures

We describe a new specimen of a supposed Paleozoic tetrapod body impression from the Lower Permian Słupiec Formation in the Intra-Sudetic Basin, Poland. The size, integument morphology of belly and part of tail imprints, and the morphology of a well-preserved pes track diagnose the specimen and readily distinguish it from other described specimens of body impressions of Paleozoic tetrapods. The eupelycosaur identity of this new specimen is based on the identification of the footprint Dimetropus leisnerianus (Geinitz, 186310. Geinitz , H. B. 1863. Beiträge zur Kenntnis der organischen Überreste in der Dyas (oder permischen Formation zum Theil) und über den Namen Dyas. Neues Jahrbuch für Mineralogie, Geologie und Paläontologie, : 385–398. View all references), which is connected with the inferred body imprint. The morphology of integument impressions indicates the presence of the various-sized square or rectangular-shaped scales on the bottom part of the belly and tail of this eupelycosaurid trackmaker.
Content may be subject to copyright.
This article was downloaded by: [Maciej Bojanowski]
On: 30 August 2012, At: 07:19
Publisher: Taylor & Francis
Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House,
37-41 Mortimer Street, London W1T 3JH, UK
Ichnos: An International Journal for Plant and Animal
Publication details, including instructions for authors and subscription information:
A Supposed Eupelycosaur Body Impression from the
Early Permian of the Intra-Sudetic Basin, Poland
Grzegorz Niedźwiedzki a b & Maciej Bojanowski c
a Department of Organismal Biology, Uppsala University, Uppsala, Sweden
b Faculty of Biology, University of Warsaw, Warszawa, Poland
c Faculty of Geology, University of Warsaw, Warszawa, Poland
Version of record first published: 29 Aug 2012
To cite this article: Grzegorz Niedźwiedzki & Maciej Bojanowski (2012): A Supposed Eupelycosaur Body Impression from the
Early Permian of the Intra-Sudetic Basin, Poland, Ichnos: An International Journal for Plant and Animal Traces, 19:3, 150-155
To link to this article:
Full terms and conditions of use:
This article may be used for research, teaching, and private study purposes. Any substantial or systematic
reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to
anyone is expressly forbidden.
The publisher does not give any warranty express or implied or make any representation that the contents
will be complete or accurate or up to date. The accuracy of any instructions, formulae, and drug doses should
be independently verified with primary sources. The publisher shall not be liable for any loss, actions, claims,
proceedings, demand, or costs or damages whatsoever or howsoever caused arising directly or indirectly in
connection with or arising out of the use of this material.
Ichnos, 19:150–155, 2012
Copyright c
Taylor & Francis Group, LLC
ISSN: 1042-0940 print / 1563-5236 online
DOI: 10.1080/10420940.2012.702549
A Supposed Eupelycosaur Body Impression from the Early
Permian of the Intra-Sudetic Basin, Poland
Grzegorz Nied´
zwiedzki1,2 and Maciej Bojanowski3
1Department of Organismal Biology, Uppsala University, Uppsala, Sweden
2Faculty of Biology, University of Warsaw, Warszawa, Poland
3Faculty of Geology, University of Warsaw, Warszawa, Poland
We describe a new specimen of a supposed Paleozoic tetrapod
body impression from the Lower Permian Słupiec Formation in
the Intra-Sudetic Basin, Poland. The size, integument morphology
of belly and part of tail imprints, and the morphology of a well-
preserved pes track diagnose the specimen and readily distinguish
it from other described specimens of body impressions of Paleozoic
tetrapods. The eupelycosaur identity of this new specimen is
based on the identification of the footprint Dimetropus leisnerianus
(Geinitz, 1863), which is connected with the inferred body imprint.
The morphology of integument impressions indicates the presence
of the various-sized square or rectangular-shaped scales on the
bottom part of the belly and tail of this eupelycosaurid trackmaker.
Keywords Dimetropus, Tetrapod tracks, Body impression, Early
Permian, Poland
Body impressions of tetrapods are very rare in the fossil
record (Lucas et al., 2010). Most such findings pertain to “am-
phibians” and have been reported only from the Carboniferous-
Permian of Europe and North America (Langiaux et al., 1974;
Langiaux, 1980; Walter and Werneburg, 1988; Martens, 1991;
Lucas et al.).
Here we describe a supposed partially preserved body
impression of a large eupelycosaurian synapsid from an Early
Permian tracksite in Poland (Figs. 1 and 2). The described
specimen was found, initially recognized, and collected by
the second author of this communication and provides direct
evidence of the nature of the integument of a 295-million-year-
old amniote.
The integument is a critically important structure be-
cause it mediates between an organism and its environment.
Address correspondence to Grzegorz Nied´
zwiedzki, Subdepart-
ment of Evolution and Development, Department of Organismal
Biology, Uppsala University, Norbyv¨
agen 18A, 752 36 Uppsala,
Sweden. E-mail:
Accordingly, integumentary structures, morphologies, and func-
tions are very diverse among extant amniote. The integument
propere includes the epidermis and dermis; integumentary
derivatives includes scales, hairs, feathers, claws, and spikes.
As they adapted to subaerial conditions, early reptiles acquired
protection against excessive evaporative water loss through the
body surface lipidization and cornification of their epidermis.
Protection against mechanical trauma was achieved by the
formation of cornified scales that are separated by a less-
cornified epidermis to provide a tough yet flexible body surface.
The body fossil record of early amniotes (Caseasauria, Eu-
pelycosauria, Araeoscelidia) begins in the Late Carboniferous,
about 318–315 Ma (Clack, 2002; Carroll, 2009). Their trace
fossil record begins in the early part of the Late Carboniferous
(Falcon-Lang et al., 2007, 2010, but see also Keighley et al.,
2008). Numerous amniote footprints are widely known from
the upper part of the Upper Carboniferous and Lower Permian
red beds across Pangea, and they sometimes preserve skin
impressions or tail-drag marks (Voigt, 2005; Minter and Braddy,
2006). Little is known about what kinds of scales covered the
bodies of Paleozoic amniotes. Newly discovered trace fossils,
including those described herein, provide a great opportunity to
study the integumentary morphology of large, eupelycosaurian
The isolated inferred body impression is preserved in a
fine-grained, subarkosic sandstone slab. From the same layer,
numerous tetrapod tracks, invertebrate traces, and plant remains
were also collected. The slab was discovered in May 2007
during a field excursion with geology students and is stored
in the Geological Museum, Faculty of Geology, University
of Warsaw (specimen MWG 009709). Assignation of the
footprints associated with the body impression was performed
following an ichnotaxonomic concept that is entirely based on
anatomically controlled characters of the imprint morphology
(Haubold, 1996).
Downloaded by [Maciej Bojanowski] at 07:19 30 August 2012
FIG. 1. Simplified geological map of the Intra-Sudetic Basin (ISB) (modified from Awdankiewicz et al., 2003) with outline map of central Europe, showing its
Vertebrate footprints from the Sudetic area have been
reported for more than 150 years, beginning with H. R. Goeppert
in 1861 (to the Schlesische Gesellschaft f¨
ur Vaterl¨
Kultur) and in 1864–65 (see Czy˙
zewska, 1955). The vertebrate
footprints from the Early Permian of Intra-Sudetic Basin
were later described by Pabst (1908), Czy˙
zewska (1955), and,
more recently, Ptaszy´
nski and Nied´
zwiedzki (2004) and Voigt
et al. (2012).
The specimen described herein came from a site very rich
in trace fossils (mainly vertebrate tracks): the Rotliegendes at
the old Tłumacz´
ow Quarry, Intra-Sudetic Basin, southwestern
Poland (Nemec, 1981; Mastalerz et al., 1995; Awdankiewicz
et al., 2003). The Intra-Sudetic Basin is situated at the northern
margin of the Bohemian Massif in the West Sudetes and is one
of the larger intramontane basins that are widespread along
the Variscan belt of Europe. It is filled with a Lower Car-
boniferous to Lower Permian volcano sedimentary succession
(Awdankiewicz et al., 2003).
The sedimentary rocks exposed in the Tłumacz´
ow quarry
consist of reddish-brown mudstones and sandstones that belong
to the Słupiec Formation (Nemec, 1981; Mastalerz et al.,
1995; Awdankiewicz et al., 2003). The deposits at Tłumacz´
accumulated on a flat, mud-sand plain and the shore of a shallow,
ephemeral lake (Mastalerz et al., 1995; Awdankiewicz et al.,
2003); they preserve wavy ripples, mud crack polygons, surfaces
with rain drops, and microbially induced structures. They also
commonly show bioturbation, of both invertebrate and verte-
brate origin, and numerous Walchia twigs and fructifications
(Mastalerz et al., 1995).
The tetrapod ichnoassemblage from Tłumacz´
ow, presently
undergoing detailed study, is very rich containing the ichnotaxa
Downloaded by [Maciej Bojanowski] at 07:19 30 August 2012
152 G. NIED´
FIG. 2. A supposed eupelycosaurian body impression from the Tłumacz´
ow tracksite: m – manus imprint; p – pes imprint; dc – dessication cracks. A, Photograph
of the specimen (MWG 009709) and outlines of the resting traces. B and C, Details of integument morphology from the belly and tail portions of the body
impression. Scale bar: 5 cm.
Amphisauropus kablikae (Geinitz and Deichm¨
uller, 1882),
Ichniotherium cottae (Pohlig, 1885), Dimetropus leisnerianus
(Geinitz, 1863), and Dromopus lacertoides (Geinitz, 1861),
which can be referred to seymouriamorph, diadectomorph,
“pelycosaurian”-grade synapsid, and early sauropsid trackmak-
ers, respectively. The cosmopolitan ichnofauna recorded at
ow corroborates the widely regarded hypothesis of
global vertebrate faunal uniformity during the Early Permian
(Lucas and Hunt, 2006; Voigt et al., 2012). Furthermore,
these footprints represent the easternmost occurrence of typical
Rotliegendes tetrapod tracks on the paleogeographic map of the
Central European Early Permian (see Voigt, 2005).
The inferred body impression is preserved in convex hypore-
lief on the underside of a bed. Three tracks, an association of
the left manus and pes and a deformed probably left pes (all
tracks probably were left by the same individual), are located
in association with the inferred body impression and are quite
clearly visible (Fig. 2A). The supposed imprint of the body is
not located between the left and right pes or manus imprints and
all three tracks are imprints of left limbs. The manus and pes are
pentadactyl, plantigrade, and show clear sole impressions. One
of the pes imprints is well preserved and demonstrates distinctive
features of Dimetropus (Fig. 3), which are interpreted as tracks
Downloaded by [Maciej Bojanowski] at 07:19 30 August 2012
FIG. 3. Close up of Dimetropus leisnerianus (Geinitz, 1863), left manus
and pes imprints associated with the body trace specimen (MWG 009709)
discovered at the Tłumacz´
ow Quarry: m – manus imprint; p – pes imprint. Scale
bar: 3 cm.
of the carnivorous members of Eupelycosauria (Haubold, 1971,
2000; Haubold et al., 1995; Voigt, 2005). Dimetropus (Geinitz,
1863) tracks are characterized by an extended sole, pronounced
and circular metatarsophalangeal pads, slender digits, and
tapered claws. There is no other tetrapod tracks on the described
Dimetropus is a widely distributed ichnotaxon in Upper
Carboniferous and Lower Permian red beds (Haubold, 1971,
1984, 1996; Haubold et al., 1995; Voigt, 2005, 2007). Three
ichnospecies have been distinguished: Dimetropus leisnerianus
(Geinitz, 1863), D. bereae (Tilton, 1931), and D. nicolasi (Gand
and Haubold, 1984). The new footrpints described here are
similar to the ichnospecies Dimetropus leisnerianus (Geinitz,
1863). The specimen described herein exhibits only a few
anatomically controlled details of the imprint morphology.
Without well-preserved material, a separation of different
ichnospecies is not reasonable for this ichnogenus (Voigt, 2005).
We refer the analyzed specimen to D.leisnerianus (Geinitz,
1863) as the first available ichnospecies. This is unsurprising
given that the holotype of D. leisnerianus, which is housed in the
Staatliches Museum f¨
ur Mineralogie, Dresden, Germany, comes
from Wambierzyce (Haubold, 1971). The new finding from
ow corresponds in age and ichnofaunal composition to
the track assocation described from Wambierzyce (Pabst, 1908;
zewska, 1955; Voigt et al., 2012).
The new specimen of a supposed synapsid body trace was
left in a rather awkward position during one of the phases of
locomotion of the trackmaker. The left pelvic limb registered
trace spanning almost the entirety of its length, including
impressions of parts of the thigh and lower leg. Imprints of the
left set of the manus and pes are preserved with fine anatomical
details. In both tracks, traces of claws are perfectly visible. In
the pes, all four circiular metatarsophalangeal pads and the full
lengths of the digits are deeply imprinted (Fig. 3). The pes
partially covers the caudal margin of the manus trace. This is
typical for synapsid tracks and may be a function of their specific
mode of locomotion.
The supposed imprints of the part of abdomen and the
proximal part of the tail are located between the traces of
the left manus-pes association and the left limb impression.
The wider part of the imprint probably pertains to the belly
of the animal. Impressions of individual square or rectangular
scales 1–3 cm long from the ventral surface of the animal
are well preserved within the supposed abdominal and tail
traces (Figs. 2B, C); in some places, they are heavily distorted,
irregular, or S-shaped, indicating that the trackmaker moved
while the trace was registered. Nevertheless, all the scale traces
are similar in size and shape.
Dimetropus is considered to be the track morphotype of
many Late Carboniferous and Early Permian non-therapsid
synapsids such as caseids, edaphosaurids, ophiacodontids,
and sphenacodonts (Haubold, 2000; Voigt, 2005). A potential
trackmaker is represented by the sphenacodontia Haptodus (=
Datheosaurus)macrourus (Schroeder, 1905) known from body
fossils from the Late Carboniferous Ludwikowice Formation
near Nowa Ruda (see Reisz, 1986).
A supposed body impression associated with well-preserved
Dimetropus leisnerianus (Geinitz, 1863) tracks (manus and pes
imprints with anatomical details) from the Słupiec Formation
(Lower Permian) represents the first trace fossil evidence
of the integumentary structure of the ventral surfaces of a
eupelycosaurian synapsid. Eupelycosaurian were among the
dominant terrestral vertebarte animals from the Carboniferous
to the end of the Early Permian. They were common from
Downloaded by [Maciej Bojanowski] at 07:19 30 August 2012
154 G. NIED´
their first appearance became progressively larger during the
Early Permian (Reisz, 1986). Skin impressions, or other skin
fossils, are rare in fossil record, but scales are rarely preserved
in pelycosaurs. These structures are liable to discovery only in
exceptional cases. They have been found partially disarticulated
in three specimens of ophiacodonts and one specimen of
Dimetrodon milleri (see Reisz, 1975). The best preserved set of
scales was described from Archaeothyris, latest Carboniferous
genus of Ophiacodontidae from the Nova Scotia (Reisz, 1975).
The oldest record of amniote scales (transverse scale
impressions preserved on digits) comes from the basal part of
the Pennsylvanian Grande Anse Formation, New Brunswick,
Canada (Falcon-Lang et al., 2007, 2010). Scales are considered
as a fundamental amniote synapomorphy. Many groups of
tetrapods from Carboniferous and Permian, both amphibians
and amniotes bore osteoderms (Dias and Richter, 2002), but
keratinous (with beta-keratin) scales appear to be a sauropsid
FIG. 4. Close up of the 1–3 cm long scale imprints organized in similar,
transverse rows. Scale bar: 3 cm.
feature, present in Chelonia, Lepidosauria, Crocodylia, and
Aves (Zug et al., 2001; see also Falcon-Lang et al., 2007).
Eupelycosaurids, however, being outside Sauropsida, would not
be expected to have scales of beta-keratin. Extant mammals do
not have such scales, but that does not mean that basal synapsids
did not and that they were lost somewhere along the line to
therapsids. In extant lizards (monitor lizards) and crocodylians
(alligators and crocodiles), scales are organized in different ways
on the legs, head, and lower and upper parts of the body, but
usually the belly has a characteristic, transverse pattern of square
or rectangular scales (Zug et al., 2001).
The new trace fossil specimen from the Early Permian of the
Intra-Sudetic Basin shows that the ventral surfaces of the bellies
and tails of eupelycosaurian synapsid were covered by square
or rectangular-shaped, 1–3 cm long scales organized in similar,
transverse rows (Fig. 4). A similar distribution of scales was
also identified in the much smaller tetrapod body impression
from the Early Permian of Turyngia (Martens, 1991).
We thank Jerry D. Harris, Spencer G. Lucas, and Martin
G. Lockley for their constructive reviews and Sebastian Voigt
of Freiberg, Germany, for his comments about the specimen
decribed in this paper. We are also grateful to the students for
carrying the specimen out of the quarry and to the colleagues
who helped in transporting it to Warsaw.
Awdankiewicz, M., Kurowski, L., Mastalerz, K., and Raczy´
nski, P. 2003. The
Intra-Sudetic Basin – a record of sedimentary and volcanic processes in late-
to post-orogenic tectonic setting. GeoLines, 16: 165–183.
Carroll, R. L. 2009. The Rise of the Amphibians 365 Million Years of Evolution.
Johns Hopkins University, Baltimore, MD, 366 p.
Clack, J. A. 2002. Gaining Ground: The Origin and Evolution of Tetrapods.
Indiana University Press, Bloomington, 369 p.
zewska, T. 1955. Tropy gad´
ow permskich z Wambierzyc (Dolny ´
Acta Geologica Polonica, 5: 131–160.
Dias, E. V. and Richter, M. 2002. On the squamation of Australerpeton cosgriffi
Barberena, a temnospondyl amphibian from the Upper Permian of Brazil.
Anais da Academia Brasileira de Ciˆ
encias, 74: 477–490.
Falcon-Lang, H. J., Benton, M. J., and Stimson, M. 2007. Ecology of early
reptiles inferred from Lower Pennsylvanian trackways. Journal of the
Geological Society of London, 164: 1113–1118.
Falcon-Lang, H. J., Gibling, M. R., Benton, M. J., Miller, R. F., and Bashforth, A.
R. 2010. Diverse tetrapod trackways in the LowerPennsylvanian Tynemouth
Creek Formation, near St. Martins, southern New Brunswick, Canada.
Palaeogeography, Palaeoclimatology, Palaeoecology, 296: 1–13.
Gand, G. and Haubold, H. 1984. Traces de vert´
es t´
etrapodes du Permien du
bassin de Saint-Affrique (Description, datation, comparaison avec celles du
bassin de Lod`
eve). Revue G´
eologie m´
eenne, 11: 321–348.
Geinitz, H. B. 1861. Saurier-F¨
ahrten des Rothliegenden im Riesengebirge.
Neues Jahrbuch f¨
ur Mineralogie, Geognosie, Geologie und Petrefakten-
kunde, 65–67.
Geinitz, H. B. 1863. Beitr¨
age zur Kenntnis der organischen ¨
Uberreste in
der Dyas (oder permischen Formation zum Theil) und ¨
uber den Namen
Dyas. Neues Jahrbuch f¨
ur Mineralogie, Geologie und Pal ¨
Geinitz, H. B. and Deichm¨
uller, J. V. 1882. Die Saurier der unteren Dyas von
Sachsen. Palaeontographica, 29: 1–46.
Downloaded by [Maciej Bojanowski] at 07:19 30 August 2012
Haubold, H. 1971. Ichnia Amphibiorum et Reptiliorum fossilium. Handbuch
der Pal¨
aoherpetologie, Teil 18. Fischer, Stuttgart.
Haubold, H. 1984. Saurierf¨
ahrten. A. Ziemsen Verlag, Wittenberg Lutherstadt.
Haubold, H. 1996. Ichnotaxonomie und Klassifikation von Tetrapodenf¨
aus dem Perm. Hallesches Jahrbuch Geowissenschaften, Series B, 18:
Haubold, H. 2000. Tetrapodenf¨
ahrten aus dem Perm – Kenntnisstand
und Progress 2000. Hallesches Jahrbuch f¨
ur Geowissenschaften B, 22:
Haubold, H., Hunt, A. P., Lucas, S. G., and Lockley, M. G. 1995. Wolfcampian
(Early Permian) vertebrate tracks from Arizona and New Mexico. In Lucas,
S. G. and Heckert, A. B. (eds.), Early Permian Footprints and Facies.
New Mexico Museum of Natural History and Science Bulletin, 6: 135–
Keighley, D. G., Calder, J. H., Park, A. F., Pickerill, R. K., Waldron, J. W.
F., Falcon-Lang, H. J., and Benton, M. J. 2008. Discussion on ecology of
earliest reptiles inferred from basal Pennsylvanian trackways (Journal, Vol.
164, 2007, 1113–1118). Journal of the Geological Society of London, 165:
Langiaux, J. 1980. Premi`
eres observations a Morteru (St´
ephanien de Blanzy-
Montceau) ichnites, faune, flore. Revue P´
eriodique de Vulgarisation des
Sciences Naturelles et Historiques “La Physiophile” Montceau-le-Mines,
93: 77–88.
Langiaux, J., Parriat, H., and Sotty, D. 1974. Faune fossile du bassin de Blanzy-
Montceau acquisitions r´
ecentes. Revue P´
eriodique de Vulgarisation des
Sciences Naturelles et Historiques “La Physiophile” Montceau-le-Mines,
80: 55–67.
Lucas, S. G. and Hunt, A. P. 2006. Permian tetrapod footprints: Biostratigraphy
and biochronology. In Lucas, S. G., Cassinis, G., and Schneider, J. W.
(eds.), Non-marine Permian Biostratigraphy and Biochronology. Geological
Society, London, Special Publications, 265: 179–200.
Lucas, S. G., Fillmore, D. L., and Simpson, E. L. 2010. Amphibian body
impressions from the Mississippian of Pennsylvania, USA. Ichnos, 17:
Martens, T. 1991. Ein besonderes fossil. Pal ¨
aontologische Zeitschrift, 65:
Mastalerz, K., Prouza, V., Kurowski, L., Bossowski, A., Ihnatowicz, A.,
and Nowak, G. 1995. Sedimentary record of the Variscian orogeny
and climate—Intra-Sudetic Basin, Poland and Czech Republic. Guide
to Excursion B1. XIII International Congress on Carboniferous-Permian,
August 28–September 2, Krak´
ow, Poland. Pa´
nstwowy Instytut Geologiczny.
Minter, N. J. and Braddy, S. J. 2006. The fish and amphibian swimming traces
Undichna and Lunichnium, with examples from the Lower Permian of New
Mexico, USA. Palaeontology, 49: 1123–1142.
Nemec, W. 1981. Tectonically controlled alluvial sedimentation in the Słupiec
Formation (Lower Permian) of Intrasudetic Basin. In International Sympo-
sium Central European Permian, Jabłonna, April 27–29, 1978, 294–311.
nstwowy Instytut Geologiczny.
Pabst, W. 1908. Die Tierf¨
ahrten in dem Rotliegendes, Deutschlands. Nova Acta.
Abhandlungen des Kaiserlichen Leopoldinisch-Carolinischen Deutschen
Akademie der Naturforscher, 89: 315–482.
Pohlig, H. 1885. Fossile Thierf¨
ahrten in dem unteren Rothliegenden von
Friedrichroda. Verhandlungen des Naturhistorischen Vereins der Preußis-
chen Rheinlande und Westfalens, 42: 285–286.
nski, T. and Nied´
zwiedzki, G. 2004. New finds of vertebrate footprints
from the Lower Permian of Wambierzyce, Poland. Geological Quarterly,
48: 199–202.
Reisz, R. 1975. Pennsylvanian pelycosaurs from Linton, Ohio and N´
Czechoslovakia. Journal of Paleontology, 49: 522–527.
Reisz, R. R. 1986. Pelycosauria. Handbuch der Pal ¨
aoherpetologie, Teil 17.
Fischer, Stuttgart.
Schroeder, H. 1905. Datheosaurus macrourus nov. gen. nov. sp. aus dem
Rotliegenden von Neurode: Jahrbuch der K¨
oniglich Preussischen Geolo-
gischen Landesanstalt und Bergakademie, 25: 282–294.
Voigt, S. 2005. Die Tetrapodenichnofauna des kontinentalen Oberkarbon und
Per m im Th ¨
uringer Wald—Ichnotaxonomie, Pal¨
okologie und Biostrati-
graphie. Cuvillier, G¨
ottingen, 1–179.
Voigt, S. 2007. Auf den Spuren der Saurier: Die Tetrapodenf¨
ahrten aus dem
Oberkarbon und Unterperm des SaarNaheBeckens. In Schindler, T. and
Heidtke, U. H. C. (eds.), Kohles¨
umpfe, Seen und Halbw¨
usten. Dokumente
einer rund 300 Millionen Jahre alten Lebewelt zwischen Saarbr ¨
ucken und
Mainz. Pollichia, Bad D¨
urkheim, 288–303.
Voigt, S., Nied´
zwiedzki, G., Raczy´
nski, P., Mastakerz, K., and Ptaszy´
T. 2012. Early Permian tetrapod ichnofauna from the Intra-Sudetic Basin,
SW Poland. Palaeogeography, Palaeoclimatology, Palaeoecology, 313–314:
Walter, H. and Werneburg, R. 1988. ¨
Uber Liegespuren (Cubichnia) aquatischer
Tetrapoden (?Diplocauliden, Nectridea) aus den Rotteroder Schichten
(Rotliegendes, Th¨
uringer Wald/DDR). Freiberger Forschungsheft Reihe C,
419: 96–106.
Zug, G. R., Vitt, L. J., and Caldwell, J. P. 2001. Herpetology: An Introductory
Biology of Amphibians and Reptiles. Academic Press, New York.
Downloaded by [Maciej Bojanowski] at 07:19 30 August 2012
... Dimetropus have been described from numerous strata encompassing the upper Carboniferous and the lower Permian of the United States 63,64,70,71 , Canada 65 , United Kingdom 72 , Germany 60,62,73 , France 66,74 , Poland 75,76 , Italy 77 , Spain 78-80 , and Morocco[81][82][83] . Dimetropus tracks have been generally affiliated with representatives of various non-therapsid synapsid groups, such as early sphenacodontians, ophiacodotids, edaphosaurids, caseids, and varanopids57,60,62,64,67,73,75,76,[84][85][86][87] . Even though the phylogenetic placement of Varanopidae among amniotes has been contentious (see Refs. ...
Full-text available
Lower Permian deposits of the Boskovice Basin in the Czech Republic have long been renowned for extraordinarily abundant specimens of discosauriscid seymouriamorphs, some of which showing exceptional preservation, including widespread soft tissues. The only other tetrapods from the strata are represented by rare temnospondyls. However, recent fieldwork in the Asselian (lowermost Permian) of the Boskovice Basin has yielded a diverse assemblage of tetrapod tracks, illuminating a hidden terrestrial tetrapod diversity. Here, we describe well-preserved isolated tracks, manus-pes couples, and a slab with trackways composed of approximately 20 tracks in at least four different directions belonging to early-diverging, or ‘pelycosaur-grade’, synapsids. The material originates from three localities situated within the Letovice and Padochov formations and is assignable to the ichnotaxon Dimetropus. The best-preserved specimen further shows rare skin impressions, which have not been observed from the hands or feet of early-diverging mammal-line amniotes before. The new material adds to the scarce record of synapsids from the Carboniferous/Permian transitional interval of equatorial Pangea. At the same time, it highlights the significance of the ichnological record of the Boskovice Basin which has long been neglected despite offering evidence for the presence of diverse faunal components that have not been reported from these basinal deposits before.
... Reisz (1975) mentioned the presence of ventral "scales" in Carboniferous "pelycosaurs", but these represent ossified subcutaneous structures homologous with gastralia, not external integumentary scales. Based on preserved skin impressions, Niedźwiedzki and Bojanowski (2012) proposed that "pelycosaurs" had scales at least on the ventral surfaces of the body. This hypothesis seems to be confirmed in a spectacular fashion by the recent discovery of the exceptionally preserved varanopid Ascendonanus, the known specimens of which preserve a scaly skin covering that is at least superficially very similar to that of extant lepidosauromorphs (Spindler et al. 2018). ...
... Dimetropus is well known from Western and Central Europe (Tucker & Smith, 2004;Voigt, 2005;Voigt & Ganzelewski, 2010), Eastern Europe (Lucas, Lozovsky & Shishkin, 1999;Ptaszy nski & Niedźwiedzki, 2004;Niedźwiedzki & Bojanowski, 2012;Voigt et al., 2012), North America (Hunt, Lucas & Lockley, 2004;Sacchi et al., 2014;Voigt & Lucas, 2015), Morocco (Voigt et al., 2011a;Voigt et al., 2011b), and Argentina (Hunt & Lucas, 1998). Dimetropus tracks have been found from the middle Moscovian to the middle Kungurian (Tucker & Smith, 2004;Voigt & Lucas, 2015). ...
Full-text available
A series of sandstone slabs from Hamstead, Birmingham (West Midlands, UK), preserve an assemblage of tetrapod trackways and individual tracks from the Enville Member of the Salop Formation (late Carboniferous: late Moscovian–Kasimovian). This material has received limited previous study, despite being one of the few British sites to preserve Carboniferous tetrapod footprints. Here, we restudy and revise the taxonomy of this material, and document it using 3D models produced using photogrammetry. The assemblage is dominated by large tracks assigned to Limnopus isp., which were made by early amphibians (temnospondyls). A number of similar but smaller tracks are assigned to Batrachichnus salamandroides (also made by temnospondyls). Dimetropus leisnerianus (made by early synapsids) and Dromopus lacertoides (made by lizard-like sauropsids such as araeoscelids) are also present. This ichnofauna contrasts with a slightly stratigraphically older, more extensive and better-studied assemblage from Alveley (Shropshire), which is dominated by small amphibians with relatively rare reptiliomorphs, but which lacks Dromopus tracks. The presence of Dromopus lacertoides at Hamstead is consistent with the trend towards increasing aridity through the late Carboniferous. It is possible that the assemblage is the stratigraphically oldest occurrence of this important amniote ichnotaxon.
... far only very few remains of pelycosaur-grade synapsids are known from the Intra-Sudetic Basin. The exact type locality of Datheosaurus is unknown, but nevertheless this taxon is a possible producer of a track reported by Niedźwiedzki and Bojanowski (2012). These tracks, consisting of imprints of the ventral body side along with footprints, were assigned to a member of the Eupelycosauria, with large caseids being excluded based on their advanced foot type. ...
Full-text available
This study represents a reinvestigation of two historical fossil discoveries, Callibrachion gaudryi (Artinskian of France) and Datheosaurus macrourus (Gzhelian of Poland), that were originally classified as haptodontine-grade sphenacodontians and have been lately treated as nomina dubia. Both taxa are here identified as basal caseasaurs based on their overall proportions as well as dental and osteological characteristics that differentiate them from any other major synapsid subclade. As a result of poor preservation, no distinct autapomorphies can be recognized. However, our detailed investigations of the virtually complete skeletons in the light of recent progress in basal synapsid research allow a novel interpretation of their phylogenetic positions. Datheosaurus might represent an eothyridid or basal caseid. Callibrachion shares some similarities with the more derived North American genus Casea. These new observations on Datheosaurus and Callibrachion provide new insights into the early diversification of caseasaurs, reflecting an evolutionary stage that lacks spatulate teeth and broadened phalanges that are typical for other caseid species. Along with Eocasea, the former ghost lineage to the late Pennsylvanian origin of Caseasauria is further closed. For the first time, the presence of basal caseasaurs in Europe is documented.
Full-text available
All modern mammals are descendants of the paraphyletic non-mammaliaform Synapsida, colloquially referred to as the “mammal-like reptiles.” It has long been assumed that these mammalian ancestors were essentially reptile-like in their morphology, biology, and behavior, i.e., they had a small brain, displayed simple behavior, and their sensory organs were unrefined compared to those of modern mammals. Recent works have, however, revealed that neurological, sensory, and behavioral traits previously considered typically mammalian, such as whiskers, enhanced olfaction, nocturnality, parental care, and complex social interactions evolved before the origin of Mammaliaformes, among the early-diverging “mammal-like reptiles.” In contrast, an enlarged brain did not evolve immediately after the origin of mammaliaforms. As such, in terms of paleoneurology, the last “mammal-like reptiles” were not significantly different from the earliest mammaliaforms. The abundant data and literature published in the last 10 years no longer supports the “three pulses” scenario of synapsid brain evolution proposed by Rowe and colleagues in 2011, but supports the new “outside-in” model of Rodrigues and colleagues proposed in 2018, instead. As Mesozoic reptiles were becoming the dominant taxa within terrestrial ecosystems, synapsids gradually adapted to smaller body sizes and nocturnality. This resulted in a sensory revolution in synapsids as olfaction, audition, and somatosensation compensated for the loss of visual cues. This altered sensory input is aligned with changes in the brain, the most significant of which was an increase in relative brain size.
Full-text available
Briefly reports the results of a recent sedimentological study concentrated on one particular sedimentary sequence of the basin-fill - the Lower Permian Slupiec Formation in the syntectic Intrasudetic Basin. This sequence forms a large sandstone body representing, one of the major basin-filling episodes. Details of depositional environment, and inferences about the possible style of tectonic control on sedimentation are presented.-Author
La description et l'étude statistique d'empreintes de vertébrés récoltées dans le Permien du bassin de St-Affrique montrent qu'elles se rapportent pour la plupart d'entre elles à neuf ichnogenres déjà connus et récoltés également dans l'Autunien du Lodévois. Une ichnoespèce nouvelle est décrite : Dimetropus nicolasi. Ces traces précisent l'existence dans de grands bassins marécageux d'Amphibiens parmi lesquels de lourds Rachitomes, de Captorhinomorphes, d'agiles Eosuchiens à aspect de lézard, d'assez grands Pélycosauriens. L'association palichnologique récoltée en deux endroits différents du bassin de St-Affrique, comparée à l'échelle palichnostratigraphique établie pour le Permien mondial montre que les niveaux de Calmels et Le Viala, ainsi que ceux situés à l'Est du Dourdou, pourraient appartenir à l'Autunien supérieur.