Quaternary Research

Copyright © University of Washington. Published by Cambridge University Press, 2019.

doi:10.1017/qua.2019.40

New fossil sea turtle trackway morphotypes from the Pleistocene of

South Africa highlight role of ichnology in turtle paleobiology

Martin G. Lockleya*, Hayley C. Cawthrab,c, Jan C. De Vynckb, Charles W. Helmb,d, Richard T. McCread, Ronel Nele
a
Dinosaur Trackers Research Group, Campus Box 144, University of Colorado Denver, P.O. Box 173364, Denver, Colorado 80217-3364, USA
b
African Centre for Coastal Palaeoscience, P.O. Box 77000, Nelson Mandela University, Port Elizabeth 6031, South Africa
c
Marine Geoscience Unit, Council for Geoscience, P.O. Box 572, Bellville 7535, South Africa
d
Peace Region Palaeontology Research Centre, Box 1540, Tumbler Ridge, British Columbia V0C 2W0, Canada
e
Department of Zoology, Nelson Mandela University, Port Elizabeth 6031, South Africa
*Corresponding author at: e-mail address: martin.lockley@ucdenver.edu (M.G. Lockley).
(RECEIVED April 4, 2019; ACCEPTED June 10, 2019)

Abstract
More than 130 late Pleistocene trackway sites from the coastal eolianites and beach deposits of the Cape south coast, South
Africa, have previously mostly yielded tracks of large mammals and birds. However, two sites east of Still Bay, and a third
near Garden Route National Park, yield distinctive trackways of hatchling sea turtles, made during the short posthatching
(postemergence) interval when the trackmakers headed for the sea. One assemblage of approximately parallel trackways indi-
cates smaller loggerhead turtle hatchlings, with alternating gaits, and contrasts with a wider trackway indicating a leatherback
turtle hatchling. These are the world’s first reports of fossil traces that document this brief “run-for the-sea” phenomenon.
They help delineate late Pleistocene sea turtle breeding ranges and indicate climatic conditions along the Cape south
coast. Ichnotaxonomically defined swim tracks of large adult sea turtles are known from a few Mesozoic sites. Likewise,
walking and swim traces of terrestrial freshwater turtles are also known from the Mesozoic and Cenozoic. However, as no
ichnotaxonomy exists for these diagnostic hatchling trails, we assign the trackways of the inferred loggerheads to the new
ichnotaxon Australochelichnus agulhasii ichnogen. et ichnosp. nov., and the inferred leatherback trackway to Marinerichnus
latus ichnogen. et ichnosp. nov.
Keywords: Pleistocene; Hatchling sea turtle; Trackways; Ichnotaxonomy; Coastal eolianites; Paleoecology; Loggerhead
turtle; Leatherback turtle

INTRODUCTION
Diverse fossil footprints are abundant and widely distributed
in late Pleistocene coastal eolianites and cemented foreshore
deposits, preserved along the Cape south coast of South
Africa (Fig. 1). These cemented paleodune deposits, which
form the ∼100,000-yr-old Waenhuiskrans Formation along
the Cape south coast, have been surveyed on foot between
2007 and 2019 by the present authors (mainly CWH) from
Arniston in the west to Robberg in the east, a distance of
∼350 km (Fig. 1).
These tracksite discoveries have generated a recent spate of
publications detailing a variety of tracks of mammals and
Cite this article: Lockley, M. G., Cawthra, H. C., De Vynck, J. C., Helm,
C. W., McCrea, R. T., Nel, R. 2019. New fossil sea turtle trackway
morphotypes from the Pleistocene of South Africa highlight role of
ichnology in turtle paleobiology. Quaternary Research 92, 626–640.
https://doi.org/10.1017/qua.2019.40
626
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other tetrapods. Most notable have been the reports of homi-
nin tracks (Helm et al., 2018b, 2019b), bird tracks (Helm
et al., 2017), and various large mammal tracks (Roberts
et al., 2008; Helm et al., 2018a, 2019a). All these studies
stress the vulnerability of track-bearing blocks because of
proximity to the ocean and the inevitable damage caused
by continuous wave action and cliff collapse.
Although it has been inferred that the Cape south coast
track record may be biased toward the tracks of larger, heavier
trackmakers, especially mammals (Helm et al., 2018a, 2018b,
2018c, 2019a, 2019b), some tracks, including those of birds
(Helm et al., 2017), small mammals (Helm et al., 2018c), and
the hatchling sea turtles reported here, show the potential for
registration and preservation of smaller tracks. The only other
possible reptile tracks, putatively made by tortoises, and
observed in small numbers at three localities (Dana Bay,
Gericke’s Point, and Goukamma), are morphologically
quite distinct from hatchling sea turtle tracks and are as yet
undescribed and will be reported elsewhere.
 New fossil sea turtle trackway morphotypes from the Pleistocene of South Africa 627

Figure 1. (color online) Locality map showing the location of track-bearing Waenhuiskrans Formation outcrops in the Still Bay area of South
Africa. Site 1 is the more westerly, and site 2 the more easterly of the two sites, marked with stars. The inset at lower right shows the
iSimangaliso Wetland Park World Heritage Site northern KwaZulu-Natal, near St. Lucia, known as a nesting area for extant sea turtles.

The purpose of this article is to describe a number of highly
distinctive and diagnostic tracks (trails) attributed to sea turtle
hatchlings. These tracks have considerable ichnological and
paleobiological significance for the following reasons: (1)
these are the first sea turtle hatchling tracks known from the
fossil record; (2) this is the first documentation of fossil reptile
tracks from South Africa’s Cape south coast, in an ichnofauna
otherwise dominated by mammal and bird tracks; (3) from a
paleobiological viewpoint these tracks evidently capture the
behavior of sea turtle hatchlings in the first few minutes of
postemergence activity, the only time they will make terres-
trial tracks at this stage of ontogenetic development; (4)
from a paleogeographic and paleoecological viewpoint the
tracks tell us about the breeding range of sea turtles in the
Pleistocene; and (5) the tracks represent two new and highly
distinctive and morphologically diagnostic categories of fos-
sil turtle tracks, which require the naming of two new ichno-
taxa in order to distinguish them from other formally named
turtle track morphotypes.
With reference to the fact that hatchling sea turtle tracks
have never previously been reported, it is pertinent to com-
pare them with other previously described tracks of sea and
terrestrial turtles from the fossil record, as well as hatchling
sea turtle tracks made on southern African beaches today.
The fossil trackways are known mostly from the Mesozoic
of the Northern Hemisphere and include evidence of both
swimming and walking behaviors of presumed mature adults.
These are associated with different paleoenvironmental set-
tings from the coastal beach-dune facies described here. How-
ever, the morphology of these various Mesozoic trackways,
reported with increased frequency in recent years (Bernier
et al., 1982; Thulborn, 1989, 1990; Gaillard et al., 2003;
Avanzini et al., 2005; Lockley et al., 2018a, 2018b, 2019),
also has ichnotaxonomic significance, which is reviewed
here and discussed in relation to a coherent classification of
the fossil tracks and trackways of turtles.
GEOLOGIC SETTING AND STRATIGRAPHIC
CONTEXT
The track-bearing Pleistocene coastal dunes and beach sands
of the Cape south coast have been described by Roberts et al.
(2012) and Cawthra et al. (2018) based on the Great Brak
River outcrops as including three types of Pleistocene facies
that can look quite similar. These can be broadly

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 628
Figure 2. Large sandstone block showing parallel sea turtle trackways (black arrows) preserved in concave epirelief on the track-bearing sur-
face. Large and small red squares correspond to line drawings in Figure 5A and C. (For interpretation of the references to color in this figure
legend, the reader is referred to the web version of this article.)
characterized as eolianite facies characteristic of the Waen-
huiskrans Formation (Malan, 1989) and foreshore and shore-
face facies that form part of the Klein Brak Formation (Malan,
1991). These are most likely dated from Marine Oxygen
Isotope Stage (MIS) 5, between ∼130 and ∼90 ka (see Helm
et al. [2018b] for discussion and the remote possibility of
older MIS 11 dates). These facies intergrade to various degrees
and are characterized locally by high-angle cross bedding,
dune sets, and low-angle to planar bedded foreshore facies.
The track-bearing surfaces are referred to as turtle tracksites
1 and 2. Turtle tracksite 1 was exposed by the fall of a large
block, about 2.5 m thick, from the cliffs above. A possible
source of the block is ∼15 m above its present position
(Fig. 2). The block came to rest in a position that, when exam-
ined in 2017, exposed the track-bearing surface in a near-
vertical orientation. The surface is on the underside of the
block where the planar bedded beach sands split along an
apparent stratigraphic plane of weakness to expose a very
flat surface with maximum dimensions of about 3.0 ×
5.6 m, representing a surface of no more than 17.0 m2, on
which multiple elongate hatchling turtle trails were registered
(Figs. 2–5). The sides of the block reveal that the sand was
horizontally bedded or part of a very low-angle sequence,
with what appear to be at least two tracks in cross section
(Fig. 3B), at a horizon separated from the track-bearing sur-
face by about 40 cm. These tracks appear as “v-shaped
depressions” with at least 6–7 cm of relief, but as they do
not indent the horizontal layers at the apex of the “v,” they
are best referred to as track or bioturbation “features” and
not depressions.
The substrate, on which the trackways were registered, was
sand, now consolidated and yellowish brown in color. Short
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M. G. Lockley et al.
dark streaks on the upper part of this surface at the junction
with the upper surface of the block reflect continuation of
the dark-brown to black coloration on the upper surface,
which is one of the sides where the block is seen in cross sec-
tion (Fig. 2). Newly exposed surfaces in this area are light yel-
lowish brown, but after prolonged exposure to weathering,
exposed surfaces become dark brown or black and, in places,
are covered in a biofilm layer, which attaches preferentially to
convex and irregular surfaces (Helm et al., 2017). This bio-
film also eventually tends to develop on flat surfaces. This
dark-brown upper layer, which implies prolonged exposure,
when viewed in the context of the source cliffs above, allows
the inference that the trackways we describe were heading
south in a seaward direction. The trails on the track-bearing
surface appear as elongate, slightly curved tripartite furrows
with the middle part a raised ridge (convex epirelief) about
one-third of trail width, with the two side furrows showing
long sequences of transverse paddle traces in concave epire-
lief. Thus, the trackways are the original natural impressions
(concave epireliefs) not the infilling casts (Figs. 2–4). Unfor-
tunately, the counterpart (convex hyporelief) of this surface
or the in situ surface from which the block fell is not exposed
or accessible for study. When revisited in 2018, the block had
split at a stratigraphic level below the track-bearing surface
and was tilted seaward away from the near-vertical inclination
to an angle of ∼60°.
Turtle tracksite 2 is situated 4.4 km east of the previously
identified site 1 (turtle tracksite 1) and consists of a subrectan-
gular, fallen sandstone slab 160 cm in maximum length,
90 cm in maximum width, and 32 cm in maximum thickness
(Fig. 6). Bedding appears parallel or subparallel, without
clear cross bedding. Bedding planes vary in thickness from
 New fossil sea turtle trackway morphotypes from the Pleistocene of South Africa 629

Figure 3. (color online) (A) Three sea turtle hatchling trackways (Australochelichnus agulhasii) from central area of slab shown in Figure 2.
Note that two trackways (right) overlap. (B) Stratigraphic section of track-bearing block, showing the track-bearing surface in vertical orien-
tation (up section to left) and presumed tracks in cross section, center, in layers just left of 15-cm scale bar.

2 to 10 mm. The track-bearing surface represents a single
bedding plane with the natural impression (concave epirelief)
of a single trackway (Figs. 6 and 7). Shell fragments or other
coarse material were not observed. The slab is located above
the intertidal splash zone and ∼ 5m from the bottom of steep
coastal cliffs. It is not possible to reliably determine from
which layer in the cliffs it originated, and the counterpart
slab has not been located. It is also not possible to determine
the original bearing of the trackway relative to the coastline.
Moreover, the slab is wedged almost vertically between two
much larger rocks and is reasonably well protected from
erosion, at least in the short term.
Tectonic activity is inferred to have been minimal on the
Southern Coastal Plain during the Pleistocene (Fleming
et al., 1998). Thus, the planar bedded strata seen throughout
the block can be inferred to represent strata previously situ-
ated in situ somewhere in the overlying cliff strata, where
the bedding planes reflected inclinations close to their
original angle of deposition.
MATERIAL AND METHODS
The tracks at sites 1 and 2 were inspected and photographed
in the field in order to obtain both two-dimensional and three-
dimensional (3D) images (Figs. 2–7). Two-dimensional
images were obtained from photographs, and tracings made
on clear acetate film. Tracings were made of three representa-
tive site 1 trackway segments (Fig. 5) after chalking the

Figure 4. Photogrammetric image of turtle trackway shown in left side of large box in Figure 2 and on left side of Figure 3. This trackway is
designated as the holotype of Australochelichnus agulhasii, and represented by a “hard copy” three-dimensional (3D) print (MGL 600 and
UCM 230.281) of the representative segment shown in red box. See text for details. (For interpretation of the references to color in this figure
legend, the reader is referred to the web version of this article.)

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 630 M. G. Lockley et al.

Figure 5. (A) Three subparallel sea turtle trackways (A. agulhasii), inferred to represent leatherback hatchlings: gray arrows show direction of
progression. Note appearance of a median ridge (small black arrows) after appearance of paddle traces. (B) Paired trackway-parallel traces
before appearance of paddle traces. (C) Detail of trackway shown in panel A to illustrate that paddle traces alternate (left and right), as
shown by plotting transverse lines (red) across trackway axis from anterior border of each paddle trace. (D) Sea turtle trackway (Marinerichnus
latus), inferred to represent a loggerhead hatchling. Note that left and right paddle traces (red lines) registered symmetrically about the trackway
midline. Note that panels C and D are shown at the same scale to show that the M. latus trackway width (TW) is much wider. (For interpretation
of the references to color in this figure legend, the reader is referred to the web version of this article.)

outlines of the main features. The tracings are reposited in the
University of Colorado Museum of Natural History as T
1781. Photogrammetric images of the trackways from turtle
tracksite 1 were obtained using a Canon PowerShot ELFPH
340 HS camera by one of us (JCDV). Point clouds and digital
terrain models were compiled (by RTM) using Agisoft Pho-
toscan Professional (v. 1.0.4), and color topographic profiles
were created with CloudCompare (v. 2.6.3.beta) to produce
an image of the trackway shown in Figure 4. Attempting to
obtain photogrammetric images of the trackways from turtle
tracksite 2 proved challenging because of the lack of camera
access in cramped space beside the trackway. However, obli-
que and close-up photos of trackway segments were obtained,
from which measurements were procured (Figs. 6–7), and a
photogrammetric model was also obtained (Fig. 6C), cour-
tesy of Carina Helm with Agisoft MetaShape Professional
(v. 1.0.4) using 27 images from Canon PowerShot ELPH
110 HS with 4608 × 3456 resolution and pixel size of
1.34 × 1.34 µm using a 4.3 mm focal length. The average
camera altitude was 0.22 m. The reprojection error of the
model is 0.71 pix.
The 3D digital image data for the site 1 trackway were used
to generate a 3D print of a segment of the trackway, shown in
Figure 4, which is reposited in the fossil footprint collections
of Moab Giants Museum (MGL), Utah, as “hard copy” rep-
lica MGL 600. A second hard copy replica is reposited at
the University of Colorado Museum of Natural History
(UCM) as specimen UCM 230.281.

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 New fossil sea turtle trackway morphotypes from the Pleistocene of South Africa 631

Figure 6. (A) The Marinerichnus latus trackway block wedged in between much larger fallen blocks. (B) Close-up view of trackway with
arcuate red lines to show curvature of trackway and approximate lengths of inner (concave) and outer (convex) margins at 600 mm and
830 mm, respectively. (C) Photogrammetric model with color vertical profile of leatherback turtle trackway east of Still Bay, courtesy of Carina
Helm (see “Material and Methods”). Vertical scale is in meters. Horizontal scale bar is 0.25 m. (For interpretation of the references to color in
this figure legend, the reader is referred to the web version of this article.)

While this article was in review, a third tracksite yielding a
hatching turtle trackway was discovered (in April 2019), in a
coastal section of the Garden Route National Park, by Andre
and Emily Brink. This site is 110 km east of sites 1 and 2 and
consists of an isolated loose slab ∼60 cm × 60 cm wide and
long and ∼20 cm thick (Fig. 8). The slab was found on a
dune surface below vegetated slopes lacking nearby visible
outcrops to suggest the stratigraphic level from which it
came. Faint horizontal bedding is evident, parallel to that of
the track-bearing surface, which appears to have been

Figure 7. (color online) (A) Close-up photograph of Marinerichnus latus trackway segment showing measurements of major morphological
features made directly form the trackway (see “Description”). (B) Line drawing of the same area showing midline ridge, margins of trackway,
and symmetrical arrangement of paddle traces (see “Description”). Compare with Figures 5D and 6.

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 632
Figure 8. (color online) Turtle trackway site 3, Garden Route National Park area. (A) View of isolated track-bearing slab on beach. (B) Detail
of slab with clear, diagnostic trackway segment (long arrow) and unclear trace (short arrow left) with opposite sense of progression. Trackway
segments occur on either side of an uneven area that might represent remains of a nest. See text for details.
exposed and weathered/eroded for some time. Nonetheless,
the natural mold of one clear turtle trackway with an alternat-
ing paddle trace gait is similar in size and morphology to the
site 1 trackways inferred to represent a hatchling loggerhead
turtle (Caretta caretta).
The diagnostic trackway segment, 25 cm long and 9 cm
wide, is preserved in the middle of the slab with evidence
of forward motion away from a disturbed (bioturbated) area
(left to right in Fig. 8) and toward a broken area (far right
in Fig. 8). On the opposite side of the uneven area from the
diagnostic trackway segment, other paddle-like traces, with
curved edges, consistent with hatchling turtle trackmaker
morphology are evident and apparently suggest motion in a
direction ∼180° opposite to that of the main trackway. This
ichnological evidence is difficult to interpret but could possi-
bly suggest at least two trackways emanating from a nest site.
Ichnologically, the trackways from sites 1 and 2 can be clas-
sified as epifaunal trails, whereas a nest would be an infaunal
trace, and potentially more easily preserved than surface
trails, and potentially amenable to study through sectioning
of the sedimentary rock comprising the slab. However,
although the site 3 specimen (Fig. 8) is portable and can be
removed to the safety of a repository for further study, it is
premature to undertake any potentially damaging analysis.
A fourth turtle trackway site was reported to us (see
“Acknowledgments”). However, details of this fourth site,
although indicated by photographs, are insufficiently
known to allow us to describe the material adequately without
further study of the field evidence.
ICHNOTAXONOMY
General ichnotaxonomic observations
According to the latest taxonomic revisions (Rhodin et al.,
2017), all tortoises, a subgroup within the turtles (Chelonia),
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M. G. Lockley et al.
are terrestrial. Thus, all tortoises are turtles, but not all turtles
are tortoises. Nontortoise turtles are mostly aquatic and
include both terrestrial, freshwater turtles and sea turtles. In
the discussion that follows, we use the term turtle to exclude
terrestrial tortoises. This is convenient when discussing the
fossil track record of chelonians, as there are no confirmed
reports of the tracks of terrestrial tortoises, even though a
few questionable Cretaceous occurrences have been sug-
gested (Fiorillo, 2005; Pascual-Arribas and Hernández-
Medrano, 2015), and we have recently observed a few as
yet undescribed tortoise tracks in the coastal dune and
beach deposits described here.
Fossil turtle tracks are known from the Mesozoic and
Cenozoic, with some evidence that the earliest known tracks,
from the Early Triassic, are older than the oldest known skel-
etal remains (Lichtig et al., 2018). The turtle track record cov-
ers a variable range of morphologies reflecting the
locomotion of different-sized marine and freshwater turtles
while either swimming or progressing on land. In most
cases there are clear-cut differences between the trackway
configurations made by swimmers and walkers, and also
between the facies associations of marine or sea turtle traces
(Fig. 9) and those associated with freshwater settings. There
are also at least four widely used ichnogenus names applied
to turtle tracks. These include Chelonichnium, originally
applied to an undiagnostic isolated track (C. vogesiacum)
from the Early Triassic of Germany (Schimper, 1850). As
reviewed by Avanzini et al. (2005), Abel (1935), and Hau-
bold (1971), this ichnotaxon was declared dubious (a
nomen dubium). This led Demathieu and Gaillard (1982, in
Bernier et al., 1982), to propose adopting (transferring) the
ichnogenus name to label large sea turtle tracks from the
Late Jurassic of France. These are preserved in a long trail
(Fig. 10), which they named Chelonichnium cerinense.
These authors used the potentially ambiguous term “turtle”
throughout their study, which we take to mean nontortoise
 New fossil sea turtle trackway morphotypes from the Pleistocene of South Africa 633

Figure 9. (color online) (A–C) Modern hatchling loggerhead turtles en route to sea, registering trackways on sandy substrates with variable
surface textures. Note scale bars in panels A and C for comparison with Figure 7. (D) Adult loggerhead digging nest. Trackway (∼1.0 m wide)
indicates onshore (up-beach) progression and shows distance from sea. Photographs in panel A (November 2015) and panel D (2013) courtesy
of Linda Harris. Photographs in panels B and C (January 2019) courtesy of Diane Le Gouvello. All photographs from Bhana Nek, in iSiman-
galiso Wetland Park World Heritage Site northern KwaZulu-Natal, South Africa.

turtles. (However, they may have wished to imply tortoise.)
They explicitly described the Chelonichnium trackway as
progressing from a drier to wetter substrate in an area with
what they described as “soft sediment slides” confirming a
substrate at least partly immersed, subaqueously, in shallow
marginal marine facies. Interestingly, as in the case noted pre-
viously where Early Triassic turtle tracks may predate the old-
est body fossil remains, so these Late Jurassic sea turtle tracks
apparently also predate the oldest known body fossil remains.
Bernier and his colleagues also named Saltosauropus latus
(Bernier et al., 1984), which they initially misinterpreted as
tracks of a hopping dinosaur. These tracks were convincingly
reinterpreted as those of large swimming marine turtles,
which provided trace evidence of large paddles used in syn-
chronous swimming strokes (Thulborn, 1989, 1990). This
interpretation has been supported by additional finds of track-
ways “attributed to giant turtles” (Gaillard et al., 2003, p. 315)
that resemble Saltosauropus (Fig. 9). Thus, in the sense used
by Bernier and his colleagues, Chelonichnium and Saltosaur-
opus respectively represent the progression of a marine turtle,
in the former case over a wet partly submerged substrate, and
in the latter case, swimming more freely, but still touching the
substrate with paddle strokes.
Because freshwater turtles, associated with fluviolacustrine
facies in terrestrial deposits, may also walk or swim, we can
expect to find trackways representing both modes of progres-
sion. This is exactly what we find in the track record. Gener-
ally, the traces of freshwater turtles have been labelled
Chelonipus (Rühle von Lilienstern, 1939), although the ich-
nogenus name Emidhypus (Fuentes Vidarte et al., 2003)
has also been used (see Avanzini et al., 2005, for review of
ichnotaxonomy). The record of Chelonipus extends from
the Early and Late Triassic of Europe (Rühle von Lilienstern,
1939, and Haubold, 1971, respectively) to the few sites in the
Jurassic of North America (Foster et al., 1999; Lockley and
Foster, 2006) and multiple tracksites in the Cretaceous of
North America (Lockley et al., 2018a, 2018b). The label
Emidhypus has been used to describe Middle Jurassic
(Klein et al., 2018) and Late Jurassic turtle tracks from
Spain (Avanzini et al., 2005) and Morocco, as well as the
original tracks described from the Early Cretaceous of
Spain (Fuentes Vidarte et al., 2003). The difference between
the type specimens of Chelonipus and Emidhypus is slight,
based on more “parallel ungual traces” in the manus (front
foot), turned slightly inward, and “apparently always away
in respect to” the pes (hind foot) (Avanzini et al., 2005,
p. 748). These authors considered that Emidhypus might be
a “junior synonym” of Chelonipus but did not opt for this
conclusion because the trackway configurations were differ-
ent. Fuentes Vidarte et al. (2003) had failed to compare
their Emidhypus with Chelonipus but were “saved” by Avan-
zini et al. (2005) who demonstrated diagnostic differences
justifying both names as valid. Note here that the differences
in trackway configuration are considered sufficient to validate
the naming of different ichnogenera.
Avanzini et al. (2005) explicitly interpreted the type mate-
rial of both ichnogenera as examples of walking behavior and
supported this conclusion by comparing the fossil trackway
with those of living turtles, including the tortoise Testudo
and other nontortoise turtle genera. Reolid et al. (2018)
described trackway evidence of what they called “semi-
aquatic locomotion” but indicated that tracks could not be
given a “concrete assignment” to either Chelonipus or Emid-
hypus. Thus, these two ichnogenera may be used as labels for
walking trackways, in the type specimens, but may elsewhere
be difficult to distinguish and may additionally indicate
semiaquatic locomotion.
Turtles are capable of subaqueous bottom walking and,
depending on substrate consistency (firmness), may register
very indistinct trackways that do not closely reflect foot

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 634 M. G. Lockley et al.

Figure 10. (color online) Sea turtle (A–D) and terrestrial turtle (E–G) trackways. (A–C) From the Jurassic of France. (A) Chelonichnium cer-
inense (after Bernier et al., 1982). (B) Saltosauropus latus (after Bernier et al., 1984). (C) Unnamed Saltosauropus-like giant turtle tracks
(modified after Gaillard et al., 2003). (D) Australochelichnus agulhasii from South Africa. (E) Chelonipus plieningeri from the Upper Triassic
of Germany (after Haubold, 1971, 1984). (F) Chelonipus isp. from the Cretaceous of Utah (after Lockley et al., 2018a). (G) Turtle swim track
assemblage T 1491 from the Cretaceous of Colorado (after Lockley et al., 2018b).

morphology (Avanzini et al., 2005), or they may register quite
regular trackway patterns that are difficult to distinguish from
those made when walking on emergent subaerial substrates
(Foster et al., 1999; Avanzini et al., 2005; Lockley et al.,
2014). In other paleoenvironmental settings, high densities
of parallel to subparallel, incomplete tracks (mostly distal
toe traces) with no discrete or recognizable trackways config-
urations have been registered (Lockley et al., 2018a, 2019).
The inability of observers to recognize trackways may be
because of multiple overprinting. These suggest swimming,
with toes touching the substrate under the influence of unidi-
rectional or preferred orientation currents, or some other pale-
oenvironmental influence on the direction of progression of
multiple individuals.
The South African tracks named here are morphologically
quite distinct from any turtle trackways previously described
but exhibit some general similarities to the turtle traces named
Chelonichnium cerinense (Demathieu and Gaillard, 1982, in
Bernier et al. 1982). For these reasons, we erect the new ich-
notaxa Australochelichnus agulhasii ichnogen. et ichnosp.
nov. and Marinerichnus latus ichnogen. et ichnosp. nov.,
which recognize the “chelonian” and “mariner” affinities of
the trackmakers and also recognize that the two trackways
registered quite different “terrestrial” gaits (i.e., alternating
in A. agulhasii and symmetrical in M. latus). As these differ-
ent gaits are unequivocally tied to loggerhead (genus Caretta)
and leatherback (genus Dermatochelys) turtles, respectively,
it is ichnotaxonomically consistent to place them in separate

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 New fossil sea turtle trackway morphotypes from the Pleistocene of South Africa
ichnogenera. However, both represent extant sea turtles, or
close relatives, and can be united in a single ichnofamily.
Because of the opinion of various ichnologists that Chelo-
nichnium is a nomen dubium (Avanzini et al., 2005, and ref-
erences therein), we resist using this ichnogenus name as the
basis of an ichnofamily. Instead, we erect the new ichnofam-
ily Marineropodidae (derived from mariner, meaning sea-
farer: Spanish/French/English).
Formal ichnotaxonmy
Ichnofamily Marineropodidae
Referred material
Australochelichnus agulhasii ichnogen. et ichnosp. nov.
MGL 600/UCM 230.281, this study. Marinerichnus latus
ichnogen. et ichnosp. nov., this study.
Narrow, elongate subsymmetrical trackways (trails) char-
acterized by a trilobed configuration consisting of a central
ridge, or groove, with serial, posteriorly divergent paddle
traces oriented postero-laterally on both sides of the trackway
midline. Paddle impressions may register as mirror image
traces, or with slight alternation on either side of the central
ridge.
Discussion of ichnofamily features
Where paddle traces are deeper than the trace of the ventral
plastron, because of the lifting of the trackmaker’s body
above the deepest paddle registration points, the central or
midline trace between the paddle marks will appear as a
raised ridge in natural impressions (a convex epirelief ridge
separated by concave lateral grooves with serial paddle traces.
A natural cast of the natural impression will show all features
in reverse—that is, an indented groove (concave hyporelief
groove separated by convex lateral ridges with serial paddle
trace casts). The height (or depth) of the central ridge
(groove) in natural impressions will vary depending on the
substrate and the ability of the trackmaker to raise its plastron
enough to reduce or minimize contact with the substrate.
Australochelichnus ichnogen. nov.
Diagnosis. Continuous subsymmetrical trails about 10 cm
wide with median groove averaging about 35–40% as wide as
whole trail separating very regular serial, posteriorly angled
rhomb-shaped “paddle” traces, with anteriorly convex mar-
gins. Anterior margins of paddle traces show an alternating
pattern. Median groove may be locally absent.
Type material. Holotype and paratype trails occur as natu-
ral impressions on a large fallen block of consolidated beach
sandstone. Holotype preserved as photo series used to create
3D image (Fig. 3).
Age of type material. Middle to late Pleistocene.
Type horizon and locality. Klein Brak Formation, east of
Still Bay, ∼4.4 km west of M. latus type locality.
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635
Derivation of ichnogenus name. “Southern turtle trail,”
Australo meaning southern, chel referring to chelonian or tur-
tle, and ichnus meaning trace or trail.
Australochelichnus agulhasii ichnosp. nov., Figures 2–5
Type material. As for ichnogenus. MGL 600/UCM
230.281.
Age of type material. As for ichnogenus.
Type horizon and locality. As for ichnogenus.
Derivation of ichnospecies name. Referring to Agulhas
current, which may have influenced turtle breeding range in
the study area.
Description. Continuous subsymmetrical trails about
10 cm wide with median ridge ranging from 25% to 50%
as wide as whole trail but averaging 35–40% of trail width.
Median ridge separating posteriorly angled rhomb-shaped
flipper or paddle traces slightly wider than long, where length
is measured parallel to trail axis. Left and right paddle traces
show regular alternating pattern, with anterior margins on
each side offset about half a paddle trace length relative to
opposite side. In cross section median ridge appears in con-
vex epirelief, whereas lateral paddle traces are subhorizontal
to slightly concave epireliefs. Anterior margins of paddle
traces are crescent to semicircular in shape and anteriorly con-
vex, with posterior margins marked by identical anteriorly
convex traces of previously registered paddle marks, which
repeat to create a highly regular series of traces on each
side. Median margins of paddle traces are angled at about
110° to direction of travel marked by trackway midline, and
curving postero-laterally to create outer trackway margin.
Some trails show segments that registered only lateral pad-
dle traces, followed continuously by segments where the
median ridge is also present. This suggests a change in loco-
motor style or substrate consistency that is reflected in differ-
ences in the configuration of the trail.
Marinerichnus ichnogen. nov.
Diagnosis. Continuous subsymmetrical trails about 14–
15 cm wide with median ridge averaging about 30% as
wide as whole trail separating very regular serial, posteriorly
angled rhomb-shaped “paddle” traces, with anteriorly convex
margins. Anterior margins of paddle traces show mirror
image configuration about trackway midline.
Type material. Holotype trails occur as natural impressions
on fallen block of consolidated beach sandstone (Figs. 5D, 6,
and 7).
Age of type material. Middle to late Pleistocene.
Type horizon and locality. Klein Brak Formation, east of
Still Bay, ∼4.4 km east of A. agulhasii type locality.
Derivation of ichnogenus name. “Seafarer turtle trail,”
mariner meaning one who goes to sea, and ichnus meaning
trace or trail.
Marinerichnus latus ichnosp. nov., Figures 6–7
Type material. As for ichnogenus.
Age of type material. As for ichnogenus.
Type horizon and locality. As for ichnogenus.
 636
Derivation of ichnospecies name. Referring to latus, the
width of the trackway.
Description. A continuous symmetrical trail about 14–
15 cm wide and ∼70 cm long as measured along curved mid-
line, with median ridge about 30% as wide as whole trail.
Median ridge separating posteriorly angled arcuate paddle
traces, longer than wide, where length is measured parallel
to trail axis. Left and right paddle traces show symmetrical
arrangement about the median ridge/trackway midline. In
cross section median ridge appears in convex epirelief
whereas lateral paddle traces are concave epireliefs, up to
15 mm deep. Anterior margins of paddle traces are arcuate
anteriorly and antero-laterally with posterior margins marked
by identical anteriorly convex traces of previously registered
paddle marks. However, the length of paddle traces measured
parallel to the trackway axis varies, averaging ∼2.5 cm (range
∼1.5–2.5 cm). Median margins of paddle traces are angled at
about 130° to direction of travel marked by trackway midline
and curve postero-laterally to create outer trackway margin.
Comparisons between A. agulhasii and M. latus
A. agulhasii and M. latus can be differentiated by a number of
distinctive morphological features of the traces, which appear
to have unequivocal ichnotaxonomic significance. For exam-
ple, M. latus is a wider (14.0–15.0 cm) trackway with larger,
more posteriorly oriented paddle traces, arranged symmetri-
cally about the trackway midline. This makes the median
ridge narrower in proportion to the trackway width (∼30%,
compared with 35–40% in A. agulhasii). This is in marked
contrast to the narrower trackway of A. agulhasii
(∼10.0 cm, thus only ∼67–71% as wide as M. latus), which
has alternating and shorter, less posteriorly oriented paddle
traces. All features of both trackways are consistent with
inferring that A. agulhasii was made by a loggerhead turtle
hatchling and M. latus by a leatherback turtle hatchling.
The M. latus trackway is curved, and there is greater variation
in the length of the paddle traces as measured parallel to the
midline. In contrast, the alternating A. agulhasii paddle traces
are very regular in size and shape.
Ichnological observations and interpretations
In order to place the A. agulhasii trails (trackways), as well as
the M. latus trackway, in their paleoenvironmental context, it
is important to note that in the case of A. agulhasii there are
multiple parallel trackways on a single block (∼5.6 x 3.0 m:
∼17.0 m2), which we interpret as a beach deposit (see Discus-
sion). Five of these trackways are marked with arrows
(Fig. 2), two are shown in Figure 3A, one (the holotype) is
shown as a 3D image (Fig. 4), and four are illustrated with
line drawings (Fig. 5). Distinctive features of these trackways
are their continuous and subsymmetrical configurations and
the very regular serial pattern created by the paddle marks
(Figs. 3A, 4, and 5). However, as noted in the description,
trackway segments where only lateral paddle traces were reg-
istered are, in places, followed by segments where a median
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M. G. Lockley et al.
ridge occurs. This suggests either (a) that the trackmaker car-
ried the animal’s central carapace clear of the substrate (with
an elevated posture) for a certain distance before lowering to
the point where the carapace left a central groove, or (b) that
substrate conditions changed subtly as the trackmaker pro-
gressed. The central ridge appears at approximately the
same place in the three parallel trackways illustrated in Fig-
ure 5A, which potentially suggests the latter interpretation
(b) is the more parsimonious.
Given this latter possibility, differences in the registration
of traces might reflect the passing of the trackmakers across
substrate areas with different physical properties (consolida-
tion, moisture content, etc.), as would be expected across a
beach, especially perpendicular to the shoreline. In this
regard, we note that the trails occur as natural impressions
on the once upward-facing surface of a large fallen block
from which the matching surface, the “counterpart” natural
casts, are lost. The block is several meters thick, and the
lower portion is composed of planar bedded sandstone of
the Klein Brak Formation. This bedding style indicates that
the block represents beach deposits rather than dune deposits
of the Waenhuiskrans Formation, which elsewhere in the for-
mation are represented by cross-bedded facies. This interpre-
tation is also consistent with the occurrence of sea turtle
hatchling trackways.
Given the subparallel trackway pattern that, as noted previ-
ously, we infer to be shore-perpendicular, it is possible to sug-
gest that the tracks were not made in, or radiating out from, the
immediate vicinity of a nest, likely in dry sand above the high
tide mark. Rather they likely represent down shore movement
from some point near the previous high-tide mark, with sand
wet enough to leave well-defined tracks. There is no unam-
biguous evidence to reveal the mechanisms that allowed
tracks to be buried and preserved without being eroded or
washed out. However, a very plausible mechanism would
be the blowing or drifting of sand over the wet surface without
the power to cause deflation. If this was the preservation
mechanism, it is possible that the covering layer of blown
sand was more susceptible to postconsolidation and postex-
humation erosion, perhaps because of being drier and less
prone to cementation. Such a scenario would help explain
why the impression surfaces are preserved and as yet no infill-
ing cast surfaces have been found—that is, the lack of casts
could be because of the preferential weathering away of the
infilling layer to more readily expose the more resistant sur-
face with the natural trackway impressions.
The Marinerichnus latus trackway is not associated with
any other recognizable tetrapod traces, although some inver-
tebrate traces are evident (Fig. 6B). The trackway appears
“suddenly” relative to the bedding plane surface of the
block. It is tempting to suggest that the trackmaker was
emerging from the sand, possibly its nest, but as noted previ-
ously, the apparent sudden appearance may represent the pas-
sage of the trackmaker from one layer of sand to another more
suitable for registering traces. Thus, the emergence possibil-
ity, although not entirely ruled out, is speculative. However,
the discovery of two sea turtle hatchling trackway
 New fossil sea turtle trackway morphotypes from the Pleistocene of South Africa
morphotypes (representing at least six individuals), within a
geographic area of paleoshoreline less than 5 km wide, sug-
gests that more sea turtle tracks and traces, including nests,
could be found. This inference is supported by the aforemen-
tioned suggestion that the site 3 specimen (Fig. 8) might rep-
resent a nest, which, as an infaunal trace, potentially has
higher if more localized preservation potential than epifaunal
surface trackways. As illustrated in Fig. 9, the shore perpen-
dicular zones of nest excavation and trackway registration
are quite localized on the scale of tens of meters. Tetrapod
nests and nesting grounds are not uncommon in the fossil
record at least as early as the Mesozoic (Carpenter et al.,
1994; Carpenter, 1999).
DISCUSSION
Neoichnology of sea turtle tracks
There seems little doubt that the A. agulhasii and M. latus
trackways are unequivocally attributable to hatchling sea tur-
tles, moving over exposed surfaces (i.e., creating epifaunal
trails). Prior to making this determination, we considered
the need to eliminate the possibility that the trackways
might conceivably represent infaunal sand-swimming traces
of a genus like Eremitalpa from the fossorial golden mole
family (Chrysochloridae) (Stuart and Stuart, 2000). However,
such traces are not compatible with the features seen here.
Less likely possibilities, including crawling penguins during
times of preterminal distress leaving tramline impressions
with their wings, are incompatible with the evidence reported
here of seven similar subparallel trackways, all too small to
permit the unlikely conjecture that they could be attributable
to distressed, beached birds. Other “tramline trackmakers”
such as crawling, rather than hopping anurans (frogs or
toads) or invertebrates such as scorpions or crabs, known to
inhabit beach and sand dune settings, are all easily discounted
because of their known trackway configurations being quite
different in size and morphology (Häntzschel, 1975, and mul-
tiple references to invertebrate trails therein). Likewise we can
discount the clawed tracks of other reptiles such as terrestrial
tortoises, terrapins, and crocodylians, which again leave dis-
tinctive trackways described from both modern and ancient
sediments (e.g., Milàn et al., 2010).
The formality of excluding the aforementioned list of
potential trackmakers for A. agulhasii and M. latus leaves
the sea turtle hatchling interpretation as the only realistic pos-
sibility. This is strongly supported by several compelling
lines of evidence. First, the tracks are morphologically highly
similar to those of extant sea turtles. Second, they occur in
coastal deposits, which represent beach environments sea-
ward of sand dunes. Third, the presence of seven parallel to
subparallel trackways, in an area less than 6.0 m wide, indi-
cates a large number of individuals moving in the same direc-
tion in a small area, evidently at the same time. Fourth, the
similarity in trackway configuration suggests they moved at
the same speeds. This is the behavior expected of hatchling
sea turtles heading across a beach toward the sea. The fact
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637
that we have two morphologically diagnostic morphotypes
that are compellingly attributed to loggerheads and leather-
backs not only supports the sea turtle interpretation generally,
but also points to the possibility of deriving paleoecological
and paleogeographic interpretations pertaining to two major
sea turtle groups.
Online images of sea turtle hatchling trackways and videos
of baby turtles scrambling to the sea after hatching are abun-
dant and easily found but pertain to a wide range of geo-
graphic localities. These include various images that show
parallel tracks and trackways that change in appearance as
the substrate the trackmakers cross also changes in consis-
tency (moisture content, firmness of sand) in a seaward direc-
tion. Many of the images show a great variety of turtle
trackway configurations including patterns similar to those
recorded here. For example, a trackway from Turtle Sanctuary
Beach, Palau Perhentian Besar, Malaysia, is practically iden-
tical to those recorded here (Fig. 9). The variation seen in
modern sea turtle hatchling trackways can be attributed to dif-
ferences in species morphology such as relative size of larger
anterior paddles and smaller hind feet.
All trackways generally share the common characteristic of
having been made on sandy substrates with anterior paddle
traces on the lateral margins. However, although the central
portions of some trackways show traces of body or carapace
drag traces, others do not, but instead show the traces of the
hind limbs or other indications that the carapace did not
touch the substrate. These differences may also be attributed
to the speed of progression of individuals, which in turn may
be influenced by substrate consistency.
Paleoichnology of sea turtle tracks
To the best of our knowledge, there are no previous reports of
sea turtle hatchling tracks from the fossil record. The only
reports of sea turtle tracks come from Jurassic limestones
from the Cerin region of France. These fall into the category
of swimming traces attributed to a large marine turtle (Thul-
born, 1989, 1990; Lockley and Meyer, 2000; Gaillard et al.,
2003; Fig. 10). These trackways were initially, and controver-
sially, interpreted as evidence of hopping dinosaurs (Bernier
et al., 1984). It was argued that the symmetrical configuration
of footprints, each with three-digit traces, supported such
inferences. So, the tracks were named Saltosauropus latus,
implying a hopping or jumping reptile (Fig. 10). These paired
tracks were subsequently reinterpreted as the result of syn-
chronous registration of the paddles of a large marine turtle
(Thulborn, 1989, 1990) swimming in shallow water over a
calcareous mud substrate. Thulborn’s interpretation was sub-
stantiated by further discoveries of swimming sea turtle tracks
by Gaillard et al. (2003), which they claimed represented the
oldest evidence of marine turtles. These Jurassic trackways
are quite different from those described here from the Pleisto-
cene Klein Brak Formation, in all relevant ichnological
details. First, the trackways are much larger, up to 1.5 m
wide. Second, they represent swimming behavior and show
only paddle or flipper marks with no central body trace.
 638
Third, they occur in very fine-grained calcareous and suba-
queously deposited substrates.
As noted previously, prior to the incorrect hopping dino-
saur interpretation proposed by Bernier et al. (1984), Bernier
et al. (1982) had described a turtle (chelonian) trackway they
formally named Chelonichnium cerinense, using the ichno-
genus name Chelonichnium originally proposed by Schimper
(1850) (see Avanzini et al., 2005, for discussion). Bernier
et al. (1982, p. 447) interpreted the C. cerinensis trackway
(Fig. 10A) not as that of a sea turtle but as a “terrestrial” che-
lonian walking over a “very wet, superficially soft, and slop-
ing” substrate interpreted as a carbonate “platform” behind a
ribbon of islands. They also considered that the animal was
progressing down a decreasingly steep slope and over an
increasingly wet substrate. Despite the suggestion that this
was a terrestrial turtle (? tortoise) the slope interpretation
begs the question: Was this in fact the trackway of a marine
turtle going to sea? Gaillard et al. (2003) hint at this possibil-
ity without explicitly reinterpreting the Chelonichnium track-
way. They note that large marine turtles likely came into very
shallow water and may have nested in the area.
Literature on sea turtle locomotion discusses the develop-
ment of the hypertrophied, hydrofoil- or wing-like front
limbs (paddles or flippers) and the modified coordination
of the four limbs as part of the secondary adaptation of land-
based tortoises (chelonians) to aquatic marine and freshwater
environments (Davenport et al., 1984; Renous et al., 1989).
Of interest here are the differences between the alternating
(left-right) gaits of walkers on land, the perhaps alternating
strokes made by some freshwater swimmers, and the synchro-
nous strokes made by large marine turtles when swimming
(and also on beaches), as inferred for the large Jurassic
swim trackways (Thulborn, 1989, 1990; Lockley and
Meyer, 2000; Gaillard et al., 2003). Smaller modern sea tur-
tles may swim with alternating strokes, but given that all fossil
trackways attributable to marine turtles are Jurassic in age,
conjectures on the swimming styles of all fossil turtles are
necessarily speculative.
Paleoenvironmental implications
The Pleistocene beaches represented by the late Pleistocene
Klein Brak Formation sediments would have provided suit-
able nesting sites for sea turtles. Basic requirements for suc-
cessful nesting incubation of extant sea turtles are as
follows: beach sand above the high-tide mark that is deep
enough (50–80 cm) to allow for burial of eggs; sufficient
sand moisture so that the leathery eggs do not dry out; sand
that is not so wet as to drown eggs by prohibiting gas
exchange; temperatures between 25°C and 35°C to allow
for successful incubation. Sand temperatures below the piv-
otal temperature of ∼29.3°C generate mostly male hatchlings,
and above this pivotal temperature mostly females (Maxwell
et al., 1988; Davenport, 1997). Compared with other extant
sea turtles, loggerhead and leatherback turtles are able to
nest successfully in subtropical, rather than tropical regions.
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M. G. Lockley et al.
In southern Africa, these conditions exist most consistently
more than 1200 km to the northeast along the beaches of
northern KwaZulu-Natal, as a result of the Agulhas Current
generating warm conditions along the shore for successful
incubation. The rookeries of loggerhead and leatherback tur-
tles on the KwaZulu-Natal coastline, in the iSimangaliso
Wetland Park World Heritage Site just south of the border
with Mozambique, represent the southernmost regular sea
turtle breeding sites in the world (see St. Lucia area: Fig. 1
inset). However, occasionally nests of either loggerhead or
leatherbacks are reported alongshore between St. Lucia to
Cape Town, either at the beginning (October) or end (January
or February) of the nesting season, suggesting that female tur-
tles had an early or late clutch still to lay on the migration
between the foraging and nesting grounds. Of these nests,
one near Alexandria (500 km east of the Pleistocene site
described here) was reported to have incubated successfully.
Depending on the seasonal temperature and assuming the
clutch is not deposited in a dune shade or under vegetation,
with 55–80 days of warm weather available (i.e., the incuba-
tion period for turtle nests), occasional turtle nests could con-
ceivably incubate along most parts of the coast east of Cape
Agulhas. However, this assumes near-ideal incubation
conditions.
We are aware of two anecdotal reports of recent nesting tur-
tles (Dermochelys) on the Cape south coast at Still Bay in
2005 (du Plessis, J., personal communication, 2018) and at
Buffelsbaai in 2007 (Combrink, X., Baker, N., personal com-
munication, 2018), as well as reports and photos of tracks and
clutch of eggs from Nautilus Bay in 2011 (Scholtz 2011;
Heyns, K., personal communication, 2018) and a report of
a leatherback turtle at Kleinemonde in January 2018.
Although these reports indicated failed nests, they are rele-
vant to the evidence for fossil nesting just east of Still Bay,
because they indicate the overlap, at least marginally, if not
extensively with the paleogeographic and geographic regions
where extant sea turtle nesting occurs. The A. agulhasii track-
ways reported here appear consistent with the alternating gait
tracks of extant loggerhead hatchlings (Caretta caretta) (per-
sonal observation by RN; see Fig. 9). The occurrence of M.
latus trackways representing hatchlings, in the same region,
confirms that leatherbacks also nested here in the Pleistocene.
MIS 5e was characterized by warmer global temperatures,
with sea levels 5–6 m higher than present levels. These
warmer temperatures could conceivably have allowed turtles
to nest on the Cape south coast. The existence of both logger-
head and leatherback hatchling turtle trackways within a 5 km
stretch of coastline on the Cape south coast suggests a similar
phenomenon to that which currently occurs in the iSimanga-
liso Wetland Park World Heritage Site near St. Lucia (Figs. 1
and 9), with the two species nesting in proximity to one
another.
CONCLUSIONS
Fossil trackways of hatchling sea turtles have never previ-
ously been reported from the ancient track record. The
 New fossil sea turtle trackway morphotypes from the Pleistocene of South Africa
multiple trackways reported here from the Klein Brak Forma-
tion and named Australochelichnus agulhasii ichnogen. and
ichnosp. nov. are therefore a novel and highly distinctive
addition to the tetrapod paleoichnological record. The track-
ways are unequivocally diagnostic of hatchling loggerhead
turtles, which left traces that are morphologically quite dis-
tinct from formally named fossil trackways of large adult
sea turtles, currently known only from the Jurassic of the
Northern Hemisphere. The occurrence of multiple subparallel
trackways, all representing similarly sized hatchlings, is also
of behavioral significance in indicating that they were made
postemergence from their nests, during a downshore “run
for the sea.” The finding of a trackway (Marinerichnus
latus) consistent with those of hatchling leatherback turtles,
less than 5 km from the loggerhead turtle tracksite, appears
to duplicate the current nesting situation in a subtropical
coastal environment farther to the northeast, where the two
species breed in proximity to one another. Their occurrence
in Pleistocene deposits of the Cape south coast, likely dated
from the MIS 5 stratigraphic interval, is the first direct evi-
dence of the antiquity of sea turtle nesting in this region
and is consistent with an extension of the present-day range
of successful sea turtle nesting sites. The occurrence of a
third site with an Australochelichnus-like trackway associated
with a possible nest trace serves to underscore the importance
of this area for the study of Pleistocene sea turtle paleobiol-
ogy, also suggesting that their traces may be more common
than previously supposed. As such traces of hatchling sea tur-
tle trackways have both paleobiological and paleogeographic
significance and utility in comparison with the traces of their
extant descendants.
ACKNOWLEDGMENTS
We acknowledge the support of Xander Combrink, Guy Gardner,
Linda Helm, Kei Heyns, Johan Huisamen, Christina Mars, Gerrit
Mars, Peter Todd, and Boy van Rensburg. We thank Andre and
Emily Brink for finding and reporting the trackway slab herein
referred to as site 3 (Garden Route National Park area). Mark
Dixon also reported a fourth turtle tracksite quite close to sites 1
and 2, for which he provided supporting photographic evidence.
However, these tracks are at a difficult-to-access location and cannot
be described here in detail without further field investigations. Linda
Harris and Diane Le Gouvello kindly provided the photographs used
in Figure 9. The photogrammetric image in Figure 6C was processed
and provided courtesy of Carina Helm. The three-dimensional print
of the A. agulhasii holotype (MGL 600/UCM 230.281) was made
with the help of Luis Sanchez Vega (University of Colorado Den-
ver), Neffra Matthews, National Operations Center (Denver, Colo-
rado), Dinosaur Designs LLC (Golden, Colorado), and the offices
of the University of Colorado Denver Dinosaur Trackers Research
Group.
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