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THE ECOLOGY OF MEIOLANIA PLATYCEPS, A PLEISTOCENE TURTLE FROM

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 Lucas, S.G. and Sullivan, R.M., eds., 2018, Fossil Record 6. New Mexico Museum of Natural History and Science Bulletin 79.

363
THE ECOLOGY OF MEIOLANIA PLATYCEPS,
A PLEISTOCENE TURTLE FROM AUSTRALIA

ASHER J. LICHTIG and SPENCER G. LUCAS

New Mexico Museum of Natural History, 1801 Mountain Road NW, Albuquerque, New Mexico 87104; -email: ajlichtig@gmail.com

Abstract—The Australian Pleistocene turtle Meiolania platyceps was subjected to quantitative methods
of inference of paleohabitus. These are based on the shell, forelimb, and femur morphology. All methods
indicate that Meiolania was aquatic. This is further supported by the presence of a plastral fontanelle,
known only from one extant terrestrial turtle– Malacochersus torieri– and no fossil turtles infered to
be terrestrial, but known in many fossil turtles inferred to be aquatic. Further corroboration comes from
the shape of the femoral head and the chelydrid-like distribution of armor. This analysis serves as an
important reminder that qualitative analogies are no substitute for quantitative analysis of paleohabitus.

INTRODUCTION on qualitative analogy rather than quantitative comparisons to

Meiolania platyceps (Fig. 1) is a large turtle from the
Pleistocene of Lord Howe Island off the coast of Australia (Sterli,
2015). Meiolanids are a very basal group of turtles that survived
almost to the present with extensive unique specializations that
render paleoecologic interpretation problematic. Sterli (2015)
recognizes three valid species of Meiolania, but this paper only
addresses the morphology and habitus of Meiolania platyceps.
This turtle has been inferred to be terrestrial in habitus, but its
habitus has never been analyzed by quantitative methods (Sterli,
2015). This paper seeks to assess the paleoecology of Meiolania
based on several quantitative methods.
HISTORY OF STUDY
The first meiolaniid fossils were described by Owen (1880)
and, at that time, referred to the large varanid lizard Megalania
prisca. Owen (1886) later revised this based on the material
discovered on Lord Howe Island, naming Meiolania platyceps.
Huxley (1887) was the first to argue that Meiolania is a cryptodire
turtle rather than a lizard. Shortly after this, Boulenger (1887)
suggested that Meiolania platyceps is a pleurodire. After this,
Baur (1889) concluded that it is a cryptodire, for the first time
listing synapomorphies linking the two: (1), descending process
of the prefrontals reach vomer; and (2), processus trochlearis
present. This was followed and expanded upon by Gaffney
(1979a, 1981, 1983, 1996; Gaffney et al., 2007), stating that
meiolaniids are on the stem of Cryptodira. Conversely, Joyce
(2007) and Sterli (2015) suggested that meiolaniids are stem to
both pleurodires and cryptodires. This difference of opinion was
greatly debated, but ultimately remains questionable. In either
case, meiolaniids are a very basal group of turtles that survived
almost to the present with extensive unique specializations.
The inferred ecology of Meiolania platyceps has largely
been based on inference from where it was found and qualitative
assessments of its morphology (Anderson, 1925, 1930; Sterli,
2015). The first assessment based on quantitative data was
presented by Lichtig and Lucas (2018), which found it to be
aquatic, counter to almost all other authors. A recent description
of the ecology of Meiolania platyceps and the reasoning for
such inferences was by Sterli (2015), which provided the
most detailed explanation of the reasoning of any author. She
listed several reasons to consider Meiolania a terrestrial turtle,
including the presence of limb and tail osteoderms, the shape of
the humerus, the presence of a tail “club,” and the shape of the
ungual phalanges. This is similar to the assertions of Anderson
(1930), who suggested the terrestrial nature of Meiolania based
on limb proportions, the structure of the phalanges and its
heavily armored condition. Both of these analyses were based
extant turtles.
Sterli (2015), citing Gaffney (1996), pointed out several
similarities between Meiolania platyceps and testudinids,
suggesting they indicated meiolanids were terrestrial. Several
of these supposed similarities are open to question, as Gaffney
(1996) stated that Meiolania platyceps is similar to testudinids
in these regards, but also to other turtles, including the aquatic
chelonioids, Chelydra and Glyptops. The features of similarity
include: the possession of short coracoids, a large angle between
the dorsal process of the scapula and the acromion, similarly
shaped olenacron and sigmoid notches, spherical femoral
heads, limb osteoderms, a reduced manus digital formula of
2-2-2-2-2, short phalanges and a vaulted carapace. Of these, the
short coracoid, large angle between the dorsal process of the
scapula and the acromion, and the spherical femoral heads were
noted by Gaffney (1996) to be present in aquatic turtles. Thus,
Gaffney (1996) makes no ecological assertions based on these
comparisons. Conversely, Sterli (2015) took these as strong
indications of the terrestrial habitus of Meiolania platyceps.
QUANTITATIVE METHODS
Previous analyses of the habitus of meiolaniids have been
qualitative. But, several quantitative methods of inference of
turtle paleohabitats have been used. Joyce and Gauthier (2004)
proposed a method based on the relative lengths of the humerus,
ulna and manus of a turtle. They concluded that shorter handed
turtles are more terrestrial. Benson et al. (2010) proposed a
method of paleoecological inference based on the curvature of
the carapace. Lichtig and Lucas (2018) introduced a method
of paleohabitus inference based on two ratios of the shell
dimensions. These are the length-to-height and the maximum
carapace-width-to-plastron-width at the abdominal-femoral
sulcus. Lichtig and Lucas (2018) concluded that turtles with
higher length-to-height ratios are more aquatic and turtles with
higher carapace-width-to-plastron-width measurements also are
more aquatic turtles.
Zug (1971) discussed several patterns of variation in the
femora of extant testudines. One is that the intertrochanteric
fossa of aquatic turtles tends to be deeper than that of terrestrial
turtles (Fig. 2). This was used to create the femur-based method
of Lichtig and Lucas (2018). In turtles that primarily walk, rather
than swim, including bottom walking, the femoral head tends to
be more circular than elongate.
Scheyer (2007) examined the histology of Meiolania sp.
and suggested this indicates that it was terrestrial. Scheyer
(2007) was not clear which species of Meiolania was studied,
so his observations may be related to one of the other species
364
FIGURE 1. Reconstruction of Meiolania platyceps, modified from the reconstruction of Gaffney (1996), in left lateral (bottom) and
ventral (top) views.
of Meiolania. The details of Scheyer’s (2007) reasoning are
not described, but the specimen lacks the internal cortex and
is superficially similar to some semi-aquatic turtles such as
Platysternon. So, lacking further evidence we consider this
assessment doubtful. In this paper, we attempt to use the
reconstructions of Gaffney (1996) and images provided by
Juliana Sterli to assess the paleohabitus of Meiolania platyceps.
METHODS
To assess the paleohabitus of Meiolania platyceps the
limb proportions described in Joyce and Gauthier (2004) were
measured from the reconstruction of Gaffney (1996). Working
from reconstructions is not ideal, and an inherent result of
this is the dependence of our results on the accuracy of the
used reconstructions. As a result we also address how far the
measurements can vary before the results would change. These
data are then plotted on the graph of Joyce and Gauthier (2004)
for comparison (Fig. 3). The same images are used to calculate
the shell ratios of Meiolania platyceps as described in Lichtig
and Lucas (2018). These are both plotted on the graph of Lichtig
and Lucas (2018) and entered into their equation (Fig. 4). In
addition, the method modified from the observations of Zug
(1971) in Lichtig and Lucas (2018) is used to infer its habitus
based on the morphology of the femur (Fig. 2). This is then
compared to living turtles to indicate the most similar extant
turtles.
RESULTS
The forelimbs of Meiolania platyceps are 34.6% humerus,
30.8% ulna, and 34.6% hand (Fig. 3). This coincides with the
aquatic range of Joyce and Gauthier (2004). The closest match
of Meiolania platyceps among the species included in Joyce and
Gauthier (2004) is Podocnemis sextuberculata, an aquatic turtle
from northern South America. To move into the terrestrial range
these values would need to change at least 10%, more than can
be explained by the angle of photography. Meiolania platyceps
is also similar in hand proportions to Macrochelys (Fig. 3) and
Platysternon. The femur-length-to-intertrochanteric-fossa-depth
ratio of Meiolania is 17.2; this is closest to the large, bottom-
walking turtle Macrochelys temminckii.
 365

FIGURE 2. Femur drawings of Zug (1971) showing the variation in the morphology of the proximal end of the femur that he linked
to their mode of locomotion and habitus. On right are Gaffney’s (1996) line drawing of the dorsal view of Meiolania platyceps
femur and below is a ventral view of the femur, MMF 13825 (photograph by Juliana Sterli). post. = posterior.
The shell ratios of Meiolania platyceps are carapace-width- aquatic, even though it was long considered a terrestrial turtle.
to-plastron-width 2.54, and carapace-length to total-height 2.00 Nevertheless, we consider this reasonable when reexamining
(Fig. 4). This carapace-width-to-plastron-width ratio is higher some of the arguments for the purported terrestrial habitus.
than in any extant terrestrial turtle by 0.72, or ~28% more 1. As suggested by Lichtig and Lucas (2018), the
(Lichtig and Lucas, 2018). This is a large difference and unlikely correlation of limb and tail osteoderms of turtles to a terrestrial
to be changed by any issues with the reconstruction. The length- habitus (Joyce, 2015; Sterli, 2015) has been overstated. The
to-height ratio is in the mid-range for either aquatic or terrestrial extant tortoises that bear the osteoderms this comparison is
turtles, not clearly providing an indication one way or another. based on have armor on the limbs and, in rare cases, only on
The discriminant function described in Lichtig and Lucas (2018) the dorsal side of the tail. This distribution of osteoderms results
results in a score for Meiolania platyceps of 1.73, well above the from a change in their function between defense while passive
dividing line between aquatic and terrestrial (Fig. 4). (limb osteoderms) and defense while mobile (tail osteoderms).
DISCUSSION Tortoise limb armor is used to defend soft body parts pulled
into the shell while immobilized. Chelydra’s spines provide
Our analysis indicates that Meiolania platyceps was defense while fully mobile and provide the benefits of a long,
366
FIGURE 3. Graph modified from Joyce and Gauthier (2004) showing the placement of Meiolania platyceps forelimb ratios relative
to the turtles examined by Joyce and Gauthier (2004).
hydrodynamically efficient tail while alleviating some of the
predation risk this imposes. The distribution of limb osteoderms
in Meiolania platyceps is unclear. But, given the large number
of specimens that show restriction of osteoderms to the manus
and pes, we think an extensive covering of limb osteoderms is
unlikely. Furthermore, the porous, lightly built structure of these
osteoderms is not what would be expected in armor. Another
use of osteoderms in reptiles is as nutrient storage (Klein et al.,
2009), which we consider a more probable function of these
limb osteoderms in Meiolania platyceps.
2. The presence of a tail club has been a recurring factor
in suggestions that fossil turtles are terrestrial (e.g., Joyce, 2015;
Sterli, 2015). The assumption is that this structure is swung like
a flail at an enemy, as has been suggested for ankylosaurian
dinosaurs (Arbour and Zanno, 2018). This assumes spiny
protrusions of the tail, such as those in Chelydra serpentina,
which are made of keratin, are a separate issue, though these
would function in the same manner if made of bone. However,
such swinging would not work in the water, as the resistance
of the water would prevent the accumulation of momentum
before impact. Nevertheless, the notion that these “clubs” in
turtles would be swung into an adversary is questionable. This
is because the spines present on these tails would impede the
“club” being swung into anything because they would hit an
object first, softening the blow. We suggest, as did Lichtig and
Lucas (2018), that these “clubs” are better interpreted as passive
defense (armor) to prevent injury to the posterior of the tail,
which could not be retracted and was thus vulnerable to attack.
3. The shape of the ungual phalanges was claimed
to be similar to those of testudinids to argue that Meiolania
platyceps is a terrestrial turtle. This does not appear to be the
case, as testudinids usually have blunt, very rounded unguals
(Hay, 1908), but those of Meiolania platyceps come to a well-
developed point, giving them a more elongate shape similar to
Chelydra. The dorsal side of the unguals of Meiolania platyceps
bears a flexor tubercle that would be associated with walking,
either terrestrially or as a subaqueous bottom walker.
4. The short manus formula with only two phalanges
per digit has also been suggested to indicate that Meiolania
platyceps was terrestrial. Conversely, the ungual phalanges of
Meiolania platyceps are more elongate than in extant testudinids
(Crumly and Sánchez-Villagra, 2004), resulting in the relatively
long hand noted above. This results in a hand similar in length-
to-width ratio to Trachemys but with thicker digits reducing
space for intra-digital webbing. This may have limited its fully
buoyant mobility, but if it is a bottom walker like Chelydra
serpentina this would have little impact as it would rarely be
fully buoyant. Instead it would be slightly negatively buoyant,
walking against the substrate much like a terrestrial animal. This
elongation of the ultimate phalanges, rather than an increased
phalangeal count, may suggest that this is an aquatic animal
derived from a terrestrial animal that had lost these additional
phalanges, present in basal turtles, earlier in evolution. In other
words, Meiolania has a long hand made out of fewer bones than
extant, long-handed turtles.
5. The heads of the humeri of Meiolania platyceps and
testudinids are similar in their morphology. They have the
rounded shape, also seen in Chrysemys and Apalone. But,
the general form of the humerus more closely corresponds to
that of the bottom-walking turtles Chelydra and Kinosternon
 367

FIGURE 4. Graph of the ratios of carapace width to plastron width on the horizontal axis and shell length to height on the vertical
axis based on 250 measured specimens in 94 modern species (from Lichtig and Lucas, 2017). Each dot on the graph represents a
specimen measured; the dot for Meiolania is labelled to the right. Outlines are minimum convex polygons for aquatic and terrestrial
turtles. Shaded area indicates overlap of these polygons. As in Lichtig and Lucas (2017), Homopus boulengeri and Malacochersus
tornieri were excluded from the terrestrial minimum convex polygon based on their unusual morphology. Line indicates the
discriminant function resulting from the analysis.
(Olsen, 1968). These are similar in their more robust shaft than Finally, some features of Meiolania platyceps not discussed
in Gopherus, or Chrysemys, and in the increased size of the previously in reference to paleohabitus, are suggestive of
anterior process of the humerus. an aquatic habitus. The most visually obvious of these is the
6. The morphology of the pectoral girdle of Meiolania plastral fontanelle in the adult Meiolania platyceps (Fig. 1),
platyceps was suggested to indicate terrestrial behavior (Sterli, which is only otherwise seen in adult sinemydids, Otwayemys
2015). Conversely, Gaffney (1996, p. 39) compared it to cunicularius, chelonioids, trionychids, thalassochelydians, and
testudinids and the paracryptodire Glyptops, a taxon generally platychelyids, universally suggested to be of aquatic habitus
inferred to be aquatic (Gaffney, 1979b). However, the presence (Brinkman, 2001) and Malacochersis tornieri, an extant
of similar pectoral morphology in both aquatic and terrestrial terrestrial turtle specialized for squeezing between rocks with
turtles excludes its use as a means of inference of paleohabitus. its thin, flexible shell (Mautner et al., 2017). Furthermore, in a
The large angle between the two processes of the scapula in terrestrial turtle a plastral fontanelle would be a vulnerability to
Meiolania platyceps was also suggested to be indicative of a predators, particularly if overturned. In addition, the relatively
terrestrial habitus (Sterli, 2015), but, as Gaffney (1996) pointed long tail of Meiolania platyceps is unlike that of any living
out, this is also seen in the bottom-walking chelydroids. Thus, it terrestrial turtle. Terrestrial turtles generally have a short tail
is not an indicator of terrestrial habitus. that can be sheltered by the carapace. Long, exposed tails
7. Spherical (or more accurately subspherical) femoral are generally seen in aquatic forms, including Chelydra spp.,
heads suggested by Sterli (2015), in her comparisons with Platysternon megacephalum, and Mauremys spp.
testudinids, to be indicative of terrestrial habitus were addressed This brings up an important question: if Meiolania platyceps
at length by Zug (1971), who she did not cite. Zug (1971) was a large aquatic herbivore, where was it living on the small
concluded that this was related to walking either terrestrially or Lord Howe Island? This is confounded by the fact that most
aquatic bottom walking, as in chelydrids (Fig. 2). Furthermore, existing streams on the island are low lying and would have
both Chelydra and Meiolania platyceps have the slightly less been under water at the time the fossils were buried (Woodroffe
perfectly circular shape of the head of the femur that Zug (1971) et al., 1995). As such, the sub-areal nearshore localities where
associated with bottom-walking turtles. Meiolania platyceps are excavated today were subaqueous when
8. It was long stated that Meiolania has a carapace the animals were living. This may indicate that Anderson’s (1925)
that is particularly high domed (Fig. 1), but it can be seen by interpretation that these turtles were nearshore marine in habitus
quantitative comparison with other turtles in our sample that it may not be so far from the truth. Meiolania platyceps likely took
is only modestly domed (Fig. 4). Thus, the “vaulted” carapace advantage of the shallow lagoons off either side of Lord Howe
of Meiolania platyceps (Gaffney, 1996; Sterli, 2015) can be Island, much as Amblyrhynchus cristatus (Marine Iguanas) do
seen from the length-to-height ratio (2.00) of this turtle to be today in the Galapagos islands. This may also explain the cause
not especially high domed or vaulted (Figs. 1, 4). Instead, it is of their abrupt extinction as the falling sea levels in more recent
average, not giving an indication of habitus one way or another. times would have caused a shrinking of these offshore lagoons
368
and the food stocks they provided. Conversely, if these animals
were fully terrestrial at this time their habitat would have likely
been expanding as the sea levels dropped and more land became
available. Also of note is the nasomaxillary sinus identified
by Gaffney (1983). This is a large sinus adjacent to the nasal
passages. We speculate that this may have held a salt excreting
organ as seen in other marine reptiles where the salt could be
excreted into the nasal effluent and disposed of.
In conclusion, three methods of inference based on three
separate anatomical regions indicate Meiolania platyceps was
aquatic. Furthermore, these analyses found Meiolania platyceps
close to the chelydrids, which suggest that these animals share
a similar ecology.
This analysis serves as a reminder of the importance of
quantitative assessments. For more than 100 years, the only
inferences on Meiolania platyceps’s habitus were based
on where it was found, a general sense of the doming of the
carapace, or vague qualitative comparisons to testudinids.
To accurately assess the shape and paleohabitus of turtles,
quantitative measurements are necessary (Joyce and Gauthier,
2004; Lichtig and Lucas, 2018).
CONCLUSIONS
1. Meiolania platyceps was an aquatic turtle.
2. Meiolania platyceps was likely a bottom walker, as is the
modern Chelydra.
3. Quantitative analysis of turtle paleohabitus is important, as
simple, qualitative observation can lead to false conclusions.
ACKNOWLEDGMENTS
We thank Juliana Sterli for providing images of the femur
of Meiolania platyceps. We thank Tomasz Szczygielski and Ren
Hirayama for their thorough and thought-provoking reviews.
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