THE PRINCIPAL MORPHS OF LOWER PERMIAN TETRAPOD
TRACKS
HARTMUT HAUBOLD
SEBASTIAN VOIGT
Institute of Geological Sciences and Geiseltalmuseum, Martin-Luther-University
Halle-Wittenberg, Domstrasse 5, 06108 Halle, Germany
<haubold@geologie.uni-halle.de> <voigt@geologie.uni-halle.de>
1
The principal morphs of Lower Permian tetrapod tracks
Introduction
Recent knowledge of the Lower Permian terrestrial vertebrate fauna is based on
osteological and ichnological data. Skeletal remains as well as tetrapod tracks of upper
Paleozoic red beds have been known since the 19th century (Case 1907; Cope 1878;
Cotta 1848; Credner 1881; Geinitz 1861; Pabst 1908; Williston 1911). The
investigation of these originated in North America and Europe due to the prevalence
and early recovery of corresponding fossiliferous sediments. However, differences in
ancient facies and present habitat in connection with geographic factors have resulted
in isolated and partly divergent tendencies of study on each side of the Atlantic. Fossil
skeletons and bones have been especially known from the red beds of the United States
(Dunkard Basin, Western Interior, Colorado Plateau). Although described by Gilmore
(1926-28) and Moodie (1929-30), for example, tracks could not gain attention in the
same way. In contrast to that, for a long time the record of Permocarboniferous fossils
of some central and western European basins was dominated by tracks and traces.
Discoveries of terrestrial tetrapods remained rarely and fragmentary. In the past two
decades, accentuated from about 1990 the bias could be contemporaneously attenuated
by extensive trackway discoveries in New Mexico (Haubold et al. 1995; Hunt et al.
1993, 1995; Hunt and Lucas 1998; Lucas et al. 2001), and by the recovery of important
skeletal remains in Germany (Berman and Martens 1993, Berman et al. 1998-2001;
Boy and Martens 1991). The completion and intensified exchange of data is of
considerable interest with respect to phylogeny, biostratigraphy, and taxonomy. Based
on new facts, now it is possible to determine the anatomically controlled taxa of the
2
Lower Permian tetrapod ichnofauna. Additionally, the reflection of evolutionary trends
of the locomotory apparatus by tracks can be used to establish the relationship to
potential trackmakers.
Lower Permian Terrestrial Tetrapods And Their Locomotion – Evidence From
Bone Record
The terrestrial vertebrate fauna of Lower Permian Red Beds is dominated by
Temnospondyli, Seymouriamorpha, Diadectomorpha, ‚Pelycosauria’,
Captorhinomorpha, and Araeoscelidia (Berman et al. 1997; Lucas 1998; fig. 1). Among
these, the amniotes indicate a first significant divergence of locomotory abilities (Olsen
1955; Romer 1935; Sumida 1997). This is due to the amniote radiation in the
Carboniferous, from which a still incompletely known diversity of terrestrial tetrapods
arose in Early Permian time.
<FIGURE 1 NEAR HERE>
Character transformations of the appendicular skeleton which will be expressed
in tracks first of all concern the structure of tarsus and carpus as well as the number,
form and orientation of phalanges. These anatomical features determine the lengthwidth ratio of the autopodium as well as the relative length and the arrangement of
digits. The tarsus of Lower Permian amniotes is characterized by an increasing
consolidation accompanied by a stronger ossification of tarsal and carpal elements. Due
to the fusion of tibiale, intermedium and proximal centrale, including the fibulare, two
modified protarsal elements result, astragalus and calcaneum. They are considered as
amniote synapomorphies (Gauthier et al. 1988; Rieppel 1993; Sumida 1997). Changes
of posture, gait and maneuverability of the body are additionally based on
3
modifications of the stylo- and zeugopodium. For example, there is an evolutionary
trend toward a flexible acetabulo-femoral articulation and gracile long bones. The
locomotorally quite progressive araeoscelids indicate a prolongation of epi- and
propodial extremity segments and the development of a rather sigmoid shaped femur
(Sumida 1997). These are adaptations for compensation of the increasingly dynamic
load of the skeleton, which is accompanying the trend to enhanced agility. The
diadectomorhs show modifications in the knee and ankle joint, that are functionally
comparable to analogous structures of recent Lacertilia, and which allow a parasagittal
orientation of meta- and autopodium (Berman and Henrici 2003). These changes are
considered as an adaptation of tetrapods of sprawling type posture and gait to maximize
the efficiency of parasagittal thrust caused by the hind limbs. Autopods that are setting
down near the body and in the direction of motion induce a higher pace angulation, an
enhanced stride, and generally greater maneuverability of the body during locomotion.
In contrast, the semiterrestrial to terrestrial representatives of amphibian-like or
anamniote Temnospondyli show a quite conservative formation of the locomotory
apparatus. The structure of their autopodia seems to have been optimised already
during an early stage of the temnospondyl radiation. Apparently after that, relative
length, position, and orientation of the digits have hardly changed during the late
Paleozoic time span. Presumably due to temporally extended early ontogenetic phases
of restriction to water bodies, the carpus and tarsus of these animals maintained a more
flexible bone mosaic in comparison to the corresponding structures of amniotes.
Early Permian Tetrapod Ichnofauna
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As showed above, divergent evolutionary tendencies in the anatomy of extremities and
of locomotory abilities are documented in body fossils of Lower Permian terrestrial
vertebrates. These developments are reflected by the trackway record. The range of
pentadactyl amniote track morphologies is extended from that of the Upper
Carboniferous. In addition, new types of trackway pattern appear within that group. In
contrast to that, tetradactyl ichnotaxa, which are referred to temnospondyls, pass the
system boundary without variation.
There are six principal morphs of Lower Permian tetrapod tracks, which can be
differentiated by means of imprint morphology (fig. 2). As a result of revision and
proceeding collections at numerous sites in Europe and North America these six
morphs are represented by the following ichnogenera: (1) Batrachichnus Woodworth,
1900, Limnopus Marsh, 1894 (2) Amphisauropus Haubold, 1970, (3) Ichniotherium
Pohlig, 1892, (4) Dimetropus Romer & Price, 1940, (5) Hyloidichnus Gilmore, 1927,
Varanopus Moodie, 1929, Gilmoreichnus Haubold, 1970, Erpetopus Moodie, 1929, (6)
Dromopus Marsh, 1894, Tambachichnium Müller, 1954. The six principal morphs will
be subsequently explained and characterized.
(1) Batrachichnus - Limnopus
Batrachichnus and Limnopus are interpreted as tracks of terrestrially adapted
temnospondyls of different body size (Haubold 1970, 1971, 1996, 2000; Hunt et al.
1995). Their tracks differ from all other mentioned ichnogenera by the tetradactyl
manus imprint. Batrachichnus has been described by Woodworth (1900) from the
Conemaugh Series of the Upper Carboniferous of Massachusetts. The genus is one of
the predominant taxa of the Permocarboniferous ichnofauna. There are only two formal
5
and geographically distinguishable Permian ichnospecies: Batrachichnus delicatulus
(Lull, 1918), e. g., from the Wolfcampian Hueco Group of New Mexico (Haubold et al.
1995), and Batrachichnus salamandroides (Geinitz, 1861), which is widely distributed
in the central European Rotliegend (Haubold 1996). Both are probably synonyms of
Batrachichnus plainvillensis Woodworth, 1900.
Marsh (1894) has established the taxon Limnopus for tracks from the Howard
Limestone Formation, Virgil Series, of the Upper Pennsylvanian of Osage, Osage
County, Kansas (Baird 1952). Limnopus is attributed to larger imprints than
Batrachichnus. Major differences concern the width of pace, the length width ratio of
the imprints, and the digit lengths of the manus imprint, in particular.
(2) Amphisauropus and (3) Ichniotherium
The ichnotaxa Amphisauropus and Ichniotherium are interpreted as Seymouriamorpha
and Diadectomorpha, respectively. The tracks of both genera show a considerable
similarity in the arrangement of the digits. However, the pes imprints differ in respect
to the length of digit V and the shape of sole. In Ichniotherium the last is characterized
by an ovate, transversally extended, and often sharply outlined heel pad (Voigt &
Haubold 2000). In contrast, the sole and digits form nearly a structural unit in the
Amphisauropus imprint.
<FIGURE 2 NEAR HERE>
The Diadectidae record a step wise consolidation of the proximal tarsus,
resulting in the formation of the astragalus and calcaneum (Berman and Henrici 2003).
Whereas basal family members are still showing sutures of the tripartite origin of these
tarsal bones, sutures are lost in more advanced relatives. Finally, the most progressive
6
of the known Diadectes species indicate a fusion of the astragalus and calcaneum to
create an astragalo-calcaneum complex. It is a prominent, mediolaterally extended
element of the proximal tarsus, which convincingly fits the oval heel imprint of
Ichniotherium cottae (Pohlig, 1885). In Limnoscelidae and Tseajaiidae, rather primitive
taxa of Diadectomorpha, the astragalus and calcaneum are still absent (Berman and
Henrici 2003). Consequently, in these forms a similar structure of the sole imprint can
not be expected. That’s why apart from seymouriamorphs, basal diadectomorphs have
to be taken into account as potential trackmakers of Amphisauropus.
The locus typicus of Amphisauropus is situated in the Goldlauter Formation of
the Thuringian Forest, Germany (Haubold 1970). Numerous specimens of the
ichnogenus come from Europe (Haubold 1996; Haubold and Stapf 1998), and, recently,
in increasing extent also from North America (Lucas et al. 2001). The ichnotaxon is
limited to Permian sediments as yet. Ichniotherium is an ubiquitous element of the
Thuringian Forest Rotliegend ichnofauna. The best record comes qualitatively as well
as quantitatively from the type locality of the Tambach Formation (Pabst 1908;
Haubold 1998; Voigt 2001, 2002). With the exception of the holotype of Ichniotherium
willsi Haubold and Sarjeant, 1973 from the Upper Carboniferous of England, all other
Ichniotherium specimens from outside of the Thuringian Rotliegend are only preserved
as single imprints (Gand 1987; Haubold 1971, 1973; Haubold and Stapf 1998; Hunt et
al. 1995). Due to incompleteness of the record and preservation their identification still
remains uncertain.
7
(4) Dimetropus
Dimetropus imprints, which are interpreted as tracks of carnivorous Eupelycosauria,
are characterized by an extended sole, stressed and circular shaped metatarsal
phalangeal pads, slender digits, and tapered claws (Haubold 1971-2000; Haubold et al.
1995). The characteristics of imprint morphology will rapidly be lost in the common
undertrack preservation by decreasing clarity of outlines. In that case, the imprints
indicate a proximodistally extended, narrow sole and short, and slender digits.
Dimetropus is a widely distributed ichnotaxon of Permocarboniferous red beds
(Haubold 1996; Haubold et al. 1995). Three ichnospecies are differentiated:
Dimetropus leisnerianus (Geinitz, 1863), D. bereae (Tilton, 1931), and D. nicolasi
Gand and Haubold, 1984. Criteria for species separation still remain under discussion.
(5) Hyloidichnus, Glimoreichnus, Varanopus, and Erpetopus
The representatives of these four genera are collected as one basic morph due to the
similarity of the digit arrangement and proportions, respectively, in manus and pes
imprint. Both kind of imprints are characterized by a moderate increase of digit length
from I to IV, remarkably acute angled hypex, inward curved prints of claws as well as a
laterally oriented digit V. The imprints of digit tips often show an extramorphologically
caused bifurcation. Referred trackmakers are “pelycosaurs” to captorhinomorphs. Up to
now a more detailed interpretation is not possible due to the nearly uniform proportions
of phalanges and digits of known skeletons (Haubold 2000). The differentiation of the
four ichnotaxa is limited in a similar manner. Only in the case of well recorded
preservation there is a character for separation by the relative length and orientation of
digit V on the manus and pes imprints. However, due to various conditions in substrate
8
and gait many transitions in preservation of imprints and trackway patterns exist. They
complicate differentiation and even sometimes identification of the forms. The Hermit
Shale of the Grand Canyon (Gilmore 1926, 1927; Haubold 1973) is the stratum
typicum of type species of Hyloidichnus and Gilmoreichnus, H. bifurcatus Gilmore,
1927, and G. (Hylopus) hermitanus (Gilmore, 1927), respectively. Further specimens
of the taxa come from south-western France (Gand 1987) and New Mexico (Haubold et
al. 1995). Varanopus and Erpetopus has been described by Moodie (1929) based on the
type genera V. curvidactylus and E. willistoni from the Choza Formation, Texas. A
reexamination and collecting of additional material at the type locality has confirmed
the state of both ichnogenera as valid taxa (Haubold and Lucas 2001). Varanopus is
also known from the Lower Permian of New Mexico, France and Germany. There are
presumptive Erpetopus specimens in France and Italy (Haubold and Lucas 2001).
(6) Dromopus and Tambachichnium
This basic morphotype unites imprints of so called lacertoid or lizard-like shape. That
means the manus and pes present long and slender, claw-bearing digits, which are
distally curved inwards. Digit length increases from I to IV. Digit V is straight,
conspicuously shorter than IV and straddled away from the last in a different manner.
Dromopus and Tambachichnium are interpreted as tracks of araeoscelids (Haubold
2000). That interpretation is quite convincing and generally accepted. Support comes
from both the proportions of the body and extremities and by known skeletons of the
manus and pes. Dromopus has been described by Marsh (1894) based on collections
from the Upper Carboniferous of Kansas. Aside from Batrachichnus, the ichnogenus is
the most common element of the Upper Carboniferous to Lower Permian ichnofauna.
9
Dromopus (Saurichnites) lacertoides (Geinitz, 1863) in Europe and D. agilis Marsh,
1894 in North America are regionally applied species names. Haubold (1996) takes the
view that both species are identical to one another. As yet, a separation could not be
explained. In a strong sense, the differentiation between D. lacertoides / agilis and D.
palmatus (Moodie, 1929) = D. didactylus (Moodie, 1930) still remains an unresolved
problem. Digit III and IV are the most prominent and even often the only elements of
D. palmatus imprints. In addition, digit V seems to be abducted farther backwards.
Accordingly, it could be concluded that the trackmaker has longer metacarpalia and
metatarsalia in comparision to that of D. lacertoides (Haubold and Lucas 2001).
In contrast to Dromopus, Tambachichnium presents straight digits of smaller
divarications and presumably a more variable trackway pattern. Until now the
ichnotaxon is only known from the Tambach Formation in Thuringia by few
specimens, one trackway segment and three isolated and disputable imprints (Müller
1954; Haubold 1998).
<FIGURE 3 NEAR HERE>
The trackway pattern
The trackway pattern is determined by body proportions of the trackmaker, especially
the coupling value. This is the ratio of the gleno-acetabular distance to the length of
extremities. Variation of the trackway pattern of Lower Permian tetrapod ichnotaxa is
comparatively uniform. This means that the body proportions of associated
trackmakers could not have had great differences. If at all existing, in most cases these
differences were not prominent enough to appear in the trackway. Thus, the trackway
pattern of upper Paleozoic tetrapod ichnia is only a character of secondary
ichnotaxonomic importance (Haubold 1996). Nevertheless, there is a spread of
10
different trackway patterns within each single ichnospecies (fig. 3). That range reflects
variable speed of trackmaker locomotion (Voigt and Haubold 2000; Voigt 2001).
Among the Lower Permian ichnotaxa, low speed of gait is recorded by trackways, in
that in one the manus imprint of the left side is faced to the pes of the right side, and
vice versa. With increasing speed of gait, manus and pes of each single set from left
and right side, respectively, approach one another. The result is a primary overstepping
with a partial covering of imprints. Erpetopus, Tambachichnium and Dromopus
represent deviation from that spread of trackway pattern. Only from specimens of these
three ichnogenera an optional, completely primary overstepping of manus imprint by
the pes is known (Haubold 1971, 1973; Haubold and Lucas 2001; fig. 3). In addition,
the lacertoid tracks, Dromopus and Tambachichnium, show different width of pace on
the hind and the front feet. The larger width of pace on the hind limb as well as an
enhanced variability of trackway pattern correlate with the anatomically recorded
transformations of appendicular skeletons of the animals as mentioned above. In some
instances, the different orientation of manus and pes within the trackway gains
ichnotaxonomic importance. Amphisauropus is characterized by a significant
divergence of manus and pes axis with respect to the direction of motion. Manus
imprint is generally oriented inwards, whereas the pes points outwards. The axis of
both are separated by up to 50°. That character can take on importance as a criterion for
differentiation between tracks of Amphisauropus and Batrachichnus/ Limnopus. In
particular, it applies to doubtful cases of extramorphologically distorted tracks, where
the real number of digits on the fore feet can not be clarified.
Summary
11
The described imprint morphologies and trackway patterns can be considered as
representative forms of the Lower Permian tetrapod ichnofauna. The result is supported
by numerous conceptual integrative specimens from widely distributed equatorial to
subequatorial areas of Pangaea (fig. 4). The ichnotaxa, characterized in the previous
paragraphs, reflect the spectrum of terrestrial tetrapods of this time period that are
distinguishable by imprint morphology and trackway pattern. Tetrapod tracks from the
Permian are represented in scientific papers by 150 names of genera and about 280
species names (Haubold 2000). The majority of these are referred to Lower Permian
ichnotaxa. All up to now separated and named Lower Permian ichnotaxa can be
principally referred to the six basic morphs discussed in this paper. Four of these basic
morphs are already known from specimens from the Upper Carboniferous. These are
(1) Batrachichnus/ Limnopus, (2) Ichniotherium, (3) Dimetropus and (4) Dromopus.
The diversity is extended to six basic morphs with 11 ichnogenera altogether in the
Lower Permian due to evolutionary progress of the locomotory apparatus of synapsids
and eureptiles. The majority of the ichnofauna and occurrences, respectively, are
dominated by Batrachichnus, Dimetropus and Dromopus. They represent, presumably,
tracks of temnospondyls, sphenacodontid “pelycosaurs” and araeoscelids.
<FIGURE 4 NEAR HERE>
The example of microsaurs shows that possibly not all osteologicallyestablished taxa can be recorded by ichnological specimens. Terrestrially adapted
tuditanomorph microsaurs range from the Upper Carboniferous to the Early Permian.
They have shorter extremities in relation to the glenoid-acetabulum distance than do
temnospondyls. The differences in body proportions should be reflected in the
trackway pattern by secondary overstepping. However, separation of trackways with
12
complete secondary overstepping from such with primary or without overstepping is
generally difficult. Therefore, it is doubtful to identify only from trackway pattern
distinct trackmakers without any significant character deviation in imprint morphology.
The majority of microsaurs have a tetradactyl manus, a character they share with
temnospondyls. Therefore, as yet it is impossible to document microsaur tracks in the
Lower Permian Red beds or to separate them from the Batrachichnus/ Limnopus track
morph. Parallelisms or convergent developments inducing almost identical or
functionally homologous extremity morphologies can limit the specification of
osteological trackway interpretation in some cases. In addition, the postcranium is of
less taxonomic importance than the anatomy of the cranium, which represents the base
of the tetrapod evolutionary system in the Permian Period, too. However, in particular,
the structure of the distal extremities of Lower Permian terrestrial vertebrates are of
sufficient significance to allow the differentiation of at least six principal track morphs.
It establishes an interpretation of fossil tracks and trackways that is based on
anatomical characters of the trackmakers. In that way, ichnia tetrapodorum are another
kind of tool for stratigraphic and faunistic conclusions, which should be considered as
supplementary and controlling factors of osteological data.
Acknowledgment. Special thanks to Dr. Spencer Lucas for revising the English.
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Williston, S. W. 1911. American Permian Vertebrates, 145 pp. Chicago: University
Press.
Woodworth, J. B. 1900. Vertebrate footprints on carboniferous shales of Plainville,
Massachusetts. Bulletin of the Geological Society of America 11: 449-454.
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FIGURE LEGENDS
FIGURE 1-Temporal distribution of tetrapod taxa with potential terrestrial
representatives during the Lower Permian based on Lucas (1998). Most of the tetrapod
track-bearing strata mentioned in the text, the Goldlauter and the Tambach Formation,
the Hermit Shale, and the Choza Formation, for example, are considered to belong to
the Cisuralian – Lower Permian – time series.
FIGURE 2-Representatives of the six significant Lower Permian track morphs and
osteological taxa of inferred trackmakers. In all cases, the manus imprint is in front of
the pes. The photograph of Dromopus only shows the pes imprint. In the related
drawing below the outline of the manus print is added. Line drawings are not scaled.
FIGURE 3-Typical Lower Permian trackway patterns (modified after: Haubold 1971).
The majority of studied trackways are spanning a range between 1 and 3. Striking
overstepping of pattern number 4 is only recorded in specimens of Erpetopus,
Dromopus, and Tambachichnium as a result of higher speed of gait.
FIGURE 4-Lower Permian – Cisuralian - Pangaea (based on: Paleogeographic Atlas
Project, University of Chicago) showing position of the most important Lower Permian
tetrapod track locality areas of North America and Europe which both were arranged in
equatorial latitudes during that period. The tetrapod ichnofossil assemblages, the bone
record, and the paleogeography confirm the supposition of an extended uniform habitat
across the central part of the supercontinent.
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