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A review of Australasian ichthyosaursMaria Zammit
a
aSchool of Earth and Environmental Sciences, North Terrace
Campus, University of Adelaide, Adelaide, South Australia, 5005,Australia
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: http://dx.doi.org/10.1080/03115511003663939
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A review of Australasian ichthyosaurs
MARIA ZAMMIT
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 identified: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, Platypterygiuscontains 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 ofP. australis vs P. longmanidemonstrates that they represent
the same taxon. Thus, P. longmanishould 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 [[email protected]], 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: McGo-
wan & 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 (Call-
away & 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 confidently attributed to a family
or genus (Fleming et al. 1971). However,
remains from the Cretaceous of Australia
have diagnostic features, although their
taxonomic status is still controversial (Wade
1990, McGowan & Motani 2003, Kear
2005a).
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, Geologi-
cal Survey, Wellington, New Zealand; MV,
Museum Victoria, Melbourne, Australia;
WAM, Western Australian Museum, Perth,
Australia.
Regional setting
Eastern Australia, New Zealand and NewCaledonia were situated in the southeast
corner of Gondwana throughout the early
Mesozoic (Audley-Charles 1978, Fleming
ISSN 0311-5518 (print)/ISSN 1752-0754 (online) 2010 Association of Australasian PalaeontologistsDOI: 10.1080/03115511003663939
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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 duringthis time (Aitchisonet al. 1995, Landiset 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 (Schellartet al. 2006), post-
dating the initiation of separation fromAustralia (Aitchison et al. 1995). Rifting
associated with the anticlockwise break-up
of eastern Gondwana initiated in the
Fig. 1. A, Map of ichthyosaur fossil localities in Australia, modified from Kear (2003). B, Map of ichthyosaur fossil
localities in New Zealand, modified from Fordyce (1982).C, Map of ichthyosaur fossil localities in New Caledonia,modified 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.
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Jurassic of northwestern Australia (Veevers
et al. 1991) and resulted in separation of
New Zealand from Australia by the earlyCenozoic (Schellart et al . 2006; but see
Laird & Bradshaw 2004 for an alternative
view). In contrast, little separation oc-
curred between Timor and Australia
throughout the Mesozoic since the forma-
tion of Timor on the passive Tethyan
margin of Gondwana (Audley-Charles
1978). Large areas of western, southern,
and eastern Australia were flooded by
shallow epicontinental seas in the mid-Cretaceous (Frakes et al. 1987), and the
region has occupied mid- to high-latitude
settings throughout much of the Mesozoic
(Embleton 1984).
Review of Australasianichthyosaur taxa
Triassic specimens
New Zealand has most Triassic specimens,
with isolated ichthyosaur bones occurring
on both the North and South islands (Fig.
1B). The first reported finds originated
from the Mount Potts region near Canter-
bury, 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 vertebralcentra 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 Shoni-
saurus sikanniensis, an animal well known
as a giant (Nicholls & Manabe 2004).
Unfortunately, these particular specimens
can not be located, and are presumed tohave been either sent to the US with other
marine reptile bones that have now van-
ished (Cox 1991), or alternatively lost on
the transport ship Matoaka, which disap-
peared en route to London (Hector 1874,
Brazier et al. 1990). Hector (1874) diag-nosed 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 Mixosaur-
idae by Campbell (1965), but was laterregarded 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 unidentifi-
able ichthyosaur (Fleming et al . 1971).
Further Triassic ichthyosaur specimens from
South Island, New Zealand, include unde-
scribed 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 (For-
dyce 1982). Putative ichthyosaurian teethare known from Nugget Point and the
Wairoa district, but they exhibit labyrintho-
dont 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 (Camp-
bell 1984), but have been referred to
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Formation
Location
Age
Taxonomicassignment
ToolebucFormation
Queensland,Australia1,
2,
3,
4
late
Albian5,
6
P.
australis7(currentlyvalid),
P.
longmani3(juniorsynonym)
AllaruMu
dstone
Queensland,Australia1,
2,
3,
4
mid
-lateAlbian8
P.
australis7(currentlyvalid),
P.
longmani3(juniorsynonym)
WallumbillaFormation
Queensland1,
2,
3,
4
andNewSouth
earlyAptianlateAlbian10
P.
australis7(currentlyvalid),
Wales9,Au
stralia
P.
longmani3(juniorsynonym)
Doncaster
Member
NewSouthWales,
Apt
ian12,
13,
14,
15
P.
australis7(currentlyvalid),
(WallumbillaFormation)
Australia1
1
P.
longmani3(juniorsynonym)
BulldogSh
ale
SouthAus
tralia16
Apt
ian12,
13,
14,
15
P.
australis7(currentlyvalid),
P.
longmani3(juniorsynonym)
BirdrongS
andstone
WesternA
ustralia17
Hau
terivianBarremian11,
18
P.
australis7(currentlyvalid),
P.
longmani3(juniorsynonym)
AlingaFormation
WesternA
ustralia17
late
AlbianCenomanian19
P.
australis7(currentlyvalid),
P.
longmani3(juniorsynonym)
MolecapG
reensand
WesternA
ustralia20
Cen
omanianTuronian3
P.
australis7(currentlyvalid),
P.
longmani3(juniorsynonym)
DarwinFo
rmation
NorthernTerritory,
late
Aptian/Albian14,
22
P.
australis7(currentlyvalid),
Australia3
,21
P.
longmani3(juniorsynonym)
Makirikiri
Formation
NewZeala
nd23
Cretaceous23
Ichthyosauria23
Unituncer
tain
MountPotts,NewZealand24
Mid
dleTriassic25
Ichthyosaurusaustralis24
(nomendubium),
I.hectori26,Ichthyosauria23
AratauraF
ormation
Kawhia,N
ewZealand27
HettangianSinemurian27
Ichthyosauria27
OuamouiFormation
NewCaled
onia28
earlyNorian29
Sh
onisaurus30
Lepredour
Shellbeds
NewCaled
onia28
earlyNorian29
Ichthyosauria30
AitutuFormation
Timor31
Lad
inianNorian32
M
ixosaurustimorensis31
(nomendu
bium),
Mixosaurussp.3
3
Table1.D
istributionofichthyosaurmate
rialintheAustralasianregion.
Seefootnotesforsourcetexts
included.Currentstatusoftaxonomic
assignmentinbold
1Molnar(1
982),
2Wade(1984),
3Wade(199
0),
4Kear(2002b),5Mooreetal.
(1986),
6McMinn&Burger(1986),7McCoy(1867a),8Krieg&R
odgers
(1995),
9Etheridge(1904),
10Helbyetal.(19
87),11Kear(2003),
12Ludbrook(1966),
13Johns(1968),
14Day(1969),15Hendersonetal.(2000),
16
Alley&
Pledge(2000),17Choo(1999),
18McLoughlinetal.(1995),
19Siverson(1999),
20Teichert&Matheson(1944),
21Kear(2002a),22Henderson
(1998),
23Flemingetal.(1971),
24Hector(1874),25Campbell&Warren(1965),
2
6Lydekker(1889),
27Sachs&Gr
ant-Mackie(2003),
28Campbell
(1984),
29Callaway
&Massare(1989),
30Mazin(1985),31Broili(1931),
32Audley-Charles(1968),
33Mazin(1983).
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Shonisaurus spp. by Mazin (1985), based
on the crown ornamentation and morpho-
logy of the teeth. Long bones and vertebraehave also been recorded from the lower
Norian Lepre dour Shellbeds (Campbell
1984).
Eight vertebrae and a basioccipital
from the Middle Triassic (Callaway &
Massare 1989), possibly from the Ladi-
nianNorian (Table 1) shallow-marine
Aitutu Formation (Audley-Charles 1968),
Timor, were originally described as Mix-
osaurus 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 incon-
gruous report by von Huene (1936), which
identified 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 attribu-
ted to Mixosaurus by Broili (1931), Cym-
bospondylus is here considered not
confidently recorded from the Australasian
region.
Jurassic specimens
Only one putative Jurassic ichthyosaurspecimen is currently known from the
Australasian region. This comprises an
isolated centrum from Kawhia, New Zeal-
and (Sachs & Grant-Mackie 2003), from the
HettangianSinemurian Arataura Forma-
tion (Sachs & Grant-Mackie 2003). It is
not yet formally described.
Cretaceous specimensA 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 identified as ichthyosaurianbut 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 ichthyo-
saurs are known from numerous deposits
(Table 1), including: the Toolebuc Forma-
tion, Allaru Mudstone and Wallumbilla
Formation of Queensland (Molnar 1982,
Wade 1984, 1990, Kear 2002b); WallumbillaFormation (Etheridge 1904) and, more
specifically, 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 Terri-
tory (Wade 1990, Kear 2002a). Kear (2003)
recently summarized Australian ichthyo-
saur remains, including diagnostic speci-
mens. 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 WesternAustralia is more similar to the Eurasian
species, P. campylodon or P. kiprijanoffi,
than the recognized Australian form (Choo
1999).
Non-ichthyosaurian remains
Two records of Australasian ichthyosaurs
are now considered non-ichthyosaurian. A
presumed ichthyosaur jaw from NewZealand (Benham 1936) has since been
referred to a squaladontid cetacean (Camp
1942). The only proposed Triassic ichthyo-
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saur from Australia (Cosgriff & Garbutt
1972) has not been studied or figured.
Its ichthyosaurian affinity 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 misidentified amphi-
bian, so this specimen requires further study.
Systematics of the AustralianmaterialSubclass DIAPSIDA Osborn, 1903
Superorder ICHTHYOPTERYGIA Owen,
1840
Order ICHTHYOSAURIA de Blainville,
1835
Family OPHTHALMOSAURIDAE Baur,1887
Platypterygiusvon 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)
1867Ichthyosaurus australisMcCoy, p. 356.
[1867a]
1888 Ichthyosaurus marathonensis Etheridge,
p. 408, pl. 1, figs 1 3.
1922 Myopterygius marathonensis von
Huene, pp. 96, 98.
1944 Myopterygius australis Teichert &Matheson, p. 169, figs 1 3.
1972 Platypterygius australis McGowan,
p. 17, pls 3 4.
1990Platypterygius longmaniWade, p. 120,
figs 1 6.
Neotype. MV P12989 (Fig. 2AC), desig-
nated herein; the original holotype compris-
ing numerous centra is lost.
Type locality, formation and age. Flinders
River, north central Queensland; Allaru
Mudstone; Early Cretaceous (late Albian).
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. Max-illa 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 well-
developed anterior foramen and one or more
foramina present (in nasal) posterodorsal to
external bony nasal opening. Parietal con-
tributes to the facet for the paroccipital
process of the opisthotic on the supratempor-
al. 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 bifurca-
tion occurring in the primary axis (digit IV).
Neural spines of at least neck and anteriortrunk 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 australismaterial 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-axatlasaxis complex; basbasioccipital;
bnabony nasal aperture; lac lacrimal;maxmaxilla; nanasal; snf supernarial foramen. Scale bars10 cm.
3
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locality, though collected one year later
(Wade 1990). MV P12989 (Fig. 2AB) in
particular, an incomplete skull and atlasaxis complex, has a combination of three
characters differentiating it from other
ichthyosaur taxa (listed here to satisfy
ICZN Article 75.3.2): (1) reduced extracon-
dylar 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 poster-
odorsal to the bony nasal aperture (Fig. 2A;
B. Kear, pers. comm. 2009). The firstcharacter listed is diagnostic of Platypter-
ygius, whereas the other two simultaneously
differentiate the Australian Platypterygius
species and demonstrate that P. longmani
and P. australis are the same taxon,
relegatingP. longmanito 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 scientific institution (satisfy-
ing 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.
AcknowledgementsMany thanks to the curatorial staff and
museums for providing information and/or
access to specimens: Neville Hudson (Auck-
land 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 fortheir comments on the manuscript. Use of
information contained within the New
Zealand Fossil Record File is acknowledged.
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