Revision of the ?Permian-Triassic Tetrapod Ichnogenus Procolophonichnium Nopcsa 1923 with Description of the New Ichnospecies P. lockleyi

Abstract

Procolophonichnium is a globally distributed but rare tetrapod ichnogenus ranging from the
?Late Permian/Early Triassic through the Late Triassic. A Permian age of the holotype from the
Karoo of South Africa, as has been proclaimed by some workers, is doubtful. Descriptions lack
coordinates of the type locality but suggest instead a position in Lower Triassic strata.
Procolophonichnium material from North and South Africa, central Europe, North and South America is re-evaluated. Furthermore, the new ichnospecies Procolophonichnium lockleyi from the Upper Triassic of the Germanic Basin is introduced, based on material with well-preserved trackways that show distinct morphological features, such as an extended “heel” behind digit V. Five ichnospecies are considered as valid: P. nopcsai (type ichnospecies), P. haarmuehlensis, P. nectouxi (new comb.) and P. polonicum as Procolophonichnium tracks from ?Upper Permian, Lower and Middle Triassic strata, and P. lockleyi ichnosp. nov. from the Upper Triassic. Trackmakers of Procolophonichnium were most likely therapsids based on the digit configuration and the phalangeal formula derived from preserved pads. A prococolophonid affinity, as was generally proclaimed, or the attribution to other groups such as captorhinids, is less probable.

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Ichnos
An International Journal for Plant and Animal Traces
ISSN: 1042-0940 (Print) 1563-5236 (Online) Journal homepage: http://www.tandfonline.com/loi/gich20
Revision of the ?Permian-Triassic Tetrapod
Ichnogenus Procolophonichnium Nopcsa 1923
with Description of the New Ichnospecies P.
lockleyi
Hendrik Klein, Spencer G. Lucas & Sebastian Voigt
To cite this article: Hendrik Klein, Spencer G. Lucas & Sebastian Voigt (2015) Revision
of the ?Permian-Triassic Tetrapod Ichnogenus Procolophonichnium Nopcsa 1923
with Description of the New Ichnospecies P. lockleyi, Ichnos, 22:3-4, 155-176, DOI:
10.1080/10420940.2015.1063490
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Revision of the ?Permian-Triassic Tetrapod Ichnogenus
Procolophonichnium Nopcsa 1923 with Description of
the New Ichnospecies P. lockleyi
Hendrik Klein
1
, Spencer G. Lucas
2
and Sebastian Voigt
3
1
Saurierwelt Pal€
aontologisches Museum, Neumarkt, Germany
2
New Mexico Museum of Natural History, Albuquerque, New Mexico, USA
3
Urweltmuseum GEOSKOP, Burg Lichtenberg, Thallichtenberg, Germany
Procolophonichnium is a globally distributed but rare tetrapod
ichnogenus ranging from the ?Late Permian/Early Triassic
through the Late Triassic. A Permian age of the holotype from the
Karoo of South Africa, as has been proclaimed by some workers, is
doubtful. Descriptions lack coordinates of the type locality
but suggest instead a position in Lower Triassic strata.
Procolophonichnium material from North and South Africa,
central Europe, North and South America is re-evaluated.
Furthermore, the new ichnospecies Procolophonichnium lockleyi
from the Upper Triassic of the Germanic Basin is introduced,
based on material with well-preserved trackways that show
distinct morphological features, such as an extended “heel” behind
digit V. Five ichnospecies are considered as valid: P. nopcsai (type
ichnospecies), P. haarmuehlensis, P. nectouxi (new comb.) and P.
polonicum as Procolophonichnium tracks from ?Upper Permian,
Lower and Middle Triassic strata, and P. lockleyi ichnosp. nov.
from the Upper Triassic. Trackmakers of Procolophonichnium
were most likely therapsids based on the digit configuration and
the phalangeal formula derived from preserved pads. A
prococolophonid affinity, as was generally proclaimed, or the
attribution to other groups such as captorhinids, is less probable.
Keywords Tetrapod footprints, Permian, Triassic, Procolophonich-
nium, Procolophonids, Therapsids
INTRODUCTION
The ichnogenus Procolophonichnium was established in
1923 by Nopcsa based on a single pes-manus set on a small
slab from the Lower Triassic? of Middelburg in the Karoo
basin of South Africa. The exact geographic and stratigraphic
position of the location is uncertain, and relevant information
from the literature is discussed below. Originally this material
was described and illustrated by Seeley (1904, 1905) under the
body fossil taxonomic name Procolophon. Seeley (1904,
1905) suggested an affinity to procolophonids and to the osteo-
logical taxon Procolophon, which is known from the Lower
Triassic of South Africa and Antarctica (deBraga, 2003). Since
the first description, Procolophonichnium has been identified
in different Lower-Middle Triassic deposits of Europe and
even in the Upper Triassic strata of the Newark Supergroup of
North America (Fig. 1). This wide stratigraphic range matches
the distribution of procolophonids, which encompasses the
Upper Permian through the Upper Triassic. Despite the fact
that the type material from South Africa was never re-studied,
and probably since Seeley’s time no ichnologist has seen it
again, many footprints and trackways, mostly from the Trias-
sic, have been referred to Procolophonichnium. However, the
correctness of these assignments has never been tested, and it
appeared that Procolophonichnium might be merely a waste-
basket ichnotaxon for small amniote footprints with a rela-
tively short digit IV. Here we review the Procolophonichnium
track record reported in the literature from Africa, Europe,
North and South America.
Whereas a Permian record of this kind of tracks is question-
able, Procolophonichnium is well-known from Lower-Middle
Triassic strata in central Europe, especially from the Buntsand-
stein and some marginal marine Muschelkalk deposits of The
Netherlands and Germany (Haubold, 1971a; Demathieu and
M€
uller, 1978; Demathieu and Oosterink, 1983, 1988; Die-
drich, 2002b).
A problem with the re-assessment of all this material is that
the holotype of the type species, Procolophonichnium nopcsai
from South Africa, which was originally housed in the Bavar-
ian State Collection for Palaeontology and Geology of Munich
since 1880 (Seeley, 1904), is lost (O. Rauhut, pers. com.).
However, a plaster cast was given to the Natural History
Museum of London by K. A. v. Zittel (Seeley, 1904, 1905),
and this specimen could be re-located by Sandra Chapman in
the museum collections. Thus, photos of the cast are the only
reference for a comparison of the holotype with other
Address correspondence to Hendrik Klein, Saurierwelt
Pal€
aontologisches Museum, Alte Richt 7, Neumarkt D-92318,
Germany. E-mail: Hendrik.Klein@combyphone.eu
155
Ichnos, 22:155–176, 2015
Copyright ÓTaylor & Francis Group, LLC
ISSN: 1042-0940 print / 1563-5236 online
DOI: 10.1080/10420940.2015.1063490
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footprints referred to Procolophonichnium besides the draw-
ings of Seeley (1904) (Fig. 1). Based on ICZN rules (Art. 75),
an artificial (man-made) cast cannot be approved as a holo-
type, and a so-called “plastotype” as used by different authors
in the ichnological literature is taxonomically without official
status. In this case a neotype should be selected, but this should
come from the same stratum and locality as the (lost) type.
However, as far as we can determine, there is no available
(topotypical) material of P. nopcsai from the type locality.
We admit that this is not an ideal basis for a comprehensive
revision of the ichnogenus. However, given a background of an
increasingly well-preserved record, some of which is published
here for the first time, we started to re-evaluate the ichnogenus
based on a global overview because earlier workers have failed
to do this and were mostly dealing with local occurrences. We
are aware of the fact that the isolated material of the generotype
from South Africa may give rise to doubts about the validity of
the ichnogenus. However, in the literature, Procolophonichnium
is a widely used name. So, we propose to keep this name and the
ichnogenus based on an emended diagnosis given here. The
question of establishing a neotype can be only resolved by the
collection of new material from the stratum typicum in South
Africa in the future. Here we describe also a new Procolopho-
nichnium ichnospecies from the Upper Triassic of Germany
based on well-preserved long trackways, a circumstance that is
rare in the global record.
Institutional and collection abbreviations. BG and
NMS DNaturmuseum, Bozen, Italy; CDUE DDepartement
of Earth Sciences, Chouaib Doukkali University, El Jadida,
Morocco; HOW DPrivate collection Henk Oosterink, Winter-
swijk, The Netherlands; IPB DDivision of Palaeontology,
Steinmann Institute, University of Bonn, Germany; KR D
Kazimierz Rdzanek private collection, Warsaw, Poland;
MCN.PIC. DMuseu de Ci^
encias Naturais, Rio Grande do Sul;
MFW DMuseum Freriks, Winterswijk, The Netherlands;
MHNA DMus
eum d’Histoire Naturelle d’Autun, France; Cg
DCollection G. Gand; NHMUK DNatural History Museum,
London, Great Britain; SKF DSaurierwelt Pal€
aontologisches
Museum, Collection H. Klein, Neumarkt, Germany; UCMP D
University of California Museum of Paleontology, Berkeley,
USA; YPM DYale Peabody Museum, Yale, USA; ZPAL R D
Institute of Palaeobiology of the Polish Academy of Science,
Warsaw, Poland.
MATERIAL AND METHODS
Material in the CDUE, NMS, SKF, UCMP and ZPAL has
been examined directly. Measurements were taken from avail-
able specimens according to standard procedures in Haubold
(1971a, b) and Leonardi (1987) (Tables 1, 2), and others come
from different studies as specified (Table 2). Outline drawings
of tracks and trackways were made on transparency film and
FIG. 1. Paleogeography of the Triassic with distribution of Procolophonichnium footprint localities in the ?Upper Permian/Lower–Upper Triassic. 1 DKaroo
Basin, South Africa; 2 DSanta Maria Formation (Upper Triassic), Brazil; 3 DShnabkaib and Wupatki formations of Moenkopi Group (Lower Triassic, Olene-
kian), Utah, Arizona; 4 DPassaic Formation of Newark Supergroup (Upper Triassic, Norian) of New Jersey; 5 DTimezgadiouine Formation (T4, Middle Trias-
sic), Argana Basin, Morocco, 6 DLower – Upper Triassic of France, The Netherlands and Germany; 7 DWi
ory Formation (Lower Triassic, Olenekian), Poland;
8DMiddle Triassic (Anisian) of northern Italy.
156 H. KLEIN ET AL.
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digitized with a vector-based drawing software. All photo-
graphs were taken under natural light conditions.
GEOGRAPHIC AND STRATIGRAPHIC DISTRIBUTION
OF FOOTPRINTS REFERRED TO
PROCOLOPHONICHNIUM
Type Locality of Procolophonichnium and its Geological
Age
The only data available on the type locality of Procolopho-
nichnium nopcsai are provided by Seeley (1904, p. 287), who
states that the slab of the holotype “is labelled ‘Middelburg,’
presumably the well-known locality in the north-east of Cape
Colony, and with it is a small new theriodont skull from the
same locality which, when the matrix is removed, may prove
to be allied to Hyorhynchus.” Kuhn (1963, p. 43), Haubold
(1971a, 1971b, p. 31) and subsequent authors have assumed
this is an Upper Permian locality, probably largely because
“Hyorhynchus” is a synonym of Permian Pristerognathus
(which, incidentally, is a taxon of Middle, not Late Permian
age).
However, Middelburg is a town in the eastern Karoo basin
built on strata of the Dicynodon (Late Permian) and Lystrosau-
TABLE 2
Track and trackway parameters of different Procolophonichnium ichnospecies
P. haarmuehlensis P. nectouxi P. polon. P. nop.
Ichnospecies
Specimen
MFW
D36
MFW
D52
MFW
D63
MFW
D80
P.
“jageri”
Cg
54 Piste A
Cg
54 Piste B
ZPAL
R7/27Z PAL R7/1–9
NHMU
KR3173
pl 31 24 26 35 28–34 17 16 32 30
pw 36 26 30 42 26–31 14 – 30 28
ml 21 18 21 19 19–24 11 14 – –
mw 24 21 25 22 22–27 14 18 – –
p I–IV 6095756045–55–– – 64

p II–IV 50605045–15
12–41

p I–V 90120108 100–55
––92

PL 107 – 100 96 90–113 55 60 – –
SL 146 – 145 145 130–156 76 89 250 –
TW 128 – – 93 – 48–25 ?45–18 78 –
PA 80–90
10075–959095130–
Measurements in millimeters and degrees; some longer trackways allowed defining ranges, all other values are from short sequences only; pl Dpes length; pw D
pes width; ml Dmanus length; mw Dmanus width; p I–IV, p II–IV, p I–V Ddivarication between digits; PL Dpace length; SL Dstride length; TW Dtrackway
width; PA Dpace angulation. Data from Demathieu and Oosterink (1983) and Demathieu and M€
uller (1978) for P. haarmuehlensis, Gand (1977) for P. nectouxi
and Klein and Nied
zwiedzki (2012) for P. polonicum.
TABLE 1
Track and trackway parameters of P. lockleyi ichnosp. nov.
Specimen SKF 130 I SKF 130 II
pl 30* 30*
pw 25* 25*
ml 20* 20*
mw 20* 20*
p I–IV 21*28
*
p II–IV 15*20
*
p I–V 37*36
*
PL 90, 105, 110, 110, 115, 115, 115 120, 110, 110, 110, 120
SL 150, 155, 190, 180, 185, 195, 195 180, 150, 150, 180
TW 70, 65, 68, 67, 74, 73 67, 73
PA 95.4, 109.5, 105.4, 109.6, 107.9, 110.3104.5, 91.3
Measurements in millimeters and degrees; * Dmean values; pl Dpes length; pw Dpes width; ml Dmanus length; mw Dmanus width; p I–IV, p II–IV, p I–V D
divarication between digits; PL Dpace length; SL Dstride length; TW Dtrackway width; PA Dpace angulation.
REVISION OF THE ICHNOGENUS PROCOLOPHONICHNIUM 157
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rus (Early Triassic) zones, and most of the vertebrate fossil
localities near Middelburg are of Lystrosaurus zone age (e.g.,
Kitching, 1977; Rubidge et al., 1995, fig. 1). Indeed, Kitching
(1977, p. 95) lists “Middelburg district” as an “unreliable
locality” in the Lystrosaurus zone, but only lists the taxon Pro-
lacerta from this locality. Furthermore, Procolophon is a char-
acteristic taxon of the Lower Triassic Lystrosaurus zone, and
the fact that Seeley (1904) originally identified the tracks as
Procolophon implies a Triassic age. Seeley (1904, p. 287) also
remarks that the find demonstrates “an interesting difference
from the absence of bones with footprints in the Trias of this
country.” Thus, it seems very likely that the type locality of P.
nopcsai is Early Triassic in age.
Like Seeley (1904), Nopcsa (1923), when erecting the new
ichnogenus Procolophonichnium based on the Middelburg
material, considered a relationship with the Early Triassic par-
areptile Procolophon. He noted also the presence of
“Procolophon zone” strata near Middelburg (Nopcsa, 1923, p.
138). Kuhn (1958, p. 47, pl. IX) lists Procolophonichnium
from South Africa, obviously Seeley’s tracks, under Triassic
forms and the Middle Beaufort beds as the provenance. Baird
(1986, p.132) gives a Lower Triassic (“Procolophon zone”)
position for this material. Smith (1993, p. 340) mentions the
footprints described by Seeley (1904), together with other
records from the Karoo, but without referring to their exact
stratigraphic position.
Global Distribution
The ichnogenus Procolophonichnium is used for tracks of
Late Permian to Late Triassic age. Supposed Permian records
of the ichnotaxon include the Eisleben Formation of the Saale
Basin, Germany (M€
uller, 1970), the Rio do Rasto Formation
of the Paran
a Basin, Brazil (Silva et al., 2012), as well as
unpublished tracks from the uppermost part of the Ikakern For-
mation, Argana Basin, Morocco (S. Voigt, pers. observation).
However, none of the referred Permian material is well-
enough preserved to confirm the ichnotaxonomic assignment.
The holotype of the type ichnospecies (Seeley, 1904, 1905)
may come from the uppermost Balfourt Formation (Upper
Permian) or the Lower Triassic Katberg Formation, both
Karoo Basin, South Africa, however, the latter is more likely
(see above).
Unequivocal Mesozoic records of Procolophonichnium
include: (1) Lower Triassic (Wi
ory Formation, Olenekian) of
the Holy Cross Mountains, Poland (Ptaszy
nski, 2000; Klein
and Nied
zwiedzki, 2012); (2) Lower-Middle Triassic (Bunt-
sandstein, Olenekian-Anisian) of Germany (Haubold, 1971a;
Demathieu and M€
uller, 1978); (3) Lower Triassic (Moenkopi
Group, Shnabkaib and Wupatki formations, Olenekian) of
Utah and Arizona (Klein and Lucas, 2010b; Thomson, 2013);
(4) Middle Triassic (Anisian-Ladinian) of France (Demathieu,
1977; Gand, 1977); (5) Middle Triassic (Muschelkalk, Vos-
senveld Formation, Anisian-Ladinian) of Winterswijk, The
Netherlands (Demathieu and Oosterink, 1983, 1988); (6) Mid-
dle Triassic (Buntsandstein–Muschelkalk, R€
ot–Karlstadt for-
mations, Anisian) of Germany (Diedrich, 2000, 2002a–c); (7)
Middle Triassic (Timezgadiouine Formation, T4) of the
Argana Basin, Morocco (Klein et al., 2011; Voigt et al.,
2011); (8) Upper Triassic (Santa Maria Formation) of Brazil
(Silva et al., 2008; this article); (9) Upper Triassic (Hassberge
Formation, Coburger Sandstein, Carnian) of northern Bavaria,
Germany (this article); and (10) Upper Triassic (Passaic For-
mation, Norian) of New Jersey, USA (Silvestri and Szajna,
1993; Baird, 1986) (Fig. 1).
Occurrences of Procolophonichnium in the Permian are
rare and not definitive (see below). The well-established strati-
graphic range of the ichnogenus encompasses almost the entire
Triassic, ranging from the earliest Triassic (Induan DLoots-
bergian land-vertebrate faunachron) through the Late Triassic
(Norian DRevueltian land-vertebrate faunachron) (cf. Klein
and Lucas, 2010a). The ichnogenus is best known from exten-
sive European records that encompass the Early—Middle Tri-
assic (Olenekian-Anisian) boundary. Records in the Early and
Late Triassic are much less common (Fig. 1).
SYSTEMATIC PALEOICHNOLOGY
Remarks on Procolophonichnium Ichnotaxobases
Features relevant for the ichnotaxobase of the ichnogenus
are: (1) overall digit proportions, shape and orientation, (2)
development and relative size (length) of the sole. Features rel-
evant for the ichnotaxobases of the ichnospecies are (1) details
of the shape and orientation of digits, (2) digit divarication, (3)
proximal margin of the sole and presence/absence of a “heel”
trace, (4) the relative length of digit V, and (5) the trackway
pattern with relative stride length, pace angulation and rotation
of imprints (Table 3).
Ichnogenus Procolophonichnium Nopcsa 1923
1904 Procolophon: Seeley, p. 287–289, fig. 1 A–C
1923 Procolophonichnium: Nopcsa, p. 138, pl. VI, fig. 6
non 1954 “Procolophonichnium”?: M€
uller, p. 190, fig. 1,
pl. 16 (1, 2), tab. 1.
non 1958 Procolophonichnium (?): Kuhn, p. 34, pl. III,
fig. 21
non 1962 Procolophonichnium:M
€
uller, p. 22, pl. V–VII.
1955 Procolophonichnium: Lessertisseur, p. 105, fig. 58e
1958 Procolophonichnium: Kuhn, p. 46, pl. IX, fig. 2
1963 Procolophonichnium: Kuhn, p. 145, pl. 1, fig. 7, p.
147, pl. 2, fig. 5, p. 155, pl. 6, fig. 36
1970 Rhynchosauroides: Holst et al., figs. 2, 7, pl. I, fig. 3,
pl. II, figs. 4–6
1971a Procolophonichnium: Haubold, pl. III
1971b Procolophonichnium: Haubold, p. 30, fig. 19.1.(7)
158 H. KLEIN ET AL.
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1977 Procolophonichnium: Demathieu, p. 357, fig. 3, pl. 2
(2)
1977 Circapalmichnus: Gand, p. 20, 22, pl. D, figs. 1–2,
pls. 3–4
1978 Procolophonichnium: Demathieu and M€
uller, p. 157–
163, figs. 2–4
1983 Procolophonichnium: Demathieu and Oosterink, p.
13, 39, 41–42, 44–46, figs. 8, 33–50
1983 Phenacopus: Demathieu and Oosterink, p. 15, fig. 10,
p. 16, fig. 12, p. 47, figs. 51–53
1984 Rhynchosauroides: Haubold, p. 147, fig. 98 (13)
1984 Procolophonipus: Haubold, p. 149, fig. 100(5)
1984 Circapalmichnus: Haubold, p. 149, fig. 100(6)
1986 Procolophonichnium: Baird, p. 133, figs. 5–6B.
1988 Procolophonichnium: Demathieu and Oosterink, p.
13, fig. 6
1988 Phenacopus: Demathieu and Oosterink, p. 14,
fig. 7A–B
1990 Rhynchosauroides: Fuglewicz et al., figs. 9(5–8), 10
(1), pls. 8(1–4), 10(1)
2000 Procolophonichnium: Ptaszy
nski, p. 178, fig. 20
2000 Procolophonichnium: Diedrich, p. 385, fig.6A–D
2002a Procolophonichnium: Diedrich, p. 82, fig. 3D, p. 85,
fig. 4
2002b Procolophonichnium: Diedrich, p. 45, figs. 8–9
2002c Procolophonichnium: Diedrich, pl. II, fig. 9, fig. 7,
2007 Procolophonichnium: Valdiserri and Avanzini, p.
113, figs. 8–9
2007: Circapalmichnus: Gand et al., p. 16, fig. 6T, pl. 2O
2008 Dicynodontipus: Silva et al., p. 102, fig. 3, p. 103,
fig. 4, p. 104, fig. 5, p. 105, fig. 6
2010 Procolophonichnium: Klein and Lucas, p. 5–6, p. 11,
fig. 9A–C
2011 Procolophonichnium: Todesco and Bernardi, p. 207, pl. 1
2011 Procolophonichnium: Klein et al., p. 227, 9A–D
2012 Procolophonichnium: Klein and Nied
zwiezki, p. 51,
fig. 50C, F, I–J
Type ichnospecies: Procolophonichnium nopcsai Kuhn,
1963, holotype specimen figured in Seeley (1904, p. 287–289,
fig. 1 A–C) and Nopcsa (1923, p. 138, pl. VI, fig. 6) (Fig. 2).
Stratigraphic and geographic distribution: ?Late Permian/
Early Triassic to Late Triassic of Africa (Morocco, South
Africa), Europe (France, Germany, Netherlands, Italy,
Poland), North America (Arizona, Utah, New Jersey), South
America (Brazil).
Revised diagnosis (partially based on Haubold,1971a, b):
Pentadactyl and semi-plantigrade to plantigrade asymmetric
footprints of small quadrupeds with digits increasing in length
from I–IV; digit IV subequal in length with digit III; digit V
subequal in length with digit II. Manus similar in shape but
smaller than pes and positioned anterior to or being slightly
overstepped by the latter posteriorly. Differs from the most
similar tracks of other ichnogenera in the following features:
(1) from Rhynchosauroides and Dromopus by digit propor-
tions, with minor differences of length and relatively longer
digit V, and (2) from Rhynchosauroides, Dromopus and Vara-
nopus by proportionately shorter and stouter digits, by
(mostly) straight to outward curved digits vs. the strong inward
curvature of digits in the latter, and by semi-plantigrade to
plantigrade imprints vs. digitigrade to semi-plantigrade
imprints observed in these latter ichnotaxa.
TABLE 3
Diagnostic features of Procolophonichnium ichnospecies
Diagnostic features of Procolophonichnium ichnospecies
P. nopcsai P. haarmuehlensis P. nectouxi P. polonicum P. lockleyi
Digits straight,
tapering
Digits I–II straight, III–IV
curved outward with
margins parallel or
distally expanded
Digits straight or curved
inward with margins
parallel or distally
expanded
Digits straight to curved
outward, tapering
Digits straight with
margins parallel or
distally expanded
Large digit
divarication
Large digit divarication Small digit divarication ? Small digit divarication
Proximal margin of
sole straight
Proximal margin of soles
traight or posteriorly
convex
Proximal margin of sole
straight or posteriorly
convex
? Proximal margin of sole
with elongated heel of
digit V
Digit V moderately
long
Digit V moderately long Digit V (if present) very
long
Digit V moderately long Digit V very long
——— Short strides Short strides Long strides Long strides
——— Low pace angulation Low pace angulation High pace angulation High pace angulation
——— Outward rotation of pes Inward rotation of pes Outward rotation of pes Inward rotation of pes
REVISION OF THE ICHNOGENUS PROCOLOPHONICHNIUM 159
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Procolophonichnium nopcsai Kuhn 1963 (Type
ichnospecies) (Fig. 2)
1904 Procolophon: Seeley, p. 287–289, fig. 1 A–C
1923 Procolophonichnium: Nopcsa, p. 138, pl. VI, fig. 6
1958 Procolophonichnium: Kuhn, p. 46, pl. IX, fig. 2
1963 Procolophonichnium: Kuhn, p. 145, pl 1, fig. 7
1971b Procolophonichnium nopcsai: Haubold, p. 30,
fig. 19.1.(7)
Holotype: An isolated pes-manus set from the ?Lower Trias-
sic near Middelburg, Karoo basin, South Africa illustrated in
Seeley (1904, p. 287–289, fig. 1 A–C) and Nopcsa (1923, p.
138, pl. VI, fig. 6) (Fig. 2). The original slab without a specimen
number was, based on Seeley (1904), deposited in the Bavarian
State Collection for Palaeontology and Geology, Munich, Bava-
ria, Germany, and is lost as it was destroyed during WWII. An
artificial (plaster) cast is housed in the Natural History Museum,
London and catalogued as NHMUK R3173 (Figs. 2A–B).
FIG. 2. Photographs and sketches of Procolophonichnium nopcsai. A–B, overview and detail of holotype plaster replica (NHMUK R3173) in collections of the
Natural History Museum, London. C, sketch of holotype specimen with left pes-manus set, redrawn from Seeley (1904). D. Reinterpreted drawing of holotype
specimen in this paper. Photographs by the Natural History Museum, London and published by courtesy of this institution.
160 H. KLEIN ET AL.
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Type horizon and locality. ?Katberg Formation (Lower Tri-
assic) near Middelburg, Karoo basin, South Africa (see Seeley,
1904).
Stratigraphic and geographic distribution: Type horizon
and locality only.
Revised diagnosis: Small, pentadactyl imprints with
long and slender digits and large oval plantar surface. Dif-
ferent from (1) P. haarmuehlensis and P. polonicum by the
more extensive sole surface and straighter digits, from (2)
P. lockleyi ichnosp. nov. by the tapering shape of digits,
the much larger digit divarication (92vs. 37for digits I–
V), the shorter digit V, and the lack of a distinct “heel”
behind digit V, and from (3) P. nectouxi by tapering digits
that show a wider divarication (92vs. 55for digits I–V)
and the shorter digit V.
Description: Pes-manus couple on a small slab. Both are
subequal in size. Manus positioned anterior to the pes. The
digits in the pes and manus are moderately spread. In the pes,
digit IV is subequal to digit III in length and digit V is subeq-
ual to digit II in length. Digits I–V are long, straight and rela-
tively slender, with indistinctly preserved, tapering distal ends.
The plantar area is broad, its length comprises about 35 % of
the total imprint. The corresponding area in the manus is not
distinctly preserved. Also, the manus lacks the impression of
digit V. The trackway pattern is unknown. Seeley (1904) iden-
tified a tail drag mark, but we cannot verify that from the cast
of the specimen he described.
Discussion: Because the holotype of the type ichnospe-
cies Procolophonichnium nopcsai is apparently lost, and
the specimen consisted of only two imprints, the diagnosis
of the ichnogenus is based on relatively weak grounds.
However, the presence of a cast in the NHMUK collections
allowed us to study overall-shape and digit proportions,
and to make a comparison with other tracks described
under Procolophonichnium from different localities. The
general shape with digits increasing in length from I–IV,
with IV being subequal with III, V being relatively long
and subequal with II, and the distinct plantar/palmar sur-
face is observed in all other ichnospecies assigned to Pro-
colophonichnium as revised here.
Seeley (1904, p. 289) illustrated the manus and the pes of P.
nopcsai separately as “A” and “B” and not in the correct rela-
tive position, which is the former impressed anterior to the lat-
ter. This author published only drawings and no photographs,
but correctly stated that “the impression of the hind foot imme-
diately follows the fore foot” (Seeley, 1904, p. 288). Gener-
ally, Procolophonichnium shows a strict anterior, antero-
medial or rarely (in the new ichnospecies) antero-lateral posi-
tion of the manus relative to the pes. The drawings of Seeley
also show a pedal digit V that is extremely long and subequal
to digit IV. Based on our studies, however, pedal digit V is
much shorter and subequal to digit II. Also, the manual digit V
added in Seeley’s drawing could not be identified by us from
the cast (Fig. 2).
Procolophonichnium haarmuehlensis (Holst et al., 1970)
(Figs. 3A–B, D–G, J, 4–8)
1970 Rhynchosauroides haarm€
uhlensis: Holst et al., figs. 2,
7, pl. I, fig. 3, pl. II, figs. 4–6
1971a Procolophonichnium sp.: Haubold, pl. III
1977 Procolophonichnium sp.: Demathieu, p. 357, fig. 3,
pl. 2 (2)
1978 Procolophonichnium jageri: Demathieu and M€
uller,
p. 157–163, figs. 2–4
1983 Procolophonichnium winterswijkense: Demathieu and
Oosterink, p. 13, fig. 8, p. 41 figs. 33–35, p. 42, figs. 36–38,
p. 39, figs. 39–41, p. 44, figs. 42–44, p. 45, figs. 45–47, p.
46, figs. 48–50
1983 Phenacopus faberi: Demathieu and Oosterink, p. 15,
fig. 10, p. 47, figs. 51–52
1983 Phenacopus agilis: Demathieu and Oosterink, p. 16,
fig. 12, p. 47, fig. 53
1984 Rhynchosauroides haarm€
uhlensis: Haubold, p. 147,
fig. 98 (13)
1984 Procolophonipus jaegeri: Haubold, p. 149, fig. 100(5)
1986 Procolophonichnium winterswijkense: Baird, p. 133,
figs. 6A.
1988 Procolophonichnium winterswijkense:: Demathieu
and Oosterink, p. 13, fig. 6
1988 Phenacopus faberi: Demathieu and Oosterink, p. 14,
fig. 7A
1988 Phenacopus agilis: Demathieu and Oosterink, p. 14,
fig. 7B
2000 Procolophonichnium winterswijkense: Diedrich, p.
385, fig.6A–D
2002a Procolophonichnium haarmuehlensis: Diedrich, p.
82, fig. 3D, p. 85, fig. 4
2002b Procolophonichnium haarmuehlensis: Diedrich, p.
45, figs. 8–9
2002c Procolophonichnium haarmuehlensis: Diedrich, pl.
II, fig. 9, fig. 7,
2007 Procolophonichnium isp.: Valdiserri and Avanzini, p.
113, figs. 8–9
2007: Procolophonichnium sp.: Gand et al., p. 16, fig. 6U
2010 Procolophonichnium isp.: Klein and Lucas, p. 5–6, p.
11, fig. 9A–C
2011 Procolophonichnium isp.: Todesco and Bernardi, p.
207, pl. 1
2011 Procolophonichnium isp.: Klein et al., p. 227, 9A–D
Lectotype: Trackway with 5 pes-manus sets and tail trace
on a slab illustrated in Holst et al. (1970) on plate I, fig. 3,
uncatalogued.
Paralectotype: Partial trackway of two pes-manus sets and
tail trace on a slab illustrated in Holst et al. (1970) on plate II,
fig. 4, uncatalogued.
Additional material: Slabs with isolated tracks and track-
ways in the HOW collection figured in Demathieu and
REVISION OF THE ICHNOGENUS PROCOLOPHONICHNIUM 161
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FIG. 3. Overview with sketches of different Procolophonichnium specimens documented from the Lower – Upper Triassic. A–B, D–G, J are assigned here to P.
haarmuehlensis (A DP.“winterswijkense”; B D“Phenacopus faberi”; D DP. “isp.”; E DP. haarmuehlensis holotype; F D“Phenacopus agilis”; G DP. “sp.”;
JDP.“jageri”). C, P. ispp. H, P. polonicum holotype. I, P. nectouxi (“Circapalmichnus”nextouxi) holotype. A–B, E–F, Muschelkalk (Middle Triassic, Anisian-
Ladinian), Winterswijk (The Netherlands) and Germany. D, Timezgadiouine Formation (T4, Middle Triassic), Morocco. G, I, Middle Triassic (Anisian-Ladi-
nian), France. H, Wi
ory Formation (Lower Triassic, Olenekian) Poland. J, Buntsandstein (Lower-Middle Triassic), Germany. From Holst et al. (1970), Dema-
thieu (1977), Demathieu and M€
uller (1978), Demathieu and Oosterink (1983), Klein et al. (2011) (A–B, D–G, J), Baird (1986)(C), Klein and Nied
zwiedzki
(2012)(H), Gand (1977)(I). Note that E represents two pes imprints of different trackways, originally misinterpreted as a pes-manus set by Holst et al. (1970).
162 H. KLEIN ET AL.
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Oosterink (1983) under D22, D41a,b, D52a, b, D34, D36, D61,
D63a, b, D64, D67, D69, D75, D89, D113, D117, D121, D139
from the Middle Triassic of Winterswijk locality, The
Netherlands; slabs with isolated tracks and trackways in
the IPB listed and figured in Sumpf (2014) under
“Procolophonichnium R525” and “Procolophonichnium 1–7”
from the Middle Triassic of Winterswijk locality, The Nether-
lands; several isolated pes and manus imprints CDUE 223–
226, CDUE 228 from the Timezgadiouine Formation (T4),
Argana Basin, Morocco; several isolated pes and manus
imprints on slabs NMS 3.55b and BG.E 126-II.4 from the Mid-
dle Triassic of the Southern Alps, Italy.
Remarks: Holst et al. (1970) did not designate a holotype.
Therefore, we consider the trackway and a partial trackway
illustrated by these authors as syntypes, listing them here as
the lectotype and the paralectotype (ICZN rules, Art. 74.1).
Type horizon and locality: Muschelkalk (Middle Triassic,
Anisian), Haarm€
uhle, 3 km west of Alst€
atte, Westphalia,
Germany.
Stratigraphic and geographic distribution: Lower-Middle
Triassic (Olenekian-Anisian-Ladinian); Buntsandstein and
Muschelkalk of the Germanic Basin in Germany and The
Netherlands, Middle Triassic deposits of France and northern
Italy, Moenkopi Group (Wupatki and Shnabkaib formations)
of Arizona and Utah, Timezgadiouine Formation (T4) of the
Argana Basin (Morocco).
Revised diagnosis (emended after Holst et al., 1970, Dema-
thieu and Oosterink, 1983): Broad trackways, pace angulation
between 80and 100. Stride/trackway width ratio »1. Pes
imprints with slight outward rotation or parallel to midline.
Pes and manus pentadactyl, semiplantigrade to plantigrade.
Digit traces straight except in digits III and IV, which are
sometimes curved outward, sometimes also slightly inward.
Divarication angles between digits large (90–120for digits
I–V). Proximal margin of imprints straight or posteriorly con-
vex. Tapering claw marks that are often curved outward. Dis-
tinguished from: (1) P. nopcsai by the less extensive sole
surface and by the different shapes of the digits, which are
tapering distally in P. nopcsai but in P. haarmuehlensis show
mostly parallel margins with rounded pads and knobby distal
ends and often outward curved claws; (2) P. lockleyi isp. nov.
by the shorter digit V, the wider digit divarication, the lack of
an elongated heel behind digit V, the relatively shorter strides,
and by the outward orientation of pes imprints vs. the inward
rotation in P. lockleyi ichnosp. nov.; (3) P. nectouxi by the
wider digit divarication, the shorter digit V and the outward
rotation of the pes; and (4) P. polonicum by the shorter strides
and smaller pace angulation, and the parallel margins of digits
that are tapering in the latter.
Description: In well-preserved specimens, such as those
from the Muschelkalk (Anisian-Ladinian) of the Haarm€
uhle
(type) andWinterswijk localities, and from the Middle Triassic
of Italy (NMS 3.55b; BG. E 126-II.4; Fig. 7A–B), imprints
show long and slender to moderately robust digits, sometimes
with distinct, rounded phalangeal pads and scale impressions.
Often, the distal ends are transversely extended and knobby,
showing outward pointing claws due to the dynamics of the
foot. The number of preserved digits can vary between five
FIG. 4. Partial trackway (A) and trackway (B) of Procolophonichnium haarmuehlensis (HOW uncatalogued) from the Muschelkalk (Middle Triassic, Anisian-
Ladinian) of the Winterswijk locality (The Netherlands). Photos courtesy of Henk Oosterink, Winterswijk.
REVISION OF THE ICHNOGENUS PROCOLOPHONICHNIUM 163
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and three due to substrate conditions. Trackways show a rela-
tively broad pattern and often sinuous tail drag marks; the
manus is generally placed anterior to the pes. Imprints show
variable orientation from parallel to slightly outward relative
to the midline.
Discussion: Thus far, the material from the Muschelkalk
(Middle Triassic, Anisian-Ladinian) of Winterswijk has been
considered the most representative Procolophonichnium
sample in the global record (Demathieu and Oosterink (1983,
1988) (Figs. 3A, 4–6). Complete trackways and imprints, that
partly show details such as skin impressions, are well known.
Despite the fact that trackway parameters cannot be compared,
because the holotype from South Africa consists of isolated
tracks only, the plantigrade to semi-plantigrade imprints, and
the digit proportions with the relatively long digit V are diag-
nostic of Procolophonichnium. The inward or outward
FIG. 5. Trackways of Procolophonichnium haarmuehlensis from the Muschelkalk (Middle Triassic, Anisian-Ladinian) of the Winterswijk locality (The Nether-
lands) (A–B DIPB “Procolophonichnium 1”; C–D DIPB R525). A, C, photographs. B, D, sketches. All from Sumpf (unpubl. Master Thesis, 2014). Notice sinu-
ous tail drag marks.
164 H. KLEIN ET AL.
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curvature of digits often seen in the Winterswijk material, as
well as the “knobby” distal digit ends that are transversely
extended with laterally curved claw traces, are particularly dif-
ferent from the South African holotype. These peculiar fea-
tures justify a differentiation at the ichnospecies level and the
assignment to Procolophonichnium haarmuehlensis is con-
firmed here.
The ichnospecies was originally described under Rhyncho-
sauroides haarm€
uhlensis by Holst et al. (1970) based on mate-
rial from the Muschelkalk of Germany (Fig. 3E). Besides the
incorrect, non-Latinized spelling, the holotype illustrated by
Holst et al. (1970) is Procolophonichnium, not Rhynchosaur-
oides, the latter being a widespread lacertoid footprint with
digit IV longest and a short digit V (Diedrich, 2002a–c).
Because it is similar in all dimensions to the coeval Winter-
swijk material later published under P. winterwijkense (Dema-
thieu and Oosterink, 1983, 1988), the name introduced by
Holst et al. (1970) has priority, and all Winterswijk
Procolophonichnium must be referred to P. haarmuehlensis
with the correct spelling, as is done here.
Demathieu and Oosterink (1983, 1988) described another
new ichnogenus, Phenacopus, with the two ichnospecies Ph.
faberi and Ph. agilis from the Winterswijk locality (Figs. 3B,
F). However, these are so similar to P. haarmuehlensis in over-
all-shape that we consider them as synonyms. Purported dif-
ferences in shape mentioned by Demathieu and Oosterink
(1983, 1988) are extramorphological features or substrate-
related.
Diedrich (2000, 2002a–c) referred different imprints from
the Muschelkalk of Germany to Procolophonichnium winter-
swijkense and P. haarmuehlensis. The latter assignment is con-
firmed here for this material. Shape and digit proportions
identical to those of P. haarmuehlensis are seen in other track-
ways and isolated tracks from the Buntsandstein (Lower-Mid-
dle Triassic) of Germany (Haubold, 1971a; Demathieu and
M€
uller, 1978), the Middle Triassic of France (Demathieu,
1977), Italy (Valdiserri and Avanzini, 2007; Todesco and Ber-
nardi, 2011), the Timezgadiouine Formation of the Argana
Basin, Morocco (Klein et al., 2011; Voigt et al., 2011) and the
Lower Triassic of the Moenkopi Group of Arizona and Utah
(Klein and Lucas, 2010b; Thomson, 2014) (Figs. 3D, G, J, 7,
8). Therefore, they are all referred to here as P.
haarmuehlensis.
Haubold (1971a, pl. III) illustrated Procolophonichnium sp.
from the Lower Triassic Buntsandstein of Germany. No tail
traces are observed. These imprints are referred here to P.
haarmuehlensis based on their similar overall shape and digit
proportions.
FIG. 6. Isolated pes and manus imprints of Procolophonichnium haarmueh-
lensis from the Muschelkalk (Middle Triassic, Anisian-Ladinian) of the Win-
terswijk locality (The Netherlands)(B, D DIPB “Procolophonichnium 3”; C–
EDIPB “Procolophonichnium 5”; A DIPB “Procolophonichnium 7”. A–C
photographs, D–E sketches corresponding to B–C. All from Sumpf (unpubl.
Master Thesis, 2014). Notice skin impressions, claw marks and scratches.
FIG. 7. Pes and manus imprints of Procolophonichnium haarmuehlensis from
the Middle Triassic (Anisian) of the Southern Alps (northern Italy). A, speci-
men NMS 3.55b. B, specimen BG.E 126-II.4.
REVISION OF THE ICHNOGENUS PROCOLOPHONICHNIUM 165
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Demathieu and M€
uller (1978) described relatively broad
trackways with a moderately outward turned pes and with a
variable orientation of the smaller manus, which is positioned
anterior to the pes, but not overstepped by the latter (Fig. 3J).
They referred this material to the new ichnospecies P. jageri.
In this ichnospecies digits increase in length from I–IV with
digit III (in some cases digit IV) projecting farthest anteriorly
along the long axis of the imprint. They are moderately spread
and straight, slightly inward curved (I–IV) or straight or out-
ward curved (V), in some cases with tiny claws. The palmar/
plantar surface is subtriangular to broadly oval. The manus is
smaller, positioned anterior to the pes and has a relatively long
digit V that is outward curved. No tail drag mark is observed.
All these features are observed also in P. haarmuehlensis,so
we synonymize here P. jageri with P. haarmuehlensis, which
has priority according to ICZN rules (Art. 23).
Klein and Lucas (2010b) illustrated isolated imprints and
trackways on a slab with a trampled surface from the Wupatki
Formation (Lower Triassic) of the Moenkopi Group that they
assigned to Procolophonichnium isp. (Fig. 8). Our re-
FIG. 8. Trackways and numerous isolated pes and manus imprints of Procolophonichnium haarmuehlensis from the Wupatki Formation of the Moenkopi Group
(Olenekian) of Arizona, USA (UCMP uncatalogued). A–B, closeup view of slab. C, overview of slab with associated pes and manus imprints of Chirotherium
sickleri. From Klein and Lucas (2010b). Notice tail drag marks. Scale bars in cm.
166 H. KLEIN ET AL.
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evaluation of the material during a detailed analysis of the ich-
nogenus and the referred ichnospecies suggests an assignment
to P. haarmuehlensis based on the following characters: (1)
digit proportions with I–IV increasing in length, IV subequal
to III, and V as long as II; (2) digits I and II straight, III–V
sometimes slightly curved outward, V laterally spread; (3) pes
and manus parallel or turned slightly outward with respect to
the midline; and (4) presence of slightly undulating, thin tail
traces.
Demathieu (1977) illustrated a partial trackway with a tail
trace from the Middle Triassic of France that he assigned to
Procolophonichnium sp. (Fig. 3G). As in the examples above,
this specimen is identical in morphology to P. haarmuehlensis
and therefore referred here to that ichnospecies.
Procolophonichnium lockleyi ichnosp. nov. (Figs. 9–13)
Holotype: SKF 130 I (a and b), trackway with 9 pes-manus
sets on slab and counterslab (Figs. 10–12).
Paratype: SKF 130 II (a and b), trackway consisting of 5
pes-manus sets and 1 manus imprint (Fig. 13). The specimen
was positioned in close proximity to the holotype, showing
only a slight change of orientation relative to the latter, but
was separated by deep erosional features. It is likely that it
belongs to the same trackway.
Additional material: Indistinct isolated imprints on slabs
SKF 130 I and II.
Type horizon and locality: Coburger Sandstein, Hassberge
Formation (Carnian). Gleussner Sandstone quarry, about
1.5 km WSW of Neubrunn, Hassberge, NW N€
urnberg, north-
ern Bavaria, Germany (5001056.6900 N; 1040008.9200 E)
(Fig. 9).
Stratigraphic and geographic range: Type horizon and
locality only.
Diagnosis: Small, broad, plantigrade to semiplantigrade,
pentadactyl pes and manus imprints of similar shape. Pes up to
30 mm in length, manus slightly smaller. Digits with minor
differences of length, digit proportions III IV >II DV>I..
Extensive plantar/palmar surface with digit V being proxi-
mally elongated into a heel that reaches up to 50% of total
footprint length. Digits straight, with slight divarication only,
subparallel, digit V sometimes more strongly everted, all with
robust rounded pads, sometimes showing tiny tapering claws.
Trackways moderately broad, with relatively long strides
(150 mm–195 mm) and high pace angulation (91.3–110.3).
Manus positioned anterior to the pes, mostly with slight poste-
rior overstep by the latter. Pes and manus with distinct inward
rotation relative to midline by up to 18in the pes and up to
20in the manus. Distinguished from: (1) P. nopcsai and P.
haarmuehlensis by the smaller digit divarication, the propor-
tionately longer digit V and the presence of an elongated
“heel” behind digit V; (2) P. haarmuehlensis by the inward
rotation of the pes and the longer relative strides; (3) P. nec-
touxi by the longer relative strides and the continuous presence
of a digit V with an elongated “heel”; and (4) P. polonicum by
the straight digits with parallel margins, whereas in the latter
they are curved outward and tapering, by the inward rotaton of
the pes and by the presence of the “heel.”
FIG. 9. Geographic and stratigraphic position of Procolophonichnium lockleyi ichnosp. nov. A, map with the type locality in northern Bavaria, Germany. B,
Map with distribution of Triassic-Jurassic strata and the type locality in the Hassberge area northwest of Bamberg (N€
urnberg area). C, The type horizon in the
Coburger Sandstein of the Hassberge Formation (Upper Triassic, Carnian). Geological map in B from Karl and Haubold (1998), stratigraphy in C after Bachmann
and Kozur (2004).
REVISION OF THE ICHNOGENUS PROCOLOPHONICHNIUM 167
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Etymology: In honor of Martin G. Lockley, leading expert
and researcher on vertebrate ichnology, and a principal con-
tributor to the present understanding of fossil vertebrate foot-
prints and their formation.
Description: SKF 130 I is a slab 90 £50 cm in length and
width consisting of three pieces. It shows a trackway with nine
pes-manus sets preserved as natural molds and casts (a and b).
Originally, the trackway sequence on the illustrated slab SKF
130 I b (Fig. 12A) consisted of 10 sets, however, the second is
missing along a broken edge of the slab. This set is still present
on the less well-preserved counterslab (SKF 130 I a), which is
not illustrated here. It shows a constant walking pattern in
which the manus is overstepped by the pes at the proximal
margin. Only the first and the fourth sets (referring to the origi-
nal count of 10 sets) show a medial overstep of the manus by
the pes.
SKF 130 II (Fig. 13) is a slab 60 £35 cm in length and
width consisting of three pieces. It shows a trackway with five
pes-manus sets and one manus imprint preserved as a natural
mold and cast (a and b). It was found a short distance from
SKF 130 I and had the same general orientation. Because it
shows the same preservation and pattern as the latter it might
belong to the same trackway. However, this cannot be deter-
mined with certainty. Features of imprints and trackway pat-
tern are as listed in the diagnosis.
Discussion:ForProcolophonichnium, there are only a few
trackways and reliable data available from the global record,
whereas most specimens are isolated imprints or trampled sur-
faces where trackways are difficult to identify. Therefore, the
newly described Procolophonichnium trackways from the
Hassberge Formation can at best be compared with the mate-
rial from the Middle Triassic Muschelkalk of Winterswijk,
The Netherlands, which shows well-preserved trackways (P.
haarmuehlensis; Demathieu and Oosterink, 1983, 1988;
Sumpf, 2014). Other complete trackways come from the
Middle Triassic of France (P. nectouxi; Gand, 1977), the
FIG. 10. Slab SKF 130 Ib showing holotype trackway with pes-manus sets of
Procolophonichnium lockleyi ichnosp. nov. from the Coburger Sandstein,
Hassberge Formation (Upper Triassic, Carnian) of Germany, preserved as nat-
ural cast. Notice posterior overstep of manus by the pes and inward rotation of
imprints.
FIG. 11. Detail of holotype trackway SKF 130 Ib of Procolophonichnium
lockleyi ichnosp. nov. in Fig. 13 with different pes-manus sets. Notice digit V
being proportionately long and also the imprint posteriorly elongated into a
distinct heel.
168 H. KLEIN ET AL.
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Lower Triassic of Poland (P. polonicum; Klein and
Nied
zwiedzki, 2012), and the Lower–Middle Triassic of Ger-
many (P. haarmuehlensis; Demathieu and M€
uller, 1978). All
data from this material in the following comparison were taken
from these studies. For trackway parameters such as stride,
trackway width and pace angulation, only pes imprints were
considered, because manus imprints are positioned anteriorly
and close to the pes in nearly all examples, and therefore val-
ues of these, besides their rotation, differ only slightly. Param-
eters of P. lockleyi are listed in Table 1, and of other
Procolophonichnium ichnospecies in Table 2.
Features of SKF 130 useful in the comparison with other
Procolophonichnium are: (1) The relative stride/pes length
ratio is larger than in Procolophonichnium haarmuehlensis
from Winterswijk, which means 6:1 on average compared
with 5.5:1, 4.7:1, and 4.1 : 1 in the latter. In P. nectouxi, this
value reaches 4.50 and 5.50, and in P. haarmuehlensis from
the Buntsandstein of Germany it is 4.77; however, in P. polo-
nicum it is 7.8, which is very high. (2) The stride length/track-
way width ratio is much higher in the new ichnospecies (2.14–
2.79) vs. »1 in the Winterswijk material; in P. nectouxi this
value ranges 1.58–4.9 and in P. polonicum it is 3.2. (3) The
pace angulation of the pes ranges 91.3–110.3in the new ich-
nospecies; in the Winterswijk material the pace angulation
ranges 80–100, values of P. nectouxi range 90–95and in
Procolophonichnium from the Buntsandstein of Germany
these are 80–95. (4) Pes and manus are turned signifi-
cantlyinwardbyupto18
and 20, respectively; this fea-
ture is elsewhere only present in P. nectouxi,inwhichthe
rotation ranges from 8in the pes to 29in the manus. In
contrast, all other known Procolophonichnium trackways
show an outward rotation or parallel orientation of the pes
and manus relative to the midline. (5) The digit divarica-
tion angles in the new ichnotaxon are much smaller com-
pared with the Winterswijk material. The smallest value
for digits II–IV in the pes is 15. , whereas in the Winter-
swijk specimens it is 45; other angles are I–IV D21vs.
60and I–V D36vs. 90;inP. nectouxi values of divari-
cation angles II–IV range 12–15.
Obviously, the greatest similarity to the new ichnospecies
from the Upper Triassic of Germany is with P. nectouxi from
the Middle Triassic of France. This ichnospecies, as originally
assigned to the distinct ichnogenus Circapalmichnus by Gand
(1977), is referred here to Procolophonichnium in a new com-
bination (see below). It shows some features similar to the
new ichnospecies, such as the inward rotation of the pes and
manus relative to the midline, which is larger in the manus
compared with the pes. Furthermore, the stride/trackway width
ratio and the interdigital angle II–IV in P. nectouxi show a
variability that overlaps the values measured in the new
FIG. 12. Sketches of Procolophonichnium lockleyi ichnosp. nov. A, holotype trackway SKF 130 Ib with nine pes-manus sets. Notice that the second set at lower
right is missing. B., detail demarcated in A by quadrangle.
REVISION OF THE ICHNOGENUS PROCOLOPHONICHNIUM 169
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ichnospecies. Differences of P. lockleyi from P. nectouxi are
1) the higher stride/pes length ratio in the former, 2) the rela-
tive projection of digits with digit III projecting farthest ante-
rior in P. lockleyi, whereas in P. nectouxi digit IV shows the
longest projection, 3) the development of a distinct elongated
heel behind digit V in P. lockleyi not present in P. nectouxi.
The stride/pes length ratio in P. lockleyi is higher than in all
other Procolophonichnium, except P. polonicum, which shows
the highest value of all. The relatively high pace angulation of
P. lockleyi overlaps ranges of the Winterswijk material but dif-
fers from all other trackways in the higher values, except of P.
polonicum, which shows the largest angle. Some of these
parameters might also reflect a large variability due to differ-
ent gait, velocity and substrate conditions.
Unique features of the new ichnospecies are present in the
morphology of the pes and manus imprints: (1) digit propor-
tions with a very long digit V and subequal length of all digits;
(2) orientation of digit V parallel to other digits, whereas in all
other Procolophonichnium it is more strongly abducted; (3)
digits I–V straight, whereas in all other Procolophonichnium
(except P. nopcsai) digits I–IV are curved inward or outward,
and digit V is straight or curved outward; (4) posterior elonga-
tion of digit V into a massive heel that reaches 50% of the pes
and manus length, whereas in the similar P. nectouxi and all
other Procolophonichnium the posterior margin is straight,
convex or slightly concave; (5) the continuous presence of dis-
tinct digits I and V impressions in all tracks, whereas, espe-
cially the specimens of P. nectouxi and P haarmuehlensis
from Winterswijk often lack these, sometimes being tetradac-
tyl to tridactyl. The fact that these pecularities in the morphol-
ogy are continuous along the trackways suggests that they are
anatomically based and therefore justify the erection of a new
ichnospecies. Also, the imprints are not very deeply
impressed, so the continuous presence of a heel and all five
digits is obviously not related to substrate conditions.
Procolophonichnium nectouxi (Gand 1977) comb. nov.
(Fig. 3I)
1977 Circapalmichnus nectouxi: Gand, p. 20, 22, pl. D,
figs. 1–2, pls. 3–4
1984 Circapalmichnus nectouxi: Haubold, p. 149, fig. 100
(6)
2007 Circapalmichnus nectouxi: Gand et al., p. 16, fig. 6T,
pl. 2O
2007 Circapalmichnus sp.: Gand et al., p. 27, pl. 4, p. 28
Lectotype: Trackway B labelled “dalle Cg 54 piste B” in
Gand (1977, pl. D, fig. 1 [in part] and fig. 2 [photographs], pl.
3 right [sketch]) and deposited in the MHNA (Fig. 3I).
Paralectotype: Trackway A labelled “dalle Cg 54 piste A”
in Gand (1977, pl. D, fig. 1 [in part; photograph], pl. 3 left
[sketch]) and deposited in the MHNA.
FIG. 13. Sketch of paratype trackway SKF 130 IIb of Procolophonichnium
lockleyi ichnosp. nov. with five pes-manus sets and one manus imprint. Long
arrows indicate relative position and orientation of SKF 130 I (holotype) and
SKF 130 II (paratype).
170 H. KLEIN ET AL.
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Referred material: Trackways on slab labelled “dalle Cg
13” and figured in Gand (1977, pl. 4), deposited in the MHNA.
Remarks: Gand (1977) designates two trackways on a sin-
gle slab as the holotype. These are considered here as syntypes
based on ICZN rules (Art. 74.1). Here we designate the better
preserved trackway “Cg 54 piste B” as the lectotype, and “Cg
54 piste A” as the paralectotype of Procolophonichnium
nectouxi.
Type horizon and locality: Middle Triassic (Anisian-Ladi-
nian) of Culles-les Roches (S^
aone-et-Loire), France.
Stratigraphic and geographic distribution: Type horizon
and locality only.
Revised diagnosis (after Gand, 1977 emended): Trackways
of pentadactyl tetrapods with digitigrade to semiplantigrade
imprints. Manus slightly smaller than pes, wider than long,
rotated more strongly inward relative to midline compared
with the pes. If preserved, digit V in the manus is slightly lon-
ger, in the pes substantially longer than digit I and subequal to
digits II and III in length. Interdigital angle II–IV distinctly
larger in the manus (42on average) compared with the pes
(12on average). Distinguished from: (1) P. nopcsai by the
smaller digit divarication and the shape of digits that show par-
allel margins or distally expanded ends, whereas in the latter
they are tapering, by the proportionately longer digit V and by
the smaller digit divarication; (2) P. haarmuehlensis by the
smaller digit divarication, the proportionately longer digit V
and the rotation of imprints toward the midline; (3) P. lockleyi
by the frequent lack of digit V in the pes imprint, the absence
of a “heel” behind digit V, the longer projection of digit IV rel-
ative to digit III, and by the shorter relative strides; and (4) P.
polonicum by the straighter digits, the shape of digits with par-
allel margins and sometimes expanding distal ends, whereas in
the latter they are tapering, the proportionately longer digit V,
the shorter stride, smaller pace angulation and by the inward
rotation of the imprints.
Description: Based on Gand (1977). Trackways are rela-
tively broad with occasional, posterolateral overstep of the
manus by the pes. After the measurements of Gand (1977), the
stride length/pes length ratio is 4.5 and 5.5, and the pace angu-
lation of the pes slightly larger than that of the manus, ranging
90–96. Digits IV and III in the pes and manus are subequal
in length, and, in the pes, digit IV is mostly projecting farthest
forward along the long axis of the imprint. Digits I–IV are
curved inward, and digit V is straight or curved outward.
The interdigital angle II–IV is distinctly larger in the manus
(42on average) compared with the pes (12on average)
(Gand, 1977). Digits show well-developed, rounded phalan-
geal pads and, in some specimens, tiny claws. The phalangeal
pad formula is 2-3-3-3-?3. Digits are more deeply impressed
at their distal and proximal portions, whereas the middle part
is less deep. The plantar/ palmar impression is broad, often
with a straight posterior margin.
Discussion: Several trackways from the Middle Triassic of
Culles-les Roches (S^
aone-et-Loire), France (Fig. 3I) were
described under the new ichnogenus and ichnospecies Circa-
palmichnus nectouxi and Circapalmichnus isp., respectively,
by Gand (1977). They are referred here to the new combina-
tion Procolophonichnium nectouxi based on the overall shape,
which is identical to Procolophonichnium. Also, purported dif-
ferences of C. isp. from C. nectouxi (Gand, 1977) are consid-
ered here as extramorphological features rather than
anatomically based. P. nectouxi is different from P. haarmueh-
lensis, especially in the inward rotation of the pes and the
manus, whereas in the latter, the imprints are oriented parallel
to the midline or are slightly outward rotated. Furthermore,
digit divarication angles are smaller in P. nectouxi compared
with P. haarmuehlensis, in which the digits are more spread.
In these features, P. nectouxi resembles the new ichnospecies
P. lockleyi described here. The morphology of P. nectouxi
(“Circapalmichnus”nectouxi) is within the range of variation
of Procolophonichnium, and trackway parameters given by
Gand (1977) match those of typical Procolophonichnium, for
example the sample from Winterswijk. However, the observed
peculiar characters justify the retention of the ichnospecies P.
nectouxi.
Procolophonichnium polonicum (Ptaszy
nski 1990)
(Fig. 3H)
1990 Rhynchosauroides polonicus Ptaszy
nski sp. n.: Fugle-
wicz et al., figs. 9 (5–8), 10 (1); pls. 8 (1–4), 9 (1–2), 10 (1).
1999 Rhynchosauroides: Lockley and Meyer, fig. 3.14
(bottom),
2000 Procolophonichnium polonicum: Ptaszy
nski, fig-
s. 18G–H, 20A–G.
2012 Procolophonichnium: Klein and Nied
zwiezki, p. 51,
fig. 50C, F, I–J
Holotype: KR 11: 4.1,3 (plaster cast ZPAL R.7/40: 1; Klein
and Nied
zwiedzki, 2012, fig. 50C, J) (Fig. 3H).
Referred material: ZPAL R.7/22, (Klein and
Nied
zwiedzki, 2012, fig. 50I), trackway; ZPAL R.7/23, (Klein
and Nied
zwiedzki, 2012, fig. 50F), several pes and/or manus
imprints; ZPAL R.7/26, isolated imprint; ZPAL R.7/27 1, 2,
isolated imprints and tail trace; ZPAL R. 7/46, left set.
Type horizon and locality: Outcrops at right bank of the

Swi
slina River, Wi
ory, Holy Cross Mountains, Poland. Wi
ory
Formation (Lower Triassic, late Olenekian).
Stratigraphic and geographic distribution: Type horizon
and locality only.
Revised diagnosis (emended after Ptaszy
nski in Fugle-
wicz et al., 1990;Klein and Nied
zwiedzki,2012): Large Pro-
colophonichnium with a relatively short digit group I–IV and
digits III and IV nearly equally long. Tip of digit V at the level
of half the length of digit IV. Distinguished from: (1) P. nopc-
sai by the less extensive sole and outward curvature of digits;
(2) P. haarmuehlensis, P. lockleyi and P. nectouxi by the
REVISION OF THE ICHNOGENUS PROCOLOPHONICHNIUM 171
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tapering shape of digits, the longer relative stride and larger
pace angulation; and (3) P. lockleyi by the proportionately
shorter digit V, the outward rotation of imprints vs. the inward
rotation in the latter.
Description: These are pentadactyl imprints and trackways
of a quadruped, in which the pes is larger than the manus. The
digits are slender to moderately broad and straight or curved
outward distally and increasing in length from I to IV. Digit I
is extremely short, and digit V is positioned posterolateral to
the digit group I–IV, being relatively long compared with the
conditions in Rhynchosauroides. Small triangular claws are
visible on digits I–IV. The manus is positioned anterior or
antero-medial to the pes. It is often incompletely impressed,
with digits I–IV or II–IV being preserved. Imprints show an
outward rotation relative to the midline.
Discussion: Fuglewicz et al. (1990) and Ptaszy
nski (2000)
established a new ichnospecies Procolophonichnium poloni-
cum, originally described as Rhynchosauroides polonicus
(Ptaszy
nski, 1990), from the Wi
ory Formation (Lower Trias-
sic, Olenekian) of the Holy Cross Mountains, Poland
(Fig. 3H). The ichnospecies was confirmed by Klein and
Nied
zwiedzki (2012), in spite of the poorly-preserved mate-
rial. Because of some pecularities such as the relatively long
stride and large pace angulation angle compared with other
known Procolophonichnium (see above), we tentatively con-
sider the ichnospecies as valid, emphasizing that more and bet-
ter-preserved material may possibly change this view.
Procolophonichnium ispp. (Figs. 3C, 14)
1986 Procolophonichnium sp.: Baird, p. 133, figs. 5–6B.
2008 Dicynodontipus protherioides: Silva et al., p. 102,
fig. 3, p. 103, fig. 4, p. 104, fig. 5, p. 105, fig. 6
Referred material: New Jersey: YPM (PU) 21754, manus
and pes tracks in convex hyporelief (Baird, 1986, figs. 5, 6B);
Brazil: MCN.PIC.001–004, MCN.PIC.016–019, (Figs. 3C,
14).
Stratigraphic and geographic distribution: Passaic Forma-
tion of Newark Supergroup (Upper Triassic, Norian) of New
Jersey, USA; Santa Maria Formation (Upper Triassic, Car-
nian) of Brazil.
Description: The imprints from the Upper Triassic of New
Jersey illustrated by Baird (1986) obviously represent a set
with a larger pes and a slightly smaller manus anterior to the
pes, about 2.5–3 cm in length (Figs. 3C, 14). Both are penta-
dactyl. Digits are robust and have broad, “knobby” distal ends
and claw marks that in digit IV and V of the pes point outward.
They are moderately spread with digit V in the pes being more
strongly curved outward. Imprints show a larger plantar and
palmar surface, which is posteriorly convex at the proximal
margin, its length reaching about 50% of the total footprint
length. Digit proportions with increasing lengths of digits I–
IV, digit III being subequal to IV, and digit V being subequal
in length to II.
FIG. 14. Pes and manus imprints of Procolophonichnium ispp. YPM (PU) 21754 in convex hyporelief from the Passaic Formation (Upper Triassic, Norian) of
New Jersey, USA. Original in Baird (1986, fig. 5). A, photograph. B, sketch. Photograph by Sebastian Dalman, sketch from Baird (1986).
172 H. KLEIN ET AL.
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Small (»1 cm length) footprints from the Santa Maria For-
mation (Upper Triassic) of Brazil described by Silva et al.
(2008) as a new ichnospecies of Dicynodontipus and re-
assigned here to Procolophonichnium (see below) consist of
relatively broad trackways with pentadactyl pes and manus
imprints. They are mostly poorly preserved. More distinct
imprints show digit proportions with I–IV increasing in length,
digit IV being subequal in length with digit III or slightly
shorter, and digit V being subequal with digit II or digit I.
Imprints show a more or less extensive sole surface reaching
»50% of footprint length, and with a concave proximal mar-
gin. The manus is only slightly smaller than the pes and posi-
tioned anterior to the latter. Trackways are characterized by
short strides, low pace angulation and parallel orientation of
imprints relative to midline. A slightly sinuous tail trace is
present.
Discussion: Procolophonichnium from the Late Triassic is
scarcely documented from the global record. Besides the
tracks from New Jersey and Brazil, the new ichnospecies P.
lockleyi described above in detail is the third record of the ich-
nogenus from this time interval. The first was described by
Baird (1986) from the Passaic Formation of the Newark Super-
group of New Jersey as Procolophonichnium sp. and com-
pared with the Winterswijk form (Figs. 3C, 14). Baird’s
material is an isolated pes-manus set that shows the character-
istic overall shape and digit proportions with relatively long
digit V and extended plantar surface.
Typically, the manus is rotated more strongly inward rela-
tive to the pes and along digit III. The stratigraphic position
suggests a comparison with the new ichnospecies P. lockleyi.
However, the material from the Newark Supergroup is too
fragmentary. Digit proportions with the relatively long digit V
are slightly similar to P. lockleyi. Distinct is the sole of the
track that shows a straight or convex margin, whereas P. lock-
leyi has a distinctly elongated heel behind digit V. Neverthe-
less, we are aware of the fact that the Newark specimen is an
isolated and incompletely preserved record, and new discover-
ies with well preserved trackways could possibly result in a re-
evaluation of the Newark material.
Silva et al. (2008) introduced the new ichnospecies Dicyno-
dontipus protherioides from the Santa Maria Formation
(Upper Triassic, Carnian) of Brazil. Dicynodontipus is a the-
rapsid trackway that was introduced by R€
uhle v. Lilienstern
(1944) based on material from the Lower-Middle Triassic
Buntsandstein of Germany. However, the morphology of the
imprints, as well as the tail trace, illustrated by Silva et al.
(2008) suggest an affinity with Procolophonichnium. Tail
traces of Dicynodontipus are unknown in all other occur-
rences, especially from the Lower-Middle Triassic of the Ger-
manic Basin of Europe. Furthermore, the material from South
America shows relatively long digits with lengths that increase
from I to III in the pes and from I–IV in the manus. This is in
contrast with Dicynodontipus, which characteristically shows
short digits that are subequal in length and often separated
from the large sole. Also, the concave proximal margin of the
imprints can sometimes be observed in Procolophonichnium,
whereas in Dicynodontipus it is always convex. Therefore, we
re-assign these tracks from Brazil to Procolophonichnium
ispp., noting that the preservation allows no concrete ichnospe-
cific determination.
Ichnotaxa with dubious affinities to Procolophonichnium
Different ichnogenera and ichnospecies have been
described in the literature and variously synonymized with
Procolophonichnium, especially by Haubold (see 1971a, b,
1984).
Procolophonipus triadicus and acutus were described from
the Buntsandstein (Olenekian-Anisian) of Germany by R€
uhle
v. Lilienstern (1939). Procolophonipus muelleri was estab-
lished as a new ichnospecies based on material from the same
unit by Haubold (1971a). All three ichnotaxa are tetradactyl.
This may be a preservational feature or anatomically based.
However it is constantly present in all imprints of these track-
ways. Haubold (1971b) re-assigned them to Procolophonich-
nium and “Diverse Trias indet.,” respectively, together with
Procolophonipus italicus originally described by v. Huene
(1941) from the Upper Triassic of Italy (Haubold, 1971b, p.
31, p. 100, fig. 62). Haubold (1984) again listed all these ich-
notaxa under Procolophonipus (Haubold, 1984, p. 149,
fig. 100 [3–4]). Procolophonipus vonhuenei described by
Bock from the Newark Supergroup (Upper Triassic) of New
Jersey (Bock 1952, pl. 46, fig. 3) encompasses groups of (up
to four) elongated marks or scratches that can be interpreted as
swim traces. They are otherwise indeterminate. Other similar,
but tridactyl imprints and broad trackways were described
from the Buntsandstein of Germany under Ruecklinichnium
tridactylum by Kuhn (1958) (see also Haubold, 1971a, p. 513,
fig. 34a).
Silva et al. (2012, p. 36, fig. 5A, p. 37, fig. 6B) illustrated
isolated imprints from the Upper Permian Rio do Rasto For-
mation of Brazil that they assign to Procolophonichnium isp.
Based on overall-shape and digit proportions, these show
some similarities to the ichnogenus by digits increasing in
length from I–IV, a relatively long digit V, and a distinct plan-
tar/palmar portion. However, the material is too poorly pre-
served for a definite determination. Therefore we consider this
assignment by Silva et al. (2012) as dubious. This is also true
for the Permian record from Germany (M€
uller, 1970) (see
above).
Footprints from the Lower Triassic of the Sydney Basin,
Australia, were described and illustrated by Retallack (1996),
who assigned them to Dicynodontipus and the new ichnospe-
cies D. bellambiensis. They show some similarities to the Pro-
colophonichnium nopcsai holotype from South Africa and are
likely of the same geological age. Congruent features concern
the proportions, shape and orientation of digits and the exten-
sive sole surface. However, the Australian material is too
REVISION OF THE ICHNOGENUS PROCOLOPHONICHNIUM 173
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poorly preserved for a concrete assignment. Nevertheless, it is
different from Dicynodontipus, which is characterized by
relatively short and subequal digits that are often distinctly
separated from the sole (R. v. Lilienstern, 1944; Haubold,
1971a, b).
DISCUSSION OF TRACKMAKERS
In earlier studies, Procolophonichnium tracks and trackways
were attributed to procolophonids (Seeley, 1904, 1905; Nopcsa,
1923; Haubold, 1971a, b; Baird, 1986), “Cotylosauria” and tur-
tles (Kuhn, 1958, 1963), and/or captorhinids (Haubold, 1984).
More recent interpretations consider “primitive amniotes”
(Klein and Lucas, 2010b) or procolophonids and therapsids
(Klein et al., 2011; Voigt et al., 2011) as trackmaker candi-
dates. More complete pes skeletons of these groups and their
reconstructions are illustrated in Figure 15 for comparison. The
digit proportions of Procolophonichnium, a relatively short
digit IV compared with Rhynchosauroides and Dromopus,and
a relatively long digit V compared with Rhynchosauroides,
Dromopus and Varanopus, are seen in procolophonid, captorhi-
nid and some therapsid pes skeletons. The manus and pes skel-
etons of the procolophonid Procolophon from the Early
Triassic (Figs. 15A, E) show a slightly longer digit IV in the
manus (relative to digit III), whereas in the pes it is subequal in
length to digit III. Also, digit V and digit II are subequal in
length. In captorhinids, the manus and pes skeletons of the
Early Permian taxa Labidosaurus and Captorhinus,respec-
tively, both have a relatively long digit IV (Figs. 15B, F). Only
in the pes, digit V is subequal in length to digit II, but in the
manus, it is proportionately shorter.
The proportions of the therapsid pes are quite different.
Thrinaxodon, a cynodont from the Middle Triassic (Figs. 15C,
G), has a pes in which digit IV is longest, while digits V and II
are subequal in length. However, in the manus, digits IV and
III are subequal in length, digit II is only slightly shorter, and
digit V is shorter than II. In the manus and pes of the Late
Permian therocephalians Aelurosuchus and Whaitsia (Fig-
s. 15D, H), respectively, digit III is longest in the manus, IV
slightly shorter, II is shorter than IV and subequal to V; digits
III, IV and V are subequal in length in the pes, digit II is
shorter, followed by I.
Based on digit proportions, the best match between skeletal
material and Procolophonichnium is seen between: (1) thero-
cephalian (therapsid) and Procolophonichnium lockleyi; also,
P. nectouxi is more or less similar in the relative digit lengths,
and (2) procolophonid Procolophon and Procolophonichnium
haarmuehlensis, P. polonicum and P. nopcsai. Captorhinids
and the therapsid Thrinaxodon have a minor overlap of digit
proportions. Captorhinids show a distinctly longest digit IV in
both the manus and the pes, and Thrinaxodon in the pes. For
P. lockleyi, the orientation of the long digit V being subparallel
FIG. 15. Autopodia of different tetrapods from the Permian and Triassic for comparison. A–D manus skeletons. E–H pes skeletons. A, E, Procolophon (Pararep-
tilia), Early Triassic. B, Labidosaurus (captorhinomorph), Early Permian. F, Captorhinus (captorhinomorph), Early Permian. C, G, Thrinaxodon (cynodont the-
rapsid), Middle Triassic. D, Aelurosuchus (therocephalian therapsid), Permian. H, Whaitsia (therocephalian therapsid), Late Permian. Notice reduction in the
number of phalanges in therapsids. All drawings from Romer (1956). Abbreviations are: a Dastragalus, c Dcentrale, ca Dcalcaneous, f Dfibulare, F Dfibula,
iDintermedium, p Dpisiform, pc Dproximal centrale, r Dradiale, R Dradius, t Dtibiale, T Dtibia, U Dulna.
174 H. KLEIN ET AL.
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to digit IV matches the conditions in the therapsid autopodia.
In contrast, a shorter and laterally spread digit V is widely dis-
tributed in other Procolophonichnium ichnospecies congruent
with pes and manus skeletons of procolophonids and
captorhinids.
Of additional importance to comparison is the phalangeal
formula, which can be derived from preserved pads in the foot-
prints, especially in the Procolophonichnium haarmuehlensis
material of central Europe, but also in the new ichnotaxon P.
lockleyi. After Sumpf (2014), the derived phalangeal formula
in the pes of P. haarmuehlensis is 2-3-3-3-x, corresponding to
the general condition in Therapsida (Figs. 15G, H), and indi-
cating a therapsid as the likely maker of P. haarmuehlensis
tracks, instead of a procolophonid, although in the therapsid
Thrinaxodon strongly reduced additional phalanges still seem
to be present (Fig. 15G).
A similar phalangeal formula is derived from the preserved
pads in the new ichnospecies Procolophonichnium lockleyi,
with 2-3-3-3-?3 for the pes and the manus. Also, based on the
illustrations in Gand (1977), P. nectouxi has a similar count. In
contrast, the phalangeal formula in the autopodia of procolo-
phonids and captorhinids is 2-3-4-5-4 for the pes and 3-3-4-5-3
and 2-3-4-5-3 for the manus, respectively (Figs. 15A, B, E, F).
Therefore, following Sumpf (2014), we consider therapsids as
the most likely trackmakers of at least some Procolophonich-
nium ichnospecies such as P. haarmuehlensis, P. nectouxi and
P. lockleyi. Another feature observed in the new ichnospecies
P. lockleyi also supports this: Impressions of the distal and
proximal phalangeal pads are more deeply impressed com-
pared to the middle portion of the digits. This is possibly due
to the so-called digital arcade (K€
ummell and Frey, 2012),
which is formed by digits bent dorsally between their distal
and proximal ends, a feature characteristic of Synapsida. Tak-
ing into account that the type ichnospecies P. nopcsai lacks
preserved phalangeal pad traces, in this case we cannot
completely exclude other tetrapod groups as the trackmaker.
CONCLUSIONS
The ichnogenus Procolophonichnium Nopcsa 1923 is a
widely distributed tetrapod footprint from ?Permian/Lower
through Upper Triassic deposits of South Africa, North Africa,
Europe and North America. A Permian age of the holotype
from South Africa, as erroneously proclaimed by some work-
ers, is less probable, and is obviously based on a misinterpreta-
tion of the statements of Seeley (1904), Nopcsa (1923) and
others. The holotype of P. nopcsai is most likely of Early Tri-
assic age.
Five ichnospecies—Procolophonichnium nopcsai (type
ichnospecies), P. haarmuehlensis, P. nectouxi, P. polonicum,
and P. lockleyi—are considered as valid. P. nectouxi is a new
combination, P. lockleyi a new ichnospecies.
The new ichnospecies Procolophonichnium lockleyi is
introduced from the German Upper Triassic based on well-
preserved long trackways and imprints with features not
observed in any other Procolophonichnium ichnospecies.
Trackmakers of Procolophonichnium were therapsids
rather than procolophonids or captorhinids considering digit
configuration and phalangeal formula derived from preserved
pads.
ACKNOWLEDGMENTS
The authors thank Sandra Chapman and the Natural History
Museum, London for making photos of the cast of the Proco-
lophonichnium nopcsai holotype and for permission to publish
these. Franziska Sumpf, and Martin Sander, Bonn helped with
figures from a master thesis that could be used here. Sebastian
Dalman provided photographs of the YPM specimen of Proco-
lophonichnium. Evelyn Kustatscher and the “Naturmuseum
Bozen” is thanked for access to Procolophonichnium speci-
mens from the Middle Triassic of the Southern Alps. We are
grateful for the critical comments and constructive review by
Ricardo Melchor, Santa Rosa, Argentina, that helped to
improve the manuscript.
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