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A review of Australasian ichthyosaurs

Maria Zammit

School of Earth and Environmental Sciences, North Terrace

Campus, University of Adelaide, Adelaide, South Australia, 5005,
Published online: 01 Jun 2010.

To cite this article: Maria Zammit (2010) A review of Australasian ichthyosaurs, Alcheringa: An
Australasian Journal of Palaeontology, 34:3, 281-292, DOI: 10.1080/03115511003663939
To link to this article:


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A review of Australasian ichthyosaurs


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Zammit, M., September, 2010. A review of Australasian ichthyosaurs. Alcheringa 34, 281292. ISSN 0311-5518.
Ichthyosaur fossils have been recorded from four landmasses in the Australasian regionAustralia, New Zealand,
New Caledonia and Timorand occur in all three systems of the Mesozoic. Most of the remains are non-diagnostic,
but at least three genera have been identied: Mixosaurus, from the Middle Triassic of Timor; Shonisaurus, from the
Upper Triassic of New Caledonia; and Platypterygius, from the Lower Cretaceous of Australia and New Zealand. Of
these, Platypterygius contains the only material that can be diagnosed to species level. However, current taxonomy of
the specimens is controversial, with two synonyms, P. australis and P. longmani, persisting in the literature. An
examination of cranial traits in the quasi-holotype of P. australis vs P. longmani demonstrates that they represent
the same taxon. Thus, P. longmani should be regarded as the junior synonym. A neotype is also here designated for
P. australis to replace the original, which is presumed lost.
Maria Zammit [], School of Earth and Environmental Sciences, North Terrace
Campus, University of Adelaide, Adelaide, South Australia 5005, Australia. Received 14.12.2009, revised 21.1.2010,
accepted 26.1.2010.
Key words: Ichthyosauria, New Zealand, New Caledonia, Timor, Australia, high-latitude, Mesozoic.

ICHTHYOSAURS were a group of extinct,

dolphin-like marine reptiles that ranged
from the Early Triassic (Spathian) until
the Late Cretaceous (Cenomanian: McGowan & Motani 2003). In Australasia, their
remains span most of this range (Fig. 1),
with material from the Triassic of New
Zealand, Timor and New Caledonia (Callaway & Massare 1989), Lower Jurassic of
New Zealand (Sachs & Grant-Mackie
2003), and LowerUpper Cretaceous of
Australia (Kear 2003) and New Zealand
(Fleming et al. 1971).
The earliest published accounts of these
ichthyosaurs come from Australia (McCoy
1867a) and New Zealand (Haast 1871,
Hector 1874), with later records from Timor
(Broili 1931) and New Caledonia (Campbell
1984). Much of the described material
can not be condently attributed to a family
or genus (Fleming et al. 1971). However,
ISSN 0311-5518 (print)/ISSN 1752-0754 (online)
2010 Association of Australasian Palaeontologists
DOI: 10.1080/03115511003663939

remains from the Cretaceous of Australia

have diagnostic features, although their
taxonomic status is still controversial (Wade
1990, McGowan & Motani 2003, Kear
This review has two primary aims: (1) to
summarize the record of ichthyosaur fossils
from the Australasian region, including
previously unpublished occurrences; and
(2) to discuss and resolve the taxonomy of
the Australian Cretaceous material.
Institutional abbreviations: AM, Australian
Museum, Sydney, Australia; GS, Geological Survey, Wellington, New Zealand; MV,
Museum Victoria, Melbourne, Australia;
WAM, Western Australian Museum, Perth,

Regional setting
Eastern Australia, New Zealand and New
Caledonia were situated in the southeast
corner of Gondwana throughout the early
Mesozoic (Audley-Charles 1978, Fleming

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Fig. 1. A, Map of ichthyosaur fossil localities in Australia, modied from Kear (2003). B, Map of ichthyosaur fossil
localities in New Zealand, modied from Fordyce (1982). C, Map of ichthyosaur fossil localities in New Caledonia,
modied from Campbell (1979). D (inset), Scale map of Australia, New Caledonia, New Zealand, and Timor
showing relative size and position of the landmasses. Symbols: triangle, Triassic specimen; square, Jurassic specimen;
circle, Cretaceous specimen.

1979) and thus experienced a convergent

tectonic regime until the split of the
supercontinent. New Zealand and New
Caledonia are composite landmasses that
assembled from disparate terranes during
this time (Aitchison et al. 1995, Landis et al.
1999) but have elements of a common
(convergent margin) geological history

(Laird & Bradshaw 2004). The two regions

may have retained connections until the
opening of the New Caledonia Basin in the
Late Cretaceous (Schellart et al. 2006), postdating the initiation of separation from
Australia (Aitchison et al. 1995). Rifting
associated with the anticlockwise break-up
of eastern Gondwana initiated in the

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Jurassic of northwestern Australia (Veevers

et al. 1991) and resulted in separation of
New Zealand from Australia by the early
Cenozoic (Schellart et al. 2006; but see
Laird & Bradshaw 2004 for an alternative
view). In contrast, little separation occurred between Timor and Australia
throughout the Mesozoic since the formation of Timor on the passive Tethyan
margin of Gondwana (Audley-Charles
1978). Large areas of western, southern,
and eastern Australia were ooded by
shallow epicontinental seas in the midCretaceous (Frakes et al. 1987), and the
region has occupied mid- to high-latitude
settings throughout much of the Mesozoic
(Embleton 1984).

Review of Australasian
ichthyosaur taxa
Triassic specimens
New Zealand has most Triassic specimens,
with isolated ichthyosaur bones occurring
on both the North and South islands (Fig.
1B). The rst reported nds originated
from the Mount Potts region near Canterbury, South Island (Hector 1874, 1879),
and have since been referred to the Middle
Triassic (Table 1) Daonella Zone (lower
Carnian; Campbell & Warren 1965). These
remains comprise indeterminate vertebral
centra of extremely large dimensions
(457 mm in diameter, Hector 1878) that
appear to represent one of the largest
ichthyosaurs in the world (Campbell 1965,
Fleming et al. 1971). This is almost double
the diameter of centra known from Shonisaurus sikanniensis, an animal well known
as a giant (Nicholls & Manabe 2004).
Unfortunately, these particular specimens
can not be located, and are presumed to
have been either sent to the US with other
marine reptile bones that have now vanished (Cox 1991), or alternatively lost on


the transport ship Matoaka, which disappeared en route to London (Hector 1874,
Brazier et al. 1990). Hector (1874) diagnosed this material, together with several
smaller vertebral centra from Rocky
Gully, near Mount Potts, as Ichthyosaurus
australis (a name pre-occupied by an
Australian species). Replacement names
include I. hectori (Lydekker 1889, Chapman
1914) and I. pottsi (see Fleming et al. 1971),
though none is currently considered valid.
This material was assigned to the Mixosauridae by Campbell (1965), but was later
regarded as simply ichthyosaurian due to a
lack of diagnostic features (Fleming et al.
1971). Campbell (1965) assigned a partial
rostrum with teeth from the upper Carnian
(Triassic) Mandeville Sandstone (Fordyce
2003) of Otamita Stream, South Island, to
the Shastasauridae, but this was also later
considered to be a taxonomically unidentiable ichthyosaur (Fleming et al. 1971).
Further Triassic ichthyosaur specimens from
South Island, New Zealand, include undescribed partial ribs and a partial humerus
from the Kiritehere coast (latest Carnian to
early Norian; Sachs & Grant-Mackie 2003),
a series of vertebral centra from Roaring
Bay, South Otago (Carnian), and teeth and
a series of vertebral centra from Etal Creek,
Southland (Anisian; Fordyce 1991)the
latter possibly constitute the oldest known
ichthyosaur material from Australasia (Fordyce 1982). Putative ichthyosaurian teeth
are known from Nugget Point and the
Wairoa district, but they exhibit labyrinthodont characters (Hector 1878, 1879, Worley
1894) and may derive from amphibians
(Fordyce 1982).
Triassic (Norian) ichthyosaur remains
have also been reported from New
Caledonia (Mazin 1985) and Timor (Mazin
1983). The jaw fragments from the
Norian Ouamoui Formation (Callaway &
Massare 1989) of New Caledonia were
originally considered plesiosaurian (Campbell 1984), but have been referred to

early Aptianlate Albian10

late AlbianCenomanian19

Middle Triassic25
early Norian29
early Norian29

Queensland1,2,3,4 and New South

Wales9, Australia
New South Wales,
South Australia16

Western Australia17

Western Australia17

Western Australia20

Northern Territory,
New Zealand23
Mount Potts, New Zealand24

Kawhia, New Zealand27

New Caledonia28
New Caledonia28

Wallumbilla Formation

Doncaster Member
(Wallumbilla Formation)
Bulldog Shale

Birdrong Sandstone

Alinga Formation

Molecap Greensand

Darwin Formation

Makirikiri Formation
Unit uncertain

Arataura Formation
Ouamoui Formation
Lepredour Shellbeds
Aitutu Formation

P. australis7 (currently valid),

P. longmani3 (junior synonym)
P. australis7 (currently valid),
P. longmani3 (junior synonym)
P. australis7 (currently valid),
P. longmani3 (junior synonym)
P. australis7 (currently valid),
P. longmani3 (junior synonym)
P. australis7 (currently valid),
P. longmani3 (junior synonym)
P. australis7 (currently valid),
P. longmani3 (junior synonym)
P. australis7 (currently valid),
P. longmani3 (junior synonym)
P. australis7 (currently valid),
P. longmani3 (junior synonym)
P. australis7 (currently valid),
P. longmani3 (junior synonym)
Ichthyosaurus australis24 (nomen dubium),
I. hectori26, Ichthyosauria23
Mixosaurus timorensis31 (nomen dubium),
Mixosaurus sp.33

Taxonomic assignment

Molnar (1982), 2Wade (1984), 3Wade (1990), 4Kear (2002b), 5Moore et al. (1986), 6McMinn & Burger (1986), 7McCoy (1867a), 8Krieg & Rodgers
(1995), 9Etheridge (1904), 10Helby et al. (1987), 11Kear (2003), 12Ludbrook (1966), 13Johns (1968), 14Day (1969), 15Henderson et al. (2000), 16Alley &
Pledge (2000), 17Choo (1999), 18McLoughlin et al. (1995), 19Siverson (1999), 20Teichert & Matheson (1944), 21Kear (2002a), 22Henderson (1998),
Fleming et al. (1971), 24Hector (1874), 25Campbell & Warren (1965), 26Lydekker (1889), 27Sachs & Grant-Mackie (2003), 28Campbell (1984),
Callaway & Massare (1989), 30Mazin (1985), 31Broili (1931), 32Audley-Charles (1968), 33Mazin (1983).


Table 1. Distribution of ichthyosaur material in the Australasian region. See footnotes for source texts included. Current status of taxonomic
assignment in bold

late Aptian/Albian14,22


mid-late Albian8

Queensland, Australia1,2,3,4

late Albian


Allaru Mudstone


Queensland, Australia


Toolebuc Formation


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Shonisaurus spp. by Mazin (1985), based

on the crown ornamentation and morphology of the teeth. Long bones and vertebrae
have also been recorded from the lower
Norian Lepredour Shellbeds (Campbell
Eight vertebrae and a basioccipital
from the Middle Triassic (Callaway &
Massare 1989), possibly from the LadinianNorian (Table 1) shallow-marine
Aitutu Formation (Audley-Charles 1968),
Timor, were originally described as Mixosaurus timorensis (Broili 1931), but Mazin
(1983) considered the material too poor to
identify to a species and reassigned it to
Mixosaurus sp. Sander (1992) referred to a
Cymbospondylus specimen from Timor
consisting of ten anterior caudal centra.
This was apparently based on an incongruous report by von Huene (1936), which
identied only three centra (one a cervical)
as being broadly similar to those of both
Mixosaurus and Cymbospondylus. Because
of this inconsistency, and the possibility
that Sander (1992) might have mistakenly
referenced the nine caudal centra attributed to Mixosaurus by Broili (1931), Cymbospondylus is here considered not
condently recorded from the Australasian

Jurassic specimens
Only one putative Jurassic ichthyosaur
specimen is currently known from the
Australasian region. This comprises an
isolated centrum from Kawhia, New Zealand (Sachs & Grant-Mackie 2003), from the
HettangianSinemurian Arataura Formation (Sachs & Grant-Mackie 2003). It is
not yet formally described.

Cretaceous specimens
A rich record of Cretaceous ichthyosaurs is
known from New Zealand and Australia.
The New Zealand material comprises sev-


eral cervical centra from the upper Albian

(Cretaceous) Makirikiri Formation (Table
1) that can be identied as ichthyosaurian
but not diagnosed any further (Fleming
et al. 1971), and a partial rostrum that is
very similar to the Australian Platypterygius
species (Sachs & Grant-Mackie 2003). In
contrast, Australian Cretaceous ichthyosaurs are known from numerous deposits
(Table 1), including: the Toolebuc Formation, Allaru Mudstone and Wallumbilla
Formation of Queensland (Molnar 1982,
Wade 1984, 1990, Kear 2002b); Wallumbilla
Formation (Etheridge 1904) and, more
specically, the Doncaster Member (Kear
2005b) of New South Wales; Bulldog Shale
of South Australia (Alley & Pledge 2000,
Kear 2006); Birdrong Sandstone (Choo
1999), Alinga Formation (Choo 1999) and
Molecap Greensand (Teichert & Matheson
1944) of Western Australia; and the
Darwin Formation of the Northern Territory (Wade 1990, Kear 2002a). Kear (2003)
recently summarized Australian ichthyosaur remains, including diagnostic specimens. Hence, this study only reassesses the
taxonomic status of the material. All
Australian specimens are referred to the
cosmopolitan genus Platypterygius based
on 17 character states (McGowan &
Motani 2003, pp. 118 119). Much of this
material is referred to a single species,
although a humerus (WAM 94.7.3) from
the HauterivianBarremian of Western
Australia is more similar to the Eurasian
species, P. campylodon or P. kiprijano,
than the recognized Australian form (Choo

Non-ichthyosaurian remains
Two records of Australasian ichthyosaurs
are now considered non-ichthyosaurian. A
presumed ichthyosaur jaw from New
Zealand (Benham 1936) has since been
referred to a squaladontid cetacean (Camp
1942). The only proposed Triassic ichthyo-



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saur from Australia (Cosgri & Garbutt

1972) has not been studied or gured.
Its ichthyosaurian anity was questioned


by Callaway & Massare (1989), and

was omitted from Mazins (1986) review
of Triassic ichthyosaurs. Kear (2004)

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considered it to be a misidentied amphibian, so this specimen requires further study.

Neotype. MV P12989 (Fig. 2AC), designated herein; the original holotype comprising numerous centra is lost.

Systematics of the Australian


Type locality, formation and age. Flinders

River, north central Queensland; Allaru
Mudstone; Early Cretaceous (late Albian).

Subclass DIAPSIDA Osborn, 1903

Order ICHTHYOSAURIA de Blainville,
Platypterygius von Huene, 1922
Type species. Platypterygius platydactylus
(Broili, 1907).
Age and distribution. EarlyLate Cretaceous
of North America, South America, Europe,
Russia, Australia and New Zealand.
Platypterygius australis McCoy, 1867[a]
(Fig. 2AM)
1867 Ichthyosaurus australis McCoy, p. 356.
1888 Ichthyosaurus marathonensis Etheridge,
p. 408, pl. 1, gs 1 3.
1922 Myopterygius marathonensis von
Huene, pp. 96, 98.
1944 Myopterygius australis Teichert &
Matheson, p. 169, gs 1 3.
1972 Platypterygius australis McGowan,
p. 17, pls 3 4.
1990 Platypterygius longmani Wade, p. 120,
gs 1 6.

Diagnosis [following Kear (2005a)]. Large

ichthyosaur, around 7 m long. Maxilla with
extensive external exposure; forms the entire
ventral portion of both the anterior maxillary
foramen and the bony nasal aperture. Maxilla also has a minor internal contact with the
prefrontal via its posterodorsal surface.
Lacrimal does not contribute to the border
of the bony nasal aperture. Prefrontal with
minor internal contribution to the bony nasal
aperture. External naris subdivided with welldeveloped anterior foramen and one or more
foramina present (in nasal) posterodorsal to
external bony nasal opening. Parietal contributes to the facet for the paroccipital
process of the opisthotic on the supratemporal. Humerus bearing tapered crest-like dorsal
trochanter and three distinct distal facets for
articulation with the ulna, radius and an
anterior zeugopodial element. Fourth distal
facet sporadically present on humerus for
articulation with pisiform. Three preaxial
accessory digits and three postaxial accessory
digits present in forelimb with digital bifurcation occurring in the primary axis (digit IV).
Neural spines of at least neck and anterior
trunk vertebrae divided into anterior and
posterior peaks by an asymmetric V-shaped
apical notch. Caudal centra from tail stock
region may bear weakly developed hemal
arch facets.

Fig. 2. Platypterygius australis material collected from the Allaru Mudstone at Flinders River, Queensland. Neotype
material (AC), and vertebral material (DM) of P. australis. Specimens include: A, MV P12989 cranial element,
right lateral view; B, MV P12989 basioccipital (right) and atlasaxis complex, dorsal view; C, MV P12989
basioccipital (left) and atlasaxis complex, left lateral view; D, MV P12992, lateral view; E, MV P22657,
anteroposterior view; F, MV P22655, lateral view; G, MV P22653, lateral view; H, MV P22656 anteroposterior view;
I, MV P22658, lateral view; J, MV P22654, lateral view; K, MV P22659, anteroposterior view; L, MV P22660,
anteroposterior view; M, MV P22661, anteroposterior view. at-ax atlasaxis complex; bas basioccipital;
bna bony nasal aperture; lac lacrimal; max maxilla; na nasal; snf supernarial foramen. Scale bars 10 cm.

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Remarks. Virtually all Cretaceous ichthyosaur material in Australia is referrable to

Platypterygius (Kear 2003). Despite the
volume of material discovered in Australia, only specimens from Queensland can
be diagnosed to species. Ichthyosaur remains from Queensland lack taxonomic
dierences and have been referred provisionally to a single species (Kear 2003).
However, there is some confusion regarding nomenclature, with both specic
epithets australis (e.g. McGowan & Motani 2003) and longmani (Kear 2003, 2005a)
in use.
Australian ichthyosaurs were rst recorded in the 1860s (McCoy 1867a,b).
McCoy (1867a) erected Ichthyosaurus (now
Platypterygius) australis on numerous
centra (McCoy 1867a, p. 355), without
giving any indication that cranial material
was available (Wade 1984). Vertebral
characters, particularly for isolated centra,
(McGowan & Motani 2003). Furthermore,
the earliest descriptions lacked detail,
leading Etheridge (1888) to require casts
(AM F125427 and AM F135426) for
comparison. Later workers also had diculty identifying the type (Hell 2001,
Wade 1984). The holotype material could
not be identied by Wade (1985) in the
Museum Victoria collections and may be
lost, but she did note that quasi-holotype
specimens MV P12989 (Fig. 2AC), MV
P12992 (Fig. 2D), MV P22653 4 (Fig.
2G, J) and MV P22656 61 (Fig. 2E, H, I,
KM) were available (Wade 1984). MV
P22656 (Fig. 2H) comprises the only
centra Wade (1985) identied as having
the same measurements as those listed by
McCoy (1867a). The cranial remains, MV
P12989 and MV P12992, were not mentioned in the original description, though
they originated from the same locality
and formation, and were referred to
P. australis in a later publication by
McCoy (1869). Limb elements were also


described in this publication, but were

only illustrated later by Chapman (1914).
Additional species now referred to P.
australis include Ichthyosaurus marathonensis (Etheridge 1888) and Platypterygius
longmani (Wade 1990). The former was
erected on a partial snout (Etheridge
1888), probably out of a sense of frustration and despair at trying to compare this
material with McCoys (McGowan 1972,
p. 15). It was later considered a junior
(McGowan 1972). Platypterygius longmani
was established on an almost complete
skull, most of a vertebral column and
pectoral girdle, both humeri, and a partial
forelimb (QM F2453), and nominated as the
holotype of P. longmani, a replacement
species for P. australis, since the holotype
of P. australis was non-diagnostic and
presumed lost (Wade 1990). However, as
the original description of P. australis
satised Articles 11 and 12 of the ICZN
guidelines (McGowan & Motani 2003, Kear
2004), P. longmani has recently been proposed as invalid. Additionally, under ICZN
guideline 75.5, QM F2453 can not replace
the holotype of P. australis on the grounds
that it was non-diagnostic without an
ocial application to the ICZNa procedure that was not undertaken. Further, QM
F2453 does not originate from the same
locality or formation as the lost holotype
and, therefore, does not full ICZN guideline 75.3.6. Nevertheless, the original holotype of P. australis still can not be identied.
Thus, following ICZN Article 75.1, under
which a lost holotype and taxonomic
uncertainty are stated as justiable grounds
for establishing a neotype, the remains
collected from the original locality and
formation as the lost holotype were reexamined (satisfying Article 75.3.6). These
specimens were given the same eld number
as the holotype by the collectorbased on
correspondence quoted by Wade (1985,
p.137) the remains derive from the same

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locality, though collected one year later

(Wade 1990). MV P12989 (Fig. 2AB) in
particular, an incomplete skull and atlas
axis complex, has a combination of three
characters dierentiating it from other
ichthyosaur taxa (listed here to satisfy
ICZN Article 75.3.2): (1) reduced extracondylar area on the basioccipital (Fig. 2B); (2)
lacrimal not contributing to the border of
the bony nasal aperture (Fig. 2A); and (3)
presence of at least one foramina posterodorsal to the bony nasal aperture (Fig. 2A;
B. Kear, pers. comm. 2009). The rst
character listed is diagnostic of Platypterygius, whereas the other two simultaneously
dierentiate the Australian Platypterygius
species and demonstrate that P. longmani
and P. australis are the same taxon,
relegating P. longmani to a junior synonym.
MV P12989 is thus designated the
neotype of P. australis with the expressed
purpose of clarifying the taxonomic status
of the Australian Platypterygius species.
MV P12989 originates from the same
locality as the holotype of P. australis
(satisfying ICZN Article 75.3.6), and is held
in a recognized scientic institution (satisfying ICZN Article 75.3.7). Lastly, it is
entirely possible that MV P12989 is part of
the same individual designated as the
original holotype of P. australis (Wade
1984), and is thus consistent with ICZN
Article 75.3.5.

Many thanks to the curatorial sta and
museums for providing information and/or
access to specimens: Neville Hudson (Auckland University); David Pickering (Museum
Victoria); and Scott Hocknull (Queensland
Museum). Thanks also to Ben Kear for
photographing specimens. Ben Kear, Rachel
Norris and two reviewers are thanked for
their comments on the manuscript. Use of
information contained within the New
Zealand Fossil Record File is acknowledged.


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