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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

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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.

282 MARIA ZAMMIT ALCHERINGA


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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

ALCHERINGA AUSTRALASIAN ICHTHYOSAURS 283


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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.


P.


WallumbillaFormation

Queensland1,

2,

3,

4

andNewSouth

earlyAptianlateAlbian10

P.


Wales9,Au

stralia

P.


Doncaster

Member

NewSouthWales,

Apt

ian12,

13,

14,

15

P.


(WallumbillaFormation)

Australia1

1

P.


BulldogSh

ale

SouthAus

tralia16

Apt

ian12,

13,

14,

15

P.


P.


BirdrongS

andstone

WesternA

ustralia17

Hau

terivianBarremian11,

18

P.


P.


AlingaFormation

WesternA

ustralia17

late

AlbianCenomanian19

P.


P.


MolecapG

reensand

WesternA

ustralia20

Cen

omanianTuronian3

P.


P.


DarwinFo

rmation

NorthernTerritory,

late

Aptian/Albian14,

22

P.


Australia3

,21

P.


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).



6/13

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-



7/13

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)



8/13

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



9/13


10/13

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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