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 Thomas H.V. Rich and Patricia V. Rich Articles

Dinosaurs and
Biotas of the Early -

Cretaceous of
Southeastern Australia
Southern polar biotas of the Early Cretaceous Valanginian-early Al-
bian) of south-central Victoria, Australia, contain > 150 taxa of
plants, invertebrates, and vertebrates, includingdinosaurs. The domi-
nant dinosaurs are small- to medium-sized hypsilophodontids (Leael-
lynasaura amicagraphicagen. et sp. nov., Atlascopcosaurus loadsigen.
et sp. nov., Fulgurotherium australe) -not usually dominant elements
in much larger thanatoceonoses elsewhere in the world. Vertebrates
also includef s h , possibly a labyrinthodontamphibian, turtles, lepido-
saurs, pterosaurs, freshwater plesiosaurs, theropods, and birds, but
thusfar no crocodiles or mammals. Thefossil-producingsedimentsare
the Obvay and Strzeleccki Groups, which some authors combine into one
unit, the Korumburra Group. These rock units have beendated by their
palynomorphs and independently by fssion-track techniques. They
represent volcanogenic clays, silts, and sands deposited in several d F
ferent fmies of braided river systems that occupied a down-dropped
graben created when Australia and Antarctica began to separate in the
late Mesozoic. Periodicity in the sedimentation regime may have been
related to meltwaterfloods in the austral spring. Invertebrates, plants,
and geochemistry sugqest that a cool, humid climate prevailed. Paleo-
magnetic studies on the Otway Group indicate that southern Victoria
w m situated at a high latitude, between 70 and 85" south, in the late
Early Cretaceous. The relict lAUosaurus sp., a possible labyrinthodont,
flora), endemic f f h e s , dinosaurs) nature of the biota, and the domi-
nance of small- to medium-sized dinosaurs may rejlect a polar 'kafe
area" isolated perhaps more by latitude and its sympathetic climate
than by geographic discordance. The Early Cretaceous occurrence of
hypsilophodontidsand theropods in polar Victoria and of hadrosaurs
and theropods in Late Cretaceous Alaska clearly demonstrate that di-
nosaurs used high latitudesfor a considerable period of time and that
their residence there was not a casual dalliance.

Dinosaurs are not common in Australia. Thus, the recovery of a moder-

ately varied late Mesozoic reptilian fauna from several localities along
the south coast of Victoria has added significantly to the understanding
of antipodean dinosaurs. Even more important is the fact that this Vic-
torian fauna lived well inside the Antarctic Circle of the time, and per- Thomas H. Rich, Curator of Verte-
haps as far south as 80 to 85' (Einbleton G1 McElhinny 1982, Idnurm brate Palaeontolo@, Museum of
1985). These dinosaurs somehow managed to survive at least three Victoria, Melbourne, Victoria 3000
Australia; and Patricia V. Rich,
months of winter darkness, either by annual migration, hibernation, or Senior Lecturer, Monash Uni~~ersity,
remaining active. Only one other place in the world, the north slope of Clayton, Victoria 3168 Australia.

NATIONAL GEOGRAPHIC RESEARCH 5(1):15-53 (1989)

 Alaska (Brouwers et al. 19871, has yielded high-latitude dinosaurs.
The Early Cretaceous vertebrate faunas of Victoria are unusual in be-
ing dominated by small ornithopod dinosaurs. Nowhere else in the
world is this the case except perhaps in the Late Triassic-Early Jurassic
of southern Africa. The small Australian ornithopods are endemic at the
generic level, but clearJy belong within the Hypsilophodontidae, indi-
cating that some form ofisolating barrier-be it climatic, geographic, or
topographic-was in effect. This idea is reinforced by the possible latest
occurrence in the world of a labyrinthodont amphibian, a group that
had become extinct elsewhere in the world by the Late Jurassic.
Other faunal elements associated with the hypsiloplzodontidsinclude
rare occurrences of carnivorous dinosaurs (Allosaurus sp. and a small-
er theropod), fi-eshwaterplesiosaurs, flying reptiles, turtles, birds, and a
variety of fishes. A varied insect fauna consisting primarily of Hemip-
tera, Coleoptera, and Diptera is also associated wit11 a flora dominated
by gymnosperms, lycopods, and ferns. The total biota suggests humid,
cool, temperate, not tropical conditions.
Recent work (Gregoly et al. in press) on 180/1G0 ratios for concretions
in the Otway and Strzelecki Groups- the rock unit from which the ma-
jority ofVictorian dinosaurs have been recovered-suggests that surface
air temperatures may have had an annual mean close to 0°C. Sedimento-
logical a~zalysisindicates that the enclosing muds, silts, sands, and con-
glo~nerateswere a typical braided stream sequence deposited under
conditions of highly varying energy. Higher energy deposition may have
occurred as meltwater floods in the springtime. Ifthe streams were melt-
water fed, the source may have been volcanic mountains located to the
east or on the shoulders of the enclosing graben. Seasonal ice buildup
has been, likewise, suggested for parts of Australia in the Early Creta-
ceous (Frakes & Francis 1988, Waldman 1971).
At the time this biota inhabited southern Australia, about 120 million
to 106 million years ago, ihe continent was still firmly attached to Ant-
arctica. Topographically this area was a down-dropped graben, a
broadvalleywl~ichreflected the beginnings of continentalbreakup, well
separated fi-om a marine influence (Frakes et al. 1987). It was thus a
"weedy," disturbed environment, a change from the stable, cratonic re-
gime of the past. The flora was changing; angiosperms were making
their first appearance, but were rare. The southern part ofAustralia was
very much a part ofAntarctica at this time, quite separate from other ter-
restrial areas now recognized as Australia. Seaways drowned much of
the remainder of the continent, leaving only large islands off to the north
of southern Victoria, and Antarctoaustralia.
The high latitude environments present in southern Victoria in the
Early Cretaceous have no modern counterpart because factors such as
cold winters and several months of annual darkness arejuxtaposed with
a relatively diverse biota.
Previous Work
At the turn of the century, William Hamilton Ferguson was mapping the
rock exposures on the Victorian coast of southeastern Australia near the
town of Inverloch, 150km southeast of Melbourne (Figure I).While do-
ing so, he found a lungfish tooth and the claw of a dinosaur. These were
subsequently reported by A. Smith Woodward (1906). The claw of the
dinosaur was allocated to the theropod Megalosaurus.
In the subsequent three quarters of a century, the remains of a turtle
(Warren 1969), a few isolated feathers (Talent et al. 1966), and a possi-
ble varanid limb bone (Molnar 1980) were the sum total of the tetrapod
1
RICH (SI RICH
48
Bi
 material recovered from the Victorian Cretaceous deposits. Aquatic ver-
tebrates were more common (Waldlnan 1971).
In 1978, Timothy Flannery and Jolm Long returned to the area near
Eagles Nest where Ferepon had discovered the two original specimens.
They found an additional specimen allnost immediately. For the bal-
ance of 1978, Flannery continued searching the Early Cretaceous coastal
outcrops in tlie area. He collected - 30 bones, including an astragalus re-
ferred lo the theropod dinosuar Allosaui-us (Molnar et al. 1981, 1985;
Welles 1983) and a fragment of the j a w of a possible temnospondyl am-
phibian (Jupp & Warren 1986).
Flannely confined his attention to the coastal outcrops of the Sirze-
lecki Group in the Gippsland Basin, although the unit occurs widely in-
land. Unfortunately, the inland occurrences tend io be poorly exposed
and have frecluently suffered extensive chemical alteration, so that even
if fossil bones once were present in the rocks within tens of meters of the
soil-covered surface, they would have been leached away.
In 1979 and 1980 a program was mounted to search for fossil verte-
brates in similar coastal outcrops of the Early Cretaceous Otway Group
that occur in the Otway Basin 200 km to the west on the flanks of ihe Ot-
way Range. This illvolved a group ofworkers from several museums and
universities: Frank Bussat, Michael Archer, Timothy Flannery, Ralph
Molnar, Patricia Rich, and Thomas Rich. Besides isolated occurrences
offossil bones, four localized areas were discovered that were relatively
rich in fossils. These were, from east to west, Point Lewis, Point Frank-
lin, Eric the Red, and Dinosaur Cove (Figure I).
Most subsequent effort has been directed at Dinosaur Cove (Fi91re 2),
the richest of the four. Here i n <400 m, three major sites were found.
East to west, and in ascending stratigraphic order, these are Slippery ,
Rock, Dinosaur Cove East, and Dinosaur Cove West. In each case, fossils
had accumulated in the lithified sediments of former stream channels.
About a dozen specimens were collected from Dinosaur Cove East be- ,
tween 1980 and 1982, the only site then recognized. More work was fu-
iile at that time because the available tools were inadequate to follow the
paleostream channels into the cliffs formed of indurated sandstone.
At the urging and with the sponsorship of the Friends ofthe Museum
of Victoria, an attempt was made in 1984 to tunnel into the sandstone
and follow the paleostrearn channel at Dinosaur Cove East. This re-
quired air-driven rock drills and breakers (provided by Atlas Copco).
Because of monetary tonsil-aints, only 16 days were available for this
first trial. Still 65 volunteers managed lo recover 85 bone fragments and
a dinosaur tooth. Work continued at Dinosaur Cove during the subse-
quent austral summers of 1985 to 1987 and resulted in the discovery of
two other productive sites plus the recovery of >2000 bone fragments,
bones, isolated teeth, toot11 rows, a n d a partial skeleton of a dinosaur.
Geology
Dlinnan 63 Chambers (1986) and more recently Douglas et al. (1988)
have summarized the Early Cretaceous geology of southern Victoria. In
brief, two Early Cretaceous basins occur along the southern coast of Vic-
toria and extend inland. These are the Otway Basin to the west and the
Gippsland Basin to the east. The two are separated by the Mornington
High, a few tens of kilometers wide. Because of this separation, the
names Otway Group and Strzelecki Group have been applied to the Ear-
ly Cretaceous rocks in these two respective basins (Douglas et al. 1988))
a convention followed here. Elsewhere, in recognition ofthe great likeli-
hood thai the rocks of these two units resulted from the same geologic

RICH& IUCH u
 event, the name Korumburra Group has been used as an alternative for
the Early Cretaceous rocks of both basins (e.g., Brown et al. 1968). Both
of these basins and the now offshore Bass Basin were initiated by Early
Cretaceous times with a north-northeast-south-southwest extention,
evidenced by "widespread shallow to moderately dipping rotational
*orma1 faults of do~ninostyle considered to sole out in major detach-
rne~ltfaults"(Etheridge et al. in press). The three basins were formed as
a of the onset of rifting between Australia and Antarctica between
- 110 million years ago (Mutter et al. 1985) and 95 million years ago (ve-
evers 1984) when 5 3 krn of primarily fluviatile volcanoclastic arkosic
sands, muds, and coals were emplaced. The sediments appear to indi-
cate pel-iodic times of high flow alternating with lower energy regimes
[Lees 1987, Wagstaff 1983). A volcanic province to the east associated
with the initiation of formation of the Tasman Sea may have been a ma-
jor source of these deposits.
Age estimates of the Otway and Strzelecki Groups have had a convo-
luted history (Drinnan & Chambers 1986). The present view was first
stated by Dettmann (1963), who placed these rocks in the Early Creta-
ceous on the basis of palynological evidence. Her work was expanded
upon in Detlmann [1986), Dettmann G3 Douglas (1976), and Dettmann
& Playford (1969). Subsequently, using fission-lllackdating, Gleadow G.1
Duddy (1980) regarded the Otway Group as late Early Cretaceous.
Wagstaff G3 McEwen Mason (this volume, p. 54) have palynologically
dated most ofthe tetrapod-bearing sites in the Otway and Gippsland Ba-
sins. Accordingly, the Otway Basin coastal sites are latest Aptian-early
Albian, and the Gippsland Basin sites are Valanginian-Aptian. But, a
hrther subdivision of the sites in Gippsland is possible. The two most
westerly coastal localities, Punch Bowl and Kilcunda, are ?Barremian-
Aptian. Where palynologically sampled, the remaining Gippsland coast-
al localities are Valanginian-Barremian in age.
Localities
The geographical and geoloeJca1 data for each site follows. Wit11 the ex-
ception of Lightning Ridge, all are locaied in south-cenkal Victoria,
Australia.
The Arch, Kilcuinda, 38'33' S, 145' 29' E, Strzelecki Group, ?Barre-
mian-Aptian
Cape Paterson, 38'40' S, 145'37' E, Strzelecki Group, Early Cretaceous
Dinosaur Cove East, Dinosaur Cove, 38'46' 5 3 4 1"S, 143"24' 14*1" E,
Otway Group, latest Aptian-early Albian
Dinosaur Cove West, Dinosaur Cove, 38'46' 53 + 1" S, 143'24' 9+1" E,
Otway Group, latest Aptian-early Albian
Eagles Nest, 38'40.5' S, 145'40.25' E, Swzelecki Group, Valanginian-
Barremian
Lightning Ridge, New South Wales, 29.5' S, 148' E, Griman CreekFor-
mation, Early Cretaceous
Point Lewis, 38'50'9 to 12'' S, 143'34'53 to 57" E, Otway Group, latest
Aptian-early Albian.
Punch Bowl, 3€i032'21"S, 145O24'9 to 13" E, Strzelecki Group, ?Barre-
mian-Aptian
SlippelyRock, Dinosaur Cove, 38"4G154+1"S, 143'24' 15.t1"E, Otway
Group, latest Aptian-early Albian
Specimens are housed at: Australian Museum, Sydney (AM); British
Museum, London [BMNH); Humboldt Museum f i r ~aturkinde,East
Berlin (HMN); Museum of Victoria, Melbourne (NMV); ~ueensland
Museum (QM); Southern Methodist University, Dallas [SMU).

- DINOSAURS OF AUSTMI.TA
 Svstematics -

The classificatioil of or~liihischiandinosaurs is currently i n a state of flux

(see Sereno 1986 a n d references therein). As a framework for this paper,
Cooper's (1985) cladistic classification of the o r n i t l ~ i s c l ~ i ais
n sfollowed
wit11 one exception: the a ~ ~ t l l ouse
r s Fabrosauridae Galton 1972, rather'
than Nanosauridae Marsh 1878.

Order Ornithopoda DIAGNOSIS: Upper cheek teeth recessed beneath a lateral projection of the
Marsh 1881 maxilla rather than marginal as in fabrosaurids. Lower cheek teeth in a mas-
Suborder Hypsilophodontia sive, rather than slender jaw, as in fabrosaurids. Unlike heterodontosaurids, no
Cooper 1985 caniniform teeth present. Femora distinguished from those of fabrosaurids by
Superfamily Hypsilopho- the lesser trochanter's being smaller than the greater trochanter. Femora distin-
dontia Cooper 1985 guished from those of heterodontosaurids by the presence of a posterior inter-
Family Hypsilophodont5dae condylar groove, a cleft between the lesser trochanter and the femoral shaft, and
Dollo 1882 the transverse axis ofthe distal femoral articular surface being horizontal rather
than inclined medially. Femora distinguished from those of iguanodontids and
camptosauridsby being less massive, the fourth trochanter's being proximal of
the midpoint of the shaft, the anterior intercondylar groove's being quite shal-
low, if present at all, rather than prominent.
DISCUSSION: The above diagnosis for the Hypsilophodontidae does not distin-
guish their cheek teeth from those of iguanodontids and camptosaurids. Cooper
(1985) does differentiate the upper cheek teeth ofthe Hypsilophodontidae from
those of the Camptosauridae and Iguanodontidae by their lacking a median
ridge (primary ridge, as used below). I11 the case of Hypsilophodon, it is cer-
tainly true that on the upper cheek teeth no ridge can be described as a median
or primary one. However, D~yosaul-usand Otlznielia, which are also called
hypsilophodontidsby Cooper (1985), may have a prominent median or prima-
ry ridge on their upper cheek teeth (see Dodson 1980).
Cooper 11985) provides no other character of the cheek teeth that is usehl in
distinguishing those of l~ypsilopl~odontids from those of iguanodontids and
camptosaurids. Given the grouping of taxa in the hypsilophodontids as he allo-
cates them, there may be none except for the smaller size of hypsilophodontid
teeth. To dojustice to Cooper, he does provide a suite of character states for oth-
er parts of the skeleton that characterize the l~ypsilopl~odontids,iguanodontids,
and camptosaurids.
Unworn upper cheek teeth o f D ~ y o s a u r may
~ ~ sbe characterized as lozenge-
shaped, because the apex of the tooth forms a horizontal blade between the an-
terior and posterior edges of the tooth that converges toward either end fioin the
widest part of the crown. This pattern is typical of iguanodontids and campto-
saurids. It contrasts markedly wit11 the condition in fabrosaurids, lleterodonto-
saurids, such llypsilophodoniids as Hypsiloplzodon and Othnielia (Galton
1974a, 1983), and the new Victorian 1~ypsilophodontidsdescribed below,
where the apex of the crown comes to a point. Thus, the teeth of tlle two new
genera are allocated to the Hypsilophodontidae because, like other primitive
taxa within the family, they are clearly not iguanodontids or camptosaurids and
yet are more derived than fabrosaurids and heterodontosaurids in the location
of the teeth in the maxilla and tlle pattern of toot11 wear. Altllough these non-
iguanodontid and non-camptosaurid dental features are shared with some
members of the Ilypsilophodo~~tidae, they do not characterize the family as a
whole because other l~ypsilopl~odoatids have a more derived condition.

teaellynasaura gen. nov. KNOWN DISTRIBUTION: Latest Aptian-early Albian (Early Cretaceous), south-
ern Victoria, Australia.
DLAGNOSIS: Distinguished from all other llypsilophodontids by a more antero-
posteriorly compressed distal end of the femur [Molnar G3 Galton 1986: fig. 4)
and from all except Othnielia, by the presence of ridges on both sides ofunworn
upper cheek teeth. Upper cheek teeth disting'uished from those of Atlascopco-

20 RICH & RICH

saul-us gen. nov. by lacking two discrete sizes of ridges, and the equal develop-
ment of ridges on both sides of unworn teeth. Distinguishedfrom the cheek teeth
ofAlocodon, Nanosnurus, and Plzyllodon (Galton 1983) by possessing ridges
that cover the entire internal and external surfaces rather than denticles restrict-
ed to the margin of the crown. Distinguished from the upper cheek teeth of
Catizptosaza-us (Galton & Powell 1980), Dryosaurus (Galton 1983), Mochlo-
don (Nopsca 1904, cited in Bartholomai & Molnar 1981), Atlascopcosaurus
gen. nov., Thescelosaurus (Galton 1974b, Morris 1976), and the Lightning
Ridge isolated hypsilophodontid cheek tooth figured by Molnar (1980: fig. 5,
QM F9505) in lacking a single primary ridge that is markedly stronger than all
other ridges. Distinguished from the upper cheek teeth of Atlascopcosauru.s
gen. nov., Otlznielia (Galton 1983), Parlcsosaurus (Galton 1973), Thescelo-
saurus (Galton 197413, Morris 1976), Zephyrosaurus (Sues 1980), and the
Lightning Ridge isolated hypsilophodontid'cheek tooth figured by MoInar
(1980: fig. 5, QM F950.5) by possessi~lga maximum of five ridges rather than a Figure 2. Dirrosaur Cove, site of the
reco_y of most of the dinosaurfas-
minimum of eight. sils fourld in Victoria, Australia. The
ETYMOLOGY: "Leaellyn" in honor of Leaellyn Rich, a Victorian schoolgirl shore platform is formed of the &no-
who participated in the discovery; "saura," Greek for '‘lizard," feminine. saur-bearing Otway Group.

Frilnk CoWl

HOLOTYPE: NMV Pi85 991-skull fragment including the left maxilla with Leaellynasaura
eight cheek teeth and alveoli for four more (perhaps five) anterior to those, left amicagraphica sp. nov-
pterygoid, left jugal, and possibly vomer and left premaxilla (Figure 3).
TYPE LOCALITY AND STRATIGRAPHIC POSITION: Slippery Rock, Dinosaur
Cove, Otway Group, latest Aptian-early Albian.
DIAGNOSIS: That of genus until other species are described.

DINOSAURS OF AUSTRALIA
 ETYMOLOGY: LLAmi~a," Latin for "friend"; "graphics," Latin for "writings,'"
feminine. In honor of the Friends of the Museum of Victoria and the National
Geographic Society, for their great assistance with the work on the Cretaceous
tetrapods of Victoria, Australia.
REFERRED SPECIMENS
Slippeiy Rock. NMVPi85 990: left and right nasals, prefrontals, frontals, and
parietals (Figure 4). NMV PI85 992: articulated proximal caudal vertebrae
plus approximately the distal seven eighths of the femur, the proximal three
fourths of the tibia and fibula, and two articulated digits, one with two and the
other-with three phalanges in articulation. NMV P185993: articulated distal
caudal vertebrae. All three specirne~lswere quite likelypart ofthe same individ-
ual as the holotype. Both skull fragments, NMV PI85991 and NMV P185990,
were found the same day -20 cm apart. Otherwise tetrapod skull material is as
yet unknown at Dinosaur Cove. The two specimens are the correct size to be
part of the same individual; both appear to have belonged to an immature ani-

I/ Figure 3. Leaelly1asaul.a arnicagra-

! phica holotype, NW PI859.91:
A (left), luternl view of dn~titiort1x2);
! A (right), r?~edinlvinv of detltitiorz
j (X2); B, nredictl view of skullfr.ctg-
merit (X1.33).
I
I

Vorner?

-Maxilla

- Jugal
Quadrate

mal; and elements between them do not overlap. The two sections of articulated
vertebrac, NMV Pi85 992 and NMV PI85 993 (the former with a partial llind
limb associated), were found within 80 cm of the two skull fragments the fol-
lowing day. Their size is that expected for an ornithopod of the dimensions of
the skull fragments.
Dinosaur Cove East. NMV P179561: right femur slightly crushed (Figure 5).
NMV PI79 564: left femur slightly crushed, condyles missing (Fipure 6). NMV
P182968: right femur crushed and missing head, proximal part of lesser tro-
chanter, and distal tip of fourth trochanter. NMV P185867: right femur crushed
and missing medial condyle; NMV P185 980: right femur missing head, proxi-

RICH 13RICH
 rnal tip of lesser trochanter, and medial condyle (Figure 7).
Dinosaur Cove West. NMV P181681: crushed left femur missing internal
condyle, fourth and lesser trocharlters.
STAGE OF GROWTH OF THE HOLOTYPE INDIVIDUAL: The two sk~tllfrag-
ments (NMV PI85990 and NMV P185991) each display a feature that suggests
an immature individual. Anterior to the eight fu~lctiol~al o r partially erupted
cheek teeth of the holotype, NMV P185991, are four undoubted alveoli in toot11
positions 1 to 4. Another alveolus may be present anterior to tooth position 1.I11
toot11 positions 2 and 4, the tips of a few serrations along the occlusal crcst of
otherwise unexposed cheek teeth are visible. The other two alveoli (tooth posi-
tions 1and 3) sllow no sign of teeth whatsoever. This pattern of a number of
empty alveoli in front of a contilluous series of partially or k l l y erupted teeth in
the maxilla is found in the young of modern crocodiles. The posterior boundaly
oftlle parietal (part of NMV P185990))which in life articulated wit11 the miss-

Cerebral
hemisphere'
/ I

.~ a r i i t a l
body
Optic lobe cerebellum

ing supraoccipital, is complete. Equally complete are the lateral boundaries of Figure 4. Leaellynasaura amicagra-
this bone where the missingpostorbitals and laterosphenoids articulated. This phica, NDlV P185990: p~.esc~rnczbly
suggests that these bones easily separated after death, as if no fusion of the su- this speci~nenis part of the sanle in-
tures between them had taken place. This is to be expected i n juveniles. dividual as the holotype, NMV
The largest referred femur ofL, amicag~aphicu,NMV PI79 564, is about 1.45 PI85991 (Figure 3). A (left, center),
times longer than that of the one likely to be from the same individual as the ho- ~antralview of rear of thefiorltals
plus the pal-ietal in nori)zal and r r -
lotype specimen, NMV PI85 992. As 1.45~=3,this indicates that at a maximum, verse; A (center, right), ster~*oscopic
the holotype individual was no Inore than one-third full grown. view, showing thefo~,nzof the erido-
DENTITION: No trace of ally preincvrillaryteeth has been found on the holo- cast in dorsal view; B, dorsal vim) of
type, the only specimen of this species with teeth preserved; but 12 or 13 cheek- the anterior- of the skzcll fragrlerzt,
tooth positions are evident. The doubtful one is a sinall alveolus(?) a t the and, behind it, the anterior of the
anterior end of the cheek-tooth row. In the discussion that follows, the cheek- natural erldocast. (X 1)
tooth positions are numbered I throug1~12,anterior lo posterior, assuming [hat
the doubtfbl alveolus is not an alveolus at all.
Tooth positions 1 and 3 are empty alveoli, and positions 2 and 4contain uner-
upted teeth with the tips of only a few denticles visible. Positions 5 , 7, 9, a n d 11
have klly erupted teeth. Position 6 has a fully erupted tooth with its root partial-
ly resorbed and the successor tooth visible. Position 8 has a partially erupted
tooth. Positions 10 and 12 bave teeth that are almost but not completely erupted.
This back-to-front replacement ofalternativeteeth with the anterior, more in-
completely emplaced teeth progressively less fully erupted is consistent wit11 a
Zahnreihen spacing of somewhat > 2.0 and significantly < 3.0; i.e., a typical
value for reptiles (DeMar 1972). The length of the erupted cheek teeth varies
from 2.2 mm for cheek tooth 5, to 3.0 mm for cheek tooth 10.

DINOSAURS OF AUSTRALIA
 Unlike At1ascopcosau1- is loadsi gen. et sp. nov., the ridges on an ullworn
upper cheek tooth are of ecpal strength on the buccal and lingual sides. With
wear, the ridges on the linlgual side of the uppers become obliterated. The stron-
gest ridge is near the rear of the tooth, and the ridges become progressively
weaker anteriorly. Also, unlike A. lorndsi, the ridges are not obviously divided
into two quite different sizes, a prominent primary ridge and secondary ones;
all are equally much weaker.
Thai these differences from A. Zoadsi are not simply related to ontogenetic
changes is s~~pported by the existence of NMV P181677, a partial upper cheek
tooth ofA. loadsi. This specilnen is quite similar to the holotype ofL. arnicapa-
phica in size but quite unlike that species in ~norphologyand virtually indistin-
guishable in form from the other, larger upper cheek teeth of A. loadsi.
BRAIN ENDOCAST: A n excellent endocast of the dorsal aspect of the cerebral
hemispheres, optic lobes, parietal body, and possibly the anterior part of the
cerebellum is preserved on NMV PI85 990 (Figure 4). The condition of the ol-

Figure 5. Right fenlur of Leaell]

saurn amicagraphica, NMV
P1795Gl:A (left), anterior view
( x i ) ;A (right), posterior view
( X I); B (left), proxinzal vi m
(x 1.5); B (rigllt), distal v i m
( X 1.5).

factory tracts is uncertain because the ventral surface of the frontals anterior to
the cerebral hemispheres is unexposed.
Between the left and right cerebral hemisphere, a broad depression clearly
separates them from one another. The length of the cerebral hemispheres is - 13
mm and their maximum combined width, 18 mm. On the ventral surface of the
frontal bones is a series of closely spaced, fine, parallel grooves. These grooves
are so numerous and nearly straight that they apparently do not reflect the ex-
pected course of sinuous blood vessels as are seen on the cerebellar regions of
some fossil reptiles such as Rromiceiomimus brevitertius (Hopson 1979: fig.
1 3 ~ )Rather,
. the dorsal surface of the endocast seems to have been smoothly
rounded with no marked furrows or ridges developed. The frontal-parietal su-
ture is readily visible near the rear of the cerebral hemispheres.
Just behind the frontal-parietal suture, a broad depression separates the ce-
rebral hemispheres from the optic lobes. The optic lobes are rounded protuber-
ances -4 mm in diameter at the lateral margins of the endocast. Between them
on the midline is a smaller protuberance -1.7 mm in diameter, interpreted as a
parietal body. The width across the endocast at this point is 16 mm.

RICH & RICH

 Figure 6. L@femur of Le;
saura amicagraphica, NIMV
P179564:A (left), anterior view;
A(right), posterior view; BI (lefi)
medial view; B (right), proxinzal
view. ( X I )

-
---il=." - m z s w w*eu*-,,aw.3 " U S " ---r.n. il4.i" .-..,..,n ,,,,
,
u #,.,A*, .J
ri*i*"u.

DINOSAURS OF AUSTRALIA 25
 Galton (1974a:fig. 6B) illustrates, but does not dcscribe, what appears to be a
small pit near the posterior edge of the ventral surface of the parietal ofH,,usilo-
phodonfoxii. Presulnably because of the relatively posterior location ofthis pit,
it was for a structure other than a parietal body.
Hopson (1979) points out that in modern Caiman, swellings similar to those
interpreted here as the optic lobes occur as a result of the presence of lateral
branches of the longitudinal venous sir~us.However, he accepts the interpreta-
tion by Russell (1969:92) for the presence of optic lobes on the endocast of tlle
theropod dinosaur Stenonyclzosau~~zrs inequalis because of ". . . their position
immediately adjacent to the cerebral hemispheres and their broadly rounded
."
form . . . The presumed optic lobes on the endocast ofL. arrzicag?uphica ap-
pear to meet these criteria. Currie (1987) regards Stenonychosaurus as ajunior
syllonym of Troodon.
Behind the optic lobes, the width across the parietals for the endocasi space
contracts to -7.5 mm. The cerebellum was located in this area. The width occu-

Figure 7. Rightfernur of Leaelly~a-

saura amicagsaphica, NMV
P185980: left, anterior view ( X i ) ;
center osterior vim (xi); right,
'.P
distal v1.m (X1.51.

pied by the cerebellum probably would have been wider, as the walls of the la-
terosphenoid and postorbital nlay have bulged outward somewhat. Hopson
.
(1979:47-481 points out that in modern Caiinan " . . the precise position, size,
and shape of the cerebellunl are totally obscured by the presence in life of the
thick, broad, dorsalvei~ouss i n ~ ~ sIn
. "light ofthese remarks, it is possible only to
make a minimum estimate of the width of the cerebellum, 7.5 mm, but not the
depth. Because the supraoccipital is not preserved, only a minimum estimate
for the anteroposterior extent of the cerebellum can be made: the distance from
the parietal body to the poster*ioredge of the parietal along the midline, 5.4 mm.
As the posterolateral corners of the parietal are a further 5.4 mm posterior than
the border of this bone at the midline, the cerebelleum likely continued that
much farther posteriorly under the supraoccipital.
Endocasts of small ornithopods have not been described. Endocasts oflarger
ornitl~ischiansdiffer markedly in that the brain is "flexed" so that the cerebel-
lum is below the level of the cerebral hemispheres, and the optic lobes are not
evident (Hopson 1979). Comparison of the endocast of L. amicagraphica with
that ofthe small theropod dinosaur Stenorzychosaurus inequalis (Hopson 1979:
fig. 13b, Russell 1969: fig. 3) shows a closer similarity. In both, the cerebral
hemispheres were smooth and somewhat wider than long. L. alnicagraphica
does not show any trace of the anterolateral swelling on the surface of the cere-
bral hemisp1lereinoted on S. inequalis. In both, the-midline separating the two
henlispheres from one another is markedly depressed. The suture between the
parietal and frontal crosses the rear part of the cerebral hemispheres on both.
The optic lobes are prominent and we11 set off from the remainder of the endo-
cast by depressions along their anterior and medial borders. S. inequalis lacks
any indication of the presence of a parietal body such as is found in L. amica-

RICH & RICH

 graphica. However, another small theropod, Dromiceiomimus breuitertius
(Hopson 1979: fig. 13c, Russell 1972: fig. 61, does have such a pineal body, so
this feature does not distinguish small theropods from ornithopods.
Jerison (1973) surveys the relationship between brain volume and body size
for as wide a spectrum of living vertebrates as he could obtain data. He uses the
heaviest individual recorded for each modern species or an average for that spe-
cies when information on individuals was not available.
Jerison's plot of these data on a log-log scale for body versus brain weight
clearly shows that modern birds and mammals (higher vertebrates) form one
cluster and modern amphibians, reptiles, and fishes (exceptsharks) form a sec-
ond cluster (lower vertebrates), not overlapping the first. On average, higher
vertebrates of a given body weight have brains 10 times as massive as lower ver-
tebrates of similar body size. To make this point more apparent,Jerison drew
polygons linking the extreme menlbers of these two groups so that all members
of each group were either on the edge or within one of these convex polygons.
The polygons for the higher and lower vertebrates do not overlap.
Jerison (1973) concludes that vertebrate evolution is characterized by stasis
in relative size of the brain in the evolution of fishes, amphibians, and reptiles.
In contrast, because of extreme sensory demands on the ancestors of birds and
mammals, these two groups evolved brains on average 10 times as large as fish-
es, amphibians, and reptiles of similar body mass.
Jerison (1973) considers data on fossils and finds that sorllc groups fall be-
tween the two polygons. These are g~~~~~ considered to be transitional in grade
of brain development between the reptiles on the one hand and modern birds
and mainmals on the other; e.g., Archaeopte~yxlithographica, the most primi-
tive bird, and the Mesozoic and earliest Tertiary mammals.
In summarizing the large amount of data on the brain weight (E) and body
weight (P) in grams of mature individuals of numerous species of vertebrates,
Jerison (1973) finds that within vertebrate classes, these two variables can be
related to one another by the following equation. This equation gives the major
axis for the brain weight-body weight polygoas. In other words, E, is the ex-
pected brain weight for an animal with body weight (P).
E = kP" Cil
Jerison uses 'I3 as the value for "a" while other workers have employed slight-
ly different values, but nonetheless, quite close toJerison's. (k is a constant nu-
merically equal to the mass of the brain in a hypothetical species of a body
weight of I g.) For mammals, Jerison (1973) calculates a value for k of 0.12.
Jerison (1973) develops the concept of encephalization quotient (EQ). This is
a measure of the ratio of the observed size of the brain of an animal Ei to that
expected for one of its same body mass E,.
EQi= - E,
Ee C2l
Combining equations [I] and [2] and substituting2/3 for "a," one can calculate
EQ for a species of known body weight, brain weight, and constant k.
EQi= - Ei
Ww 3 C3l
Hopson (1977) continues this work, concentrating on arcliosaurs. He adds
data on dinosaurs to thoseJerison has considered. Hopson (1977) also recalcu-
lates the position of Archaeopteryx lithographica using different estimates of
body weight and brain size than given in Jerison (1973). He positions A. litho-
g r a p h i c ~at the lowermost boundary of the higher vertebrates' polygon. With
this data set, Hopson (1977) calculates k as 0.005 for archosaurs.
Because the endocast of L. amicagraphica is only visible in dorsal view, i n
order to estimate the volume, an assumption must be made concerning the
transverse cross-section. This cross section is assumed to be elliptical with a
depth half that of the transverse width. (From figures of archosaur brains, half
seems a conservative and reasonable assumption.) The anterior end of the cere-
bral hemispheres is 7 mm wide, the widest point on the endocast is 11mm pos-
teriorly near the rear of the cerebral hemispheres. A further 8 mm posteriorly
and immediately behind the optic lobes, the width is 8 mm. The volume is then

DINOSAURS OF AUSTRALLA
 calculated assuming illat the brain can be approximated as the frusta of two el-
Iiptical cones. Since the volume of the cerebellum cannot be even roughly ap-
proximated, the total volume ofthe endocast calculated in this manner, 1.3 mL,
is probably a minimum estimate. Assuming a specific gravity of 1.0 g/mL for
brain tissue, as Jerison (1973) does, gives an estimate of 1.3 g for the weight of
the brain of L. amicapaphica if the entire volume measured was in fact occu-
pied by brain tissue.
In dinosaurs, Jerison (1973) assumes that 50% of the endocast volume was
occupied by brain tissue. Hopson (1979) points out that in a modern Caiman
the anterior area of an endocast is filled almost completely with tissue of the ce-
rebrum. It is in the posterior area of the endocast, particularly above the cere-
bellum, that much of the volume is occupied by tissue other than that of the
brain. As the cerebellar area cannot be included in the volume estimate and the
area that can be measured was probably occupied almost entirely by brain tis-
sue, the estimated weight of 1.3 g is probably a reasonable minimum value for
the brain of L. anzicapaphica.
Unfortunately, very little of the skeleton of L. amicagraphica is known. The
femur ofNMV PI85 992 which almost certainlv is 1.art of the same individual as
J

the endocast, is not q~litecomplete. Its length, when complete, would have been
- 78 mm. DeBeer (1954) gives a measurement of 58 mm for the length of the
British Museum specimen ofArchaeopteryx lithographica. As the body shapes
ofL. ainicagraphi~aand A. litlzographica are roughly similar and the femur is
about a third longer. in L, amicagraphica, it is reasonable to assume that the
body weight was -1.34~ or 2.4 times that ofA, lithographica.
Hopson (1977: fig. I ) uses the bodyweight of400 gforA. lithographica in his
log-log plot of brain and body weight. On this basis, the weight of the type
specimen of L. amicapaphica would have been - I kg.
When these values are plotted- k = 0.005, body mass = 1000 g, brain
weight = 1.3 g-on Hopson 1971: fig. I (Figure 8 here) -the brain for this par-
ticular L. amicag-aphica appears to score higher than expected for the brain of
an archosaur; EQ= 2.6.
Given all the assumptions illat had to be made to estimate EQ, for the one
available endocast ofL. amicagraphica, it was desirable to develop a n alterna-
tive way to calculate EQ that would rely on a somewhat different set of assump-
tions. The approach taken was to use the fact that the endocast of L.
amicqqraphica, as far as is known, looks cluite similar to that of Stenonycho-
saw& inequalis. By comparing the areas that are well represented in both to
their body weights, a second estimate of EQ can be made.
Because this approach for estimating EQ has never been used, the derivation
of equation [4]and the basis for the constant k' are given on page 47.

In equation [4], Ai is the observed area ofthe endocastfrom the anterior tip of
the cerebral hemispheres to the posterior edge of the optic lobes; P is the esti-
mated body weight; and k t is a n empirically determined constant that presurn-
ably applies to all archosaurs with a n endocast similar in shape to those ofS. in-
equalis and L. amicagraphica. Substituting the appropriate values of Ai, k',
and P for L. amicasaphica yields a n estimate for EQ of 2.3.
This rough agreement for the estimate of EQ (2.3 and 2.6) calculated in quite
different ways gives confidence that the true value is close to, if not within, that
range. However, the single available specimen of L. amicagraphica is not an
adult. At most, it was approximately one third of its adult weight at the time of
death. As EQ is calculated for presumed adults in Jerison (1973) and Hopson
(19771, an allowance must be made for this factor before meaningfid compari-
sons can be made of the EQ ofL. amicagraphica. Equation [5] gives a n estimate
of the adult encephalization quotient EQ, of L. amicagraphica (see the deriva-
tion of this equation on page 47).
EQ;, =EQ~P'"
k~ni/~ [51
If, in fact, the adult body weight P, was three times the I kg suggested for the
 adult of the individual with the endocast P, the adult EQ for L. amicupaphica
would be in the range 1.6 to 1.8, given the EQrange estimated for the immature
specimen, 2.3 to 2.6. Hopson (1977) gives a range of 0.9 to 1.5 for the EQ of or-
nithopods. IfL. arnicagraphica had a n adult body mass of 10 kg, its EQ range
would be 1.1to 1.2, well within the ornithopod range. It should be borne in
mind that the smallest ornithopod that Hopson (1977) uses to establish the EQ
range for ornithopods was approximately two orders of magnitude greater in
body mass than L. umicagraphica. It may well prove that small ornithopods,
like small theropods, have a much larger EQ than larger ones.
ORBIT: Bccause t l ~ epostorbital bone and the ventralmost part of the lacrimal
are missing, the diameter of the orbit is uncertain. However, the minimum an-
teroposterior diameter of'the orbit measured between the top of,the dorsal pro-
cess of the jugal and the anterodorsalmost part of the anterior process of the
jugal bordering the orbit is 20 mm. Dorsally, it is 22 inm anteroposteriorlybe-
tween the posterior border ofthat part of the prefrontal, forming the anterodor-

Fi@~re8. B1~in:bodysize ~~elatiorzs

of Archaeopteryx and selected dine-
saul-s 012 minimuin convex ,LIo~}J$o~s
of extant reptiles, birds, and mam-
~r~als. The heavy line irldicutes the ex-
pectecl br-ain:bodl~~rcights for
individuals of Leaellynasaura amica-
graphica that were nzore mature
than the holotypc. The scale is log-
log. Synibols-A=hchaeopteryx; 01,-
nithopods: Art =Allatosa~~'~~s,

sal border of the orbit and the most posterior part of the orbital border
preserved on the frontal. Therefore, the original orbital diameter was probably
in the range of 22 to 25 mm. From the preserved anterior edge of the nasals to
the parietal-supraoccipital suture is 60.4 nun. The ratio of the orbital diameter
to the distance between the anterior tip of thc nasals and the yarietal-supraoc-
cipital suture is, therefore, in the range of 0.36 to 0.41. For Dryosuurus nltus,
the same ratio is 0.41 (Galton 1983: fig. 2), and for Hypsilophodo~~ fo.xii, 0.46
(Galton 1974a: figs. 4a, 5b). These recoi~structionsappear to indicate that all
thrce species had similarly rather l a ~ ~ gorbits.
e
FEMUR: In medial view, the shaft ofthe femur is concave postei-iorly, but some-
what less so in the other Victorian ornithopods. In posterior view, the medial
side is straight and the lateral side, slightly concave. At the level of the fourth tro-
chanter, the cross section of the shaft is nearly circular except for two straight
segments on the medial side.
The lesser trochanter is well below the greater trochanter. A prominent
groove separates the two trochanters on the lateral side of tlle shaft of NMV
P179564. An equally pronlinellt groove between them is visible on the posterior
side of NMV Pi85 980. Neither specimen displays any indication of the groove
the other clearly shows. No cleft is evident between the two trochanters. In dor-
sal view, ihe greater trochanter plus the head are anteroposteriorly compressed.

DINOSAURS OF AUSTRALIA
 Together the two are trapezoidal in oulline, the lateral edge of the greater tro-
chanter extending slightly farther anteriorly and posteriorly than the head.
There is no clear separation between the two, and they are of equal height. The
fourth trochanter is pendant. A depression for them. caudlfemoral longus is
well developed at the base of the fourth trochanter on NMV P185980. 011 all
other specimens, this area is too damaged to allow certainty as to whether tlle
depression was present. The fourth tochanter is located one third to two fifths
the distance from the proximal to the distal end of the femur.
A shallow but distinct intercondylar groove was present. In outline, the distal
end is roughIy an elongate rectangle 16 x 6 mm in the case of NMV PI82968
and - 12 X 6 mm in the case of NMV P185980. Both condyles are bulbous in
shape, with the medial noticeably larger than the lateral. $om both condyles,
low ridges extend proximally - 60% of the distance to the level of the fourth t o -
chanter. Such ridges are present on Fulgurotherium austrah but they do not
extend as far proximally. Lateral to the lateral condyle and the ridge extending
proximally from it is a distinct groove.
With the exception of NMV PI79564 and NMV Pi81 681, all the femora re-
Table 1. Leaellynasaura
arnicagraphic&sp. nov., ferred to L. amicagraphica appear to be from juveniles. Regardless, thesefemo-
Femur Lengths ra stand in marked contrast to those ofFulgurolherium austrule and Victorian
Ornithopod Femur Type I in that the distal ends are markedly anteroposteriorly
Specimen No. Femur Length (mm)
compressed and there is no clear division between the greater trochanter and
NMV PI79561 69 the head. That the former difference is not merely an artifact of preservation is
NMV PI79564 slightly 7113 demonstrated by NMV PI85980 which shows few, if any, signs of distortion.
NMV Pi01681 103 They stand in marked contrast to these two species and Victorian Ornithopod
NMV 1'182968 63
Femur Type 2 in that the ridges ex?endingpro&rnally from the condyles contin-
NMV PI85867 50
NMV PI85980 59
ue for 60% of the distance to the base of the fourth trochanter. They also differ
NMV PI85992 IGglestimated 781
from Victorian Ornithopod Femur Types 1and 2 in that the greater trochanter
is anteroposteriorly compressed.
The estimate of 78 mm for original length of the femur that probably belongs
to the holotype of L. amicapaphica, NMV P185992, was made by comparing
this incomplete femur with the approximately same-sized and virtually com-
plete femur NMV PI79561 (Table 1).The ratio of the distances from the dorsal
tips of the fourth trochanter to the distal end of the external condyle [NMV
P185992: 51mm; NMV PI79 561: 45 mm) was calculated and multiplied by the
length of NMV PI79 561.
COMMENTS: L, alnicagruphica was a small hypsilophodontid dinosaur. Mol-
nar & Galton (1986) compare the femora of fabrosaurids with those of hypsilo-
phodontids and find that the former have a greater irochanter not expanded
anteriorly, posteriorly, and dorsally beyond the head; an anteroposteriorly flat-
tened head; and a larger lesser trochanter relative to the greater trochanter. The
femora ofL. amicagraphica are fabrosau~idlikein the first two characters, al-
though the greater trochanter is slightly more anteroposteriorlyexpanded than
the head. The association of this femoral form with a skull in which the cheek
teeth are recessed beneath a lateral projection of the maxilla as in hypsilopho-
dontids-rather than being marginal as in fabrosaurids-is the primary reason
for regarding L. amicapaphica as a hypsilophodontid rather than a fabro-
saurid, despite the form of the femora.
The holotype individual appears to be a juvenile no more than one third fully
grown. Its body weight was estimated at I kg and the brain mass at 1.3 g. The
endocast of the brain is more similar to that of the small theropod Stenonycho-
saurus inequalis than to that of larger ornithopods. However, endocasts of
small ornithopods have not been previously reported. Thus, the similarity may
in part be owing to the fact that the endocasts of L. amicagraphicu and S. irze-
qualis are those of relatively small dinosaurs.
' One feature ofL. amicag'aphica is comparable across a spectrum of hypsilo-
phodontids: the proportions of the nasals. They are relatively long and narrow
in Hypsilophodon foxii (Galton 1974a), Zephyrosaurus schafi (Sues 19801,
and possibly Parksosau?-uswan-eni (Galton 1973), as well as in L. arnicapa-
phica. In contrast stands Dryosaurus (Galton 1983), which has broader and
shorter nasals. Although this feature is comparable across a number ofhypsilo-
phodontids, its phylogenetic significance is uncertain.

RICH & RICH

 KNOWN DISTRIBUTION: Latest Aptian-early Albian (Early Cretaceous), gen. nov.
Atlascopcosau~~us
southern Victoria, Australia.
DIAGNOSIS: Distinguished from all other l~ypsilophodontidsexcept Zeplg~ro-
saul-us by location of the apex of the upper cheek teeth close to tlle rear of the
tooth rather than about midway between the anterior and posterior borders.
Distinguisl~edfrom the upper cheek teetll of Leaellynasau7.a gen. nov. by the
presence of a ~ninimumof eight rather than a maximum of five ridges; ridges of
two distinct sizes rather than a gradation between ridges of maximum and
minimum strength; and ridges confined to the buccal surface rather than equal-
ly developed on both sides of the unworn teeth. Cheek teeth distinguished from
those ofMochlodon (Nopsca 1904, cited in Bartllolonlai & Mohlar 1981) by
having a weaker primary ridge. Upper cheek teeth further distinguished by the
position ofthe prilnaly ridge near the posterior margin instead of central on the
buccal surface of the crown. Lower cheek teetll distinguished from those of
Kanpasaurus (Cooper 1985) by the much weaker primary ridge. Cheek teeth
distinguished from those of Dryosa~lrus(Galton 1983) and Valdosaurus?
(Galton 63 Taquet 1982) by lacking a lozenge-shaped crown and by possessing
stronger secondary ridges and ridges that cover all the buccal surface ofthe up-
pers and lingual surface of tlle lowers rather than confined to the toot11 nmrgins.
Cheek teeth distinguishedfrom those of Calnptosaurus (Galton 69 Powell 1980)
and Tenontosaurus (Galton 1974b, Morris 1976) by same criteria as Dye-
sazlrus and Valdosaurus plus being lllarkedly smaller. Cheek teeth distin-
guished from tl~oseof Otknielia (Galton 1983) by a well-defined primary ridge
with secondary ridges stronger and extending nearly to the base of the crown.
Cheek teeth distinguished from those ofPlzyllodon, Nanosatirus, and Alocodon
(Galton 1983)by being markedly larger, and by possessing well-definedprima-
ly and secondary ridges rather than marginal denticles only. Upper cheek teeth
distinguished from those ofHypsilophodon (Galton 1974a) by having a distinct
primary ridge and more numerous and prominent secondary ridges. Cheek
teeth distinguished from those of Parlcsosau~n~s (Galton 1973) by a distinct,
markedly larger primary ridge. Cheek teeth distinguished from those of Ze-
phyrosaurus (Sues 1980) by the stronger primary ridge and the greater differ-
ence between it and the other ridges. Upper cheek teeth distinguished from the
Lightning Ridge, isolated l~ypsilopl~odontid cheek toot11 figured by Molnar
(1980: fig. 5, QM F9505) by location of the primary ridge near the posterior
margin of the buccal surface of the toot11 rather than near the midpoint. Lower
cheek teeth distinguished from the Lightning Ridge specimen by ventral mar-
gins of the crown that diverge from one another proceeding apically from the
root at an angle of -70" rather than 45".
ETYMOLOGY: Named in honor of the Atlas Copco Corporation, which sup-
plied the pneumatic tools and accessory equipment that enabled the excava-
tions at Dinosaur Cove.
HOLOTYPE: NMV P166409, left maxilla fragment with one erupted cheek Atlascopcosuul.us loadsi
tooth and three unerupted cheek teeth wllicl~have been exposed by removal of sp. nov.
tlle enclosing bone (Figure 9).
TYPE LOCALITY AND STRATIGRAPHIC POSITION: Point Lewis, Otway
Group, latest Aptian-early Albian.
DIAGNOSIS: That of genus until other species are described.
ETYMOLOGY: In honor of William Loads in appreciation of his unfailing faith
in and support for the Dinosaur Cove project.
REFERRED SPECIMENS
Point Lewis.NMV P157390: left maxilla with seven cheek teeth. Same local-
ity and stratigraphic position as Ilolotype (Figure 10).
Dinosaur Cove West. NMV P177934: isolated, left lower cheek tooth.
Dinosau7- Cove East. NMV P181679: isolated, left upper cheek tooth. NMV
PI82962 left dentary fragment with I1 empty alveoli along the buccal margin
and three unerupted teeth lingual to them (Figure 11). The position of the most
anterior tooth in this dentary is uncertain owing to damage sustained by the

DINOSAURS OF AUSTRALIA
 specimen when it was uncovered wit11 a pneumatic rock breaker. NMV
P185970: left maxilla fragment with 10 empty alveoli along the buccal margin
and three unerupted teeth lingual to them.
SlipperyRock. NMVP181677: a n isolated, left upper cheek tooth of ajuvenile
missing the crown posterior to the primary ridge. NMV PI86 003: isolated, left
lower cheek tooth. NMV P186847: right anterior, dentary fragnlent with tips of
three unerupted teeth visible. NMV P187116: isolated, right upper cheek tooth.
DENTITION: Based on the alveoli, the holotype ~ossessedat least 10 upper
cheek teeth. Except for the one or two most posterior of these, all adult speci-
mens are approximately the same size when unworn; the crown having a maxi-
mum anteroposterior dimension of - 7 mm, the smaner ones being -5 min.
Alveoli in the most complete mandible available, NMV P182967, indicate the
presence of at least 11 functionaI teeth. The alveoli are largest in the middle of
this series and decrease toward both ends.
I
! Figlli'e 9. A t l ~ ~ s c o p c o s a laadsi
u~~~s
: la1G1i~1U.9: (left),
J I O ~ O ? J ; I C ,NMV
i ~ ~ I I I T C tlr1~1.11pted
I lnter~nl1 1 i ~ qof
cltCch-tnntlr it1 cc r)~n~.il!a Jis~rgrnrr~i
(Xi); A (~*iglit), 1t1~~1iol I ~ ~ AoJ'tltc
IJ
rtrn.rillnS,.t~,q)rrcrr~( X U ; H (left),
lntr~r.nl1~ie111of tire p~sfer'i~r.tr~ost
rtrlc~.rcpt~d11pperc l ~ e ~loot11k
(Xl.SG); IJ (riglit), Intc~.al vie^^^ 01'
'

.i e1.11ptec1~rppet.d r ~ c ktootli W?).

NMV PI57390 suggests that i n the upper cheek teeth, the length of the Zahn-
reihen or anterior-to-posterior waves of tooth replacemellt was slightly < 3.0.
Ridges are developed on both sides of the unerupted cheek teeth. On the buc-
cal side of the upper cheek teeth and on the lingual side of the lowers, a single
primary and seven to 14markedly weaker secondary ridges extend vertically or
nearly so from the occlusal crest ofthe tooth to the base of the crown. The anteri-
or and posterior margins of the crown, from root to widest point, is bounded by
a distinct ridge about as strong as the secondary ridges, except for the one on the
anterior edge of the upper cheek teeth. That ridge projects noticeably farther
away from the body of the tooth than do the secondary ridges.
On the opposite sides of the unworn cheek teeth, lingual in the uppers and
buccal in the lowers, a distinct primary ridge is absent, and faint secondary
ridges extend from the crest of the tooth less than one fourth the distance to the
base. It is these surfaces that contacted one another when the teeth were func-
tional. Conseguently, on the teeth that were functional,the secondary ridges oil
the occlusaI surfaces were obliterated.
The buccal surface of an upper cheek tooth may be immediately distin-
guished from the lingual surface of a lower cheektoothby three criteria. First, in
the uppers, the primary ridge is located close to the rear ofthe buccal surface. In
contrast, the primary ridge on the lower cheek teeth is approximately at the
midpoint of the lingual surface, giving a symmetrical appearance to the tooth.
Second, the outline ofthe occlusal crest in unworn teeth in buccal view is round-
ed on the lowers while being two straight segments that meet at an angle of - 80"
at the apex of the primary ridge on the uppers. Third, the anteriormost ridge on

RICH & RICH

 Figure 10. Atlascopcosaurus
Ioadsi, N .P157390, maxilla
with upper cheek teeth: left, lat-
eral view; right, medial view.
(X 11

Figure 11. Atlascopcosauius loadsi:

A (left), NiMV Pi82967n, r7lnr1dible
with two unerupted cheek teeth, Eat-
era1 vim (X O.(iG); A (right), NMV
P182967a, rnar~diblewit11two 11 ner-
upted cheek teetlz, ?nedinl view
(X 1.33); B (left), NMV Pi829670
(sair~einclivid~ialas NMV P182967a),
Eo~vei.cheek teetlz, lutcr-a2 v i w 1x21;
B (right), NMV P I 77,934, lower
cheek teeth, lateral view 1x2).

the upper cheek teeth is prominent, as mentioned above.

In passing from the crest of the upper cheek teeth toward the base of the
crown, the one or two seconda~yridges closest to the primary ridge converge
toward it. The secondary ridges farther from the prima~yridge curve slightly
and run parallel to the primary ridge as they approach the base of the crown.
On the two known specimens with worn upper cheek teeth (NMV PI57390
and NMV PI81677), one of two wear facets is present on the lingual side of each
functional tooth. The facets appear to be flat or slightly concave in the frontal
plane. Adjacent teeth may share a common facet. With wear, the occlusal mar-
gin ofthe cheek teeth is a straight line or nearly so. The single worn, lower cheek
tooth known (NMV P177934), has a single flat wear facet, and the occlusal
margin of that wear facet has serrations formed by secondary ridges as also oc-
cur on the upper cheek teeth.
NMV PI81677 is the one juvenile specimen ofA. loadsi known. Moderately
worn near the tip on the lillgual side, it is clearly an upper cheek tooth, because
the most anterior secondary ridge projects farther from the body of the tooth
than do the other secondary ridges, and the profile of the occlusal edge is
straight, not rounded. The anterior edge of this tooth is 1.8 mm from the center
of the primary ridge. This contrasts with a range from 4.2 to 5.6 mm for the sim-
ilarmeasurements on all the other cheek teeth ofA. Zoadsi. Other than its dimin-

DINOSAURS OF AUSTRALIA
 utive size, no feature distinguishes it from the other specimens of A. loadsi. It
has a well-developed primaiyridge and five secondary ridges anterior to it. The
secondary ridges on the lingual side of the lower cheek teeth, anterior to the pri-
mary ridge, converge on it toward the root of the iooth at a slightly greater angle
than the ones posterior to the primary ridge, which are almosi parallel to ii.
MAXILLA: None of the three maxilla fragments is complete. However, among
them. much of this element can be reconstructed.
Behind the most posterior cheek tooth, the maxilla expands laterally to form a
horizontal plate. On the dorsal surface of this plate are a series of parallel striae.
This was the area of contact with thejugal. Similar parallel striac are developed
in the posterodorsal region of the internal surface of the maxilla, ihe region
where it contacted the palatine.
The external surface of the maxilla immediately above the cheek teeth is in-
clined dorsolaterally for 7 to 10 m m and then turns 90' to a dorsomedial inclina-
tion. On the dorsolaterally inclined surface are as many as four foramina.
On the dorsal surface of the maxilla, beginning above -thefourth most posteri-
or cheek tooth and extending antericrly is a deep groove. The groove extends
forward from this point parallel to the row of alveoli for the finctional cheek
teeth. In one of the two specimens where the groove was visible, NMV PI66409
(the other being NMV P182967)) a prominent foramen occurs behind the pos-
terior end of the groove. This groove and the prominent foraillen behind it
(when present) were probably the location of the dental lamina.
Galtoil (1983) reported horizontal foramina on the medial surfaces of tlie
dentary and maxilla of onc specimen ofDryosaurus altus. No such foramina
are visible on either the dentary or n~~vrillae of A. loadsi.
DENTARY: The mandible is represented by two specimens. NMV PI82 967 is a
left dentary, missing the anterior moiegr and slightly crushed on its internal
side. Even thougl~the areas of contact with the splenial and prearticular are
preserved, damage to the mandible is so extensive in this area that the actual
surface of contact cannot be seen. Only the point of contact wit11 part of the ven-
tral border of the surangular is preserved. Tlie transverse cross section of the
dentary is a nearly perfect, outwardly convex semicircle. The depth of the den-
i a ~ beneath
y the alveoli of the cheek teeth- -20 mm-is nearly uniform along
the preserved section of the dentaiy.
NMV PI86847 is tlle second mandibular specimen, a right syrnphysial re-
gion with the seven most anterior alveoli and three unerupted cheek teeth pre-
served. The Mecklian canal is a well-defined groove running along the ventral
border of the median surface. Anterior to that is the region of symphysial con-
tact with the opposing dex~tary.
COMMENTS: Although the association of the upper and lower dentitions here
referred to A. loadsi is not a certainty, it is highly probable. Both occur at two
sites, Dinosuar Cove East and Slippery Rock. The form of the primary and sec-
ondary ridges is quite similar, and the uppers and lowers are about the same
size. Although the three differences between the upper and lower cheek teeth
listed above (position of the primary ridge, presence or absence of a noticeably
stronger anteriormost secondary ridge, and outline of the crown) are consistent
and readily recognized, none ofthe differences suggests that these various spec-
imens belong to different species.
The presence of a primary ridge and secondary ridges that extend the entire
distance from the occlusal margin ofthe tooth to the base ofthe crown separates
A. loadsi from known fabrosaurids and heterodontosaurids. Further distin-
guishing A. loadsi from fabrosaurids, the upper cheek teeth are recessed be-
neath a lateral expansion of the maxilla, rather than being marginal, and the
lower cheek teeth are in massive rather than narrow jaws. A strong median
ridge characterizes iguanodontids. Because the primary ridge is not medially
located on the upper cheek teeth of A. loadsi but is displaced posteriorly, and
because of its small size, it is not regarded here as an iguanodontid.
Within the Hypsilophodontidae, A. loadsi is about equally close dentally to
Kangnasaurus coetzeei (Cooper 1985)) Thescelosaurus neglectus (Galton
1974b1, Hypsilophodon foxii (Galton 1974a), and Zephyrosaurus scha$

RICH & RICH

 [Sues 1980). Lower cheek teeth ofA, loadsi have a much weaker primary ridge
than that on the one cheek tooth known of K . coetzeei. Otheiwise, the lower
cheek teeth ofA. loadsi are quite similar to those of the K. coetzeei specimen.
Both A. loadsi and K. coet~eeihave a single prominent primary ridge and a se-
ries of weaker secondary ridges that are subparallel to the primary one.
A. loadsi differs from T,neglectus (Galton 1974b, Morris 1976)principally in
that the primary ridge and apex of the upper cheek teeth are close to the posteri-
or border of the crown rather than approximately midway between the anterior
and posterior borders.
In the case of the upper cheek teeth of H.foxii (Galton 1974a), although the
apex is shifted somewhat posteriorly, approaching the more extreme displace-
ment seen in A. loadsi, the primary ridge in H, foxii is not as prominent.
The teeth of 2. schaJjt (Sues 1980), with their posteriorly located apexes on
the upper cheek teeth, resemble unworn teeth of A. loadsi. The much weaker
developnlent of the primary ridge on 2, schafl sets it apart.
The intrafamilial relationships of the l~ypsilopl~odontidsknown from partial
skulls and skeletons is poorly understood; e.g., Cooper (1985), Galton (1983),
Sues (1980). Given the fragmentary nature of the material, little more can be
said concerning A. loadsi except that it belongs within the family.
MATERIAL Fulgurother.iurn australe
Eagles Nest. NMV PI50 054: right femur lacking only the condyles and proxi- von Huene 1932
ma1 regions of the head and greater and lesser trochanters (Figure 12). NMV
P185948: anterointernal third of midshaft with base of capitulum of right fe-
mur. NMV P187094: partial left femur with capitulum, partial fourth trochan-
ter, and distal end; shaft between capitulum and fourth trochanter missing.
The Arch near Kilkunda. NMV P164998: left femur lackinghead and medial
half of greater and lesser trochanters (Figure 13).
Dinosaur Cove East. NMV P177935: right femur lacking the distal fourth of
the bone with much of the capitulum abraded away (Figure 14). NMV
PI85 961: left femur fragment with capitulum, greater and lesser trochanters
abraded, missing fourth trochanter and all of the bone distal to that point.
DESCRIPTION: I11medial view, the shaft is anteriorly convex. In posterior view,
the medial border is slightly concave and the lateral border, virtually straight.
From the midshaft proximally, the shaft has a roughly triangular cross section
and distally the shaft is more nearly oval in cross section.
The rodlike lesser trochanter is markedly smaller than the greater trochanter
from which it is separated by a shallow, narrow cleft. On the lateral side of the
femur, a well-developed groove begins at the base ofthis cleft and continues dis-
tally halfway to the base of the fourth trochanter. No corresponding groove is
present on the medial side of the femur. The lesser trochanter was almost cer-
tainly well below the top of the greater trochanter. The greater trochanter is ex-
panded somewhat anteriorly and posteriorly beyond the head and probably
was higher than the head as well.
The fourth trochanter is large, pendant, and located about one third the dis-
tance from the proximal to the distal end of the bone. A faint scar, presumably
the site of attachment of the M , caud~emora2i.slongus, is present on NMV
PI64998 at the base of the fourth trochanter, but not on NMV Pi50054.
An extremely sl~allowanterior intercondylar groove is present at the distal
end of NMV PI64 998, but no such groove is evident on NMV Pi50 054. The me-
dial condyle has a strikingly flat medial surface. The lateral condyle is a medio-
laterally compressed blade in contrast to the much wider medial condyle. Both
extend about the same distance posteriorlyfrom the shaft. There is a well-devel-
oped groove on the external side of the lateral condyle. In distal view, the lateral
condyle is aligned slightly posterolaterally from the shaft. But the inclination is
<. 30' that Molnar 63 Galton (1986) report for the F. australe material from
Lightning Ridge. The lateral condyle continues proximally as a low ridge that
disappears about halfivay to the base of the fourth trochanter. No similar ridge
is associated with the medial condyle. In distal view, the outline of the condyles
is roughly rectangular, being 32 mm wide and 25 mm deep on NMV P164998.
The shaft immediatelyproximal to the condyles is 27 mm mediolaterally and 16

DINOSAURS OF AUSTRALIA
mm anteroposteriorly on NMV Pi64998. On NMV P150054, the same mea-
surements of the shaft are 29 mm and 18 mm.
COMMENTS: Molnar & Galton 11986) give a number of criteria for assigning
the femora off. australe to the Hypsilophodontidae. The same criteria distin-
guish the Victorian femora of this species. These femora are distinguished from
iguanodontids by the fourth trocl~anter'slocation on the proximal half of the
shaft. From fabrosaurids, they are distinguished by the greater trochanter's ex-
pansion anteriorly, posteriorly, and dorsally from the head and separationfrom
it by a shallow sulcus. They are further distinguished by the lesser trochanter's
being smaller relative to the greater trochanter and the greater trochanter's not
being a~lteroposteriorlycompressed.
Molllar 63 Galton (1986) widely compare the Lightning Ridge femora they re-
fer to F. australe with the femora of other hypsilophodontids. With minor ex-
ceptions, the Victorian specimens display the same differentiating features.
Molnas & Galton (1986) consider F. australe from Lightning Ridge to be most

similar to fiypsiloplzodon and Othnielia. But, like the Lightning Ridge femora,
the F. australe femora from Victoria differ from Hypsilophodon and Othnielia
by having a thin, sheetlike lateral condyle, and a lesser trochanter that is lower
than the greater trochanter rather than subequal to it in height. Furthermore,
both the Lightning Ridge and the Victorian F. australe materials differ from
Hypsilophodonby having a shallow, rather than a deep, sulcus 011 the posterior
face of the head; and a triangular, rather than a cpadrangular, cross section of
the shaft immediately above the fourth trochanter.
Because NMV PI77935 lacks the distal end, this specimen from the Otway
Basin was allocated to the same species as the other femora here referred to F.
australe from the Gippsland Basin, with some hesitation. In the features avail-
able, it displays all the differentiating character states that set apart the femora
referred to F. australe from the Lightning Ridge and the Gippsland Basin from
other ornithopods. It is noticeably larger than all the Gippsland Basin speci-
mens. If complete, it would have had a length of - 200 mm, while the Gippsland
-
Basin specimens would all have been 150 lnm long. However, this larger size
does not preclude it from F. australe, as AM F66764 from Lightning Ridge
would have been about the same size, and the other Lightning Ridge specimens
referred to this species are noticeably smaller than those from the GippslandBa-
sin [Molnar & Galton 1986).

RICH & RICH

 - -A" < >a-

Figure 13. Left fen^ of FulgU

eriuln australe, NMI 'I64998:
Oeft), anterior vimq (right),
tcrior i~iplv;B (left),edial v i n
(right), lateral vim?, ,' distal 1
cx0.n

s*_/

-
MATERIAL Victoria11Hypsilophodorat i cl
Dirzosaur- Cove E a s t N M V Pl.85976: left femur missing condyles. NMV Pernur Type 1
P185995: left fernur missing condyles. NMV P185999: left femur missing only
distal tip of internal condyle and most posterior part of capitulum (Figure 15).
Slippev Rock. N M V PI86 004:right femur missing most of the condyles and
proximal tips of greater trochanter and capitulum.
Slippely Rock or Dinosaur Cove East. N M V P187l.15: distal half of the left
femur.
DESCRIPTION: In medial view, the shaft has a pronounced curve, convex ante-
riorly. In anterior view, the sides of the shaft are straigllt and parallel except at
the distal end where the two borders diverge slightly from one another. In cross

DINOSAURS OF AUSTRALIA
 section at and slightly above the fourth trochanter, the posterior, lateral, and
medial sides are each a straight line. The lateral and medial sizes arc approxi-
mately parallel to one another or converge slightly anteriorly. The posterior side
varies 5 20" from being perpendicular to both. The anterior side is a nearlyper-
fect semicircle in cross section.
The greater trochanter is a mediolaterally compressed blade that extends
both anterior and posterior to the base ofthe capitulum. The lesser trochanter is
a mediolaterally cbmpressed blade that terminates dorsally below the greater
trochanter. The two are separated dorsally by a shallow cleft. Laterally, a well-
developed groove continues distally between the greater and lesser trochanters.
The fourth trochanter is enda ant and located about two fifths the wav from the
proximal to the distal ends of the femur.
The depression for the M. caudifemoralis longus is strong on NMV Pi85976
and alnlost undetectable on NMV Pi85995. The de~ressionis on the medial
surface of the shaft immediately adjacent to the base &fthe fourth trochanter. A
very shallow anterior intercondylar groove at the distal end of the femur con-
trasts markedly with the narrow, deep posterior intercondylar groove. The me-
Table 2, Victorian
Wypsilophodontid Femur dial condyle is wider mediolaterally and projects much farther posteriorly than
T y p e 1,Lengths the lateral condyle. The outline of the distal end of the shaft is a parallelogram
slightly wider than long. The angles of the parallelogram vary considerably
Specimen No. Femur Length (mm)
from one specimen to the next. For example, the internal angle of the anteroex-
N M V PI85976 83 ternal corner varies from 60 to 120".
NMV PI85995 >77
NMV PI85999 89
COMMENTS: The marked anteroposterior expansion of the greater trochanter
N M V PI86004 >46 and relatively small size ofthe lesser trochanter readily distinguish these femora
from those of fabrosaurids. As in all the Victorian hypsilophodontids, the posi-
tion of the fourth trochanter proximal to the midpoint of the shaft sets these fem-
ora apart from those of iguanodontids.
The distal end differs markedlv from the femora of Leaellvnasaura amica-
@a.phica in that it is not anteroposteriorly compressed to the same degree and
the ridges fiom the condyles do not extend as far proximally along the shaft. The
distal end differs from that of Ful~rotheriumaustrale in that the medial con-
V

dyle projects much farther posteriorly from the shaft than does the lateral con-
dyle rather than the two being subequal in this regard, and there is no groove
external to the lateral condyle. The proximal ends differ from those of all other
l~psilophodontids(Hypsilophodon and Dlyosaurus [Galton 19751, Kangna-
saurus [Cooper 19851, Othnielia [Galton G3 Jensen 19731, Valdosaurus [Gal-
ton & Taquet 19821, and Yandrlsaurr~s[We & Cai 19841) in that the greater and
lesser trochanters are markedly expanded anteroposteriorly and compressed
mediolaterally so that it has a bladelike structure (Table 2).

Victorian Hypsilophodontid MATERIAL

Femur Type 2 Cape Paterson. NMV P156980: right femur missing the head, much of the
greater trochanter, tip of the lesser trochanter, and the condyles (Figure 16).
DESCRIPTION: In medial view, the shaft is concave posteriorly. In posterior
view, both the medial and lateral sides are straight. The lateral corner of the
shaft becomes abruptly acute midway between the bases ofthe lesser and fourth
trochanters. From that point distally as far as the shaft is preserved, the cross
section is sharply triangular. The surface at the distal end of the femur where
the condyles would be located ifthe bone were complete, does not face approxi-
mately posterior as in the other Victorian hypsilophodontids but rather, poster-
ointernally. Measured with respect to the position of the fourth trochanter, the
difference in orientation is -45".
Tlie lesser trochanter is separated from the greater trochanter by a deep
groove on the lateral surface and a shallow one on the anteromedial side. The
greater trochanter extends for some distance posterior to the lesser and was
probably mediolaterally compressed, but the state ofpreservation is inadequate
to be certain. The fourth trochanter is damaged on its dorsal side, giving the
false impression that it is directed posteriorly rather than pendant, the normal
condition. The fourth trochanter is located - 45% ofthe distance from the proxi-
mal to the preserved distal end, thus, clearly proximally located. However, the

RICH 61 RICH
 condyles are not preserved, so the fourth trochanter was located even farther
prolrimally when the bone was complete. The area where a depression for the
M , caud$ernoralis lorzgus would be expected is damaged.
The posteromedial surface of the femur has no well-defined proximodistally
directed ridges that might be a continuation of the condyles. The 1.egio11of the
shaft immediately abovc the condyles is roughly rectangular in cross section
wit11 an anteromedial length of 60 mm and anterolatel-a1width of -31, mm.
The preserved length ofthis femur is 290 mm. As the condyles are missing, it
was clearly longer than this when complete. On a referred Victorian specimen
ofF. australe (NMV Pi64 998) the proxirnodistal di~nensionsof the condyles is
one sixth the total length of the bones. Therefore, a reaso~lableestimate for the
length of this femur when it was complete is 350 mm.
COMMENTS: The relatively small size of the lesser trochanter relative to the
greater trochanter sets this femur apart from those offabrosaurids. The position
of the fourth trochanter proxinlal to the midpoint of the shaft distinguishes it
fiom those of ca~nptosauridsand iguanodontids.
Not only is Victorian Hypsilophodo~ltidFemur Type 2 markedly larger than
all other Victorian hypsilophodontids, but it differs from them morphologically
in two ways. First, the distal end is "twisted" by 45". This twist so that the sur-
face once bearing the condyles faces postero~nediallyinstead ofposteliorly does
not appear to be an artifact caused by distortion. Although the surface of the
specimen shows cracks in many places, there are no i~ldicaiionsof compressio~l
or any other form of distortion. Second, the lateral corner ofthe shaft abruptly,
rather than gradually, changes from a smooth, rounded surface in the vicinity of
the greater and lesscr trochanters to a cluite sharp one midway between those
trochanters and the fourth trochallter.
Furthermore, it differs from the femora of'Leaellynasaur.aamicagraphica in
that the shaft is not anteroposterio1.1y compressed. The rodlike character of the
lesser trochanter is markedly different from the mediolaterally compressed na-
ture of this feature 01.1 the Victorian Hypsilophodontid Femur Type I.
Of the other three Victorian ornithopod femoral groups recognized here, the
closest resemblance is with Fulgurotherir~maustrale. Were it not for the twist
in orientation of the distal end, this specimen would be included in the group
allocated to that species. However, in the small numbers of ornithopod femora
available, this amount of rotation of the distal end appears greater than to be ex-
pected in a single species.
The single specimen ofthe Victoria11Femur Type 2 is so damaged that further
comparisons between it and previously published descriptions of other hypsilo-
phodontids were not feasible.

Discussion
Owing to the nature of the holotype ofFulgurotherium australe, it is not
certain that the other specimens referred to this species definitely belong
there, However, the name is available and, for the present a t least, it
makes a convenient label for this assemblage of femora which probably
in whole o r in part a t least are generically distinct from other hypsilo-
phodontids. The range of variation of the specimens referred here to F.
australe is comparabIe with that which Molnar 63 Galton (1986) al-
lowed in their allocation of femora assigned to this species from Light-
ning Ridge, New South Wales, Australia. The amount of variation
observed in the F. australe femora from both Lightning Ridge a n d Vic-
toria is noticeably greater than that seen i n either Leaellynasaur.~ami-
cagraphica o r the Victorian Hypsilophodontid Femur Type 1sampIes.
However, in all cases, samples are few, less than a dozen in every case.
The Victorian Hypsilophodontid Femur Type 2 may b e an extreme
variant of those referred to F. australe. The twist of the distal end of the
single specimen of Type 2 clearly sets it apart from the other specimens

DINOSAURS OF AUSTRALIA
wv-M
*a#-&a,n
r. -* --'rru "uw n

Figure 14. Rightfenzur of Fu1gul7-

otheriunl australe, NMV P I 77935:
A (left), arzter'ior-view;A (right),
posterior view; B (left), medial
view; B (right), lateral view; C,
proximal view. (xO.51

referred to F. australe. However, the amount ofintraspecific variation in

the orientation of the proximal and distal ends of hypsilophodontid fem-
ora has never been quantified, and so few samples of actual femora are
available that the possibility of this specimen's being an extreme variant
of F, australe cannot be ruled out. The single specimen of Type 2 is so
battered that the presence or absence of the diagnostic features ofF. am-
trale cannot be determined. However, enough of it is preserved for confi-
dence that it is neither L.amicagraphica nor Type I.
The highly anteroposteriorly compressed distal and proximal ends of

RICH 61 RICH
 Figure 15. Victol-innHypsilopho-
dontid Femur v p e 1,NMV
P185999: A Oefi), a.nterior view;
A (right), poster-ior v i m ; B Cleft),
medial view; B (right), lateral vim
C (left), pr-oxinzal vim< C (right),
distal view. 1x 21

the femora ofL. amicagraphica and the mediolaterally flattened and an-
teroposteriorlyexpanded greater and lesser trochanters of the Victorian
HypsilophodontidFemur Type I clearly separate these two groups from
one another and all other hypsilophodontids as well.
Among the Victorian hypsilopl~odontids,the only reasonably confi-
dent association of teeth and postcranial elements, particularly femora,
in the same species is the material assigned to L. arnicagraphica. It is
represented by the partial skull and postcranial skeleton of one individ-
ual plus six referred, isolated femora. The holotype individual appears
to have had a deceptively large brain. However, this individual was aju-
venile at the time ofits death. Because brain weight increases in modern
Alligator mississippiensis (see page 47) as a function of the cubed root
ofthe increase in body weight, it appears that had the holotype ofL. ami-
cagraphica lived to adulthood, its encephalization quotient (EQ ofJeri-
son [I9731 and others) would have been lower. If as an adult it had
weighed 3 kg- the maximum size indicated for the species by the largest
referred femur- the EQ for an archosaur would have been in the range
1.6 to 1.8, slightly greater than the maximum 1.5 reported for ornitho-
pods in Hopson (1977). On the other hand, if its maximum body size

DINOSAURS OF AUSTRALIA
was 10 kg, its EQ would probably have been 1.1to 1.2, well within the
ol-nithopod range. The form of the brain in dorsal view is similar to that
of the small theropod Stenonychosaurus inequalis (Hopson 1979, Rus-
sell 1969). They differ in that S. inequalis has a higher EQ (5.8) as com-
pared with a maximum of 1.8 for L. amicag-aphica. L. amicagraphica
has a distinct pineal body while S. inequalis does not. However, other
small theropods do have such a pineal body; e.g., Dromiceiomi?nusbre- I
vitertius (see Russell 1972).
Dinosaur Cove has yielded several dozen isolated reptilian postcran-
ial elements that are clearly not turtles and, on the basis of frequency,
size, and ornithopod form, are probably parts of Atlascoposaurus
loadsi, the teeth ofwhich are found there. But no associated or articulat-
ed remains of the species demonstrate this beyond doubt. The existence
at Lightning Ridge of a small hypsilophodontid cheek tooth, unlikely to

Table 3. Distribution of Early Cretaceous Vertebrates at Principal Sites in

Victoria, Australia (Listed West to East)
Taxon present
? Taxon possibly present
- Taxon absent

Taxon Otway Basin Gippsland Basin

Dinosaur indet
Fulgurotherium australe
-- -. . . - . . . . - . - .. -- --- -
. - - - - - - - - a
Leuellynusaura
amicapaphica
~tlasco~cosaurus Ioadsi
- .. .. .
- ---.--------
Victorian Hypsilophodont
m e1 - - .. -- - - - - - - - - - -
- - - - - - - - - - - - - .
Victorian Hypsilophodont
w e2 - -
Theropoda
~llosaurussp. - - - -. -- - - - - - - - . -
- - - - - - - ? . - - - - - - -

Dinosaur footprint
Testudines -... -
- - - -
- - - - -I- - .
- - - - - - - - - -
- - I

Chelycarapookusarcuatus
Plesiosaur - . . -
- - - - - - - - -
- - - - - .
?Lepidosaur
Pterosaur
Aves
- - .
- - - - - - - - - -
? - - - - - -
- - - - - - - - - - - - - - -
- - - - - - - - - - - - - - -
Labyrinthodont
Pisces - - -..- -- -- -- --- -- -- -- --- -- - -
-- -- --- -- -- -- --- .- - - . ,
Ceratodontidae - - -
Ceratodus avus - - - - - - - - - - - - - -
Ceratodus nargun
Ceratodus sp.
Coccolepk woodwardi - - - - - - - - - -
Wadeicthys oxyops - - - - - - - - - -
Koonwarria mantfrons - - - - - - - - - -
Leptokpis koonwarri - - - - - - - - - -

be that ofA. loadsi, makes the association of the teeth ofA. loadsi with
the femora referred to F. australe from Lightning Ridge and Victoria far
from a certainty. The only other hypsilophodontid femur found at a site
which has yielded dental remains ofA. loadsi is the Victorian Hypsilo-
phodontid Femur Type 1. All Type I femora known thus far are much
too small to have come from individuals that produced most ofthe dental
material of A. loadsi. The only exception is a single, diminutive partial
upper tooth ofA. loadsi which is presumably a juvenile's. This specimen

RICH &+RICH
may have come from an individual with femora no larger than the larg-
est thus far known of the Victorian IIypsilophodontid Femur Type 1.
The teeth of L. amicagraphica and A. loadsi are clearly distinct from
one another. Both differ from the small hypsilophodontid tooth known
from Lightning Ridge, New South Wales, illustrated in Molnar (1980).
Associated Biota
The three or four hypsilophodontids (L. arnicagraphica, A. loadsi, F.
australe, and Victorian Hypsilophodontid Femur Type 1) known from
the Otway Group are all from sites dated as latest Aptian-early Albian.
The fauna associated (Table 3) with them consists of dipnoans (lung-
fishes), actinopterygians (bony fishes), testudines (turtles), pterosaurs
(flying reptiles), plesiosaurs, and small theropods. The last three goups
are known exclusively from Dinosaur Cove and are represented by only
a few isolated elements each: the pterosaurs by a ?tibiotarsus (K. Padian
and P. Wellnhoffer, personal communication), the plesiosaurs by isolat-
ed teeth. Apparently the plesiosaurs must have been freshwater forms,
for there is no other indication that the rocks of the Otway Group were
laid down in marine conditions. Although not common, freshwater ple-
siosaurs are known elsewhere in the world.
Other than dinosaurs, turtles are the most common elements in the
terrestrial fauna. They are represented by limb bones, vertebrae, and a
lowerjaw, as well as numerous shell fragments. These isolated elements
represent turtles so primitive that they cannot be assigned to either the
cryptodires or the pleurodires (E.S. Gaffney, personal communication).
One (possiblytwo) hypsilophodontid (F. australe and Victorian Hyp-
silophodontid Femur Type 2) is known from the StrzeleckiGroup in the
Gippsland Basin. The associated vertebrate fauna (Table 3) consists
solely of dipnoans, actinopterygians, testudines, a theropod (Molnar et
al. 1981), a lepidosaur [Molnar 1980), and possibly a labyrinthodont
amphibian Uupp &Warren 1986).Just as in the Otway Basin, the thero-
pod from the Gippsland Basin was small: Allosaurm sp., known from
Eagles Nest, was only -2 m high.
Birds are represented by only five feathers known from a single local-
ity in the Gippsland Basin, Koonwarra. Nothing concerning the familial
identity of these feathers can be discerned at present, despite a recent
study (Rogers 1987). The feathers do come from birds about the size of
crows, and several types of feathers are represented.
This same locality produced a variety of freshwater fishes, and many
of these are represented by complete skeletons: lungfishes [Ceratodonti-
dae), palaeonisciforms [Coccolepididae), archaeomaenids (Archaeo-
maenidae, Pholidophoriformes), koonwarriids and leptolepidids
(Koonwarriidae, Leptolepididae, both clupeiformes) (Waldman 19711.
But, thus far, not a single fossil bone of any tetrapod has been recovered
from the Koonwarra locale.
No mammalian remains have yet been found in the Victorian se-
quence, but in the Griman Creek Formation farther north at Lightning
Ridge, New South Wales, two jaw fragments of an extinct monotreme
have been found associated with dinosaur material similar to that from
the Victorian localities (Archer et al. 1985,Rich et al. in press).
Invertebrateshave, likewise, been preserved in the ~trzeleckiGroup of
the Gippsland Basin, at Koonwarra, More than 80 species have been re-
covered, most of which are insects, but an ostracod, syncarids, anostra-
cans, cladocerans [all Crustacea), spiders, possibly earthworms,
freshwater bryozoans, and unionoid bivalves were also present. Twelve
orders of insects were present, and of these Hemiptera, Coleoptera, and

DINOSAURS OF AUSTRALIA
dontid Femur v p e 2, NMV
PZ56980:A Oeft), qostel-ior vic
A (right), n.zedial v z m ; B, late]
view. (X0.25)

Diptera were the most diverse groups. Numerically dominant were im-
mature individuals of the aquatic Ephemeroptera and Diptera. Minor
elements of the inseci faunas included the Odonata, Blattodea, Plecop-
tera, Orthoptera, Psocoptera, Mercoptera, Trichoptera, Hymenoptera,
and possibly the Sipl~onaptera(Jell G3 Duncan 1986).
More than 50 floral taxa are known from the Koonwarra locality and
other contemporaneous Victorian localities, with both macroscopic and
microscopic material preserved. Most major taxonomic groups from
Bryophyta to Coniferophyta are represented (Drinnan G3 Chambeis
1986), but angiosperms are noticeably absent, or at best rare. Pterido-
phytes are the most diverse and abundant. Forests were dominated by
ginkgoes and a variety of conifers. The understory contained pentoxyla-
leans, ferns, sphenopsids, and ground or epiphytic bryophytes. Fringes
of the forest boasted sclerophyllous ferns, and more open moorlands
were covered by Lycopodium and sphagnalean mosses. Isoetales, hepa-
tics, and algae inhabited aquatic environments and indicate that circu-
lating, moderately nutrient-rich waters (Dettmann 1986).

RICH 61 RICH
 Paleoenvironment
Preliminary sedimentologicalstudies (Brown 1984, Lees 1987, Wagstaff
1983) reveal several types of depositional environments in the Gippsland
and Otway Basins during the Early Cretaceous, associated with braided
river systems: paludal, lacustrine, fluviatile, and overbank floodplain.
Most fossils have been recovered from lacustrine and fluviatile facies, al-
though a single footprint (Knowledge Creek, Otway Basin), preserved in
overbank deposits, established that dinosaurs lived in situ.
The sedimentation patterns were episodic, indicative of perhaps an-
nual flooding (Wagstaff 1983). Lees (1987) suggests that such periodici-
ty could have been associated with spring meltwater floods from distant
highlands. Oxygen isotope studies (Gregory et al. in press) in near syn-
genetic or early diagenetic concretions suggest mean annual tempera-
tures of 4f 5°C.
Environmental conditions indicated by the biota are not entirely clear.
Douglas & Williams (1982:171] and Williams 63 Douglas (1985) argue
for a "warm- to cool-temperate, relatively equable climate of moderate
seasonality with no widespread winter freezing." They further suggest
the possibility of a reduced obliquity of the ecliptic during the Mesozoic
to explain the presence of evergreen plants in the southern Australian
polar floras which had "no sign of specialized structures . . . designed to
cope with conditions of polar light and darkness" (Williams & Douglas
1985:355). But Drinnan & Chambers (1986:8-9) state that the "Koon-
warra Fossil Bed flora does not contain positive evidence for anyparticu-
lar palaeoclimate, but is consistent with the indication of the
environment suggested by the invertebrate fauna also preserved." They
suggest that a "cool climate is consistent with the two major components
of the preserved flora, Ginkgo australis and Taeniopteris daintreei.
Both taxa were large, simple, petiolate leaves, with were abscised com-
plete, and were probably from deciduous plants."
Based on the pollen content of the Strzelecki Group sediments, espe-
cially at Koonwarra, Dettmann (1986) reconstructed the Early Creta-
ceous climate of southern Australia as cool, humid, possibly seasonal,
but she was unwilling to speculate on the significance of the flora rela-
tive to annual light availability,
Based on taphonomic analysis of the fauna, particularly the verte-
brates, together with analysis of the sediments, Waldman (1971) pic-
tures Koonwarra as a cool, shallow lake covered with ice during the
winter which resulted in a "winterkill" of the fish.
Jell & Duncan's (1986:lii) work on the invertebrate fauna from the
same locality indicate that the environment of deposition was a "shal-
low, semi-isolatedbody of water marginal to a shallow freshwater lake
with periodic, probably seasonal, replenishing of the fauna from the
lake and periodic mass mortality of the isolated fauna." They suggest
that conditions could have been cool (based on the presence of siphlon-
urid mayflies and cantharid beetles) but are not willing to accept Wald-
man's (1971) evidence for winter ice. For other reasons, Wilson (1977)
and Elder & Smith (1988) also object to the evidence for winter ice.
Waldman's rather well-argued case for winterkill, however, seems, for
the moment, the best explanation for the thanatocoenose at Koonwarra.
Indeed, the biota of the southern Australian Early Cretaceous was
moderately diverse (more than 150 taxa now recognized). Forests of
both deciduous and evergreen plants were dominated by ginkgoes, po-
docarps, and araucarian conifers. Trees exhibited rings, so some kind of
seasonality occurred. Whether this was controlled by water availability
or light or some other factor is not at all clear. Both deciduous and ever-

DINOSAURS OF AUSTRALIA
 green plants were present (nearly 60 taxa) . Fossil vertebrates included
ceratodont lungfish, dinosaurs (both juvenile and adult), but, interest-
ingly, no crocodilians. All of the vertebrates, including the dinosaurs,
were of moderate to small size, the largest reaching only - 2 m in height
(a small Allosaurus). Hypsilophodont dinosaurs (including both juve-
niles and adults) were unusually diverse for this group, and dominant in
terrestrial faunas. In general, vertebrate diversity was not high, but '
wheiher this is a small sample size bias or a tsue reflection oflow diversi-
ty is at present not certain.
The authors cannot agree with a number ofstatements concerning the
paleoclimaticsignificance of the southern Victorian Early Cretaceousbi-
ota. More data are needed before definitive statements on many aspects
of these communities can be made. Nevertheless, the following can be
pointed out:
The presence of evergreen plants does not necessarily militate against
a lengthy polar night. Definitive information has yet to be published on
controlled, long-term studies of evergreen araucarians and podocarps
held under polar light cycles coincidentwith warmer-than-present con-
ditions at comparable latitudes and with higher oxygen levels predicted
for Early Cretaceous times in southern Australia (Barron 1983). Ever-
green plants, whose metabolism would have been slowed by cool to cold
conditions might well have been able to survive a long polar night that
would have been punctuated each month by a period of moonlight. Con-
trary to Douglas & Williams (19823, the authors conclude that such con-
ditions might not have been conducive to evergreens' metabolizing
themselves to death in the darkness. Perhaps long-term, controlled stud-
ies of the remnants of this Early Cretaceous assemblage would provide
some pertinent answers or eliminate some possibilities.
Modern distributions and tolerances of certain forms may be decep-
tive, as studies such as that by Archbold (1981) on Lingula have demon-
strated. Frakes & Francis (1988547) point out that 'ccycadophytes,
classically wasm temperate indicators, appear to have had a broader
adaptive range than their living counterparts and could tolerate cool
high-latitude environments." The same might be said for the ceratodont
lunpfish, whose modern survivors have a relict range in tropical
Queensland, part of Africa, and South America (Waldman 1971). Dur-
ing the Mesozoic their range extended to paleolatitudes near both the
North and South Poles (Schaeffer 1970).
The presence ofjuvenile dinosaurs in the Victorian Early Cretaceous
faunas does not necessarily indicate that dinosaurs lived year-round in
the area (contrary to the interpretation of Brouwers et al. [I9871 for a
similar situation on the North Slope of Alaska in the Late Cretaceous),
Juvenile presence only indicates that dinosaurs nurtured young in the
area at some time during a year, perhaps even using this area as a high-
energy nursery during the Antarctic summer, like many vertebrates use
the Arctic and Antarctic areas today.
Whether dinosaurs migrated, hibernated, or lived year-round in situ is
not at all resolved for the Early Cretaceous of southern Victoria, despite
arguments to the contrary (Douglas G3 Williams 1982). Migration might
well have been possible if dinosaur herds behaved like modern caribou,
which move over great distances annually. Exactly how far the nearest
light-enriched area was from the Otway and Gippsland Basins in the
Early Cretaceous has not been solidly resolved, since the paleolatitudes
for this area range from latitude 70 to 85' S.
The discussion concerningwhether dinosaurs were warm-blooded or

RICH & RICH

i?
 Basis for Estimating Encephalization
Quotient (EQI from Area of Endocasts
4.: Expected area of endocast in dorsal view from anterior
tip of cerebral hemisphere to posterior edge of
optic lobes
4: Observed area of endocast in dorsal view from anterior
tip of cerebral hemispheres to posterior edge of
optic lobes
E,: Expected brain mass
E,: Obselved brain mass
EQi: Encephalization quotient
k: Constant relating endocast volume (presumed estimate of
brain mass) to body weight
k": Constant relating endocast area to endocast volullle
P: Body mass in grams
- -
A, area of an ellipse with major
and minor axes a and b.
V, volume of an ellipsoid with axes
a, b, and c.
Because ellipsoids being compared
have same shape, each homologous
pair of axes of each ellipsoid have
constant ratio.
Same reason as [3']
From [l'] and [3'1
From 12'1 and [4'1
From [5'] and [6'1
[8'1 E = Vp By definition, the mass, E, of a vol-
ume, V, is equal to the product of
that times its density, p.
19'1 A = ( n k*/4) From [7'] and [8']
(np k*k**/6)-213E2/3
[lo'] k***= ( a k*/4) By definition.
(ap k*k**/G)-213
[ l i t ]A = ~***EWS From [9'] and [lo']
[12'] A, = kl'E,"" From [ l l ' ] if it is assumed that the
con~pleteendocast of Leaellyna-
saura arnica~aphicadoes not sig-
nificantly depart in shape from an
ellipsoid.
[13'] A, = kt%? Sallle reason as [12']
[ I ~ Jkt] = pl3kll By definition
[15'] E, = kPu3 From [3]
116'1 EQi = E,/E, From 121
[17'] EQ," = (Ei/E,)2/3
From [16']
[18'] EQi"" A i / 4 From [12'], [13'], and [17'1
[IS'] EQi = A,""/AeWL From [18']
120'1k' = (E,/PU3)u3/k" From [14'1 and [15']
[211] = E ~ 3 k " / P 4 " From [201]
[22'] = A C P m From [12'1 and [Zl']
[23'] = A, (EQi)-2nP-" From [18'] and [22']
124'1 = 4.9 m m 2 p From [23'] and based on following
estimates for Stenonychosaurus ine-
qualis from Hopson (1977) for EQ,
and P. A, measured from Hopson
(1979, fig. 13b)
DINOSAURS OF AUSTRALIA
* EQ,: 5.80
* P: 45000 g
* A; 1852 mm2
125'1 A, = k2I3kl'P From [22']
[26'1 = (4.9 mm2go") Expected endocast area for Leael-
(1000 g)" lynasaura amicapaphica from
[24'], [25'], and assuming body
weight (P) is 1000 g.
[27'] = 106 mn? From [26']
[28'] EQ, = (182 m r n 2 ) 3 ' V r o m[19'], [27'], and measure-
(106 mm"-3n ment of the area of the endocast of
Leaellynmaura arnicag~aphicafor
A, 182 mm2
[29'] = 2.3 From [28'], estimate of EQ for a ju-
venile individual of Leaellynasaura
amica~aphica,NMV PI85 990-
-
NMV PI85993
Basis for Estimating Encephalization
Quotient (Em of Adult Archosaus
when EQ ofJuvenile Known
For ontogenetic variation within a single species, the relationship
between body weight and brain weight l ~ a sthe same general
form as the variation between species; namely E=kPu where:
E =brain weight
P= body weight
k= constant
a = power function
However, the value of "ay' in such circumstances is not as it is
for interspecific variation. Rather, values near '4 have commonly
been used in the studies of such ontogenetic change in body:brain
weight. Most ofthese studies have been on mammals, particular-
ly primates. Using the data on Alligator mississippiensis from
Crile & Quirling (1940), it was found that "a" was estimated as
0.32 using a least squares best fit of the data. As this was based on
on1 five datapoints, the value 0.32 was taken to be a best estimate
P
of j3. Therefore, in the derivation given below, the estimate of
blain weight as a function of body weight within a species is as-
sumed to be as in equation [2"].
E,: Observed brain mass of juvenile in grams
En: Brain mass in grams calculated for adult
EQ,: Encephalization quotient calculated for adult of same spe-
cies with observed EQ,
EQ,: Encephalization quotient
k: Constant relating endocast volume (presumed estimate of
brain mass) to body weight
k'": Constant relating endocast volume to body weight within a
single species
P: Body mass observed in grams
P,: Body mass of adult in grams
[l"] E, = k1"p1/3 See paragraph above
[2"] EQ, = Ei/(kPu3) From [3]
[3"] = (k'"P113]/(kP2'3)
From [I"] and [Z"]
[4"] EQ, = kt1'p-lf3/k From [3"]
[5"] EQ, = k'"P,-u3/k Substitute EQ,, for EQ,, and
P, for P in [4"1
[6"] EQ,/EQ, = P'lW/P,'" From [4"] and [5"]
[TI EQ, = EQiP1"/PaBJ3 From [6"]
47
cold-blooded is not sufficiently resolved to use body temperature regula-
tion as a n argument either pro or con the presence of long polar nights
(contra Douglas & Williams 1982).
Based on available data, a moderately diverse biota appears to have
lived in the vicinity of latitude 70 to 85' S in southern Victoria under
probably cool, humid conditions that experienced some seasonality
both in energy resources and sedimentary influx. Quite possibly near-
freezing conditions occurred sometime during the year, and a polar
night of a few months affected the area. If this were the case, moonlight
would have been available for at least two weeks of eve~yfour-week cy-
cle. Dinosaurs and a variety of other vertebrates, insects, and plants oc-
cupied the area either continually or episodically-perhaps some
groups migrated, while others hibernated or became dormant during
certain periods. Clearly, many groups were permanent, nonmigratory
residents. Dinosaurs were among the inhabitants of this polar area, indi-
cating that they occupied such environments for at least 45 million years
fkom the Early until the Late Cretaceous, ifboth the Australian and Alas-
kan polar occurrences are considered (Brouwers et al. 1987).
Relictual Distributions and Endemism
Some taxa in the Early Cretaceous biota ofvictoria appear to be late sur-
vivors of groups that had become extinct elsewhere-"living fossils" of
the time. Allosaurus sp. (Molnar et al. 1981,1985; Welles 1983) and a
possible labyri~ithodontOupp G.1 Warren 1986) occur in the Strzelecki
Group of the Gippsland Basin (Eagles Nest and the Punchbowl, respec-
tively), dated as Valanginian-Aptian or Early Cretaceous.
Although some controversy concerns identification of the Victorian
Allosaurus sp. (Welles 1983), Molnar et al. (1985) point out four unique
synapomorphic characters it shared with Allosaurus Ji-agilis from
North America. A.fragiZis is known from the latestJurassic and possibly
the earliest Cretaceous of North America, but the occurrence in Austra-
lia is the youngest currently known.
The youngest occurrences of labyrinthodonts, with the exception of
the possible Victorian form, are in the Early Jurassic of Queensland,
Australia (Warren & Huchinson 1983), and the MiddleJurassic of CM-
na (Dong 1985). The Victorian specimen extends the range of the labyr-
inthodonts another 50 million years forward in time. Both appear to be
relicts that survived longer in Australia and in a polar safe area (Vermeij
1987) than elsewhere. Drinnan & Chambers (1986:7), likewise, note
that "the Victorian Early Cretaceous flora appears to be a remnant, or at
least a late developing flora, e.g. the persistence ofthe Pentoxylales, and
the apparent rarity of angiosperms." '
Thus, in the Early Cretaceous of southern Australia, parts of the biota
were late survivors or endemics, indicating at least a partial isolation of
this region of the continent from the rest of the world. However biased
the sample, the dominance of hypsilophodontids in the thanatocoenose
is noteworthy, as in other regions of the world this is not the case. This
could result from paleogeographic or paleoclimatic factors. In this polar
biota, latitude may have played a major role.

Conclusions
At least three and perhaps four species of hypsilophodontid dinosaurs
inhabited south-central Victoria, Australia, during the Early Cretaceous
.
(Valanginian-Aptian) In one case -Leaellynasaura amicagraphica ,

RICH & RICH

 gen. et sp. nov. -the teeth and femora may, with reasonable co~lfidence,
be associated. Atlascopcosaurus loadsi is defined by its cheek-tooth
morphology. No femora can be associated with reasonable certainty to
these dental remains.
The femora can be divided into three different taxa-L. arnicapa-
phica, Fulgurotherium austr-ale, Victorian Hypsilophodontid Femur
Type I-with the possibility that a fourth species is represented by a sin-
gle specimen, Victorian Hypsilophodontid Femur Type 2. ~lternatively,
this single specimen may be a n extreme variant of F. australe.
A partial endocast of the braincase of one of the hypsilophodontids, L.
amicagraphica, suggests tliat it had an encephalization quotient (EQJ
(sensu Hopson 1977,Jerison 1973) slightly greater than that or other or-
nithopods. As with other members of this group, the eyes were large,
and it is interesting to speculate that a large brain and visual acuity
might be favorable preadaptations for life in polar latitudes under a re-
gime of low light intensity for several months of the year. In this regard, it
is interesting to note that the optic lobes were not evident on any of the
other three hypsilophodontid endocasts available for direct examina-
tion. All three were from much lower paleolatitudes than the ~ ~ i s t r a l i a n
specimen: Hypsilophodon foxii BMNH R.2477 from the Isle of Wight,
England (paleolatitude 39 to 43" N; Barron 1987), Dryosauru Zettow-
vorbecki HMN dy A from Tendaguru, Tanzania (paleolatitude 28 to 33O
S; Smith et al. 1981), and a n as yet unnamed hypsilophodontid in SMU
block 3BS 2-2 from the Proctor Lake site, Texas (see Winkler et al. 1988)
(paleolatitude 33 to 37O N in Barron 1987).
Associated with these dinosaurs is a varied biota of vertebrates, inverte-
brates, and plants made u p of > 150 tam. Forests of both deciduous and
evergreen plants were dominated by gingkoes (Gingko australis), podo-
carps, and araucarian conifers, while the understory contained pentoxyla-
leans, ferns, sphenopsids, and both ground and epiphytic bryophytes.
Forest fringes contained sclerophyLlous ferns, and open areas contained
Lycopodium and sphagnalean mosses. Isoetales, hepatics, and algae dom-
inated aquatic environments and reflect that they were circulaiing and
moderately nutrient-rich. Plants and invertebrates are consistent with a
cool, humid paleoenvironmental reconstruction. Vertebrates offer little in-
formation in this regard.
Sedirnentolopical and regional studies suggest that sediments of the Ot-
way and Strzelecki Groups were deposited in a graben by braided river sys-
tems that derived from volcanic highlands. 180/160 studies suggest that air
temperatureswere cold, at times close to or below 0" C. Sediments are epi-
sodic in nature, and tree rings suggest seasonality.
Paleomagnetic determinations on the volcanogenic sediments in
southern Victoria during the Early Cretaceous place it between latitudes
70 and 85" S, well inside the Antarctic Circle. The biota that inhabited
this area must have experienced three months or more of winter dark-
ness, punctuated only by moonlight for about a fortnight each month.
This biota contained a number oflate survivors (Allosaurus sp., a labyr-
inthodont amphibian, flora), endemics (teleost fishes), and a dominant
element in the vertebrate fauna that elsewhere is a minor component
(hypsilophodontiddinosaurs). Perhaps the presence of these late survi-
vors is attributable to their having found their way to polar "safe areas"
or isolated regions. Without doubt, however, this biota successfilly used
such Mesozoic high latitudes, and dinosaurs were among the groups
that existed at such high latitudes for a considerable period of time, at a
minimum through much of the Cretaceous. Their sojourn near the poles
was not a transitory phenomenon.

DINOSAURS OF AUSTRALIA
 -
We extend our heartfelt thanks to the some 300 vol~~nteers who have excavated the rock
and found the fossils. Among the many who assisted in other ways are: James Warren,
James Bowler, Lesley Kool, Elizabeth Thompson, Natalie Schroeder, John Chessells,
Robert Edwards, Anlis Heislers, Jolm Herman, William Loads, Jack Mackenzie, Max
Manley, and Patrick O'Neill. Org~u~izations that supported the project, bothin kind and
in cash, include the National Geographic Society, Atlas Copco Australia Pty. Ltd., Aus-
tralian Research Grants Scheme, David Holdings Pty. Ltd., Earthwatch, Friends of the
Museum of Victoria, Imperial Chemicals Industries Australia Operations Pty. Ltd., Mel-
bourne Metropolitan Board of Works, Mobil Oil Australia Ltd., National Herbarium of
Victoria, National Parks Service of Victoria, SurfLife SavingAssociation of Victoria, and
Victorian Police. Mnally, for allowing access to collectionsand facilities, as well as shar-
ing ideas and information: William A. Clemens and Kevin Padian, University of Califor-
nia, Berkeley; Michael Cluver andJuri van den Heever, South African Museum; Eugene
S. Gaffney, American Museum of Natural History; Peter Galton, University of Connecti-
cut, Bridgeport; Robert Gregory, Monash University, Melbourne; Nicholas Hotton, Na-
tional Museum of Natural Histo~y,Washington, D. C.; Harry Jerison, Universily of
California, Los Angeles; Angela Milner, British Museum (Natural History); Ralph Mol-
nar, Queensland Museum, Brisbane; John Osirom, Peabody Museum, Yale University;
Albert Rich, Soft Warehouse, Ho~lolulu;Alexander Ritchie, Australian Museum, Sydney;
Arnold Suzummoto, Bishop Museum, Honolulu; William Tunibull, Field Museum of
Natural History, Chicago; Peter Wellnhofer, Bayrische Staatssammlung fir Palaontolo-
gie und Histo~ischeGeologic, Munich; Rupert Wild, Staatliches Museum fir Natur-
kuade, Stuttgart. Steve Morton photographed the specimens and Draga Gelt prepared
the diagrams. Elizabeth Thompson provided editorial assistance and Frank Coffa sup-
plied the photograph of Dinosaur Cove.

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Archer, M.; Flannery, T. F.; Ritchie, A,; et al.
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