Review of the Early Cretaceous Hazel Creek Assemblage, the Oldest Mammal Bearing Fauna in Northern Australia

Home , Cardabiodon, Notochelone, Rolling Downs Group, Wallumbilla Formation

Review of the Early Cretaceous Hazel Creek assemblage,

the oldest mammal-bearing fauna in northern Australia

Daniel Joseph Traub

A thesis in fulfilment of the requirements for the degree of

Master of Science

School of Biological, Earth and Environmental Sciences

Faculty of Science

University of New South Wales

March 2019

Thesis/Dissertation Sheet Australia's Global University

Surname/Family Name Traub Given Name/s Daniel Abbreviation fordegree as give in the University calendar MSc Faculty Science School BEES Thesis Tille Review of the Early Cretaceous Hazel Creek assemblage, the oldest mammal bearing fauna in northern Australia

Abstract 350 words maximum: (PLEASE TYPE) Situated in the Boomarra Sub-basin, Hazel Creek is a moderately sized locality in the Toolebuc Formation, defined by fossiliferous slabs of flaggy limestone. In the mid-late Albian the area was likely a shallow marine estuary, placing it uniquely close to shore. Due to this shallow depth and proximity to shore, the fauna is incredibly diverse. Among the marine invertebrates are the bivalve genera Aucellina and lnoceramus, the common belemnite genus Dimitobelus, and the ammonite genus Myloceras. Vertebrate remains are dominated by disarticulated fish bones and shark teeth (e.g. species of Cardabiodon, Echinorhinus, Richmondichthys, Pachyrhizodus, and an undescribed species of lizardfish) as well as rare fragmentary remains of chimeroids (Ptykoptychion tayyo). Marine turtles (Notochelone costata) are also common, being represented by numerous pieces of shell and limb bone. Large marine reptiles like plesiosaurs (Elasmosauridae) and ichthyosaurs (Platypterygiusaustralis) have also been recovered as isolated vertebrae, teeth, and limb elements. However, of greater significance are the previously undocumented terrestrial taxa that were washed in from the nearby shore. These elements allow us a singular glimpse into the terrestrial fauna of Australia in the mid Cretaceous. Small, fragile limb bones of birds have been preserved, although poor preservationstates limit identification and description. Of unique import, a mammal premolar has also been discovered, making Hazel Creek the only Mesozoic mammal locality known for northern Australia. This tooth represents the oldest mammal in northern Australia. This unique shallow marine locality holds promise of future discoveries which could shape our understanding previously undocumented terrestrial taxa and the Australian Cretaceous ecosystem.

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Situated in the Boomarra Sub-basin, Hazel Creek is a moderately sized locality in the Toolebuc Formation, defined by fossiliferous slabs of flaggy limestone. In the mid-late Albian, the area was likely a shallow marine estuary, placing it uniquely close to shore. Due to this shallow depth and proximity to shore, the fauna is incredibly diverse. Among the marine invertebrates are the bivalve genera Aucellina and Inoceramus, the common belemnite genus

Dimitobelus, and the ammonite genus Myloceras. Vertebrate remains are dominated by disarticulated fish bones and shark teeth (e.g. species of

Cardabiodon, Echinorhinus, Richmondichthys, Pachyrhizodus, and an undescribed species of lizardfish) as well as rare fragmentary remains of chimeroids (Ptykoptychion tayyo). Marine turtles (Notochelone costata) are also common, represented by numerous pieces of shell and limb bone. Large marine reptiles like plesiosaurs (Elasmosauridae) and ichthyosaurs (Platypterygius australis) have also been recovered as isolated vertebrae, teeth, and limb elements.

However, of greater significance are the previously undocumented terrestrial taxa that were washed in from the nearby shore. These elements allow us a singular glimpse into the terrestrial fauna of Australia in the mid-Cretaceous. Small, fragile limb bones of birds have been preserved, although poor preservation states limit identification and description. Of unique import, a mammal premolar has also been discovered, making Hazel Creek the only Mesozoic mammal locality known for northern Australia. This tooth represents the oldest mammal in northern

Australia. This unique shallow marine locality holds promise of future discoveries which could shape our understanding previously undocumented terrestrial taxa and the Australian Cretaceous ecosystem.

i TABLE OF CONTENTS

ABSTRACT….…………………………………………………………………….i

TABLE OF CONTENTS………………………………………………………….ii

LIST OF FIGURES………………………………………………………………iii

ACKNOWLEDGEMENTS……………………………………………………….v

INTRODUCTION…………………………………………………………………1

Geologic Setting………………………………………...... ….……………2

Lithostratigraphy…………………………………………………………..5

Environmental Setting……………………………………………………..7

Fossil Assemblage…………………………………………………………7

METHODS AND MATERIALS……………………………………………….....9

Institutional Abbreviations……………………………………………….11

SYSTEMATIC PALAEONTOLOGY…………………………………………...12

DISCUSSION……………………………………………………………………61

Conclusions………………………………………………………………65

Further Study……………………………………………………………..66

REFERENCES CITED…………………………………………………………..68

ii LIST OF FIGURES

Figure 1. Eromanga Basin overlaid with extent of Eromanga Sea

Figure 2. Hazel Creek Locality Map

Figure 3. Hazel Creek Geologic Map

Figure 4. Stratigraphic relations of formations present at Hazel Creek

Figure 5. Example of the Toolebuc limestone slabs present at the Hazel Creek

Locality

Figure 6. Aucellina sp. cf. A. hughendenensis Etheridge internal view of right

valve in situ.

Figure 7. Inoceramus sp. cf. I. sutherlandi McCoy fragments in situ.

Figure 8. Large Inoceramus sp. cf. I. sutherlandi McCoy fragment in situ

Figure 9. Dimitobelus sp. longitudinally bisected in situ

Figure 10. Dimitobelus sp. in situ. Longitudinal cross-section (A) and transverse

cross-section (B)

Figure 11. Myloceras ammonoides. Side view (A) and posterior view of whorl (B)

Figure 12. Pachyrhizodus sp. cf. P. grawi jaw fragments (A) and vertebrae (B)

Figure 13. Richmondichthys sweeti skull fragment (A) and scales from the flank

(C-E) and from various locations (B)

Figure 14. Gen et. sp. incertae sedis cf. Apateodus sp. jaw fragments (A) and teeth

Figure 15. Squalicorax sp. indet. from Hazel Creek Locality, lingual and labial

views

Figure 16. Squalicorax sp. indet from Bilelo (1969) (photo from Welton and

Farish, 1993)

Figure 17. Carcharias striatula teeth

Figure 18. Plicatolamna arcuata tooth, lingual view

iii Figure 19. Cardabiodon sp. indet tooth, lingual view

Figure 20. Echinorhinus australis teeth

Figure 21. Echinorhinus australis, as pictured by Kemp (1991).

Figure 22. Ptyktoptychion tayyo right mandibular tooth plate. Lingual view (A),

dorsal view (B), dorsal anterior view (C)

Figure 23. Notochelone costata jaw bones. Dorsal and ventral views (A&B),

ventral view (C)

Figure 24. Notochelone costata humeri growth series. Adult, dorsal and ventral

views, (A), sub-adult (B), and juvenile (C)

Figure 25. Notochelone costata plastron, dorsal and ventral views

Figure 26. Notochelone costata carapace peripherals, ventral view

Figure 27. Platypterygius sp. paddle bone, dorsal and ventral views

Figure 28. Platypterygius sp. vertebrae. Dorsal (A), caudal (B), indet (C)

Figure 29. Elasmosauridae tooth, labial and lingual views

Figure 30. Elasmosauridae phalange

Figure 31. Elasmosauridae vertebra anteroposterior view and profile view

Figure 32. Possible dinosaurian limb fragment

Figure 33. Hollow limb fragments from Enantiornithes

Figure 34. Hazel Creek mammal incertae sedis premolar

Figure 35. Comparative mammal premolars of the Australian Cretaceous. Hazel

Creek premolar (A), Corriebaatar marywaltersae (B) (Rich et al.

2009a), Teinolophos trusleri (Rich et al. 2016) (C), Ausktribosphenos

nyktos (Rich et al. 2016) (D), and Bishops whitmorei (D) (Rich et al.

2009b).

Figure 36. Various coprolites from Hazel Creek Locality

iv ACKNOWLEDGEMENTS

There are many people I need to thank for their part in the completion of this degree. Firstly, I need to thank my supervisors Professor Michael Archer and

Dr. Suzanne Hand. It goes without saying that none of this would have been possible without them. They have been patient and trusting and I am ever grateful for their support.

My thanks must also be given to those within the Vertebrate Palaeontology

Laboratory, University of New South Wales, Dr. Troy Myers and Dr. Anna

Gillespie without whom I would have had no specimens to work with and no place to start. I would also like to thank my committee members, Dr. Carol Oliver,

Dr. Malte Ebach, and Dr. Ian Graham, and the future reviewers of this this paper.

Thanks to Henk Godthep and Gill Goode for the previous work done with the

Hazel Creek locality.

Thanks must also be given to the organizations that supported collection for this thesis. Thank you to The Riversleigh Society and to the Palaeontology,

Geobiology and Earth Archives Research Centre (PANGEA) Research Centre.

I am indebted to the students who helped me process the large amounts of the collected material: Ashleigh Ford, Jessica Tam, Jessica Schemer, and Antonia

Parker. They waded through many a fish bit and came through the other side sane.

I am extremely grateful to those who provided insight and comparative material for the Cretaceous biota. Jacqueline Nguyen and Patrick Smith, both of the Australian Museum. Especially Patrick Smith for his fount of knowledge and positivity.

Finally, I would like to thank my friends and family who helped keep me together through this process. I could not have done this without you.

v INTRODUCTION

The mid Albian of Australia has been documented through marine deposits across the Great Australian Basin (GAB). The GAB consists of three large continental basins, the Carpentaria Basin in the north, the Surat Basin in the southeast, and the

Eromanga Basin between them (Figure 1) (Cook, 2012). The Eromanga basin, the largest of the three, possesses a number of well-documented fossil localities which have revealed the biota of Australia’s Cretaceous marine environment. However, the

Cretaceous terrestrial fauna of Australia is globally one of the least well-documented.

Within the basin, marine deposits

Figure 1. Eromanga Basin overlaid with extent of Eromanga Sea (Bell, 2018). Red point shows

location of Hazel Creek Deposit.

1 have, on occasion, produced rare terrestrial elements that had been washed out to sea.

Hazel Creek is a previously undocumented locality in north-western Queensland that has the potential to provide significant insights into the Early Cretaceous terrestrial, as well as shallow marine biotas of northern Australia. This shallow near-shore deposit has produced rare terrestrial fossils including a single mammal premolar. This tooth is the only Mesozoic mammal specimen known from the northern half of the continent. The present study has focused on taxonomic and taphonomic features of the Hazel Creek faunal assemblage.

Geologic Setting

The Eromanga Basin is the geologic remnant of the Eromanga Sea which covered much of Australia’s landmass in the Cretaceous (Bain and Draper 1997). Mid

Aptian to Albian sediments in the basin are represented by the Rolling Downs Group.

This group consists of four formations (see Figure. 4 for stratigraphic relationships): the

Wallumbilla Formation (Fm), the Toolebuc Fm, the Allaru Fm and the Normanton Fm

(Kemp 1991).

The Wallumbilla Formation is primarily a mudstone unit mid Aptian to mid

Albian (107-115 million years old) in age. These basin-wide, shallow marine deposits were deposited during the initial period of the Cretaceous marine transgression (Bain and Draper 1997; Ramsden 1983). Along with mudstone, the Wallumbilla Fm also contains siltstone, minor limestone, and glauconitic sandstone. Measurements within the

Carpentaria Basin provide an average thickness of 150-220 metres (Day et al. 1975).

The Toolebuc Formation is an interbedded limestone and black shale unit, mid to late Albian in age (97-107 million years old). It was deposited during the height of the Cretaceous marine transgression after the northern seaway was established. At this

2 time, the depositional environment was relatively clear, shallow, marine waters (Bain and Draper 1997; Kemp, 1991). According to Smart et al. (1980), the Toolebuc Fm is predominantly composed of carbonaceous and bituminous shale interbedded with bioclastic and micritic limestone deposited in an oxygen-restricted environment. There is also a calcareous oil shale often associated with a thin coquina bed within the lower sequence of this formation (Swarbrick, 1974). The upper sequence of this formation commonly contains layers of coquinite composed of fine-grained fragments of pelecypods and fossil fish (Day, 1983). Measured across a variety of sites, the Toolebuc

Fm is about 5-20 metres in thickness (Day et al., 1975).

The Allaru Formation is a marine mudstone sequence, late Albian in age. Apart from being primarily mudstone, the Allaru Fm also contains beds of glauconitic siltstone and minor limestone. On average this formation is 160-215 metres thick (Smart et al., 1980).

The Normanton Formation, the upper most formation in the Rolling Downs

Group, is primarily siltstone and sandstone of late Albian age. There has been some indication that the formation may possibly extend into the earliest Cenomanian (Smart et al., 1980). In its entirety, the Normanton Fm consists of labile glauconitic sandstone, siltstone, minor silty mudstone, and limestone beds ranging between 200-300 metres thick (Day et al., 1975).

The Hazel Creek locality is about 300km northeast of Mt Isa and 190km southwest of Normanton (Figure 2). Occupying an area of more than 40 square kilometres, the Hazel Creek locality occurs between latitudes 19’12’ and 19’18’S and longitudes 140’17’ and 140’22’E in the vicinity of the Three Ways Intersection of

Burketown and Julia Creek Normanton Roads. Preliminary and informal geologic studies were conducted in 2003 by Gill Goode in conjunction with the University of

3 Figure 2. Hazel Creek Locality Map. (Goode, 2003)

Figure 3. Hazel Creek Geologic Map. (Goode, 2003)

4 New South Wales. The Hal Creek per se and associated tributaries form a braided system. During the rainy season the creek is prone to flash floods, moving sediment (cobbles and pebbles during flooding) down slope to Dismal Creek (Smart,

1973). During the remainder of the year, the area is dry and arid. Slabs of fossiliferous limestone have been moved, overturned, and buried by flash floods as well as by road graders and other anthropogenic activities.

Lithostratigraphy

Deposits at the Hazel Creek locality are primarily those of the Toolebuc

Formation. However, there appear to also be outcrops of the Wallumbilla and Allaru

Formations, as well as relatively smaller exposures of Tertiary and Holocene deposits.

In a preliminary informal study, Gill Goode outlined the stratigraphy of the region

(Figure 3) as it is currently understood. The contact between the Toolebuc and the

Wallumbilla Formations is observable in the lower creek beds to the east of the Burke

Developmental Road. There are outcrops featuring a number of facies within the

Wallumbilla Fm toward the eastern extent of the Hazel Creek locality (calcareous cone in cone structures as well as mudstone, and siltstone).

The Toolebuc Formation provides most of the fossiliferous deposits in the Hazel

Creek area. It possesses a number of different limestone facies/structures including flaggy limestone, limestone concretions, and friable platy limestone. In some areas, fossils are abundantly evident on carbonic acid-eroded surfaces of slabs scattered across the surface of the site. However, the density of fossils varies and in some areas, fossils are completely absent. The limestone can often be pale orange in colour due to the presence of iron oxides. There is a single locality (site 15 on Figure 3) where a vertical outcrop can be observed. This should be a focus for future research into the detailed microstratigraphy of this area. Exposures of these deposits are often signalled by the 5 presence of black soils and areas of distinctive vegetation. Sites lacking limestone often contain weathered laterite deposits without superpositional shrub growth. Evaporitic gypsum crystals forming course interlocking prisms can be found in soils of this kind, often growing perpendicular to the sediment surface.

The Allaru Formation can be found in scattered surface deposits near the western extent of the Hazel Creek locality. Two deposits within the extent of the

Toolebuc (sites 013, 014, 016, and 018 on Figure 3) are also likely the Allaru

Formation. These deposits are cone in cone mudstones and concretionary limestones, the latter of which is fossiliferous.

In her study, Goode outlined Tertiary and Holocene sediments. These are represented by small exposures of conglomerate material at the southwest corner of the

Hazel Creek area. Tertiary to Quaternary colluvial and outwash sands and gravels are deposited down slope towards Dismal Creek. There are sandy creek beds to the east which contain alluvial plain deposits that have been interpreted to be Holocene in age

(Goode, 2003).

Figure 4. Stratigraphic relations of formations present at Hazel Creek. Modified from Cook (2012).

Environmental Setting

During the Cretaceous, when the Australian Continent was attached to

Antarctica in latitudes of 50 and 85 degrees south, plate tectonic movements started to break up the supercontinents. This breakup, combined with global sea level rise, flooded many of the early Cretaceous landmasses, inundation peaking about 110 million years ago (Cook and McKenzie 1997). In Australia, this resulted in a series of inland seas.

Between 116 and 98 million years ago there were four periods of marine transgression and regression (Smart et al., 1980). These marine movements deposited the Rolling

Downs Group.

The paleoenviornment at the time of deposition for the Rolling Downs Group was a large, shallow epicontinental sea where minor changes of sea level resulted in the differing depositional intervals (Day, 1983). The massive, inland Eromanga Sea, which covered about a third of the Australian Continent, was connected to open ocean waters to the north. The shallow epicontinental seas were home to abundant marine life (Cook,

2012) such as is evident in the limestones found at Hazel Creek.

Fossil Assemblage

Faunas recovered from bioclastic limestones of the Toolebuc Formation are well documented from a number of localities. These include marine invertebrates, fish, marine reptiles, and pterosaurs (Day, 1983). A large number of the previously described marine taxa are identifiable within the Hazel Creek area. These fossils make up the majority of the taxonomic review. They include aspidorhynchid teleosts and lamniform and chimeroid chondrichthyans. Invertebrates like ancyloceratid coiled heteromorph ammonites, belemnoids, and filter feeding bivalves have been recovered. Marine

7 reptiles discovered include chelid turtles, elasmosaurid plesiosaurians, and ichthyosaurs.

In some cases, however, terrestrial taxa have been found which enables an albeit limited insight into the diversity of northern Australia’s early Cretaceous terrestrial faunas.

Among terrestrial taxa so far recovered are enantiornithine birds and a single mammal premolar.

8 METHODS AND MATERIALS

Hazel Creek Fossils

The Hazel Creek locality was brought to the attention of Professor Michael

Archer and Mr. Henk Godthelp of the University of New South Wales, Sydney in the

1980’s. It has been collected periodically in the 1980s, 1990s, 2000s, and most intensively in 2017 and 2018 which provided the materials used in the present study).

Exposures of the Hazel Creek limestone are primarily broken slabs that have been eroding at ground level (Figure 5). Many have been disturbed by heavy machinery because a main road passes right through the deposit. Over the last two years, collection has involved locating and then lifting loose slabs onto a trailer or flat-bed and then transporting them to Mount Isa from where they are trucked to UNSW in Sydney. In the field, sledgehammers, rock hammers, and pry bars are used to manipulate slabs. Slabs sent to the Vertebrate Palaeontology Laboratory at the University of New South Wales are then acid-processed in a purpose-built laboratory. In previous years, fossil processing has been conducted by Dr. Anna Gillespie and Mr. Henk Godthelp. In the last two years, I have carried out preparation personally. Fossils are retrieved by repeatedly enabling weak concentrations of acetic acid (3-5%) to dissolve them. Fragile elements recovered have been consolidated using a polyvinyl resin dissolved and thinned with acetone. Once stable and isolated from the limestone matrix, fossils are sorted under a microscope and then made available for taxonomic assessment.

Following completion of this study, Hazel Creek materials will be stored in the

University of New South Wales, Sydney collections while the research continues.

Although identification mostly is done through the use of published journal articles, occasionally comparison has had to be made with other specimens. This comparative

9 material was made available by the Palaeontology Department of the Australian

Museum and School of Biological, Earth and Environmental Sciences at the University of New South Wales, Sydney.

Figure 5. Example of the Toolebuc limestone slabs present at the Hazel Creek Locality

Measurements

Osteological and dental measurements were made using a pair of electronic digital

Vernier callipers. These callipers were accurate to 0.01mm and rounded to the nearest

0.1mm. All measurements were taken at least three times. Finer measurements were made using a Leica Wild M3B binocular dissecting microscope fitted with a Wild MMS

235 digital length measuring electronic micrometre eyepiece calibrated to measure in one hundredths of a millimetre.

Photography Equipment

Photographs were taken using a Sony Alpha SLT-A58 digital camera on a fixed stand.

Detailed photographs were taken using the Scanning Electron Microscope (SEM) instrument in the Mark Wainwright Analytical Centre, UNSW Sydney. The instrument is a Hitachi EDSONS S-3400 Scanning Electron Microscope. All editing was done with Adobe Photoshop.

Institutional Abbreviations

AM, Australian Museum, Sydney;

GSQ, Geological Survey of Queensland;

MVP, Museum of Victoria, Melbourne;

QM, Queensland Museum, Brisbane;

11 SYSTEMATIC PALAEONTOLOGY

Class BIVALVIA

Subclass PTERIOMORPHIA Beurlen, 1944

Order PTERIOIDA Newell, 1965

Suborder PTERIINA Newell, 1965

Superfamily PECTINACEA Rafinesque, 1815

Family BUCHIIDAE Cox, 1953

Genus Aucellina Pompeckj, 1901

Aucellina sp. cf. A. hughendenensis Etheridge sensu

Diagnosis: As outlined by Jiang et al. (2004) ‘Orthocline to strongly prosocline, inequivalve; left valve usually convex with more or less prominently projecting umbo and prosogyous beak; right valve flat or slightly convex with its umbo scarcely projecting. The anterior auricle of the right valve in alignment with hinge, tongue- to spoon-shaped, concave inside, separated from the umbonal cavity by a ridge; subauricular notch extending almost to beak, pseudoctenolium present. The anterior auricle of the left valve is usually absent except for a very short one in some specimens.

Posterior auricles of both valves are distinct. Radial ribs occur in some species; the cardinal area consists of striated anterior and posterior ligament areas bounding a shallow to moderately deep, oblique ligament pit which extends posteriorly from just in front of the umbo; the dorsal part of the basal ear forming the anterior ligament area.’

Collected Material: A single example of the genus Aucellina have been recovered from the Hazel Creek locality. The valve remains in situ displaying an internal view of a right

12 valve (Figure 6). This specimen is 20.3mm long and 21.6mm wide. There is moderate erosional damage wearing through the valve in places. The valve is prosocline and aligns with the inequivalve nature of the genus. The anterior aurical is not visible (either by the matrix or the preservation state), while the anterior aurical in the right valve is visible in alignment with the hinge sharing the above listed characteristics. While A. hughendenensis is the commonly preserved species within the Cretaceous of Australia

(Henderson, 2004) the collected specimen is not well enough preserved for species identification.

Figure 6. Aucellina sp. cf. A. hughendenensis Etheridge internal view of right valve in situ.

Scale bar = 1cm

13 Subclass CRYPTODONTA Neumayr, 1884

Order MYALINIDA Paul, 1939

Family INOCERAMIDAE Giebel, 1852

Subfamily INOCERAMINAE Zittel, 1881

Genus Inoceramus Sowerby, 1814

Inoceramus sp. cf. I. sutherlandi McCoy, 1865

Diagnosis: As outlined by William and Box (1992) ‘Shells moderate to large in size, moderately thick and inequivalved, erect, moderately inflated, commonly having strong posterior sulci, folds, and auricles; anterior face typically flattened forming pseudolunule; hinges typically massive with medium to large resilifers. Byssal slit prominent and well defined, small pedalbyssal insertion area(s), strong mantle retractor tracks, and large, comma-shaped posterior adductor insertion area.’

Collected Material: Only fragmentary remains have been recovered and observed. The majority of which are small, seen below in Figure 7 and 8. These fragments are between

32.6mm to 12.4mm long as well as 2.2mm to 1.0mm wide. Figure 8 shows a much larger fragment on the erosional surface. These shell fragments do not survive the acid etching process and are thus unlikely to be removed from the matrix. While identification to species is impossible without more complete specimens, it is clear on size alone the assignment to the genus Inoceramus stands. According to Cook (2012),

Inoceramus sutherlandi McCoy, 1865 is the species found throughout the Toolebuc.

Given the preservation of the shells examined, there is no reason to indicate that they could not represent I. sutherlandi.

Figure 7. Inoceramus sp. cf. I. sutherlandi McCoy fragments in situ. Scale bar = 1cm

Figure 8. Large Inoceramus sp. cf. I. sutherlandi McCoy fragment in situ. Hammer provided for scale.

Class CEPHALOPODA Cuvier, 1795

Subclass COLEOIDEA Bather, 1888

Order BELEMNITIDA Zittel, 1895

Suborder BELEMNOPSEINA Jeletzky, 1965

Family DIMITOBELIDAE Whitehouse, 1924

Genus Dimitobelus Whitehouse, 1924

Dimitobelus sp. indet

Diagnosis: As outlined by Williamson (2006) ‘Guards hastate in outline, less so or cylindriconical in profile. Ventro-lateral grooves clearly inscribed are confined to the

16 alveolar region and anterior portion of the stem region. Dorso-lateral grooves lacking or rudimentary, forming weak, obscure depressions or lines. Ventro-lateral grooves are straight in the alveolar region and lie sub-parallel to the venter, but posteriorly towards the stem region they curve towards the midline of the flanks and progressively weakening. Lateral lines paired, always present in apical and stem regions, centrally placed, becoming deflected dorsally near the apex.’

Collected Material: Dimitobelus belemnites cover the surface of many of the blocks from the Hazel Creek locality. Sensitive to the acid washing process, the belemnites often dissolve during general fossil extraction. Most specimens range from 25mm to

40mm with an average diameter of 8mm. In profile, they are subhastate to cylindriconical and in transverse cross-section they are subcircular and flattened along one face (Figure 10.B). As the specimens are found shattered and eroded in half along the longitudinal axis (Figure 9&10.A), inspection of the groves cannot be conducted.

Figure 9. Dimitobelus sp. longitudinally bisected in situ. Australian dollar coin for scale

Figure 10. Dimitobelus sp. in situ. Longitudinal cross-section (A) and transverse cross-section (B). Scale bars = 1cm

18 Subclass AMMONOIDEA Zittel, 1884

Order AMMONITIDA Hyatt, 1889

Family ANISOCERATIDAE Hyatt, 1900

Subfamily LABECERATINAE Spath, 1925

Genus Myloceras Spath, 1925

Myloceras ammonoides Etheridge, 1909

Diagnosis: As outlined by NcNamara (1978) ‘Whorl section ovoid. Early whorls loosely coiled, later whorls in contact. Ribs sinuous throughout, regularly bifurcating on flanks, though rarely bifurcating at ventro-lateral tubercles; thickening across venter on early phragmocone. Ventro-lateral tubercles frequent, very small on phragmocone, become large and elongate on body chamber; present to peristome. Crozier moderately hooked, not recurved. First lateral saddle of suture bifid and asymmetrical; first lateral lobe trifid and almost symmetrical.’

Collected Material: The collected ammonite is half body fossil and half impression

(Figure 11). The phragmocone and early whorls are preserved with the body shaft left as an imprint on the matrix. All together, the specimen is 197.7mm long and 118.7mm wide. At its widest point the phragmocone is 42.2mm thick. Thickening of the shaft cannot be assessed as it is missing and the impression only represents half of the specimen. Ventro-lateral tubercles are present, visible along the outermost whorl.

Sinuous ribs are visible along the entire length and are notably bifurcating along the flanks of the shaft. This specimen is certainly Myloceras ammonoides as opposed to the very similar Myloceras auritulu outlined in McNamara 1978. Myloceras auritulu does

19 not possess ventro-lateral tubercles on the phragmocone and its ribs bifurcate at the ventro-lateral tubercles on the shaft, not along its flanks.

Figure 11. Myloceras ammonoides. Side view (A) and posterior view of whorl (B). Scale bar = 1cm.

20 Class ACTINOPTERYGII

Division TELEOSTEI Müller, 1844

Suborder PACHYRHIZODONTOIDEI Forey, 1977

Family PACHYRHIZODONTIDAE Cope, 1872

Genus Pachyrhizodus Dixon, 1850

Pachyrhizodus sp. cf. P. grawi

Diagnosis: As outlined by Forey (1977) with revisions from Bartholomai (2012).

Pachyrhizodontid fishes in which the skull roof is marked with a frontal depression.

Dermethmoid broad laterally with muted posteroventral processes, lacking definitive bone-enclosed ethmoid commissure. Dilatator fossa with a roof posteriorly, pterotic not produced into a spine. Exoccipitals meeting above and below foramen magnum.

Endochondral elements of posterior of otic region meet loosely without interdigitating sutures. Dilitator fossa present, sometimes emphasised anteriorly by a large fenestra between autosphenotic and pterotic below excavated frontal margin behind autosphenotic ‘crest’. Foramen for the orbital artery and the posterior opening of the jugular canal close together on the lateral face of prootic and enclosed within a ‘prootic cup’; pterotic roof of dilitator diminishes posteriorly to virtually disappear. Fenestration of anterior ceratohyal variably present. Posterior infraorbitals usually very broad, overlying preoperculum and often much of the operculum. Preoperculum varies from minimally expanded ventrally to significantly expanded posteroventrally and with tapered vertical limb. Preopercular sensory canal moderately branched across preopercular base. Operculum with oblique ventral margin, interoperculum longer than deep. Ventral postcleithrum expanded posteriorly, outer pectoral fin-ray large and closely articulated. Caudal fin-rays (where known) crossing hypurals at a steep angle.

21 Collected Material: Material consists primarily of fragmentary jaw bone, teeth, and vertebrae (Figure 12). These jaws are approximately 14mm to 25mm long and 1.7mm to

5.7mm deep (including the teeth). They are poorly preserved and leave room for interpretation. The teeth are small, conical, and curved lingually. They are uniform in size along the jaw fragments and tightly spaced. The collected vertebrae are not incredibly distinct, but are amphicoelous and narrow near the middle in a lateral view.

The majority of neural arches and transverse processes are missing but the surface of the vertebrae are pocketed where the attachment would have been. The centrum is circular to elliptical in shape (the latter possibly due to compression). Fine ridges and grooves run their length on the lateral surfaces. The genus Pachyrhizodus is represented in the

Cretaceous of Australia by two species: P. marathonensis Etheridge Jnr., 1905 and P. grawi Bartholomai, 2012. The difference between the two species is primarily size.

P. marathonensis is larger and more robust than the sleeker P. grawi. The slender nature of the jaw fragments would seem to indicate P. grawi, but with the lack of more complete specimens it would be impertinent to assign species at this time. It is also worth noting that none of the bones are robust enough to be considered for

Australopachycormus hurleyi Kear 2007 or large enough to be considered Cooyoo australis Woodward 1894 (Bartholomai, 1987), both of which are frequently recovered

Toolebuc species (Smith, P., personal communication, 2018).

Figure 12. Pachyrhizodus sp. cf. P. grawi jaw fragments (A) and vertebrae (B). Scale bar = 1cm

23 Family ASPIDORHYNCHIDAE Nicholson & Lydekker, 1889

Genus Richmondichthys Bartholomai, 2004

Richmondichthys sweeti Etheridge Jnr & Smith Woodward, 1891

Diagnosis: As outlined by Bartholomai (2004) ‘Very large, exceeding 1.6 metres in length. Neurocranial roof with markedly asymmetrical fronto-parietals, dermopterotico- extrascapulars and post-temporals; rostral not extending beyond maxillary; premaxillary without teeth and short, not extending anteriorly as far as the dentalosplenial; circumorbital bones are extremely well developed; maxillary without teeth, set well anterior to orbit; predentary very small, edentulate; gill filaments extremely elongated; dentalosplenial deep anteriorly and very deep posteriorly, lacking teeth; lateral line scales greatly deepened; all scales heavily covered with ganoine, raised into heavy ridges and tubercles; ornamented ganoine also covers the outer bones of the head and is present on the larger elements of the fin rays.’

Collected Material: Collected material from this site consists of preserved scales

(Figure 13.B-E). Many are fragmentary, but a number of more complete flank scales have also been recovered. These are slightly convex following the body shape. The leading edge of the scale is smooth while the trailing edge is covered in ganoine pits and ridged. A prominent ganonine ridge separates the smoothing, leading edge from the ridged edge running along the median line of the scale. The posterior margins of the flank scales are finely serrate and on at least one of them the dorsal margin is pointed

(Figure 13.C-E). The flank scales are between 99.15mm and 52.8mm long with a width between 22.9mm and 12.2mm. All the scales and scale fragments are covered in a layer of shiny ganoine. A skull fragment has also been recovered (Figure 13.A). This 24 fragment is heavily ornamented with raised bumps and ridges. It is slightly convex and

quite thin. This fragment measure 35.0mm long by 13.6mm wide.

D E

Figure 13. Richmondichthys sweeti skull fragment (A) and scales from the flank (C-E) and from

various locations (B). Scale bar = 1cm.

25 Order AULOPIFORMES Rosen, 1973

Suborder ICHTHYOTRINGOIDEI Goody, 1969

Family indet.

Gen et. sp. incertae sedis cf. Apateodus Woodward, 1901

Diagnosis: As outlined by Newbrey & Konishi (2015) ‘Snout narrow, mesethmoid pointed; long, narrow frontal bones; short, narrow, irregularly shaped parietals meeting at the midline; parietal bones not separated by the supraoccipital; supraoccipital with a short, robust crest extending beyond the posterior margin of the bone; widest areas of the neurocranium at anterior tips of sphenotics and posterior ends of pterotics; sphenotics narrow, long; sphenotic spine ventrally oriented; epiotics narrow and in contact with parietals; facet for the dorsal arm of the posttemporal situated on the epiotic, not extending past the posterior wall of the epiotic, and extending posteriorly as far as the pterotic; posterior end of pterotic moderately deep in lateral profile; pterotics medially constricted in dorsal view; posterolateral corner of pterotic with a sharp 25–35 angle; anterior part of the hyomandibular facet formed by the pterotic, prootic, and sphenotic; parasphenoid extending posteriorly to overlap the basioccipital including occipital condyle; ectopterygoid teeth pinnate and laterally compressed with anterior and posterior cutting edges; diastema between first and second dermopalatine teeth with long dentary teeth that fill (i.e., occlude) the gap; teeth of dermopalatine and ectopterygoid striated near base; length of diastema between the teeth as long as or longer than the height of the posterior tooth; length of the diastema between the posterior tooth of the dermopalatine and the anterior tooth of the ectopterygoid shorter than the height of the anterior tooth on the ectopterygoid; anterior tooth of the ectopterygoid angled posteriorly; length of the diastema anterior to the first tooth of the

26 dermopalatine shorter than the length of the base of the first tooth; coronoid process moderately high and rounded with a 90 angle; dentary-angular contact ‘V’-shaped; angular confined to posterior 42% of mandible; dentary teeth striated near base; dentary teeth greatly laterally compressed compared with palatal teeth and slightly inclined posteriorly; mandibular canal closed; cleithrum with a broad dorsal arm; narrow mesocoracoid arch present in pectoral girdle; mesocoracoid concavity deep and ‘U’- shaped.’

Collected Material: The material collected of Gen et. sp. incertae sedis cf. Apateodus sp. consists mainly of numerous teeth (Figure 14.B) and a couple jaw fragments (Figure

14.A). These teeth are all dentary teeth based on their elongated length when compared to existing Apateodus species. They are on average around 10mm to 20mm from tip to base with a base diameter of about 4.0mm. The jaw fragment pictured is 24.8mm long,

8.9mm tall (including the tooth), and 3.8mm wide. Three existing species are outlined for the Apateodus genus: A. striatus Agassiz, 1837, A. glyphodus Blake, 1863, and A. busseni Fielitz & Shimada, 2009. According to Kriwet et al. (2006), the genus

Apateodus has been found in the Antarctic fauna. Similar teeth have also been discovered in near complete specimens at the Richmond Quarry (Smith, P., personal communication, 2018). These, like the Hazel Creek specimens, have yet to be described.

27 A

Figure 14. Gen et. sp. incertae sedis cf. Apateodus sp. jaw fragments (A) and teeth (B).

Scale bar = 1cm.

28 Class CHONDRICHTHYES Huxley, 1880

Subclass ELASMOBRANCHII Bonaparte, 1838

Order LAMNIFORMES Berg, 1958

Family ANACORACIDAE Casier, 1947

Genus Squalicorax Whitley, 1939

Squalicorax sp. indet

Diagnosis: The genus Squalicorax is defined by trenchant teeth that are compressed labiolingually. The roughly triangular main cusp has no cusplets and is serrated and convex lingually. The main cusp is directed anteriorly. The root is curved but not arcuate (Wolberg, 1985).

Collected Material: The material collected from the Hazel Creek locality of the genus

Squalicorax is represented by one tooth. The tooth is small, 6.5mm wide mesiodistally and 4.2mm tall. The tooth is labiolingually compressed and lingually convex. There is a single crown with fine serrations pointing strongly distally. The root is not tall, only about 1mm. There is a species from the genus Squalicorax known in the Cretaceous of

Australia known as Squalicorax falcatus Agassiz, 1843 (Wolberg, 1985). This species is characterized by a thick root and a wide pointed crown with prominent serrations. S. falcatus is taller than it is wide. These are not characters shared with the Hazel Creek

Squalicorax sp. tooth recovered. It is shares similar characteristics with an undescribed species of the genus from Bilelo (1969). A photograph of this specimen is presented below in Figure 16. It is more akin to the Hazel Creek tooth in the shallower nature of the root, the more finely serrated edge, and in the size and orientation of the main cusp.

Figure 15. Squalicorax sp. indet. from Hazel Creek Locality, lingual and labial views

Scale bar = 2mm

Figure 16. Squalicorax sp. indet from Bilelo (1969) (photo from Welton and Farish, 1993).

Scale bar = 2mm

30 Family ODONTASPIDIDAE Mükker and Henle, 1839

Genus Carcharias Rafinesque 1810

Type Species: Carcharias taurus Rafinesque 1810

Carcharias striatula Dalinkevičius, 1935

Diagnosis: As outlined by Siverson (1997) ‘Anterior teeth are up to at least 10.2mm high. There is one pair of small- to medium-sized cusplets. Short but very course labial folds are present along the base of the crown. The lingual face of the cusps has much longer but weak flexuous folds.’

Collected material: The teeth recovered from Hazel Creek (Figure 17) vary in their preservation state, most missing the tip of primary cusps, a lobe of the root, or one or both of the cusplets. The roots are bilobate and highly arcuate. There is a prominent medial groove in the root that can be accompanied by pitting. The main cusp is thin, long, and comes to a fine point, curving lingually. The main cusp has a length to width ratio between 1:4 and 1:2. There are a pair of cusplets on either side of the main cusp.

These teeth are varied in size from approximately 15mm down to 5.5mm from the apex of their main cusp to the base of their root. These sharks are of the same genus of extant sand tiger sharks.

31 -

Figure 17. Carcharias striatula teeth. Black scale bar = 1cm, white = 2mm

32 Family CRETOXYRHINIDAE Glikman, 1958

Genus Plicatolamna Herman (fide Cappetta and Case, 1975)

Plicatolamna arcuata Woodward, 1894

Diagnosis: As outlined by Wolberg (1985) ‘Moderately large teeth with massive, arcuate, bilobed roots and a medial primary cusp. The primary cusp is triangular, externally convex, and in complete specimens is flanked by small, but well-developed, anterior and posterior denticles. Short basal plications are most pronounced externally.

In anterior teeth the primary cusp is essentially vertical, whereas in lateral teeth it is inclined. Cusps in upper lateral teeth are not as tall as in lower laterals and the secondary denticles are proportionately larger.’

Collected Material: The collected single tooth is possibly a distal lateral tooth (Figure

18). It has a large, bilobate root, a large main cusp, and two cusplets or denticles. The root is arcuate and has a short, pronounced basal plication on its lingual face. The main cusp is large, triangular, lingually convex, and (presumably) distally curved. The cusplets are small, triangular and lack any curve. The tooth is 6.5mm wide from root tip to root tip and 6.0mm tall from root base to main cusp tip. The main cusp is 3.0mm tall, the root width is 2.5mm, and the cusplets are 1.2mm tall.

Figure 18. Plicatolamna arcuata tooth, lingual view. Scale bar = 5mm

Family CARDABIODONTIDAE Siverson, 1999

Genus Cardabiodon Siverson, 1999

Cardabiodon sp. indet

Diagnosis: As outlined by Siverson (1999) ‘Four anterior and about 14 lateroposterior teeth are present in both the upper and lower jaw. The first upper anterior tooth is about half the size of the second to fourth upper anterior teeth. There are no teeth on the inferred upper intermediate bar. The first lower anterior tooth is approximately two- thirds of the size of the second and third lower anterior teeth. Lower lateroposterior teeth are greatly enlarged, with the first and second lateroposteriors exceeding all upper and lower anterior teeth in size. The fourth lower anterior tooth is markedly reduced in size compared to adjacent anterior and lateroposterior teeth. Most teeth have one pair of lateral cusplets. Anterior and distally situated lateroposterior teeth may lack cusplets on one or both sides of the cusp. There is a marked dignathic heterodonty, expressed in a

34 relatively narrow and distally directed cusp in upper jaw teeth, as opposed to a broader, more erect cusp in lower jaw teeth. Moving through the files towards the commissure, the distal inclination of cusp increases in both the upper and the lower jaw. The lingual neck is well developed medially. One or two foramina open on the lingual protuberance of the root. There is no median groove. The lobes of the root become shorter and more divergent, moving posteriorly through the files.’

Collected Material: One large tooth represents the genus Cardabiodon from the Hazel

Creek locality (Figure 19). It is quite a large tooth measuring 31.9mm in length from root base to apex and 20.3mm in width across the root. The root is dilobate and arcuate, and lacks a medial groove. The main cusp is triangular in shape and quite wide where it meets the root of the tooth. It is 14mm long and 11mm wide. There are a pair of cusplets on either mesiodistal side of the main cusp. These cusplets are significantly smaller than the main cusp (3mm by 4mm).

Figure 19. Cardabiodon sp. indet tooth, lingual view. Scale bar = 1cm

35 Order ECHINORHINIFORMES de Buen, 1926

Family ECHINORHINIDAE Gill, 1862

Genus Echinorhinus de Blainville, 1816

Echinorhinus australis Chapman, 1909

Diagnosis: This species has varied in nomenclature first as Corax australis (Chapman,

1908), then referred to Squalicorax australis by Cappetta (1987), followed by

Pseudocorax australis (Kemp, 1991) and finally reassigned to the genus Echinorhinus

(Kemp, 1996). Kemp’s (1991) photos (Figure 21) offer the best diagnosis. A labiolingually compressed, single cusped tooth with no cusplets.

Collected Material: Collected material of Echinorhinus australis consists of teeth in various states of completeness (Figure 20). The mesiodistal width of the teeth is between 5.5mm and 9mm. The crown consists of a single cusp with no additional cusplets. This cusp is highly compressed labiolingually and elongate in a distal direction. The cutting edge of the tooth is smooth across the cusp. The root is highly labiolingually compressed and has a width 1/3-1/2 of the entire tooth height. The root is simple and possesses horizontal grooves. The species E. cookei Pietschmann, 1928 and

E. brucus Bonnaterre, 1788 are modern day, extant members of the genus. They are the prickly shark and bramble shark, respectively.

Figure 20. Echinorhinus australis teeth. Black scale bar = 1cm, White = mm

Figure 21. Echinorhinus australis, as pictured by Kemp (1991).

37 Superorder HOLOCEPHALI

Order CHIMAERIFORMES

Suborder CHIMAEROIDEI Patterson, 1965

Family CALLORHYNCHIDAE Garman, 1901

Subfamily CALLINORHYNCHINAE Stahl, 1999

Genus Ptyktoptychion Lees, 1986

Ptyktoptychion tayyo Lees, 1986

Diagnosis: As outlined by Barthalomai (2008) ‘Moderate sized. Length of palatine symphyseal margin in holotype is 53mm (but is 98mm in QMF17853), while width is

38mm, giving a minimal L: W ratio of 2.6. The length of the mandibular tooth plate is

110mm while the width is 35mm, yielding a L: W ratio of 3.1. Labial margins deeply dentate especially in mandibular tooth plate. Oral surface of palatine tooth plate with three ridges. Middle ridge about three quarters as strongly developed as the innermost ridge. Grooves between all ridges deep. Four tritors present. Mandibular tooth plate with three tritors. Four ridges on oral surface at labial margin with inner three only progressively slightly weaker from innermost outwards and with three well defined grooves present.’

Collected Material: The sole specimen representing Ptyktoptychion tayyo is a rather large tooth plate. The tooth plate is likely the right mandibular plate with the posterior outer tritor and anterior outer tritor preserved. Everything below the anterior outer tritor

(symphyseal tritor, oral surface, and symphyseal margin) has not been preserved. The specimen is 66.3mm in length, 47.0mm in height, and 19.0mm in width. While the specimen is not complete (neither were those used in the descriptive study

[Bartholomai, 2008]) the length to width ratio is 3.5. The posterior outer tritor has an

38 average width of 4.7mm while the anterior outer tritor has an average width of 3.1mm.

The other comparable species is Ptyktoptychion wadeae Bartholomai, 2018. It, however, has a smaller length width ratio of 2.7 and has a total of five tritors present. The posterior inner tritor and the median tritor are not present on the Hazel Creek specimen.

Figure 22. Ptyktoptychion tayyo right mandibular tooth plate. Lingual view (A), dorsal view (B),

dorsal anterior view (C). Scale bar = 1cm 39 Order TESTUDINES Batsch, 1788

Superfamily CHELONIOIDEA Baur, 1893

Clade PANDERMOCHELYS Joyce et al., 2004

Family PROTOSTEGIDAE Cope, 1872

Genus Notochelone Lydekker, 1889

Notochelone costata Owen, 1882

Diagnosis: As outlined by Kear (2003) ‘Small protostegid turtle with plastron length <1 m. Skull with nasals present and large temporal vacuity. High lingual ridge of maxilla exposed from lateral ridge; medial process of jugal absent. Prefrontal-postorbital in contact and barely allowing the frontals to reach the orbital rims. Interorbital bridge wide. Parietal-squamosal contact absent. Foramen caroticum laterale and canalis carotici interni not noticeably larger in diameter than the foramen anterius canalis carotici interni and the medial branch of the canalis caroticus internus; canalis caroticus lateralis small in diameter compared to the canalis caroticus internus. Rod-like rostrum basisphenoidale. Possibly amphicoelous cervical vertebrae. Carapace with marked neural keel; plastron with very large central and peripheroplastral fontanelles, reducing the hyoplastral-hypoplastral contact to a narrow projection; xiphiplastra much broader than long and medially curved with a large mid-line fontanelle. Scapular angle wide.

Coracoid longer than humerus. Lateral process of humerus restricted to anterior portion of shaft and bearing a medial concavity.’

Collected Material: Many fragments of Notochelone costata were recovered from the site, the majority of which were of minimal diagnostic significance or poorly preserved.

Two jaws were recovered. One prepped out and the other remains in situ due to its

40 fragile state. The loose jaw is 26.8mm long and 27.1mm wide with a height of 9.2mm

(Figure 23.A). The in situ jaw is 52.7mm long and 37.5mm wide (Figure 23.B). The small size of these jaws could indicate juveniles of the species. The posterior end of both jaws was not preserved. Limb bones had also been recovered in what could be interpreted as a humeri growth series. The adult humerus (Figure 24.A) is broken just past halfway and missing the distal end. It measures 38.4mm in length, the head has a width of 12.2mm, and its narrowest point measures 9.4mm. The sub-adult humerus

(Figure 24.B) is mostly complete with minor erosional damage. It measures 47.1mm in length, the head has a width of 17.4mm, and its narrowest point measures 8.5mm. The juvenile humerus (Figure 24.C) is complete and lacks the large processes it would have developed later in life. It measures 31.5mm in length, the head has a width of 13.9mm, and its narrowest point measures 7.0mm. If this is indeed a growth series, it could be the first correlated for this species. Fragmented elements of the carapace peripherals and plastron have been identified as well. The peripherals (Figure 26) range between

12.1mm and 18.0mm wide and 18.9mm and 20.8mm long. The plastron fragment

(Figure 25) is 68.3mm long by 68.5mm wide with and average thickness of 6.5mm.

There are three turtles known from the Australian Cretaceous Notochelone costata,

Cratochelone berneyi, and Bouliachelys suteri (Kear and Lee, 2006). Cratochelone berneyi has a plastron length of around two metres (Kear, 2003) and Bouliachelys suteri has a plastron length of around one and a half metres (Kear and Lee, 2006), while

Notochelone costata has a plastron length of less than one metre (Kear, 2003). The fragments of shell that have been found in the Hazel Creek deposit agree in terms of size with referral to N. costata.

41 A1 A2

Figure 23. Notochelone costata jaw bones. Dorsal and ventral views (A&B), ventral view (C).

Scale bars = 1cm

42 A1

Figure 24. Notochelone costata humeri growth series. Adult, dorsal and ventral views, (A), sub-

adult (B), and juvenile (C). Scale bar = 1cm

Figure 25. Notochelone costata plastron, dorsal and ventral views. Scale bar = 1cm

Figure 26. Notochelone costata carapace peripherals, ventral view. Scale bar = 1cm

44 Order ICHTHYOPTERYGIA Owen, 1840

ICHTHYOSAURIA de Blainville, 1835

Family OPHTHALMOSAURIDAE Baur, 1887

Genus Platypterygius Wade, 1990

Platypterygius sp. cf. P. australis

Diagnosis: Large-bodied ichthyosaur up to 9 m in length. Skull low-crowned with long snout, small orbit and long postorbital region. Maxilla extremely long anteriorly.

Dentition robust with roots of teeth quadrangular in cross-section. External naris subdivided; septomaxilla well ossified. Squamosal lost. Condylus occipitalis semi- hemispherical with area extracondylaris extremely reduced. Stapes large with rounded head. Atlas-axis co-ossified with third cervical vertebra; intercentra not differentiated.

Humerus with very strong trochanter dorsalis and two or three distal facets. Anterior and posterior accessory digits of forefin well developed with all podial elements very thick, forming a close-fitting polygonal mosaic pattern. Pelvic girdle and hind limb poorly known but apparently reduced. Caudal peduncle short. (Kear, 2003)

Collected Material: Three vertebrae and a single paddle bone of Platypterygius sp. cf.

P. australis were recovered from the site. Two of the vertebrae are identifiable as dorsal and caudal, while the third remains too damaged for identification. The dorsal vertebra

(Figure 28.A) is 43.1mm wide, 37.5mm tall, and 13.7mm thick. It has rib facets just above the centre midline of the centrum and toward the ventral end of the vertebra. It is a very round pentagonal shape. The caudal vertebra (Figure 28.B) is 30.8mm wide,

29.0mm tall, and 11.3mm thick. It is a rounded teardrop with a prominent notochord- foramen. The eroded vertebra (Figure 28.C) is 57.7mm wide, 59.7mm tall, and 14.7mm

45 thick. No notable features are preserved on this specimen. The single paddle bone

(Figure 27) is roughly pentagonal in shape, 43.7mm long, 37.5mm wide, and 13.7mm thick. Platypterygius australis McCoy, 1867 is the only currently known species of the genus Platypterygius known from Australia. There is no evidence that this material does not represent Platypterygius australis.

Figure 27. Platypterygius sp. cf. P. australis paddle bone, dorsal and ventral views. Scale bar = 1cm

A B

C B

Figure 28. Platypterygius sp. vertebrae. Dorsal (A), caudal (B), indet (C). Scale bar = 1cm

47 Order SAUROPTERYGIA Owen, 1860

Suborder PLESIOSAURIA de Blainville, 1835

Superfamily PLESIOSAUROIDEA Welles, 1943

Family ELASMOSAURIDAE Cope, 1869 (sensu Ketchum & Benson, 2010)

Gen. et sp. indet.

Diagnosis: As outlined by Andrews (1910) ‘Head relatively small; neck long, in some cases excessively so. Cervical ribs with single head. Scapulae meeting in the middle line, where they join the corresponding median anterior prolongations of the coracoids, at least in fully adult individuals. Clavicles and interclavicles may both be present, but one or both are usually greatly reduced. Episodically bones much modified, being shortened up so as to resemble mesopodials.’

Collected Material: The elasmosaurid material consists of a single tooth (Figure 29), vertebra (Figure 31), and phalange (Figure 30). The vertebra is half preserved, with one face slightly crushed, while the other is worn clean away. In its current state, the vertebra is 137.6mm wide and 90.2mm tall. The centrum is elliptical bulging mediolaterally and slightly concave (the vertebrae likely being platycoelous). The centrum is 108.0mm wide and 63.2mm tall. The neural arch is present, if barely, though the dorsal and both transverse processes are missing. The neural canal is small and a rounded triangle. Both rib facets are intact shaped like bottom-heavy figure eights.

These are positioned above the centreline of the centrum. The more complete rib facet is

56.8mm tall and 25.32mm wide at its widest point. The phalange is 74.5mm long eroded at both ends. The centre diameter is 21.3mm expanding to 37.5mm and 30.7mm respectively at each end. The tooth is broken at the base and fractured near the top. It is

48 22.5mm long and 9.1mm wide at the base. It is slightly curved, slightly conical though buccolingually compressed, and has fine, thinly spaced striations running from base to tip.

Figure 29. Elasmosauridae tooth, labial and lingual views. Scale bar = 1cm

Figure 30. Elasmosauridae phalange. Scale bar = 1cm

. Scale bar 1cm=

view and profile view profile and view

anteroposterior

Elasmosauridae vertebra

31. e

Figur

50 Clade SAURIA Macartney, 1802

Clade ? DINOSAURIA Owen, 1842

Gen. et sp. indet.

Collected Material: Large fragment of dense bone approximately 65.3mm long and

24.2mm wide at widest point (Figure 32). The fragment is curved slightly. It is likely a fragment of a long bone that weathered in an exfoliative manner, fracturing along growth lines.

Remarks: While this bone may not seem remarkable, it is large for the average bone recovered from the Hazel Creek Deposit. It is also not fish, mammal, or bird due to its size and density. It is potentially plesiosaur or ichthyosaur, but seems more likely to be dinosaur. Beyond that speculative distinction, it is unlikely that further diagnoses can be made; however, it an important indicator of other potential specimens. Further research needed to look at bone microstructure is beyond the scope of this study. It should also be noted that the bone has an exfoliation-like weathering pattern which would require terrestrial based weathering.

Figure 32. Possible dinosaurian limb fragment. Scale bar = 1cm

Class AVES Linnaeus, 1758

Clade ORNITHOTHORACES Chiappe, 1995

Clade ENANTIORNITHES Walker, 1981

Family Gen et. sp. incertae sedis

Collected Material: A number of small fractured limb bones were recovered from the

Hazel Creek locality (Figure 33). These limb bones were between 7mm and 12mm long and 1.5mm and 3mm wide. They were unfortunately broken at both distal and proximal ends leaving only the centre shaft. Damaged as they were, their rounded, hollow nature, small size, and time period of occurrence provide substantial evidence for description as enantiornithean avialans. While further identification of these specimens is unlikely, the discovery of more complete elements will hopefully prove useful toward that end. The material collected is not pterosaur elements. This is clear as pterosaur bone is not

52 typically preserved with the hollow of the bone intact. Most long bones are flattened during the preservation process. Those that are intact are generally triangular or rectangular in cross-section rather than the circular nature of the collected

Enantiornithes material.

Figure 33. Hollow limb fragments from Enantiornithes. Scale bar = 1cm

53 Class MAMMALIA Linnaeus, 1758

Subclass incertae sedis

An isolated premolar represents the Class Mammalia in the Hazel Creek assemblage. This is of considerable importance as it is now the only known Mesozoic mammal from the northern half of the Australian continent, the nearest other occurrences were the Early Cretaceous Griman Formation of Lightning Ridge, New

South Wales. It is also important as it adds weight to the conclusion reached in this study that the Hazel Creek assemblage was accumulated in a shallow marine environment not far from land, or at least closer to land than the environment in which the Richmond fossil assemblage accumulated (see Discussion below).

Description: While the Hazel Creek premolar is distinctly mammalian in form, it exhibits a relatively plesiomorphic premolariform morphology which makes it difficult to assign to any particular subclass, let alone family or genus previously known.

Comparisons are made here with the only other mammals from Australia that are represented by Mesozoic taxa known from premolars: ausktribosphenids and monotremes. While it might have additional similarities to premolars of other groups not yet known from the Australian Mesozoic, further comparisons would be more useful once additional teeth, hopefully molars, have been recovered from the Hazel Creek

Locality.

This premolar is in all probability a left lower premolar of uncertain serial position. This is likely because the convex curvature of the primary blade, which is nearer the tooth margin than the concave side, the primary cuspid (protoconid) has a marked convex flank that sweeps up from the side, interpreted here to be the buccal side

54 of the crown and there is a shallow talonid basin that occupies more of what is interpreted to be the lingual flank of the talonid.

The crown is long and narrow with an attenuated triangular basal outline with the apex just anterior to the protoconid and the wider portion below the posterior part of the crown. Both tooth roots are robust and vertically deep and are at least twice the height of the crown. The posterior root is longer and more robust than the anterior root.

The saddle between the roots is substantial, only just shorter than the length of the anterior root.

There is only one principle cusp, the protoconid. At the posterior margin of the occlusal surface, there is a small posterior cuspid that provides support for the posterior end of the crown’s primary blade. This blade which descends posteriorly from the apex of the protoconid, after a concave inflection at a position above the posterior root of the tooth, then ascends to and terminates on the buccal apical flank of the small posterior cuspid. There appears to be a curved anterior vertical blade that extends from the apex of the protoconid down towards the base of the anterior end of the crown, on the lingual side. There are no cingulids. From the lingual view, the primary blade from the protoconid is capable of being honed via thegosis to maintain sharpness in the leading edge of the blade, with longitudinal hollowing subtending this blade on the lingual side.

There may be a slight subvertical ridge developed in the talonid basin of the tooth, developed on the anterior flank of the posterior cuspid.

The tooth is relatively unworn except for a slight wear facette on the trailing edge of the primary posterior blade subtending the protoconid and a small bit of apical wear at the tip of the protoconid. Given the state of preservation of the tooth, it is not entirely clear how far down or even below the swollen base of the crown the enamel extended.

Discussion: In comparison with other Australian Mesozoic mammals, the Hazel Creek premolar is unlike the posterior premolars of Bishops whitmorei and Ausktribosphenos nyctos in that these Victorian Cretaceous taxa (Rich & Vickers-Rich 2004, Rich et al.

1997, 2009b) have premolars that are not as elongate and triangulate with narrow anterior ends and wide posterior ends. Although the P1 of B. whitmorei is vaguely similar, it appears to have (Rich et al. 2009b) distinct basal cingulids which the Hazel

Creek tooth entirely lacks. Posterior premolars of both of these Victorian taxa have posterior cingulids, features entirely lacking in the Hazel Creek premolar. Further, posterior premolars of the Victorian taxa have virtually trigonid-like attributes with three primary cusps, again entirely unlike the Hazel Creek tooth. Further, none of the

Victorian taxa have such widely space roots under the crown as the Hazel Creek tooth suggesting that the latter may have been part of a more extended premolar tooth row with larger gaps between and hence less crowding of the teeth.

Comparison with early monotremes do not provide any greater argument for similarity. The oldest known monotreme, and the only one with relatively conventional premolars, is Teinolophos trussleri (Rich et al. 2016). The only known premolars are present in the most recently described jaw (ibid). While this tooth, a P4, is premolariform, the outline of the damaged crown suggests that the primary cuspid was more centrally situated in the crown with an only miniscule posterior cuspid. The outline of the crown is also unlike that of the Hazel Creek tooth in that it is wider at its anterior than its posterior end.

No other Mesozoic mammals known from Australia have premolariform premolars other than Corriebaatar marywaltersae, which has been interpreted by Rich et al. (2009a) to be a multiuberculate with a plagiaulacoid posterior premolar. The

56 identity of this taxon as a multituberculate should be in doubt given the similar structure of other pre-therian groups known from the Early Cretaceous of South America.

However, whatever its affinities, the tooth present in this Victorian Cretaceous taxon is utterly unlike that of the Hazel Creek premolar.

It must be concluded, therefore, that the Hazel Creek premolar represents an as yet unknown Mesozoic mammal from Australia. When this unique and intriguing taxon is better known, it may be possible to determine its phylogenetic relationships to other groups of Mesozoic mammals.

Figure 34. Hazel Creek mammal incertae sedis premolar. Scale bar = 1mm

Figure 35. Comparative mammal premolars of the Australian Cretaceous. Hazel Creek premolar

(A), Corriebaatar marywaltersae (B) (Rich et al. 2009a), Teinolophos trusleri (Rich et al. 2016) (C),

Ausktribosphenos nyktos (Rich et al. 2016) (D), and Bishops whitmorei (D) (Rich et al. 2009b).

(Photos not to scale)

58 Coprolites

Collected Material: The coprolites retrieved from the Hazel Creek deposits (Figure 36) come in an array of sizes from about 5cm long with a diameter of 2cm to small millimetre sized fragments (Table 1 provides a list of photographed coprolites. All mostly intact and not fragmentary). They are generally a buff colour with shades of iron oxide and chalk.

Remarks: One of the more common preserved elements recovered from the Hazel Creek locality are coprolites. These coprolites came in various sizes from 13-14mm in length and 4-5mm in diameter to coprolites 50mm in length and 19-20mm in diameter While detailed analysis of these coprolites is outside the scope of this study, they are important to recognize. It is likely that the majority of the coprolites are from fish species.

Coprolites can be useful in determining information on species diet (Qvarnström, 2019) and paleoenviornment (Bull et al., 2002). Objects preserved in the coprolite can be used to identify diet and animal of origin as well as the possibility of locating new species.

Unfortunately, thus far no objects have been located within the coprolites recovered at the Hazel Creek locality.

Figure 36. Various coprolites from Hazel Creek Locality. Scale bar = 1cm

60 DISCUSSION

Australian Cretaceous Fauna Review

Australia’s early-mid Cretaceous faunas are some of the least well-documented

Cretaceous faunas globally. The documented faunas are preserved in a number of deposits including the Surat Basin in north-central New South Wales and south-east

Queensland, the Gippsland and Otway Basins in coastal Victoria, and Eromanga Basin in central Queensland. These three faunal groups are discussed here to provide context for the Hazel Creek Locality within Australia’s Cretaceous.

The Griman Creek Formation of Lightning Ridge, in the Surat Basin (Albian–

Cenomanian) has been reviewed and described by Bell et al. (2019). The faunas of this formation are as follows: a diverse invertebrate record is present that includes crayfish, tharid, viviparid and succineid gastropods, hyriid and unionid bivalves, and rare aminiferans. Various forms of bacteria, slime moulds and fungi have also been identified. Marine teleosts are rare, evidenced by rare aspidorhynchid scales. Extremely rare lamniform chondrichthyans are also present represented by an undescribed taxon.

Dipnoans are conversely diverse, represented by numerous isolated tooth plates pertaining to a variety of ceratodontiform taxa. Shell fragments, limb elements, and vertebrae of testudines have been recorded as among the most common vertebrate remains recovered; possible chelid, meiolaniform, and meiolaniid-like turtles have been identified though few taxa have been named. At least two crocodyliforms have been recorded, a conical-toothed eusuchian and an indeterminate broad-snouted, ziphodont mesoeucrocodylian. Isolated plesiosaurian teeth are relatively abundant, which are best referred to as a species of Leptocleidia or a leptocleidid-like taxon. Anhanguerian pterosaurs are represented by two isolated tooth crowns. Dinosaurians present are

61 primarily ornithopods along with an indeterminate ankylosaurian, isolated titanosauriform sauropod teeth, and an indeterminate megaraptorid theropod. A few distal ends of femora represent enantiornithine birds. Mammals are known from this locality by two taxa: Kollikodon ritchei, an australosphenidan, and Steropodon galmani, originally described as a steropodontid but referred to Ornithorhynchidae.

The Victorian Gippsland and Otway basins (Barremian–Albian) are contained within the Strzelecki and Otway groups. In a review of the faunas of the basins, Poropat et al. (2018) outlined the taxa present as follows. The invertebrate taxa are significantly more diverse within the Strzelecki and Otway groups. There are 13 orders and more than 70 insect species currently recorded along with two specimens of araneans and a single opilionid. Only a single species of xiphosuran has been recovered. The most common invertebrates are freshwater crustaceans. There are also three groups of branchiopods, a single ostracod, and a single syncarid. A number of other invertebrates have been recorded, but not well described including relatively common annelid mesofossils, freshwater bivalves, and uncommon bryozoans. Teleosts are represented by an array of relatively common actinopterygian fishes. Three species of dipnoans have been described based on tooth plates. A number of undescribed temospondyls, all within Brachyopoidea, are recorded. Reptiles are represented by isolated teeth and bones of small-bodied ‘pliosauromorphs’ that may be leptocleidids, and relatively common chelonians, of which only two taxa have been described. There are no recorded crocodylomorphs or pterosaurs within either the Strzelecki or Otway group faunal assemblages. Within Dinosauria there are an array of non-avian theropods (none of which are described), an indeterminate relatively small armoured ankylosaur, minimal ceratopsian records, and a few ornithopods. Dinosaur ichnofossils have been recovered but, again, none described. Three feathers discovered were recovered, originally

62 presumed to represent birds, but could just as well be from a feathered theropod. An ornithuromorph bird, specifically an enantiornithian furculum, is the only concrete evidence for the presence of avians. Just under 60 mammal specimens have been found, most correlating to ausktribosphenids and monotremes.

Eromanga Basin Cretaceous assemblages (Albian–Turonian) contain the Allaru,

Mackunda, Winton, and Toolebuc Formations. In a review of the Eromanga Basin fauna, Cook (2012) outlines the basic geology and existing taxa as follows. The Winton

Fm is a terrestrial deposit and has produced undescribed insects, teleosts, and dipnoans.

Reptilian elements from this formation include a dolichosaur, crocodylomorphs, testudines and pterosaurs. Dinosaurians make up the remainder of the fauna with the putative allosauroid Australovenator, the lithostotian titanosaurian Diamantinasaurus, and the titanisauriform Wintonotitan. The Mackunda Fm is reportedly a marine deposit with a diverse benthic and nektic fauna. Bivalves consist primarily of the genera

Inoceramus and Aucellina accompanied by multiple other genera and a number of solemyiids. Belemnites are abundant, while ammonites tend to be less common.

Gastropods are present in shell beds throughout the formation as undescribed small naticiids and higher-spired gastropods. Teleosts are uncommon, while chondrichthyans are abundant. The Allaru Mudstone has a number of common ammonite genera and low bivalve diversity that includes species of Inoceramus and Aucellina. Crustacea are locally abundant, ophiuroids are rare, and monasterid seastars are known. A single faviid coral has also been recovered. From the Toolebuc Fm, an array of ammonite genera has been noted along with the a single nautiloid genus. Teuthid genera have been recovered along with species of the belemnite genera Dimitobelus. Teleosts are abundant represented by amiids and aspidorhynchids. The chimaeroid and lamnid chondrichthyans as well as dipnoans have been recovered. Plesiosaurians known from

63 this Formation include the pliosaur Kronosaurus queenslandicus and a number of indeterminate elasmosaurids. Platypterigius australis is the only ichthyosaur so far known. Three species of testudines, Cratochelone berneyi, Notochelone costata, and

Bouliachelys suteri., are well-documented (Kear and Lee, 2006). Dinosaurian elements have been recovered including Minmi sp., and Muttaburrasaurus langdoni along with as of yet undescribed sauropod elements. A small indeterminate pterosaur has been recovered along with the enantiornithean bird Nanatius eos.

Comparison of the Hazel Creek Local Fauna

The Hazel Creek Local Fauna contains two species of bivalve, a species of

Inoceramus whose fragments are relatively common and a rarer species of Aucellina. A single ancyloceratid coiled heteromorph ammonite (Myloceras ammonoides) and highly abundant belemnoids (species of Dimitobelus). Teleost remains are the most abundant and diverse group present in the deposit. Lamniforms and chimaeroids have a low abundance while echinorhiniforms have a high abundance and represent the most common chondrichthyan. The chelid Notochelone costata is abundant, represented by juveniles, sub-adults, and adults. Elasmosaurid plesiosaurians and the ichthyosaur

Platypterygius australis are abundant. Indeterminate enantiornithine birds are relatively abundant given the sample size of collected material. Mammals are represented by a single taxonomically ambiguous premolar.

On balance, the Hazel Creek Local Fauna shares most in common with local faunas from the Eromanga Basin, primarily Toolebuc Fm, collected at localities like

Richmond and Hughenden. This seems to support the assertion that Hazel Creek deposits primarily composed of the Toolebuc Fm. Abundant taxa occurring in Hazel

Creek and Eromanga Basin at large include the bivalves, chondrichthyans and teleosts,

64 marine reptiles, and enantiornithes. The Hazel Creek Local Fauna, as so far known, lacks a number of groups that are present in contemporaneous Toolebuc assemblages such as terrestrial arthropods, gastropods, dipnoans, crocodylomorphs, non-avian dinosauria, and pterosaurs. While this may be an accident of collection biases or limited sample size, genuine absence from this palaeocommunity is a distinct possibility. With more extensive searching and acid-processing of the limestone, we might discover other species of these, and other contemporaneous groups.

The major distinction between the localities comes in relative abundances. The

Hazel Creek Locality shows increased abundances of taxa that would indicate a shallower and potentially nearer-shore deposit than those of Richmond. Marine turtles, while one of the most frequently recovered tetrapods, have a greater abundance of younger individuals present with in the Hazel Creek Locality. The minute size of some of the collected specimens suggest they are likely juveniles. Juvenile marine turtles will often remain in shallower waters until reaching a sufficient size for life in open water.

For the sample size, the avian abundance is higher than what is recorded at deeper and further off-shore deposits. While birds frequently alight over water, all have to return to land at some point to roost and breed. The size of the referred avian material also seems to suggest smaller species, which would be closer tied to the land. The presence of other terrestrial taxa, such as the potential Dinosaurian fragment and the indeterminate mammal premolar, provide indication of a near-shore deposit as well, as such material would have to be washed from shore.

Conclusions

The goals of this study were to complete a preliminary taxonomic review of the

Hazel Creek Locality and set the ground work for future study. This paper also seeks to

65 stress the importance of the unique mammal premolar discovered at the site. The Hazel

Creek local fauna was found to be comparative with other Toolebuc sites within the

Eromanga basin, however, its variances indicate that the deposit may have been shallower and closer to shore than many sampled previously. The mammal premolar was found to represent an as yet unknown Mesozoic mammal from Australia when compared with currently described specimens. With this preliminary study complete, continued processing is paramount for future discoveries.

Future study

The blocks of Hazel Creek limestone that have so far been processed have provided clear indications that further work will be rewarded with better specimens of known taxa and the appearance of additional, rarer taxa that have yet to be recovered.

Gaps within the Cretaceous fauna could start to be filled with the discovery of other terrestrial taxa that are more likely to turn up in this assemblage than other deposits that appear to be sampling faunal assemblages further off-shore. Additional avian specimens and those of non-avian dinosaurians, crocodylomorphs, plesiosaurians and even other terrestrial sauropsids, such as lacertilians and serpentes, would be greatly welcomed. In particular, more of the mammal material will be of great importance given the scarceness and low diversity of Australia’s Mesozoic mammals that have been discovered to date. The comparative ease of processing the limestone blocks should aid in rapidly expanding the known species from the site.

As well, further study should also be focused on microscopic elements such as radiolarians, diatoms, pollen, fungi, and other less conspicuous organisms that would provide additional information about proximal and distal paleoenviornments and

66 potentially help to biocorrelate this assemblage with others known from the Toolebuc

Formation.

Focus on the stratigraphy of the deposit would help to clarify potential differences in microfacies. Materials collected to date have been confined to surface blocks of limestone in what appears to be a relatively flat-lying sedimentary system.

Effort to excavate below this level might reveal older faunal assemblages that have so far not been sampled further. This excavation could enable refined biocorrelation of the

Hazel Creek Local Fauna with other Toolebuc assemblages. Geologic coring should be undertaken to establish transects across the locality.

Wider surveys of the Hazel Creek locality and subsequent mapping of the relative abundances within the site could lead to possible indicators of palaeocurrent distribution. Having a more complete knowledge of this would allow future collection teams to be most efficient instead of random collection patterns.

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