pliocene crocodilians of chinchilla: identification using dental morphometrics · 2018. 6. 18. ·...

Pliocene Crocodilians

of Chinchilla:

Identification Using

Dental Morphometrics

Christina Chiotakis

Bachelor of Applied Science (Ecology)

Supervised by Dr Matthew Phillips (QUT) and Dr Scott Hocknull (Queensland

Museum)

Submitted in fulfilment of the requirements for the degree of Master of Science

(Research)

Faculty of Science and Engineering

Queensland University of Technology

2018

CHRISTINA CHIOTAKIS 08301166

1

Keywords:

Chinchilla Local Fauna, Crocodilian, crocodile, Crocodylus, Pallimnarchus,

Pliocene, Quinkana, teeth

Abstract:

The following thesis examines the Pliocene crocodilians of the Chinchilla

Local Fauna. Crocodilians are an extremely diverse group of mostly

amphibious, opportunistic hunters that include twenty-three extant species

across the globe. A number of extinct species have also been described from

Australia’s prehistory. Two species have been described from the Chinchilla

Local Fauna, Pallimnarchus pollens and Quinkana sp. Teeth fossilise

readily, and so make up the largest dataset of fossilised evidence of

crocodilian species. For this study, the teeth were grouped visually,

measured and scanned to created three-dimensional models for analysis.

When graphed on a PCA, it is possible to distinguish two distinct groups of

teeth. The two groups are differentiated based on overall shape of the tooth.

One group contains conical teeth, the other laterally compressed teeth.

Further analysis of the teeth was able to determine the approximate position

of the teeth along the tooth row. Alveoli on available jaw specimens were

also measured to determine if there was a way to match isolated teeth to

them. However, when jaw specimens were measured the site shows

potential for three different crocodilian taxa occurring within the Chinchilla

Local Fauna. Further research will need to be conducted to determine the

third potential species and correct positioning of isolated teeth along the

tooth row.


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0.0 CONTENTS PAGE

Section Page Number

0.0 Contents Page 2

0.1 List of Figures 3

0.2 List of Tables 6

0.3 List of Abbreviations 6

0.4 Glossary 6

Statement of Original Authorship 8

Acknowledgements 8

1.0 Introduction 9

1.1 Objectives 9

1.2 Significance 10

2.0 Literature Review 10

2.1 Crocodilians 10

2.2 Cenozoic Crocodilian Fossil Record 13

2.3 Plio-Pleistocene Extinction 18

2.4 Chinchilla Sands Local Fauna 20

2.5 Australian Crocodilians 25

2.5.1 Extant Australian Crocodilians 25

2.5.1.1 Crocodylus porosus 25

2.5.1.2 Crocodylus johnstoni 26

2.5.2 Extinct Australian Crocodilians 27

2.5.2.1 Pallimnarchus sp. 27

2.5.2.2 Quinkana spp. 28

2.6 Varanus priscus 29

2.7 Fossilised Teeth 30

2.8 Future Research into Extinct Crocodilians 32

2.9 Conclusion 33

3.0 Methodology 34

4.0 Ethics and Limitations 37

5.0 Results 37

5.1 Initial Dataset 37

5.1.1 Raw Data 38


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5.1.2 Principle Components Analysis 38

5.2 CT Scanned Dataset 41

5.2.1 Raw Data 41

5.2.2 Root Means Square Data Matrix 42


5.2.4 Non-Metric Multi-Dimensional Scaling 43

5.3 Caniniform and Molariform 45

5.3.1 Raw Data 46



5.4 Analysis Based Upon Ratio 52

5.4.1 Ratio of Mid-Height and Base Length versus

Width

52


5.5 Alveoli Measurements 58

5.5.1 Crocodylus sp. 58

5.5.2 Pallimnarchus sp. 59

5.5.3 Quinkana sp. 60

5.5.4 Unknown Species 61

6.0 Discussion 61

6.1 Future Research 69

7.0 Reference List 72

8.0 Appendices 87

9.0 Supplementary Figures 116

0.1 List of Figures

Figure Number and Title Page Number

1 Ranges and Estimated Population Size of Extant

Crocodilian Species

11

2 Artists Impression of an Early Triassic Crocodilian 12

3 Possible Homo sapiens migration routes 19

4 Map of Main Queensland Fossils Sites 20


4

5 Map of Crocodylus porosus Distribution across Australia 25

6 Map of Crocodylus johnstoni Distribution across

Australia

26

7 Location of Pliocene and Pleistocene Crocodilian bearing

Fossil Sites

28

8 Palaeogeography of the Giant Varanids 30

9 Examples of ziphodontid and conical Crocodilian Teeth 32

10 Diagram of Crocodilian Teeth 35

11 Principle Components Analysis with Crocodylus spp. 39

12 Principle Components Analysis without Crocodylus spp. 40

13 Principle Components Analysis with Crocodylus spp.

without Height

40

14 Principle Components Analysis without Crocodylus sp.

and Height

41

15 Principle Components Analysis of 3D Dataset 42

16 Principle Components Analysis of 3D Dataset without

Height

43

17 Non-Metric Multi-Dimensional Scaling of 3D Dataset 44

18 Non-Metric Multi-Dimensional Scaling with Teeth 45

19 Side and Base View of Teeth from Non-Metric Multi-

Dimensional Scaling Graph

45

20 Principle Components Analysis of Caniniform Teeth 46

21 Principle Components Analysis of Caniniform Teeth

without Height

47

22 Principle Components Analysis of Molariform Teeth 47

23 Principle Components Analysis of Molariform Teeth

without Height

48

24 Non-Metric Multi-Dimensional Scaling of Caniniform

Dataset

49

25 Non-Metric Multi-Dimensional Scaling with

Caniniform Teeth

50

26 Side and Base View of Caniniform Teeth from Non-

Metric Multi-Dimensional Scaling Graph

50


5

27 Non-Metric Multi-Dimensional Scaling of Molariform

Dataset

51

28 Non-Metric Multi-Dimensional Scaling with

Molariform Teeth

51

29 Side and Base View of Molariform Teeth from Non-

Metric Multi-Dimensional Scaling Graph

52

30 Histogram Based Upon the Ratio of Mid-Height Length

vs Width

53

31 Histogram Based Upon the Ratio of Crown Base Length

vs Width

53

32 Scatterplot of Crown Base Length/Crown Base Width

vs Height

54

33 Scatterplot of Mid-Height Length/Width vs Height 54

34 Scatterplot of Mid-Height Length/Width vs Crown Base

Length/Width

55

35 Non-Metric Multi-Dimensional Scaling Coloured by

Ratios

56

36 Non-Metric Multi-Dimensional Scaling Coloured by

Ratios with Teeth

57

37 Side and Base View of Teeth from Non-Metric Multi-

Dimensional Scaling Graph

57

38 Alveoli Measurements of Crocodylus spp. 58

39 Alveoli Measurements of Pallimnarchus spp. 59

40 Alveoli Measurements of Quinkana spp. 60

41 Alveoli Measurements of Unknown species 61

42 Dentary Specimens of Modern Crocodylus porosus and

Crocodylus johnstoni

66

43 Fossilised Dentary Specimens 67

44 Quinkana Tooth from Mount Etna Deposits 69

45 Pallimnarchus sp. Dentary from the Darling Downs 71

46 Crocodilian Dentary from Winton 71


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0.2 List of Tables

Table Number and Title Page Number

1 Cenozoic Crocodilian Species from Australia and

Oceania

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2 Species belonging to the Chinchilla Local Fauna 21

0.3 List of Abbreviations

PCA – Principle Components Analysis

QM – Queensland Museum

QMF – Queensland Museum Fossil

CT (Scan) – Computed Tomography (Scan)

MDS – Multi-Dimensional Scaling

RMS – Root Means Square

Mya – Million Years Ago

0.4 Glossary

Labial – referring to cheek

Lingual – referring to tongue

Ziphodont – Laterally compressed, distally curved,

serrated crowns (D’Amore and Bluenschine, 2009)

Alveoli – Crypt in the jaw in which teeth are imbedded

Osteoderm – Bony plate under the skin

Maxilla – upper jaw

Dentary – lower jaw

Mekosuchine – Distinct group of Australia-western

Pacific crocodilians belonging to the Eusuchia

Eusuchian – Clade to which all extant crocodilians

belong

Pterygoid – Bone forming part of the palate in a skull

Local Fauna – Species making up a spatially and


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temporally discrete fauna (e.g. Chinchilla Local Fauna


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Statement of Original Authorship:

The work contained in this thesis has not been previously submitted to meet

requirements for an award at this or any other higher education institution. To

the best of my knowledge and belief, the thesis contains no material previously

published or written by another person except where due reference is made.

Signature: QUT Verified Signature

Date: June 2018

Acknowledgements:

I would like to thank my supervisors’ Dr Matthew Phillips and Dr Scott

Hocknull for all their input and encouragement throughout the project. I

extend this thank you to the Vertebrate Collection Managers Dr Andrew

Amey and Dr Patrick Couper at the Queensland Museum for allowing me

access to the modern crocodilian specimens. And thanks to my family and

friends for their continued support throughout the completion of this

research. I would also like to thank everyone at Geosciences, Queensland

Museum, in particular, Kristen Spring for allowing me access to the

collection, and Nikki Newman and Rochelle Lawrence for co-ordinating CT

Scanning for the fossil teeth. A special thanks to Greenslopes Private

Hospital and QX-ray for access to the CT Scanner for data collection, and

the Phillips Lab Team, in particular, Carmelo Fruciano, for aiding with the

statistical analysis.


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1.0 Introduction:

This chapter outlines the objectives and significance of the research. This

will be followed by a literature review of current and relevant scientific

research to provide a background on the topic. The methodology will then

be outlined followed by the results of the research. Finally, there will be a

full analysis of all data collected, relating back to the relevant literature and

potential future applications of the work.

1.1 Objectives

The aim of this study is to find a way of identifying crocodilian species and

species diversity for a single Local Fauna, the Chinchilla Local Fauna, by

utilising the most abundant fossils available. Inspection of the Queensland

Museum vertebrate fossil collection has identified teeth as one of the most

common fossils found of crocodilians from the Chinchilla Local Fauna, due

to their durability and constant shedding throughout the life of the animal.

Osteoderms are the second-most common, followed by post-cranial remains

and then finally fragmentary cranial elements. Differences in tooth shape

and size relate to dietary preferences of the species that possessed them,

therefore it is likely that determining the diversity of crocodilian taxa from

within the Chinchilla Local Fauna may be possible from teeth alone.

However, at a gross level, crocodilian teeth are very similar between species

compared to the diversity of shape seen in mammalian dentition. Therefore,

a new methodology will be introduced and tested to determine whether

shape can differentiate species of crocodilian from a single Local Fauna

based only on teeth.

A secondary objective of this project was to determine whether these

abundant isolated teeth could be associated with less common fragments of

jaws recovered from the Chinchilla Local Fauna. The tooth diversity should

reflect the diversity of jaws and overall shape disparity between the isolated

teeth and will be compared to shape disparity of alveoli found within these

jaws.


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1.2 Significance

This research is important because previous identification of fossil

crocodilians from Chinchilla have been based almost entirely on

fragmentary cranial remains which may be significantly underestimating the

diversity of species present within the Local Fauna simply due to the rarity

of these fossils. Using the most abundant crocodilian fossil recovered from

Chinchilla, their teeth, provides the greatest opportunity to sample the true

diversity of crocodilian taxa. Using the methods described below provides a

more objective comparison, which can later be adapted to other fossil types

and species as required. Being able to assign isolated teeth to fragmentary

toothless jaw fragments from the same Local Fauna will test the total

species diversity from within the Local Fauna and provide additional data

for future comparisons and taxonomic description.

2.0 Literature Review:

2.1 Crocodilians

Crocodiles are the most diverse surviving group of predatory megafauna in

non-marine ecosystems across the globe. There are three recognised

families of crocodilians: the Alligatoridae, Crocodylidae and Gharialidae

(Norell, 1989Pough et al, 2009;). These families include twenty-three extant

species (Janke et al, 2005; Meganathan et al, 2010), twelve of which belong

to the genus Crocodylus (Brochu, 2000). Most species of crocodilian are

distributed throughout the tropics and subtropics of the world, with a few

species found in warm temperate regions (Brazaitis and Watanabe, 2011;

Pough et al, 2009) (Figure 1).


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Figure 1: The ranges and estimated population sizes of crocodilian species

across the world, represented on a map. (National Geographic Society,

1996-2010). Since the creation of this image, fossil evidence has been found

as far south as Antarctica (Brochu, 2003).

Extant crocodiles live a highly amphibious lifestyle (Naish, 2001) therefore

their distribution is directly linked to fresh, estuarine and saltwater (Delfino

et al., 2007). It is uncommon to see two or more species living sympatrically

today and where this does occur, the species substantially differ

morphologically (Brochu, 2001). However, the fossil record of crocodiles

shows a greater degree of sympatry and species packing (Salisbury and

Willis, 1996). Variation in the morphology of the cranium, in particular, the

shape of the snout, along with tooth shape and body size express different

degrees of ecomorphological evolution relating to the preferred mode of

prey apprehension and dietary preference of each species (Densmore and

Owen, 1989; Pough et al., 2009). Many species of crocodilian are ambush

predators (Enax et al., 2013) preying opportunistically on a large range of

prey (Erickson et al., 2012; Webb et al., 1982).


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Crocodilians first evolved during the early Triassic Period (approximately

250 million years ago) with more modern looking and amphibious

crocodilians appearing during the Jurassic Period (Bellairs, 1969; Brazaitis

and Watanabe, 2011). Based on morphological features of the skeleton it is

hypothesised that the first crocodilians were likely terrestrial predators that

later evolved the amphibious lifestyle we see today (Naish, 2001). Some of

these early features used to infer a terrestrial lifestyle include slender leg

and foot bones as well as uncompressed tail vertebrae (Naish, 2001) (Figure

2). Crocodilians have dominated the land-water interface as apex predators

since the Cretaceous Period (approximately 85 million years ago) (Erickson

et al., 2012). Fossilised crocodilians have been found across a wide

geographical range spanning as far North as Ellesmere Island, Canada, and

as far South as Antarctica (Brochu, 2003) (Figure 1).

Figure 2: Artist’s impression of an Early Triassic crocodilian,

Barberenasuchus brasiliensis (Naish, 2001).

Most fossil evidence of crocodilians comes in the form of isolated teeth and

osteoderms (bony armour plates under the skin) (Mackness et al., 2010).

Teeth have been described as having two main forms: molariform and

caniniform (Erickson et al., 2012). Caniniform teeth grow in the anterior

part of the jaw and are generally large and conically shaped to grip prey

(Enax et al, 2013). Molariform teeth are shorter and stouter, growing in the

back of the jaw and are used to crush the bones and other rigid components

of prey. Though crocodilians today have conical shaped teeth with a

rounded base, some extinct forms had ziphodont (serrated, labio-lingually

compressed) teeth. This is possibly related to a terrestrial lifestyle, as may


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be inferred from their similar morphology to both theropod dinosaurs and

extant varanids, both of which have ziphodont dentition. The only extant

reptiles with ziphodont teeth are the varanids, which includes the largest

living lizard, the Komodo dragon (Varanus komodoensis).

2.2 Cenozoic Crocodilian Fossil Record

The Australian crocodilian record occurs in both the Mesozoic and

Cenozoic, with the Cenozoic record including taxa from both the

Palaeogene and Neogene periods (Salisbury et al., 2006; Willis, 1997). The

majority of the taxa have been recorded from sites in Queensland, the

Northern Territory and South Australia (Willis, 1997). Fossil crocodiles

have been found in lacustrine, fluvial and cave deposits (Table 1). These

sites have been studied with a primary focus on the mammalian faunas, and

therefore there are gaps in the crocodilian fossil record even though

crocodilian remains have been recovered from a wide variety of localities.

Studies have shown that the majority of species recorded throughout this

time belong to a subfamily of crocodylids, the Mekosuchinae (Brochu,

2003). The earliest known of these species is Kambara from the Early

Eocene Tingamarra deposits of South-East Queensland (Willis, 1997).

There are several species of crocodilian from the Oligo-Miocene of

Australia, all of them currently being placed within the Mekosuchinae. Baru

darrowi is known from two localities, Riversleigh, Queensland, and Bullock

Creek, Northern Territory. The earliest known species of Quinkana are also

recorded from the Oligo-Miocene, although the genus was initially

discovered in late Pleistocene cave deposits in Queensland. Mekosuchus

occurs in the Miocene deposits at Riversleigh, however, Quaternary records

of this genus are known from Vanuatu and New Caledonia (Mead, et al.,

2002). Other Australian crocodilian species that existed before the Plio-

Pleistocene include Trilophosuchus rackhami, Harpacochampsa

camfieldensis, and Australosuchus clarkae (Willis, 1997) (Table 1).


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Table 1: Cenozoic Crocodilian Species from Australia and Oceania.

Age Species Locality Site

Interpretation

Author/s

Early Eocene Kambara

murgonensis

Tingamarra, near

Murgon, South East

Queensland.

Tingamarra Fauna

Lacustrine Willis,

Molnar

and

Scanlon,

1993

Early Eocene Kambara

implexidens

Main Quarry,

Tingamarra Station,

Boat Mountain Area,

Murgon, South East

Queensland.

Tingamarra Local

Fauna

Lacustrine Salisbury

and Willis,

1996

Mid-Late

Eocene

Kambara

molnari

Stuart Deposit, The

Narrows Graben,

Brick Kiln Seam,

near Gladstone,

Queensland. Rundle

Formation

Lacustrine-

Estuarine

Holt,

Salisbury

and Willis,

2005

Mid-Late

Eocene

Kambara taraina Stuart Deposit, The

Narrows Graben,

Kerosene Creek

Member, near

Gladstone,

Queensland. Rundle

Formation

Lacustrine-

Estuarine

Buchanan,

2009

Late

Oligocene

Mekosuchus

whitehunterensis

White Hunter Site,

Riversleigh,

Queensland

Lacustrine-

Riverine

Willis,

1997

Late Quinkana White Hunter Site, Lacustrine- Willis,


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Oligocene meboldi Riversleigh,

Queensland

Riverine 1997

Late

Oligocene

Baru huberi White Hunter Site,

Riversleigh,

Queensland

Lacustrine-

Riverine

Willis,

1997

Late

Oligocene

Baru wickeni White Hunter Site,

Pancake Site, Sticky

Beak Site,

Riversleigh,

Queensland

Lacustrine-

Riverine

Willis,

1997

Oligo-

Miocene

Australosuchus

clarkae

Unnamed site, Lake

Palankarinna, South

Australia. Tedford

Local Fauna and

Tedford East Local

Fauna

Lacustrine Willis and

Molnar

1991

Late

Oligocene to

Mid Miocene

Baru darrowi Blast Site, Camfield

Beds, near Camfield

Homestead, North

Central Northern

Territory. D-Site,

Riversleigh,

Queensland

Lacustrine-

Riverine

Willis,

Murray

and

Megirian

1990

Early

Miocene

Trilophosuchus

rackhami

Ringtail Site, Gag

Plateau, Riversleigh,

Queensland

Cavernous Willis,

1993

Mid Miocene Harpacochampsa

camfieldensis

Camfield Beds,

Camfield

Homestead, North

Central Northern

Territory. Bullock

Creek Local Fauna

Lacustrine-

Riverine

Megirian,

Murray

and Willis,

1991

Mid-Late Quinkana timara Bullock Creek Lacustrine- Megirian,


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Miocene Locality, Camfield

Beds, Camfield

Station, Northern

Territory

Riverine 1994

Mid-Late

Miocene

Crocodilian

unknown species

Bannockburn

Formation,

Manuherikia Group,

New Zealand

Lacustrine Molnar

and Pole,

1997

Early

Pliocene

Quinkana

babarra

Dick’s Mother Lode

Quarry, Allingham

Formation, Bluff

Downs Station,

North East

Queensland

Riverine Willis and

Mackness,

1996

Early

Pliocene

Kalthifrons

aurivellensis

Golden Fleece

Locality,

Mampuwordu Sand

Member, Tirari

Formation, Lake

Palankarinna, South

Australia.

Lacustrine-

Riverine

Yates and

Pledge,

2016

Plio-

Pleistocene

Pallimnarchus

pollens

“the Condamine

Drift”, unknown

locality, probably on

the Darling Downs,

South East

Queensland

Riverine DeVis,

1886;

Molnar,

1982

Plio-

Pleistocene

Pallimnarchus

gracilis

Terrace Site,

Riversleigh,

Queensland.

Lansdowne Station,

Western Australia.

Tara Creek, New

Riverine Willis and

Molnar,

1997


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South Wales

Pleistocene Volia

athollandersoni

Wainibuku Cave,

Wainibuku Valley,

Viti Levu, Fiji

Cavernous Molnar,

Worthy

and Willis,

2002

Pleistocene

to Present

Crocodylus

johnstoni

Terrace Site,

Riversleigh,

Queensland

Riverine Willis and

Archer,

1990

Pleistocene

to Present

Crocodylus

porosus

Allingham

Formation, West of

Emu Valley

Homestead, North

Queensland

Riverine Molnar,

1979

Middle

Pleistocene

Quinkana sp. Mount Etna Caves,

Rockhampton,

Central Eastern

Queensland

Cavernous Hocknull,

2005,

Hocknull

et al,

2007,

Hocknull,

2009

Late

Pleistocene

Quinkana

fortirostrum

Tea Tree Cave,

Chillagoe, North

Queensland

Cavernous Molnar,

1977;

Molnar,

1982

Late

Pleistocene

Quinkana sp. King Creek, West of

Pilton, Eastern

Darling Downs,

Queensland

Riverine Sobbe,

Price and

Knezour,

2013

Late

Holocene

Mekosuchus

kalpokasi

Arapus

Archaeological Site,

Efate Island,

Vanuatu

Archaeological Mead, et

al., 2002


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Currently, Australian Plio-Pleistocene species of crocodilian include the

extinct Pallimnarchus pollens, Pallimnarchus gracilis, Quinkana

fortirostrum, Quinkana babarra, two unknown species of Quinkana and the

extant Crocodylus porosus (Molnar, 1982) and Crocodylus johnstoni (Willis

and Archer, 1990). Adding to this list are the fossil crocodilians studied for

this research that belong to the Pliocene Chinchilla Local Fauna from

Chinchilla, South-East Queensland (Figure 4).

2.3 Plio-Pleistocene Extinction

Megafaunal species were once common across Australia until

approximately 50,000 - 40,000 years ago, when most became extinct. Until

that time, many reptilian species, including crocodilians, varanid lizards and

snakes attained giant body sizes, exceeding 5 metres in length (Wroe, 2002).

This large body size made some of these species amongst the largest apex

megafaunal predators across the continent. However, by the end of the

Pleistocene, all, except Crocodylus porosus, were extinct. What caused this

extinction is still widely debated by many in the scientific community.

Currently the two main hypotheses focus on anthropogenic mediated

overkill or deteriorating climate mediated extinction.

The overkill hypothesis implicates humans, in particular, Aboriginal people,

as the direct cause of the megafaunal extinction due to rapid overhunting

and alteration of the environment (Barnosky and Lindsey, 2010; Wolverton,

2010). In Australia, this appears to be a widely accepted theory, even though

evidence to support this theory is largely indirect. Even across other

continents there are few megafaunal deposits that also contain evidence of

direct human predation. The climate change hypothesis implicates either

sudden climatic change during the last glacial period (Wolverton, 2010) or

long-term deterioration of the climatic system over several hundred

thousand years (Wroe et al., 2013). They suggest that megafaunal species

were unable to cope with this sudden or long-term deterioration in the

climatic system and so this change drove the megafauna extinct. Evidence

for these major climatic shifts is supported by geological, palynological and


19

palaeoclimatic evidence, however, there remains significant gaps in this

record.

It is widely accepted that humans first evolved in Africa, before migrating

across the globe. Through dating and analysis of mitochondrial and genomic

DNA (mtDNA), several studies have concluded that human migration

followed a path through Asia, following the southern coast (Malaspinas et

al., 2016; Stanyon et al., 2009) (Figure 3). Archaeological sites have dated

human arrival in Australia to be around 50,000 to 65,000 years ago

(Barnosky and Lindsey, 2010; Clarkson et al., 2017; Pushkina and Raia,

2008). Contrary to this, a recent study found that at least one species of

Australian megafauna, Zygomaturus trilobus, didn’t go extinct until

approximately 30, 000 years ago (Westaway et al., 2017), although support

for this date is wanting. However, at least 20,000 years of overlap in which

humans and megafauna co-existed would suggest that Homo sapiens did not

play a direct role in the megafauna extinction. There are no deposits in

Australia that conclusively shown direct evidence of human hunting of

megafaunal species (Cosgrove et al., 2010), or long-term evidence that

humans hunted them to extinction.

Figure 3: Homo sapiens migration routes across the world with estimated

arrival times (Stanyon et al., 2009). The ages marked on this map have since

been revised for human arrival within Australia to 65,000 years ago

(Clarkson et al., 2017).

This lack of evidence has also been used to support the climate change

hypothesis. Approximately 50,000 years ago Australia went from being a


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cold and dry continent to a warm and dry continent. The warming of

continental Australia caused a higher rate of evaporation which in turn

caused inland water systems to become scarce (Wroe et al., 2013). As the

water systems disappeared, so too did the vegetation they supported, giving

way to the barren Australia that we know today. It is likely that this climatic

deterioration created the conditions for the extinction of Australia’s

megafauna, in particular the aquatic species of crocodilians.

2.4 Chinchilla Sands Local Fauna

The fossil crocodilians used in this study were derived primarily from the

Pliocene Chinchilla Sands, south-east Queensland (Figure 4). In particular,

the isolated teeth used for this analysis come from a single locality to the

east of the Chinchilla town centre, QML 1. The fossils derived from this

locality come predominately from the formation known as the Chinchilla

Sands (Woods, 1960). These fossils are collectively referred to as the

Chinchilla Local Fauna.

Figure 4: Map of Chinchilla fossil locality relative to other major fossil

sites throughout Queensland (Price, 2012).

A large range of species have been recorded from the Chinchilla Sands,

including fish, reptiles, birds, and most extensively, mammals (Louys and

Price, 2015) (Table 2). Of the reptiles, Quinkana sp. and Pallimnarchus

pollens are the only identified crocodilian species. A large variety of

mammalian species have been described from this area, as they have been


21

the most extensively studied (Louys and Price, 2015). The mammalian

species range from the carnivorous Thylacinus cynocephalus (Mackness et

al., 2002; Ride, 1964) and Thylacoleo crassidentatus (Bartholomai, 1962),

through to the larger herbivorous species such as Euowenia grata (De Vis,

1887) and Protemnodon chinchillaensis (Bartholomai, 1973). The extensive

variety of species from this area, has given rise to several palaeohabitat

inferences (Montanari et al., 2013).

Table 2: List of species belonging to the Chinchilla Local Fauna as listed in

Louys and Price, 2015. *taxa not considered to be valid.

Family Genus Species Author/s

Trionychidae Gen. et sp. indet. DeVis, 1894b; Gaffney and

Bartholomai, 1979

Chelidae Emydura sp. Gaffney, 1981

Crocodylidae Pallimnarchus pollens De Vis, 1886

Crocodylidae Quinkana sp. Molnar, 1982; Louys and

Price, 2015

Agamidae Gen et sp. indet. Hutchinson and Mackness,

2002

Gekkonidae Diplodactylus cf. steindachneri Hutchinson and Mackness,

2002

Scincidae Cyclodomorphus sp. Hutchinson and Mackness,

2002

Scincidae Tiliqua wilkinsonorum Hutchinson and Mackness,

2002

Varanidae Varanus komodoensis Hocknull et al, 2009

Varanidae Varanus sp. Hutchinson and Mackness,

2002

Madtsoiidae Yurlunggur sp. Mackness and Scanlon,

1999

Casuariidae Dromaius novaehollandiae Patterson and Rich, 1987

Megapodiidae Leipoa gallinacea Boles, 2008

Anatidae Biziura lobata DeVis, 1888c; Olson, 1977


22

Anatidae Anas superciliosa DeVis, 1888c, Olson, 1977

Anatidae Aythya australis DeVis, 1888c; Olson, 1977

Phalacrocoracidae Microcarbo melanoleucos DeVis, 1888c; Miller,

1966b

Pelecanidae Pelecanus proavus DeVis, 1892; Miller,

1966a; Rich and van Tets,

1981

Ciconiidae Ciconia nana DeVis, 1888c; DeVis,

1892; Boles, 2005

Accipitridae Necrastur alacer DeVis, 1892; Rich and van

Tets, 1982

Rallidae Fulica atra DeVis, 1888c; Olson, 1975

Rallidae Gallinula mortierii DeVis, 1888c; Olson, 1975

Rallidae Charadriiformes gen. et

sp. indet.

DeVis, 1888c; Olson, 1977

Charadriidae Vanellus sp. DeVis, 1892

Archizonurus securus Mahoney and Ride, 1975

Dasyuridae Dasyurus dunmalli Bartholomai, 1971; Wroe

and Mackness, 2000a

Dasyuridae Archerium chinchillaensis Wroe and Mackness, 2000b

Thylacinidae Thylacinus cynocephalus DeVis, 1884a; Ride, 1964;

Mackness et al, 2002

Peramelidae Perameles bowensis Mackness et al, 2000

Indeterminate Koalemus ingens DeVis, 1889c;

Bartholomai, 1968

Indeterminate Koobor notabilis DeVis, 1889c; Archer,

1977

Phascolarctidae Phascolarctos ?stirtoni Price, 2008; Price et al,

2009

Diprotodontidae Euowenia grata DeVis, 1887

Diprotodontidae Euryzygoma dunense DeVis, 1888b; Longman,

1921

Diprotodontidae Zygomaturus sp. Archer and Wade, 1976


23

Palorchestidae Palorchestes parvus DeVis, 1895; Woods,

1958; Price and Hocknull,

2005

Vombatidae Vombatus ursinus DeVis, 1883a; Bartholomai

and Marshall, 1973;

Thylacoleonidae Thylacoleo crassidentatus Bartholomai, 1962

Phalangeridae Phalanger procuscus* DeVis, 1889c; Mahoney

and Ride, 1975

Macropodidae Brachalletes palmeri* DeVis, 1883b; Mahoney

and Ride, 1975

Macropodidae Synaptodon aevorum* Bartholomai, 1975;

Dawson and Flannery,

1985

Macropodidae Troposodon minor Bartholomai, 1966, 1967;

Flannery and Archer, 1983

Macropodidae Troposodon gurar Flannery and Archer, 1983

Macropodidae Sthenurus notabilis Bartholomai, 1963

Macropodidae Sthenurus andersoni Prideaux, 2004

Macropodidae Simosthenurus antiquus Bartholomai, 1963; Pledge

1980; Prideaux, 2004

Macropodidae Wallabia indra DeVis, 1895; Bartholomai,

1966 Bartholomai, 1976

Macropodidae Macropus pan Bartholomai, 1975;


1985

Macropodidae Macropus agilis siva Bartholomai, 1975;


1985

Macropodidae Macropus dryas Bartholomai, 1966, 1975;


1985

Macropodidae Macropus woodsi Bartholomai, 1975;



24

The presence of forest is inferred from the presence of tree and forest

dwelling species including Phascolarctus ?stirtoni (Price, 2008; Price et al.,

2009), Koobor notabilis (Archer, 1977; De Vis, 1889c), and Protemnodon

chinchillaensis (Bartholomai, 1973). Fossils from a number of large

herbivorous marsupials have also been collected, including Euowenia grata

(De Vis, 1887) and Euryzygoma dunense (De Vis, 1888b; Longman, 1921),

implying the presence of open grasslands. Several aquatic and semiaquatic

species, including the crocodilians Quinkana sp. (Louys and Price, 2015)

and Pallimnarchus pollens (De Vis, 1886), as well as turtle remains, suggest

permanent bodies of water were also present in the area (Montanari et al.,

2013).

The mosaic of palaeohabitats may not accurately reflect the true habitats at

the time simply due to a lack of systematic excavation with reference to

stratigraphic context. Therefore, fauna may represent a diachronous

assemblage and thus represent multiple times and palaeohabitats lumped

together in a single assemblage. Due to this, for the purposes of this study,

the usage of the term ‘Local Fauna’ will be used to describe the entire

faunal assemblage from the Chinchilla Sands with the assumption that this

1985

Macropodidae Prionotemnus palankarinnicus Bartholomai, 1975

Macropodidae Protemnodon devisi Bartholomai, 1973

Macropodidae Protemnodon chinchillaensis Bartholomai, 1973

Macropodidae Bohra wilkinsonorum Dawson, 2004a; Hocknull,

2005

Macropodidae Silvaroo bila Dawson, 2004b

Macropodidae Silvaroo sp. Dawson, 2004b

Chronozoon australe* DeVis, 1883c; Mahoney

and Ride, 1975

Muridae Pseudomys vandycki Godthelp, 1989

Molossidae Mormopterus sp. Hand et al, 1999


25

assemblage is diachronous across the Pliocene and possibly into the

Pleistocene.

2.5 Australian Crocodilians

From approximately five million years ago (5mya) to the present day, three

genera of crocodilians have lived in Australia. These genera include the

extinct Quinkana and Pallimnarchus, in addition to the extant Crocodylus.

2.5.1 Extant Australian Crocodilians

2.5.1.1 Crocodylus porosus

Crocodylus porosus (Estuarine Crocodile) is the largest extant crocodilian

(Meganathan et al., 2010; Pough et al., 2009) and has the largest distribution

of all extant species (Dodson, 2003; Hanson et al., 2015; Johnson, 1973;

Magnusson, 1980). In Australia, its range covers most of the Northern

coastline from Queensland to Western Australia (Cogger, 2014; Read et al,

2004) (Figure 5). The full range of Crocodylus porosus encompasses the

Indian Ocean through to the west of the Pacific Ocean, including India

through to the Philippines (Figure 1).

Figure 5: Distribution of Crocodylus porosus in Australia (Cogger, 2014)

The large distrubution of Crocodylus porosus is likely linked to its excellent

adaptation to salt water environments (Johnson, 1973). In addition to salt

water, Crocodylus porosus has been observed in brackish, coastal and

freshwater environments (Fukuda et al., 2008; Read et al., 2004).

Crocodylus porosus is a generalist feeder of aquatic prey (Erickson et al.,

2012; Hanson et al., 2015), however they also ambush land-based prey from


26

the water’s edge. To kill large prey that ventures close to the water’s edge, a

crocodile will propel itself out of the water with its powerful tail. It then

grabs the prey with strong conical teeth backed by a high bite force (Enax et

al., 2013). The earliest known fossil material of Crocodylus porosus has

been found in the Allingham formation, Queensland. The age of these

fossils shows that Crocodylus porosus has been in Australia from at least

the middle Pliocene (Molnar, 1979). In some parts of northern Australia,

Crocodylus porosus has been observed living sympatrically with C.

johnstoni, Australia’s only other extant crocodilian species (Webb et al.,

1983).

2.5.1.2 Crocodylus johnstoni:

Crocodylus johnstoni (Freshwater Crocodile) is Australia’s only extant

endemic crocodile (Tucker, 2010; Willis, 1997). Distributed across northern

Australia (Cogger, 2014), the Freshwater Crocodile is frequently found in

freshwater streams, rivers, lagoons, and lakes (Webb and Smith, 1984)

(Figure 6).

Figure 6: Distribution of Crocodylus johnstoni in Australia (Cogger, 2014)

Crocodylus johnstoni is a small crocodile usually growing to a maximum of

3 metres long and weighing around 60-70kg (Nolch, 2001; Webb and

Smith, 1984). It has a long slender snout with sharp, pointed teeth used to

catch its prey (Brochu, 2003; Erickson et al., 2012; Willis and Molnar,

1997). The preferred prey of C. johnstoni is fish (Johnson, 1973) however

their prey comprises a wide taxonomic range including crustaceans, insects,

small mammals, small amphibians, and birds (Erickson et al., 2012; Webb

et al., 1982; Willis and Molnar, 1997). Fossils of C. johnstoni are rare with


27

the oldest known from Pleistocene deposits at Riversleigh (Willis, 1997;

Willis and Molnar, 1997).

2.5.2 Extinct Australian Crocodilians

Throughout the Plio-Pleistocene, there were three known genera of

crocodilians living in Australia, Pallimnarchus, Quinkana, and Crocodylus.

Only Crocodylus is extant. All of these genera belong to the family

Crocodylidae, however only Pallimnarchus and Quinkana belong to the

subfamily Mekosuchinae. Crocodylus belongs to the group known as

Eusuchia, or the “True Crocodiles”. The Mekosuchinae differ from the

Eusuchia based on a number of features including a reduction or loss of the

anterior process of the palatine, a wedge formed by the supraoccipital on the

skulls dorsal surface, and a great difference between the smallest and largest

alveoli (Stein et al., 2012; Willis, 1997). Most of the Mekosuchinae became

extinct globally during the Late Tertiary (Brochu, 2003). However, in

Australia they survived well into the Quaternary and in the western Pacific

into the Holocene.

2.5.2.1 Pallimnarchus spp.

Pallimnarchus is an extinct genus of Australian crocodile that evidently

lived only in freshwater environments. Currently two species of

Pallimnarchus are recognised, Pallimnarchus pollens and P. gracilis (Willis

and Molnar, 1997). Pallimnarchus fossils have been discovered in various

deposits across central and east mainland Australia (Louys and Price, 2015;

Mackness et al., 2010; Willis and Molnar, 1997). Most deposits containing

Pallimnarchus fossils are found in Queensland, although they have also

been found as far south as northern New South Wales and South Australia

and as far west as Western Australia (Willis, 1997; Willis and Molnar,

1997) (Figure 7).


28

Figure 7: Location of Pliocene and Pleistocene crocodile-bearing deposits

in Australia (Willis, 1997).

Pallimnarchus was a large, robust crocodilian (Willis, 1997) growing to

about 5 metres in length (Molnar, 2004). It had large conical teeth in a broad

snout (Molnar, 1982; Molnar 2004; Willis and Molnar, 1997) with the orbits

directed upwards, suggesting an aquatic ambush predator (Molnar, 2004).

The broad snout and conical teeth have led palaeontologists to interpret

Pallimnarchus as a generalist feeder of large prey, much like the extant

Crocodylus porosus (Willis, 1997; Willis and Molnar, 1997).

2.5.2.2 Quinkana spp.

The earliest species of Quinkana date back to the Miocene, though the first

species described were discovered in Plio-Pleistocene deposits (Willis,

1997). Quinkana was endemic to Australia (Willis, 1997) and survived well

into the Pleistocene (Molnar, 2004). Several species of Quinkana existed

throughout this time period (Table 1).

Quinkana was a mekosuchine with ziphodont (serrated, laterally

compressed) teeth (Mackness and Sutton, 2000; Molnar 1982; Sobbe et al.,

2013; Willis 1997), and may have had a terrestrial or semi-terrestrial


29

lifestyle (Mackness and Sutton, 2000; Willis, 1997). It was a small

crocodile, growing to a maximum of 3 metres in length (Molnar, 2004).

Fossilised remains of Quinkana have been found throughout Queensland,

with possible recordings as far south as the Lake Eyre Basin, South

Australia (Molnar, 1982; Willis 1997; Willis and Molnar, 1997) (Figure 7).

The majority of Quinkana fossils have been found in cave deposits,

providing additional support to the terrestrial or semi-terrestrial hypothesis.

Where Quinkana has been found in ancient water systems (Molnar 1982)

the fossils have also been found with the remains of Pallimnarchus,

suggesting the two-species lived sympatrically and so occupied different

ecological niches within the same territories (Willis and Molnar, 1997).

2.6 Varanus priscus

Varanus priscus was the largest terrestrial lizard globally and lived in

Australia during the Late Pleistocene (Erickson et al., 2003; Hocknull et al.,

2009; Molnar, 2004). It is more commonly known as Megalania prisca,

however, review of its phylogenetic position places this taxon well within

the Varanus genus (Hocknull et al., 2009). Fossil evidence of Megalania has

been found throughout eastern Australia, ranging as far west as Lake Eyre

and as far South as Melbourne (Hocknull et al, 2009; Molnar, 2004) (Figure

8). Most fossils have been found in Queensland with several localities on

the Darling Downs (Molnar 2004; Price and Sobbe, 2005). Megalania

fossils are commonly found around rivers and streams, though some fossils

have also been found in cave deposits (Molnar, 2004).


30

Figure 8: Palaeogeography and chronology of giant varanids. Schematic

diagram illustrating the proposed taxonomy, chronology and dispersal

sequence of giant varanids from mainland Australia to the Indonesian

islands of Timor, Flores and Java during the Pliocene-Pleistocene (Hocknull

et al., 2009).

2.7 Fossilised Teeth

Teeth are made of dentine covered in a layer of enamel (Dauphin and

Williams, 2007) and are one of the most durable skeletal features

(Hendrickx et al., 2015). They readily fossilise and are one of the most

common crocodilian fossils. Variation in tooth morphology can provide

valuable insight into the palaeoecology and feeding behaviours of the

animals from which the teeth belong (Brink et al., 2015; Dauphin and

Williams, 2007). Isolated teeth are the most common crocodilian fossil

recovered. Though it is difficult to determine the exact position of an

isolated crocodilian tooth on a tooth row, they still provide insight into


31

palaeoecological diversity if they can be assigned to a taxon (Hendrickx et

al., 2015).

Extant crocodilian teeth are considered to be conical in shape, though

ziphodont forms did exist (D’Amore, 2009). The most recently extinct

ziphodont crocodilian is Quinkana. Ziphodont teeth differ from conical

teeth in that they are laterally compressed, often distally curved and have

defined serrations on the crown of the tooth (D’Amore and Blumenschine,

2009). Crocodilian teeth have evolved several specific traits to minimise

damage to the teeth during predation. Throughout the life of the animal,

teeth are continually replaced through a hollow root (Enax et al., 2013).

Teeth are also arranged in such a manner that they will not collide with one

another during predation (Enax et al., 2013). In some cases, this alignment

results in wear facets on the teeth where they may rub against each other for

extended periods of time.

Crocodilians have two forms of teeth. Caniniform, or canine-like teeth, are

situated anteriorly, and molariform, or molar-like teeth, that are situated

posteriorly (Erickson et al., 2012). Molariform teeth are much shorter and

stouter than caniniform teeth and tend to be used for crushing bone and flesh

much like the molars of mammals. In contrast, caniniform teeth, in most

extant species, are taller and more conical. Most crocodilians have conical

teeth and are rounded the entire way up the tooth. All known ziphodont-

toothed crocodilians are now extinct. Ziphodont teeth are laterally

compressed and have a much less rounded base than conical teeth (Figure

9).


32

Figure 9: A ziphodontid tooth (Top) and a conical tooth (Bottom) in both

lingual and basal view.

Reptiles known to have ziphodont teeth include extinct crocodilians,

theropod dinosaurs, and some of the extant varanids, such as the Komodo

Dragon (Varanus komodoensis) (Brink et al., 2015; D’Amore et al., 2011).

Based on the similarity in ziphodont dentition, ziphodont crocodilians,

including Quinkana, are thought to have similar behavioural and feeding

ecology as the modern ziphodont varanids (D’Amore and Blumenschine,

2009).

2.8 Future Research into Extinct Crocodiles

There has been minimal research on extinct crocodilian species, such as

Quinkana and Pallimnarchus. However, there have been hypotheses put

forth about how the Plio-Pleistocene crocodilians may have interacted with

and lived with one another in particular environments. The research to be

conducted as part of this project aims to decipher how many different

species of crocodilian lived during the Plio-Pleistocene in the Chinchilla

Local Fauna. To do this, the morphological differences between different

teeth will need to be examined to determine how many different species of

crocodilian may have been living in the area at that time. Variation along

the tooth row is taken into consideration, using modern specimens as a basis

for comparison for the fossilised teeth.


33

2.9 Conclusion

Crocodilians are one of the most diverse groups of non-marine predators

globally. The twenty-three-extant species of crocodilian are found in

tropical or sub-tropical ecosystems living an obligate amphibious lifestyle

(Brazaitis and Watanabe, 2011; Janke et al., 2005; Meganathan et al., 2010;

Pough et al., 2009). Extant crocodilians rarely live in sympatry with other

species, though it appears that this was more common in prehistoric forms,

based on the number of species found in fossil deposits. Crocodilians first

appear in the fossil record about 250 million years ago with more typical

looking crocodiles appearing during the Jurassic (Bellairs, 1969; Brazaitis

and Watanabe, 2011; Naish, 2001).

In Australia, C. porosus and C. johnstoni are sometimes observed living

sympatrically (Webb et al., 1983). Both species are found in the tropics of

Australia. Crocodylus porosus is more often found in salt water or estuarine

environments, while C. johnstoni is most often found in freshwater.

Prehistoric crocodilians of Australia are numberous (Willis, 1997), though

there are three main genera from the Plio-Pleistocene, Pallimnarchus,

Quinkana, and Crocodylus. It appears that the majority of large predators in

Australia were reptiles (Willis, 1997). The crocodiles make up a number of

these predators, though Megalania was also a large, land-based predator

throughout this time period (Erickson et al, 2003; Hocknull et al., 2009;

Molnar, 2004).

The vast majority of crocodilian fossils come in the form of teeth due to

their durability (Hendrickx et al., 2015). Fossil teeth vary morphologically

from species to species. This project will use linear and 3D geometric

morphometric methods to look at the morphological difference between

fossilised teeth to determine the number of species found in deposits and

infer the palaeoecology of the species. Further use of this research has the

potential to be used for other isolated teeth groups, such as theropod

dinosaurs.


34

3.0 Methodology:

Fossil and modern teeth used for this research are housed at the Queensland

Museum Geosciences and Vertebrate Collections respectively. Fossil

specimens are provided with the prefix QMF, and modern specimens a

prefix QMJ. Bulk fossil teeth specimens were taken from the Chinchilla

Locality of the QMF collection and sorted through for specimens that were

suitable for analysis. Specimens that were broken and heavily worn were

discarded from the study. A total of 306 fossilised teeth were initially sorted

into groups by me and labelled 1-15 based on the shape of the base using a

visual inspection. Before this, there was no prior grouping of the teeth aside

from them being placed together in the bulk collection. Group 1 were most

labio-lingually compressed in cross-section, while group 15 were circular in

cross-section at the base. To take measurements by hand for each tooth in a

logical manner, each of the teeth was given a number within its group. For

example, group 1 tooth 1 was labelled 1.1 and so on (See Appendix 1 for

full list).

A total of five measurements were taken of each tooth using digital callipers

(Figure 10). Each measurement was taken three times and then averaged to

get the most accurate measurements possible. These measurements were

labelled M1-M5 and are as follows; M1: dorso-ventral height of the tooth

measures from tooth base to tip; M2: mid-crown anterior-posterior length

measured half way up the tooth; M3 mid-crown labio-lingual width

measured half way up the tooth at the same height as M2; M4: crown-base

anterior-posterior length; M5 crown-base labio-lingual width measured at

the base of the tooth.


35

Figure 10: Diagram of crocodilian tooth with anatomical terminology and

measurements M1-M5. A) Labelled tooth crown. B) Labelled tooth base.

The modern crocodilian specimens (C. porosus (QMJ48127) and C.

johnstoni (QMJ39230) were measured using the same methods for the fossil

specimens. As the teeth of these specimens were in situ with the maxillae

and dentaries, they were numbered based on their position in the tooth row

and whether they were maxillary or dentary teeth. For example, the anterior-

most in the maxilla of Crocodylus porosus was labelled CPT1, with the

anterior-most tooth of the dentary labelled CPB1 (See Appendix 3 for full

list). T refers to the top jaw (maxilla) and B to the bottom (dentary). Both

modern specimens were adults at the time of death and had been donated to

the Queensland Museum vertebrate collection.

All the measurements taken were compiled into a Microsoft Excel spread

sheet and the averages calculated to use for analysis (Appendix 1). There

were several unknown measurements where teeth had been chipped,

cracked or broken pre- or post-deposition and so the data was cleaned to

remove these specimens (Appendix 2).

The best-preserved teeth were then CT Scanned at Greenslopes Hospital

using a Siemens Dual Energy Somatom Scanner. 3D models of each tooth


36

were generated from this data. The CT Scan data was loaded into

InVersalius 3.0 (Galantucci, L.M., 2010), a CT imaging freeware to produce

3D surface models of each specimen. As the teeth had to be scanned in large

groups, Meshlab (Siotto et al., 2015), a freeware for opening and editing 3D

models, was used to render each tooth individually into its own individual

model which could then be used for direct comparison with each other.

Once each tooth had been rendered, they were culled to include only fully

preserved tooth specimens. Teeth with missing sections, cracks or large

wear facets were excluded from the group to allow for the most comparable

dataset for the analysis.

Analysis of the scanned teeth was performed using CloudCompare (Hassett

and Lewis-Bale, 2017), a freeware which can compare 3D models to one

another and provide a measure of shape similarity and fit. Each tooth was

uploaded individually to CloudCompare and compared to each other tooth.

Due to the density of the points available in the 3D models a subset of one

million points was extracted from each surface for point-point comparison.

Three landmarks were picked to align each tooth used in each comparison;

the apex of the tooth, and the anterior and posterior points on the base of the

tooth (Figure 10). Using these points, CloudCompare aligns the models

based on the subset point clouds, using millions of iterations to provide the

closest fit of shape. It reports a computed statistic fit using a Root Mean

Square (RMS) value of each point measured across each point cloud

compared. As a result, this statistic will return a size-independent

comparison of shape fit between each compared pair of teeth. The lower the

RMS statistic, the closer in overall shape the two teeth are, removing size as

a possible bias. Each pair comparison reported RMS was entered into excel

spreadsheet to create a data matrix for later analysis.

All the datum was compiled using Microsoft Excel due to ease of use in

being able to hold multiple data sets in one file. Most of the statistical

analysis was done using PAST (PAlaeontological STatistics). PAST 3.18

was used to run Principle Components Analyses (PCA) on the linear

measurement data sets. PAST was also used to analyse the Non-Metric

Multi-Dimensional Scaling (MDS) analysis of the RMS data matrix


37

generated from CloudCompare. Due to the large data sets used for this

research PCA are useful in teasing apart groups of teeth shape data that may

not be easily discernible upon visual inspection only. A Non-Metric MDS

was used on the data matrix as it is a clear way of visualising the similarities

between teeth (Hassett and Lewis-Bale, 2017).

4.0 Ethics and Limitations:

There were no ethical considerations for this research as most the specimens

used were fossils and had died approximately 5 million years ago. The

modern Crocodylus specimens were donated to the Queensland Museum

after their time of death.

There were a few limitations with the research. The dataset was limited to

fossil teeth found within the Queensland Museum Fossil collection, and then

narrowed to those found from the Chinchilla area due to the time restrictions

within a Masters of Research. Teeth that had poor preservation, appeared to

be heavily reworked, worn or damaged were also excluded from the data set

as they would not have given accurate measurements relative to better

preserved specimens. The exact provenance of the specimens would also be

questionable. The age of the crocodilians from which the teeth come is near

impossible to determine. However, though the size of crocodilian teeth

increases with age, shape does not appear to change (Poole, 1961). Modern

specimens were also limited because the Queensland Museum has only two

adult Crocodylus skulls with in situ teeth to use for comparison and costly

travel to other collections was outside the scope of this research project.

Display skulls were also excluded because they have undergone

considerable preparation work to make them safe as touch specimens for the

public. Therefore, their teeth were considerably altered and thus would not

have provided accurate data.

5.0 Results:

5.1 Initial Dataset

The initial data set was collected using measurements taken manually using

callipers. A total of five measurements were taken of each tooth specimen


38

suitable to be included in the data set. Measurements were given short hand

notations as follows: M1, M2, M3, M4 and M5. M1 is the height of the

tooth measured from the base to the tip. M2 is the anterior-posterior

measurement half way up the tooth. The height half way up the tooth was

determined by dividing the average measurement of height by two. M3 is

the labio-lingual measurement, also half way up the tooth. Measurements 4

and 5 were both taken at the base of the tooth. M4 is the anterior-posterior

measurement, and M5 is the labio-lingual measurement. The teeth were also

added into groups visually based on the compression of the base. Groups

were labelled 1 to 15, with 1 being highly compressed and 15 being almost

circular in shape.

Modern specimens were also measured in the same way from skulls stored

at the Queensland Museum. The modern skulls belonged to the two species

of crocodilian extant in Australia, C. porosus and C. johnstoni.

5.1.1 Raw Data

The raw data includes data from all teeth selected as suitable to be analysed

for the research required. Each measurement, as listed in the methodology

(Section 3.0), was taken three times and then averaged to get the most

accurate measurements possible (Appendix 1). Some of the teeth selected

had wear facets or were chipped at the base. These gaps were filled in with

“?” for the initial data set. These specimens were deleted from the data set to

have a full data set without any unknown measurements (Appendix 2). As

much of the data is from unknown extinct species, modern specimens of

known origin were also included in the data set. In Australia, there are two

extant species of crocodilian, C. porosus and C. johnstoni. They were

measured in the same manner as the fossil specimens, however it was noted

where along the tooth row the teeth sit (Appendix 3).

5.1.2 Principle Components Analysis

A Principle Components Analysis (PCA) is typically used for large data sets

to account for correlations within the data that wouldn’t usually be visible


39

due to the amount of variability between measurements. All graphs were

generated using the statistical analysis program PAST.

Figure 11: Principle Components Analysis based on the clean data set,

including modern specimens and all five measurements taken of teeth

specimens. Crocodylus porosus are plotted using Red Crosses, Crocodylus

johnstoni are plotted using Blue Squares and the fossil teeth are plotted

using Black Dots.

A Principle Components Analysis (Figures 11-14) show that fossil teeth

occupy far broader morphospace than either of the modern species samples.

These figures are based on the clean data set, i.e. there were no specimens

with unknown measurements included in the data set. The modern

crocodilian specimens tend to plot relatively close to other teeth from the

same species (Figure 11), with very few outliers. Crocodylus porosus is

more widely scattered and overlaps almost entirely with the fossil

specimens. The C. johnstoni teeth fall within narrower morphospace, mostly

just outside of that occupied by the fossil and C. porosus teeth. In figures 7

and 8, the height (M1) is included as a component, however it appears to

provide too much stress on the data set and so was later removed. The

modern specimens were removed from several of the graphs to see how the

fossil teeth would plot on a PCA without their influence (Figure 12). The

height (M1) was included here as a possible component to allow for

comparison with the other graphs in which height was removed.

M1

M2

M3

M4

M5M5

1.1

1.2

1.4

1.5

1.61.7

2.1

2.2 2.3

2.4

2.5

2.6

2.7

2.8

2.9

2.10

2.11

2.122.13

3.1

3.2

3.3

3.5

3.6

3.7

3.8

3.12

3.13

3.14

3.153.16

4.2

4.3

4.4

4.5

4.6

4.7

4.8

4.9

4.10 4.11

4.13

4.14

4.16

4.17

4.18

4.19

5.2

5.3 5.4

5.5

5.6

5.7

5.9

5.10 5.11

5.12

5.13

5.14 5.15

5.16

5.17

5.18

5.20

5.21

5.22

5.23

5.24

5.25

5.26

5.27

6.1

6.2

6.36.4

6.5 6.7

6.8

6.9

6.10 6.12

6.13

6.15

7.1

7.3

7.4

7.5

7.6

7.7

7.87.9

7.10

7.11

7.12

7.13

7.14

7.15

7.16

8.2

8.3

8.4

8.5

8.6

8.7

8.88.9

8.10

8.118.14

8.16

9.1

9.2

9.3 9.4

9.5

9.6

9.7

9.8

9.9 9.10

9.12

9.13

9.17

9.18

9.19

9.20

9.21

9.22

9.23

9.24

9.259.26

9.27

9.28

9.29

10.2

10.4

10.6

10.7

10.8

10.10

10.11

10.12

10.13

10.15

10.16

10.17

10.19

10.20

10.22

10.2310.24

10.25

10.26

10.27

10.28

10.29

10.3010.31

10.3210.33

11.2

11.3

11.4

11.5

11.6

11.7

11.8

11.9

11.10

11.11

11.12

11.13

11.14

11.16

11.17

11.19

11.20

11.21

11.22

11.23

11.24

11.2512.1

12.2

12.3

12.5

12.6

12.712.8

12.9

12.10 12.11

12.13

12.14

12.15

12.16

12.17

12.19 12.20

12.21

13.1

13.2

13.3

13.5

13.7

13.8

13.10

13.11

13.12

13.14

13.15

13.16

13.17

13.18

13.19

13.21

13.23

13.25

14.1

14.2

14.3

14.5

14.7

14.8

14.10

14.11 14.13

14.14

14.15

14.16

14.17

14.18

14.19

14.20

15.1

15.2

15.4

15.5

15.6

15.7

15.8

15.9

15.10

15.11

15.12

15.13

15.14

15.15

15.1615.17

15.19

CPT1 CPT2CPT3

CPT5

CPT6 CPT7

CPT8

CPT9

CPT10CPT11CPT12

CPT13CPT14

CPT15

CPT16

CPT17

CPB1

CPB2

CPB3

CPB4

CPB5

CPB6

CPB7

CPB8

CPB9CPB10

CPB11

CPB12

CPB13

CPB14

CPB15

CJT1

CJT2

CJT3

CJT4

CJT5CJT6

CJT7

CJT8

CJT9

CJT10CJT12CJT13

CJT14CJT15

CJT16

CJT17

CJT18

CJT19

CJB1

CJB2CJB3

CJB4

CJB5CJB6CJB7CJB8CJB9

CJB10

CJB11CJB12

CJB13CJB14

CJB15-30 -24 -18 -12 -6 6 12 18 24

Component 1

-8

-6

-4

-2

2

4

6

8

Com

ponen

t 2


40


excluding modern specimens; all five measurements taken of teeth

specimens are included in this graph.


including modern specimens, with height excluded as a component of the

analysis. Crocodylus porosus are plotted using Red Crosses, Crocodylus

johnstoni are plotted using Blue Squares and the fossil teeth are plotted

using Black Dots.

M1

M2

M3

M4

M5M5

1.1

1.2

1.4

1.5

1.61.7

2.1

2.22.3

2.4

2.5

2.62.7

2.8

2.9

2.10

2.11

2.122.13

3.1

3.2

3.3

3.5

3.6

3.7

3.8

3.12

3.13

3.14

3.15

3.16

4.2

4.34.4

4.5

4.6

4.74.8

4.9

4.10

4.11

4.13

4.14

4.16

4.17

4.18

4.19

5.2

5.3 5.4

5.5

5.6

5.7

5.9

5.10 5.11

5.12

5.13

5.14

5.15

5.16

5.17

5.18

5.20

5.21

5.22

5.23

5.24

5.25

5.26

5.27

6.1 6.2

6.36.4

6.5

6.7

6.8

6.96.106.12

6.13

6.15

7.1

7.3

7.4

7.57.6

7.7

7.8

7.9

7.10

7.11

7.12

7.13

7.14

7.15

7.16

8.2

8.3

8.4

8.5

8.6

8.7

8.8

8.9

8.10

8.11

8.14

8.16

9.1

9.2

9.3 9.4

9.5

9.6

9.7

9.8

9.99.10

9.12

9.13

9.17

9.18

9.19

9.20

9.21

9.22

9.23

9.24

9.25

9.26

9.279.28

9.29

10.2

10.4

10.6

10.7

10.810.10

10.11

10.12

10.13

10.15

10.16

10.17

10.19

10.20

10.22

10.23

10.24

10.25

10.26

10.27

10.28

10.29

10.30

10.31

10.3210.33

11.2

11.3

11.4

11.5

11.6

11.7

11.8

11.9

11.10

11.11

11.12

11.13

11.14

11.16

11.17

11.19

11.20

11.21

11.22

11.2311.24

11.2512.1

12.2

12.3

12.5

12.6

12.7

12.8

12.9

12.10

12.1112.13

12.14

12.15

12.16

12.17

12.19

12.20

12.21

13.1

13.2

13.3

13.5

13.7

13.8

13.10

13.11 13.12

13.14

13.15

13.16

13.17

13.18

13.19

13.21

13.23

13.25

14.1

14.2

14.3

14.5

14.7

14.8

14.10

14.11

14.13

14.14

14.15 14.16

14.17

14.1814.19

14.20

15.1

15.2

15.4

15.5

15.6

15.7

15.815.9 15.10

15.11

15.12

15.13

15.14

15.15

15.1615.17

15.19

-20 -15 -10 -5 5 10 15 20 25

Component 1

-6

-4

-2

2

4

6

8

10

Com

ponen

t 2

M2

M3

M4

M5

1.11.2

1.4

1.5

1.61.7

2.1

2.2

2.3

2.4

2.5

2.6

2.7

2.8

2.9

2.10

2.112.12

2.13

3.13.2

3.3

3.5

3.6

3.7

3.8

3.12

3.133.14

3.15

3.16

4.2

4.34.4

4.5

4.6

4.7

4.8

4.9

4.10

4.11

4.13

4.14

4.16

4.17

4.18

4.19

5.2

5.3

5.4

5.5

5.6

5.7

5.9

5.10

5.11

5.12

5.13

5.14

5.15

5.16

5.17

5.185.20 5.21

5.22

5.235.24

5.25

5.26

5.27

6.1 6.2

6.3

6.4

6.5

6.7

6.8

6.9

6.10

6.12

6.13

6.15

7.1

7.3

7.4

7.5

7.6

7.7 7.8

7.9

7.10

7.11

7.12

7.13

7.14

7.15

7.16

8.2

8.3

8.4

8.5

8.6

8.7

8.8

8.9

8.10

8.118.14

8.16

9.1

9.2 9.3

9.4

9.5

9.6

9.7

9.8

9.99.10

9.12

9.13

9.17

9.18

9.19

9.20

9.21

9.22

9.23

9.24

9.25

9.26

9.27

9.28

9.29

10.2

10.4

10.6

10.7

10.8

10.10

10.11

10.12

10.13

10.15

10.16

10.17

10.19

10.20

10.22

10.23

10.24

10.25

10.26

10.27

10.28

10.29

10.30

10.31

10.32

10.3311.2

11.3

11.4

11.5

11.611.7

11.8

11.911.10 11.11

11.12

11.1311.1411.16

11.17

11.19

11.20

11.21

11.22

11.23

11.24

11.2512.1

12.2

12.3

12.5

12.6

12.712.8

12.912.10

12.11

12.13

12.14

12.15

12.16

12.17

12.19

12.20

12.21

13.1

13.2

13.3

13.5

13.7

13.8

13.10

13.11

13.12

13.14

13.15

13.16

13.17

13.18

13.1913.21

13.23

13.2514.1

14.2

14.314.5

14.7

14.8

14.1014.11

14.13

14.14

14.15 14.1614.1714.18

14.19

14.20

15.1

15.2

15.4

15.5

15.6 15.7

15.8

15.9

15.10

15.11 15.12

15.13

15.14

15.15

15.16

15.17

15.19

CPT1

CPT2CPT3

CPT5

CPT6

CPT7CPT8

CPT9

CPT10

CPT11

CPT12

CPT13

CPT14

CPT15

CPT16

CPT17

CPB1

CPB2

CPB3

CPB4

CPB5CPB6

CPB7CPB8

CPB9

CPB10

CPB11

CPB12CPB13

CPB14

CPB15

CJT1

CJT2

CJT3

CJT4

CJT5

CJT6

CJT7CJT8

CJT9

CJT10

CJT12

CJT13

CJT14

CJT15

CJT16CJT17CJT18

CJT19

CJB1

CJB2

CJB3

CJB4CJB5

CJB6CJB7CJB8CJB9

CJB10CJB11CJB12CJB13

CJB14CJB15

-20 -16 -12 -8 -4 4 8 12 16

Component 1

-4.0

-3.2

-2.4

-1.6

-0.8

0.8

1.6

2.4

3.2

Com

ponen

t 2


41

Many of the teeth have similar shapes, though not similar heights. Due to

this, height was excluded as a component of the analysis, leaving only

measurements 2-5 (Figure 13 and Figure 14). With the removal of height as

a factor, it is easier to see two distinct groups in the graphs provided. Even

with the removal of height as a factor, C. johnstoni and C. porosus still plot

relatively close to each other. Crocodylus porosus still has the most overlap

with one of the groupings of the fossils, whereas C. johnstoni has very little

overlap with the fossils and C. porosus (Figure 13). With the removal of the

modern specimens, there are still two largely distinct groups showing on the

PCA within the fossils (Figure 14). To further clean up the data, only those

teeth used for the Cloud Compare analysis were kept and further analysed

with a PCA and Non-Metric MDS (Multi-Dimensional Scaling). These teeth

were CT Scanned for the analysis.


excluding modern specimens, with height also excluded as a component of

the analysis.

5.2 CT Scanned Dataset

5.2.1 Raw Data

All the teeth from the initial data set were CT Scanned. Teeth that had large

wear facets, cracks or chips were then excluded from the clean data set

M2

M3

M4

M5

1.11.2

1.4

1.5

1.61.7

2.1

2.2

2.3

2.4

2.5

2.6

2.7

2.8

2.9

2.10

2.11

2.12

2.13

3.13.2

3.3

3.5

3.6

3.7

3.8

3.12

3.133.14

3.15

3.16

4.2

4.34.4

4.5

4.6

4.7

4.8

4.9

4.10

4.11

4.13

4.14

4.16

4.17

4.18

4.19

5.2

5.3

5.4

5.5

5.6

5.7

5.9

5.10

5.11

5.12

5.13

5.14

5.15

5.16

5.17

5.185.20 5.21

5.22

5.235.24

5.25

5.26

5.27

6.1 6.2

6.3

6.4

6.5

6.7

6.8

6.9

6.10

6.12

6.13

6.15

7.1

7.3

7.4

7.5

7.6

7.7 7.8

7.9

7.10

7.11

7.12

7.13

7.14

7.15

7.16

8.2

8.3

8.4

8.5

8.6

8.7

8.8

8.9

8.10

8.118.14

8.16

9.1

9.2 9.3

9.4

9.5

9.6

9.7

9.8

9.99.10

9.12

9.13

9.17

9.18

9.19

9.20

9.21

9.22

9.23

9.24

9.25

9.26

9.27

9.28

9.29

10.2

10.4

10.6

10.7

10.8

10.10

10.11

10.12

10.13

10.15

10.16

10.17

10.19

10.20

10.22

10.23

10.24

10.25

10.26

10.27

10.28

10.29

10.30

10.31

10.32

10.3311.2

11.3

11.4

11.5

11.611.7

11.8

11.9 11.10 11.11

11.1211.13

11.1411.16

11.17

11.19

11.20

11.21

11.22

11.23

11.24

11.2512.1

12.2

12.3

12.5

12.6

12.7

12.8

12.912.10

12.11

12.13

12.14

12.15

12.16

12.17

12.19

12.20

12.21

13.1

13.2

13.3

13.5

13.7

13.8

13.10

13.11

13.12

13.14

13.15

13.16

13.17

13.18

13.1913.21

13.23

13.2514.1

14.2

14.314.5

14.7

14.8

14.1014.11

14.13

14.14

14.15 14.1614.1714.18

14.19

14.20

15.1

15.2

15.4

15.5

15.6 15.7

15.8

15.9

15.10

15.11 15.12

15.13

15.14

15.15

15.16

15.17

15.19

-20 -16 -12 -8 -4 4 8 12 16

Component 1

-4.0

-3.2

-2.4

-1.6

-0.8

0.8

1.6

2.4

3.2

Com

ponen

t 2


42

(Appendix 4). The 3D models of the teeth that were kept were then analysed

using Cloud Compare and a PCA was also repeated for these specimens.

5.2.2 Rough Means Square Data Matrix

Each of the CT Scanned teeth was compared to every other tooth using

Cloud Compare. Cloud Compare gave a Root Means Square (RMS) output

that shows how similar each of the teeth are to each other. These results

were then placed into an excel file to create a data matrix for analysis (See

Supplementary Data 1).


As done previously, the data set was culled to include only those teeth used

for the CT Scan analysis. The new data set was then used to rerun a PCA

with and without height as a component for analysis (Figure 15 and Figure

16). Once height has been removed as a factor, there are still two clear

groupings visible on the PCA. The majority of unknown specimens group at

the top of the graph. A smaller number of them group together on the

bottom section of the graph.

Figure 15: Principle Components Analysis of teeth used for comparison in

Cloud Compare based on 3D models of specimens. Height is included as a

component in this figure.

M1

M2

M3 M4

M5

1.1

1.4

1.6

2.1

2.2

2.3

2.6

2.8

2.13

3.1

3.2

3.3

3.5

3.6

3.7

3.8

3.12

3.13

3.15

4.2

4.3

4.44.9

4.11

4.13

4.14

4.17

4.18

4.19

5.35.4

5.6

5.9

5.10

5.11

5.12

5.13

5.145.17

5.18

5.23

5.24

5.27

6.1 6.2

6.5

6.12

7.57.6

7.9

7.14

7.15

8.2

8.4

8.5

8.7

8.9

8.10

9.2

9.3 9.4

9.5

9.8

9.9

9.10

9.17

9.19

9.20

9.22

9.25

9.26

9.27

9.29

10.2

10.4

10.6

10.7

10.810.10

10.11

10.12

10.13

10.19

10.22

10.23

10.24

10.25

10.26

10.27

10.28

10.31

11.2

11.4

11.8

11.9

11.10

11.11

11.14

11.19

11.20

11.21

11.22

11.25

12.1

12.3

12.7

12.9

12.10

12.11

12.14

12.15

12.16

12.19

12.21

13.2

13.7

13.8

13.10

13.11

13.14

13.18

13.19

13.21

13.23

13.25

14.1

14.7

14.11

14.13

14.14

14.16

14.19

14.20

15.2

15.4

15.5

15.6

15.7

15.8

15.9

15.11

15.12

15.16

-20 -16 -12 -8 -4 4 8 12 16

Component 1

-4.8

-3.2

-1.6

1.6

3.2

4.8

6.4

8.0

Com

ponen

t 2


43

Figure 16: Principle Components Analysis of teeth used for comparison in

Cloud Compare based on 3D models of specimens. Height is excluded as a

component in this figure.

5.2.4 Non-Metric Multi-Dimensional Scaling

Using the RMS output from the Cloud Compare analysis, the data matrix

was used to do a Non-Metric Multi-Dimensional Scaling (MDS) analysis

(Figure 17). Though it is unclear using a graph with points, as above, the

teeth plot similar to how they would sit along the tooth row (Figure 18). The

points used to anchor the Cloud Compare analysis were the apex, and the

anterior and posterior ends of the base of the tooth.

M2

M3

M4

M5

1.1

1.4

1.6

2.1

2.2

2.3

2.6

2.8

2.13

3.1 3.23.3

3.5

3.6

3.7

3.8

3.12

3.13

3.15

4.2

4.34.4

4.9

4.11

4.13

4.144.17

4.18

4.19

5.3

5.4

5.6 5.9

5.10

5.11

5.12

5.13

5.14

5.17

5.18

5.235.24

5.27

6.16.2

6.5

6.127.5

7.6

7.9

7.14

7.15

8.2

8.4

8.5

8.78.9

8.10

9.29.3

9.4

9.5

9.8

9.99.10

9.17

9.19

9.20

9.22

9.25

9.26

9.27

9.29

10.2

10.4

10.6

10.7

10.8

10.10

10.11

10.12

10.13

10.19

10.22

10.23

10.24

10.25

10.26

10.27

10.28

10.3111.2

11.4

11.8

11.911.10

11.11

11.14

11.19

11.20

11.21

11.22

11.25

12.1

12.3

12.7

12.9

12.1012.11

12.14

12.15

12.16

12.1912.21

13.2

13.713.8

13.10

13.11

13.14

13.18

13.1913.21

13.23

13.25

14.1

14.714.11

14.13

14.14

14.16

14.19

14.20

15.2

15.4

15.5

15.615.7

15.8

15.915.11 15.12

15.16

-20 -16 -12 -8 -4 4 8 12 16

Component 1

-4.0

-3.2

-2.4

-1.6

-0.8

0.8

1.6

2.4

3.2

Com

ponen

t 2


44

Figure 17: Non-Metric Multi-Dimensional Scaling based on the Root

Means Squared Output from Cloud Compare analysis of the 3D teeth

models.

Some of the fossil teeth were laid out on a print out of the graph to create a

visual depiction of how the teeth are plotting across the graph (Figure 18).

The teeth plot from caniniform (left of graph) to molariform (right of

graph). Also, the ziphodont (labio-lingually compressed) teeth plot on the

outer ring of the MDS graph, while those with a circular base plot towards

the interior or top of the graph. Figure 19 shows the bases of each of the

teeth corresponding to those seen in Figure 18.

1.1

1.4

1.6

2.1

2.2

2.3

2.6

2.8

2.13

3.1

3.2

3.3

3.5

3.6

3.7

3.8

3.12

3.13

3.15 4.2

4.3

4.4

4.9

4.11

4.13

4.144.17

4.18

4.19

5.35.4

5.6

5.9

5.10

5.11

5.12

5.13

5.14

5.17

5.18

5.23

5.24

5.27

6.1

6.2

6.5

6.12

7.5

7.6

7.9

7.14

7.15

7.17

8.28.48.5

8.7

8.9

8.10

9.2

9.3

9.4

9.5

9.8

9.9

9.109.17

9.19

9.20

9.22

9.25

9.26

9.27

9.29

10.2

10.3

10.4

10.6

10.7

10.8

10.10

10.11

10.12

10.13

10.19

10.2210.23

10.24

10.25

10.26

10.27

10.28

10.31

11.2

11.4

11.8

11.9

11.10

11.11

11.14

11.19

11.20

11.21

11.22

11.25

12.112.3

12.7

12.9

12.1012.11

12.14

12.1512.16

12.18

12.19

12.21

13.2

13.4

13.7

13.8

13.10

13.11

13.14

13.1813.19

13.21

13.23

13.25

14.1

14.7

14.11

14.13

14.14

14.16

14.19

14.20

15.2

15.415.5

15.615.7

15.8

15.9

15.11

15.12

15.16

-0.20 -0.16 -0.12 -0.08 -0.04 0.00 0.04 0.08 0.12 0.16

Coordinate 1

-0.09

-0.06

-0.03

0.00

0.03

0.06

0.09

0.12

0.15

0.18

Coord

inate

2


45

Figure 18: Depiction of how the teeth plot on the Non-Metric MDS by

matching the teeth numbers with the points on the graph. Pictured left to

right are tooth numbers 9.27, 9.8, 9.22, 12.15, 5.11, 10.12, 3.2, 13.19 and

10.31.

Figure 19: Teeth depicted on the Non-Metric MDS graph with the base of

each tooth to the right. Pictured left to right are tooth numbers 9.27, 9.8,

9.22, 12.15, 5.11, 10.12, 3.2, 13.19 and 10.31.

5.3 Caniniform and Molariform

As previously stated, crocodilian teeth can be divided into two broad

categories, caniniform and molariform. The teeth that were used for the

Cloud Compare analysis were further split into these two groups to be

analysed separately.


46

5.3.1 Raw Data

All caniniform and molariform teeth had also been previously measured.

Caniniform teeth were grouped together to create a graph of measurements

of only the caniniform teeth (Appendix 5). The same was repeated for the

molariform teeth (Appendix 6).


As previously described for the full dataset, the caniniform teeth were

analysed with a PCA. There is very little grouping visible in the PCA which

includes height as a component (Figure 20). As height has too much effect

on the data, it was excluded as a component (Figure 21). With height

excluded as a factor for the PCA on the Caniniform teeth there are again

two visible groups on the graph (Figure 21). Again, the majority of teeth are

plotting in a group at the top of the graph, while there is still a small group

plotting at the bottom of the graph. This procedure was then repeated for the

molariform teeth.

Figure 20: Principle Components Analysis for all caniniform teeth used in

the Cloud Compare Analysis. Height is included as a component in this

figure.

M1

M2

M3

M4

M5

1.1

1.62.1

2.2 2.3 2.6

2.8

2.13

3.33.5

3.6

3.7

3.8

3.13

3.15

4.3

4.4

4.9

4.11

4.13

4.144.17

4.18

5.3 5.4

5.6

5.9

5.10 5.11

5.12

5.13

5.14

5.17

5.18

5.23

5.24

5.27

6.1

6.26.5

6.12

7.5

7.9

7.14

7.15

8.2

8.4

8.5

8.7

8.10

9.2

9.5

9.8

9.17

9.19

9.22

9.25

9.26

9.27

10.2

10.4

10.6

10.7

10.8

10.10

10.12

10.19

10.22

10.23

10.24

10.25

10.26

10.28

11.2

11.4

11.8

11.10

11.11

11.19

11.20

11.21

11.25

12.1

12.3

12.7

12.9

12.1012.11

12.14

12.15

12.16

12.19

13.2

13.7

13.8

13.10

13.11

13.14

13.18

13.23

13.25

14.1

14.7

14.1114.13

14.14

14.16

14.19

14.20

15.2

15.4

15.5

15.6

15.7

15.8

15.9

15.11

15.12 15.16

-20 -16 -12 -8 -4 4 8 12 16

Component 1

-4.8

-3.6

-2.4

-1.2

1.2

2.4

3.6

4.8

6.0

Com

ponen

t 2


47

Figure 21: Principle Components Analysis for all caniniform teeth used in

the Cloud Compare Analysis. Height is excluded as a component in this

figure.

Figure 22: Principle Components Analysis for all molariform teeth used in

the Cloud Compare Analysis. Height is included as a component in this

figure.

Most of the teeth used for the analysis were caniniform. Due to this, the

molariform data is sparse and so it is difficult to see any groupings (Figure

22). It is currently unknown as to why there is a bias in the fossil record

against molariform crocodilian teeth. The number of molariform teeth in a

crocodilian’s mouth is limited, therefore it would be expected that there are

M2

M3

M4

M5

1.1

1.6

2.1

2.2

2.3

2.6

2.8

2.13

3.3

3.5

3.6

3.7

3.83.13

3.15

4.34.4

4.9

4.11

4.13

4.144.17

4.18

5.3

5.4

5.65.9

5.10

5.11

5.12

5.13

5.14

5.17

5.18

5.235.24

5.27

6.16.2

6.5

6.12

7.5

7.9

7.14

7.15

8.2

8.4

8.5

8.7

8.10

9.2

9.5

9.8

9.17

9.19

9.22

9.25

9.26

9.27

10.2

10.4

10.6

10.7 10.810.10 10.12

10.19

10.22

10.23

10.24

10.25

10.26

10.28

11.2

11.4

11.8

11.1011.11

11.19

11.20

11.21

11.25

12.1

12.3

12.712.9

12.10

12.11

12.14

12.15

12.16

12.19

13.2

13.713.8

13.10

13.11

13.14

13.18

13.23

13.25

14.1

14.714.11

14.13

14.14

14.16

14.19

14.20

15.2

15.4

15.5

15.615.7

15.8

15.915.11

15.12

15.16

-20 -16 -12 -8 -4 4 8 12 16

Component 1

-4.0

-3.2

-2.4

-1.6

-0.8

0.8

1.6

2.4

3.2

Com

ponen

t 2

M1

M2

M3

M4

M5

1.4

3.1

3.2

3.12

4.2

4.19

7.6

8.9

9.3

9.49.9

9.10

9.20

9.29

10.11

10.13

10.2710.31

11.9

11.1411.22

12.21

13.19

13.21

-10 -8 -6 -4 -2 2 4 6

Component 1

-4.8

-4.0

-3.2

-2.4

-1.6

-0.8

0.8

1.6

2.4

Com

ponen

t 2


48

few molariform teeth fossils. Even if height is excluded as a component for

the PCA, it is still very difficult to see any groupings within the data. There

may be a clustering on the left and right side of the graph, however more

molariform teeth will need to be included with a larger dataset to determine

if these apparent clusters may be biologically meaningful or just a result of

the small number of teeth in this group (Figure 23).

Figure 23: Principle Components Analysis for all molariform teeth used in

the Cloud Compare Analysis. Height is excluded as a component in this

figure.


As these teeth were all used in a Cloud Compare analysis to provide a data

matrix (See Supplementary Data 2) a Non-Metric MDS analysis was

completed for them. Without having images of the teeth on the points of the

Non-Metric MDS graph, it is difficult to see how the teeth may be arranged

(Figure 24).

M2

M3

M4

M5

1.4

3.1

3.2

3.12

4.2

4.19

7.6

8.9

9.3

9.49.9

9.10

9.20

9.29

10.11

10.13

10.27

10.31

11.9

11.14

11.2212.21

13.19

13.21

-6.4 -4.8 -3.2 -1.6 1.6 3.2 4.8

Component 1

-1.8

-1.2

-0.6

0.6

1.2

1.8

2.4

3.0

3.6

Com

ponen

t 2


49

Figure 24: Non-Metric Multi-Dimensional Scaling for Caniniform tooth

specimens

A number of the teeth were chosen to be laid out on the Caniniform Non-

Metric MDS graph (Figure 25) to visualise how the teeth are plotting based

on size and shape. Some of the teeth included in the caniniform group

appear transitionary and could have possibly been included in the

molariform group. Due to the length of the tooth compared to the width,

visually, they have been included in the caniniform group. Those teeth that

are considered transitionary are plotting on the left-hand side of the graph,

on the outer edge. Corresponding to this, the ziphodont teeth are mostly

plotting on the left and outer edge while the teeth with circular bases are

plotting on the right and towards the middle of the graph. See Figure 26 for

images of the base of the teeth.

1.11.6

2.1

2.2

2.3

2.6

2.8

2.13

3.3 3.5

3.6

3.7

3.8

3.13

3.15

4.3

4.4

4.94.11

4.13

4.144.17

4.18

5.35.4

5.6 5.9

5.10

5.11 5.12

5.13

5.14 5.17

5.18

5.23

5.24

5.27

6.1

6.2

6.5

6.127.5

7.9

7.14

7.15

7.17

8.2

8.4

8.5 8.7

8.10

9.2

9.5

9.8

9.17

9.19

9.22

9.25

9.26

9.27

10.2

10.3

10.410.6

10.7

10.8

10.10

10.12

10.19

10.22

10.23

10.24

10.25

10.26

10.2811.211.4

11.811.10

11.11

11.19

11.20

11.21

11.25

12.1

12.3

12.7

12.9

12.10

12.11

12.14

12.15

12.16

12.18

12.19

13.2

13.4 13.7

13.8

13.10

13.11

13.14

13.18

13.2313.25

14.1

14.7

14.11

14.13

14.14

14.1614.19

14.20

15.2

15.4

15.5

15.6

15.715.8

15.9 15.11

15.12

15.16

-0.20 -0.16 -0.12 -0.08 -0.04 0.00 0.04 0.08 0.12 0.16

Coordinate 1

-0.18

-0.15

-0.12

-0.09

-0.06

-0.03

0.00

0.03

0.06

Coord

inate

2


50

Figure 25: Depiction of how the caniniform teeth plot on the Non-Metric

MDS by matching the teeth numbers with the points on the graph. Pictured

left to right are tooth numbers 2.8, 14.14, 5.11, 8.4, 12.15, 9.8, 3.8, 9.22 and

9.27.

Figure 26: Caniniform teeth depicted on the Non-Metric MDS graph with

the base of each tooth to the right. Pictured left to right are tooth numbers

2.8, 14.14, 5.11, 8.4, 12.15, 9.8, 3.8, 9.22 and 9.27.

All molariform teeth were also used in a Cloud Compare analysis to provide

a RMS data matrix (See Supplementary Data 3), and so a Non-Metric MDS

graph was completed using this data (Figure 27). It is difficult to visualise

how the teeth plot on these points so a number of them were placed on a

print out of the graph (Figure 28).


51

Figure 27: Non-Metric Multi-Dimensional Scaling for Molariform tooth

specimens

There appears to be some grouping to the molariform teeth. The ziphodont

molariform teeth are plotting on the left side of the graph while those with a

circular base are grouping on the right. See figure 25 for images of the base

of the molariform teeth. However, more data will need to be added to

confirm this.

Figure 28: Depiction of how the molariform teeth plot on the Non-Metric

MDS by matching the teeth numbers with the points on the graph. Pictured

left to right are tooth numbers 1.4, 4.19, 3.12, 11.14, 13.19, 11.22, 10.31,

3.2, 10.13 and 13.21.

1.4

3.1

3.23.12

4.2

4.19

7.6

8.9

9.3

9.4

9.9

9.10

9.20 9.2910.11

10.13

10.27

10.31

11.9

11.14

11.22

12.21

13.19

13.21

-0.40 -0.32 -0.24 -0.16 -0.08 0.00 0.08 0.16 0.24

Coordinate 1

-0.20

-0.16

-0.12

-0.08

-0.04

0.00

0.04

0.08

0.12

0.16

Coord

inate

2


52

Figure 29: Molariform teeth depicted on the Non-Metric MDS graph with

the base of each tooth to the right. Pictured left to right are tooth

numbers1.4, 4.19, 3.12, 11.14, 13.19, 11.22, 10.31, 3.2, 10.13 and 13.21.

5.4 Analysis Based Upon Ratio Data

5.4.1 Ratio of Mid-Height and Base Length versus Width

The ratios of the base and mid height of the teeth were calculated and used

for further analysis of the data. To determine if there was a clear split in the

data a histogram was plotted (Figures 30 and 31). For the purpose of this

study, the teeth that plot on the left of each histogram will be considered

Group 1 and those on the right, Group 2. Though there are clearly two

groups when the ratio is calculated for M2 and M3 (Mid-Height

Length/Mid-Height Width) (Figure 30), there is less distinction between

ratios when calculated based on M4 and M5 (Crown Base Length/Crown

Base Width) (Figure 31). The teeth were then organised to depict these

points and graphed in a scatter plot based on the ratio groups depicted.


53

Figure 30: Histogram showing two distinct groups based on the ratio

calculated from M2 and M3 at 1.39.

Figure 31: Histogram showing distinct groups based on the ratio calculated

from M4 and M5 at 1.13

Each tooth was plotted using the CBL/CBW ratio compared to height

(Figure 32). There are two close but very clear groups based on this

differentiation from groups, with no overlap between them. The same was

then repeated using the ML/MW ratio (Figure 33) for comparison. Figure 30

shows a split between two groups when the ratio for Mid-Height Length to

Mid-Height Width (ML/MW) at 1.39. When using the ratio split between

ML/MW comparing it to M1 (height) there is also a clear distinction

0.84 0.96 1.08 1.20 1.32 1.44 1.56 1.68 1.80

0

10

20

30

40

50

60

70

80

90

Fre

que

ncy

0.84 0.96 1.08 1.20 1.32 1.44 1.56 1.68 1.80

0

10

20

30

40

50

60

70

80

Fre

que

ncy


54

between two groups (Figure 33). Due to the much clearer distinction on the

histogram for the ML/MW ratio (Figure 30), it was decided that there would

be greater support to use the ML/MW ratio for further analysis. Some teeth

plotted in the group 1 when organised based on ML/MW and CBL/CBW,

some plotted in group 2 both times, and others switched from group 1 to

group 2 or visa-versa.

.

Figure 32: Ratio of CBL/CBW compared to M1 (height) after teeth have

been split into group 1 (Blue) and group 2 (Black) ratio values.

Figure 33: Ratio of ML/MW compared to M1 (height) after teeth have been

split into group 1 (Blue) and group 2 (Black) ratio values.

1.11.61.7

2.3

2.4

2.6

2.8

2.9

2.10 2.11

2.12

2.13

3.5

3.6

3.7

3.8

3.13

3.14

3.15

3.16

4.5

4.6

4.7

4.9

4.11

4.13

4.14

4.16

4.17

4.185.5

5.6

5.7

5.9

5.10

5.11

5.12

5.13

5.14

5.155.16

5.17

5.18

5.21

5.22

5.23

5.24

6.3

6.4

6.5

6.7

6.9

6.10

6.126.13

6.15

7.1

7.5

7.7

7.8

7.9

7.10

7.12

7.15

8.2

8.78.8

8.10

8.11

8.14

8.16

9.6

9.7

9.8

9.12

9.17

9.19

9.21

9.22

9.24

9.27

10.8

10.22

10.24

10.25

10.26

11.2

11.1011.11

11.13

11.2512.1

12.7

12.8

12.10

12.11

12.15

12.17

12.20

13.1

13.7

13.8

13.10

13.11

13.12

13.14

13.15

13.16

13.18

13.23

14.5

14.7

14.13

14.15

14.16

14.1714.18

15.1

15.2

15.5

15.16

15.17

15.19

CPT2

CPT3

CPT9

CPB1

CPB4

1.2

1.4

1.5

2.1

2.2

2.5

2.7

3.1

3.2 3.3

3.12

4.2

4.3

4.4

4.8

4.10

4.19

5.2

5.3

5.4 5.20

5.25

5.26

5.27

6.1

6.2

6.8

7.3

7.4

7.6

7.11

7.13

7.14

7.16

8.3

8.4

8.5

8.6

8.9

9.1

9.2

9.3

9.4

9.5

9.9

9.10

9.13

9.189.20

9.23

9.25

9.26

9.289.29

10.2

10.4

10.6

10.7

10.10

10.11

10.12

10.13

10.15

10.16

10.17

10.19

10.20

10.23

10.27

10.2810.29

10.30

10.31

10.32

10.33

11.3

11.4

11.5

11.6

11.7

11.8

11.9

11.12

11.14

11.1611.17

11.19

11.2011.21

11.22

11.23

11.24

12.2

12.3

12.5

12.6

12.9 12.13

12.1412.16

12.19

12.21

13.2

13.3

13.5

13.17

13.19

13.21

13.25

14.1

14.2

14.3

14.8

14.10

14.1114.14

14.20

14.19

15.4

15.6

15.715.8

15.9

15.10

15.11 15.12

15.13

15.14

15.15

CPT1

CPT5

CPT6

CPT7

CPT8

CPT10

CPT11CPT12

CPT13

CPT14

CPT15

CPT16

CPT17

CPB2

CPB3

CPB5

CPB6CPB7

CPB8

CPB9

CPB10

CPB11

CPB12

CPB13

CPB14

CPB15

CJT1

CJT2

CJT3

CJT4

CJT5

CJT6

CJT7

CJT8

CJT9

CJT10

CJT12

CJT13

CJT14CJT15

CJT16CJT17

CJT18

CJT19

CJB1CJB2

CJB3

CJB4

CJB5

CJB6CJB7

CJB8 CJB9 CJB10

CJB11CJB12

CJB13

CJB14CJB15

0.84 0.96 1.08 1.20 1.32 1.44 1.56 1.68 1.80

CBL/CBW

8

12

16

20

24

28

32

36

40

M1

1.11.61.7

2.3

2.4

2.6

2.8

2.9

2.10 2.11

2.12

2.13

3.5

3.6

3.7

3.8

3.13

3.14

3.15

3.16

4.5

4.6

4.7

4.9

4.11

4.13

4.14

4.16

4.17

4.185.5

5.6

5.7

5.9

5.10

5.11

5.12

5.13

5.14

5.155.16

5.17

5.18

5.21

5.22

5.23

5.24

6.3

6.4

6.5

6.7

6.9

6.10

6.126.13

6.15

7.1

7.5

7.7

7.8

7.9

7.10

7.12

7.15

8.2

8.78.8

8.10

8.11

8.14

8.16

9.6

9.7

9.8

9.12

9.17

9.19

9.21

9.22

9.24

9.27

10.8

10.22

10.24

10.25

10.26

11.2

11.1011.11

11.13

11.2512.1

12.7

12.8

12.10

12.11

12.15

12.17

12.20

13.1

13.7

13.8

13.10

13.11

13.12

13.14

13.15

13.16

13.18

13.23

14.5

14.7

14.13

14.15

14.16

14.1714.18

15.1

15.2

15.5

15.16

15.17

15.19

CPT2

CPT3

CPT9

CPB1

CPB4

1.2

1.4

1.5

2.1

2.2

2.5

2.7

3.1

3.2 3.3

3.12

4.2

4.3

4.4

4.8

4.10

4.19

5.2

5.3

5.4 5.20

5.25

5.26

5.27

6.1

6.2

6.8

7.3

7.4

7.6

7.11

7.13

7.14

7.16

8.3

8.4

8.5

8.6

8.9

9.1

9.2

9.3

9.4

9.5

9.9

9.10

9.13

9.18 9.20

9.23

9.25

9.26

9.289.29

10.2

10.4

10.6

10.7

10.10

10.11

10.12

10.13

10.15

10.16

10.17

10.19

10.20

10.23

10.27

10.2810.29

10.30

10.31

10.32

10.33

11.3

11.4

11.5

11.6

11.7

11.8

11.9

11.12

11.14

11.1611.17

11.19

11.2011.21

11.22

11.23

11.24

12.2

12.3

12.5

12.6

12.9 12.13

12.1412.16

12.19

12.21

13.2

13.3

13.5

13.17

13.19

13.21

13.25

14.1

14.2

14.3

14.8

14.10

14.1114.14

14.20

14.19

15.4

15.6

15.715.8

15.9

15.10

15.11 15.12

15.13

15.14

15.15

CPT1

CPT5

CPT6

CPT7

CPT8

CPT10

CPT11CPT12

CPT13

CPT14

CPT15

CPT16

CPT17

CPB2

CPB3

CPB5

CPB6CPB7

CPB8

CPB9

CPB10

CPB11

CPB12

CPB13

CPB14

CPB15

CJT1

CJT2

CJT3

CJT4

CJT5

CJT6

CJT7

CJT8

CJT9

CJT10

CJT12

CJT13

CJT14CJT15

CJT16CJT17

CJT18

CJT19

CJB1CJB2

CJB3

CJB4

CJB5

CJB6CJB7

CJB8 CJB9CJB10

CJB11CJB12

CJB13

CJB14CJB15

0.84 0.96 1.08 1.20 1.32 1.44 1.56 1.68 1.80

ML/MW

8

12

16

20

24

28

32

36

40

M1


55

Figure 34: ML/MW plotted against CBL/CBW ratios. Teeth remaining in

group 1 for both ratios were plotted using blue, group 2 using black and

group 3 using red.

The organisation of the specimens based on the ratios, shows there are three

groups of which some teeth switch between groups 1 and 2 (plotted in red),

and some of which remain in the same group; group 1 plotted in blue and

group 2 plotted in black (Figure 34). Only one specimen overlaps from

group 3 into group 2 and one specimen from group 2 overlaps into group 3.


Figure 36 shows the same Non-Metric MDS graph as in Figure 17, however

the points have been marked depending on which of the three previously

mentioned groups the teeth fall into after being sorted by ratios. It is unclear

how the teeth plot in the above graph, therefore specimens were placed onto

the graph as a visual depiction of this (Figure 37).

1.1

1.2

1.4

1.5

1.61.7

2.1

2.22.3

2.42.5

2.6

2.7

2.8

2.9

2.10

2.11

2.12

2.13

3.5

3.6

3.8

3.12

3.13

3.14 3.15

3.16

4.5

4.6

4.7

4.8

4.94.10

4.11

4.13

4.14

4.16

4.17

4.18

4.19

5.5

5.65.7

5.9

5.10

5.11

5.12

5.13

5.14

5.155.16

5.17

5.18

5.20

5.21

5.22

5.235.24

5.25

5.27

6.3

6.4

6.5

6.7

6.9

6.10

6.12

6.13

6.15

7.5

7.7

7.8

7.9

7.10

7.12

7.15

8.8

8.108.118.14

8.169.6

9.7

9.8

9.17

9.19

9.21

9.22

9.25

9.27

CJB14

CJT19

5.3

7.1

7.3

7.4

8.2

8.3 8.69.1

9.29.4

9.13

9.23

9.26

10.2

10.4

10.7

10.10

10.11

10.13

10.1610.17

10.19

10.20

10.23

10.24

10.26

10.27

10.28

10.29

10.33

11.2

11.411.511.6

11.7

11.8

11.9

11.10

11.11 11.1211.13

11.16 11.17

11.19

11.20

11.21

11.22

11.23

11.2411.25

12.1

12.2

12.5

12.6

12.7

12.8

12.9 12.1012.11

12.13

12.14

12.1512.16

12.17 12.1912.20

12.21

13.1

13.2

13.3

13.513.7

13.813.1013.11

13.1213.14

13.15

13.16

13.17

13.18 13.1913.21

13.23

13.25

14.1

14.2

14.3

14.5

14.7

14.8

14.10

14.11

14.1314.1414.1514.1614.17

14.18

14.19

14.2015.1

15.2 15.415.5

15.6

15.7

15.8

15.9

15.10

15.11

15.12

15.1315.14

15.15

15.1615.17

15.19

CPT2CPT3

CPT13

CPT14

CPB2

CPB6

CPB8

CPB10

CJT5CJT4CJB2

CJT3

3.1

3.2

3.3

3.7

4.24.34.4

5.2

5.4

5.26

6.1

6.2 6.8

7.6

7.11 7.13

7.14

7.16

8.4

8.5

8.7

8.9

9.3

9.5

9.9

9.10

9.12

9.18

9.20

9.249.289.29

10.6

10.8

10.12

10.15

10.22

10.25

10.3010.31

10.3211.3

11.14

12.3

CPT1

CPT5 CPT6

CPT7

CPT8

CPT9

CPT10 CPT11 CPT12

CPT15CPT16

CPT17

CPB1CPB3

CPB4

CPB5

CPB7

CPB9

CPB11

CPB12

CPB13

CPB14

CPB15

CJT1

CJT2

CJT6

CJT7

CJT8

CJT9

CJT10

CJT12CJT13

CJT14CJT15

CJT16CJT17

CJT18

CJB1

CJB3

CJB4

CJB5

CJB6CJB7CJB8

CJB9CJB10

CJB11 CJB12

CJB13

CJB15

0.9 1.0 1.1 1.2 1.3 1.4 1.5 1.6 1.7

CBL/CBW

0.84

0.96

1.08

1.20

1.32

1.44

1.56

1.68

1.80

ML/M

W


56

Figure 35: Non-Metric MDS based on the Root Means Squared Output

from Cloud Compare the Cloud Compare analysis of the 3D models,

coloured by the groups from the ratio differentiation.

As done with previous MDS graphs, teeth were chosen and laid out upon a

print out of the graph to show how the teeth specimens are plotting across

the graph (Figure 36). The teeth plot from caniniform (left) to molariform

(right). Those that sort into group 2 in Figures 32 to 34 plot on the outer

edge in Figure 37. Teeth that sort into group 1 in Figure 32 to 34 plot on the

inner section of the graph (Figure 35). Teeth that switched between the two

groups (group 3) are scattered throughout the middle of the Non-Metric

MDS graph above (Figure 35). Figure 37 shows each of the teeth placed on

the Non-Metric MDS graph (Figure 36) with their corresponding base to the

right.

1.1

1.4

1.6

2.1

2.2

2.3

2.6

2.8

2.13

3.1

3.2

3.3

3.5

3.6

3.7

3.8

3.12

3.13

3.15 4.2

4.3

4.4

4.9

4.11

4.13

4.144.17

4.18

4.19

5.35.4

5.6

5.9

5.10

5.11

5.12

5.13

5.14

5.17

5.18

5.23

5.24

5.27

6.1

6.2

6.5

6.12

7.5

7.6

7.97.14

7.15

7.17

8.28.48.5

8.7

8.9

8.10

9.2

9.3

9.4

9.5

9.8

9.9

9.109.17

9.19

9.20

9.22

9.25

9.26

9.27

9.29

10.2

10.3

10.4

10.6

10.7

10.8

10.10

10.11

10.12

10.13

10.19

10.2210.23

10.24

10.25

10.26

10.27

10.28

10.31

11.2

11.4

11.8

11.911.10

11.11

11.14

11.19

11.20

11.21

11.22

11.25

12.112.3

12.7

12.9

12.1012.11

12.14

12.1512.16

12.18

12.19

12.21

13.2

13.4

13.7

13.8

13.10

13.11

13.14

13.1813.19

13.21

13.23

13.25

14.1

14.7

14.11

14.13

14.14

14.16

14.19

14.20

15.2

15.415.5

15.615.7

15.8

15.9

15.11

15.12

15.16

-0.20 -0.16 -0.12 -0.08 -0.04 0.00 0.04 0.08 0.12 0.16

Coordinate 1

-0.09

-0.06

-0.03

0.00

0.03

0.06

0.09

0.12

0.15

0.18

Coord

inate

2


57

Figure 36: Depiction of how the teeth plot on the Non-Metric MDS by

matching the teeth with the points on the graph. Pictures left to right are

tooth numbers 9.27, 9.19, 9.8, 11.25, 7.5, 12.15, 5.11, 12.9, 4.19, 2.8, 10.13

and 13.19.

Figure 37: Teeth depicted on the Non-Metric MDS graph with the base of

each tooth to the right. Picture left to right are tooth numbers 9.27, 9.19, 9.8,

11.25, 7.5, 12.15, 5.11, 12.9, 4.19, 2.8, 10.13 and 13.19.


58

5.5 Alveoli Measurements

Alveoli are an important component of jaw specimens. The size and shape

of alveoli are used as a comparison for isolated teeth, to match them to a

species or associated specimen. Alveoli were measured from several

different specimens housed in the Queensland Museum fossil collection, as

well as the two modern Crocodylus specimens. The ratio of the anterior to

posterior measurements of the alveoli was used to create the graphs depicted

in figures 38 to 41.

5.5.1 Crocodylus spp.

Figure 38: Alveoli measurements of Crocodylus porosus and Crocodylus

johnstoni fossils and modern specimens showing an upwards trend

posteriorly.

0.8

1

1.2

1.4

1.6

1.8

2

0 5 10 15 20

Alv

eo

li R

atio

Tooth Postion

Crocodylus spp.

QMF17479 QMJ39230 Top

QMJ39230 Bottom QMF9229

QMF11611 QMJ48127 Top

QMJ48127 Bottom Linear (QMF17479)

Linear (QMJ39230 Top) Linear (QMJ39230 Bottom)

Linear (QMF9229) Linear (QMF11611)

Linear (QMJ48127 Top) Linear (QMJ48127 Bottom)


59

Alveoli were measured from both modern and fossil Crocodylus specimens.

Two specimens belonged to C. johnstoni, QMF17479 and the modern

specimen QMJ39230. Three specimens belonged to C. porosus, two fossils,

QMF9229 and QMF11611 and the modern QMJ48127. Most of the fossil

specimens are fragmentary and so it is unknown what tooth position the

alveoli specifically belong to. However, when a trend line is added to each

of the specimens there is a clear upward trend posteriorly which is

supported by the modern specimens (Figure 38). Therefore, the alveoli

length to width changes in ratio along the jaw line, starting with the anterior

most alveoli being close to having an equal length to width. As the ratio

changes along the alveolar row, those that are posterior-most have alveolar

length approaching twice that of its width.

5.5.2 Pallimnarchus sp.

Figure 39: Alveoli measurements of Pallimnarchus sp. fossils showing a

uniform or slightly downward trend posteriorly.

0

0.2

0.4

0.6

0.8

1

1.2

1.4

1.6

1.8

0 2 4 6 8 10 12 14 16 18

Alv

eo

li R

atio

Tooth Position

Pallimnarchus sp.

QMF1156 QMF2025 QMF1165

QMF11612 QMF11626 PF Left

PF Right PM Linear (QMF1156)

Linear (QMF2025) Linear (QMF1165) Linear (QMF11612)

Linear (QMF11626) Linear (PF Left) Linear (PF Right)

Linear (PM)


60

Identified fossil specimens were also used to collect alveoli measurements.

The largest number of specimens housed in the collection belong to

Pallimnarchus, the extinct giant freshwater crocodile. The ratio for the

alveoli of Pallimnarchus remain relatively close to a value of 1, equal length

to width, with the posterior-most teeth slightly more uniform resulting in a

slight downwards trend (Figure 39).

5.5.3 Quinkana sp.

Figure 40: Alveolar measurements of Quinkana sp. fossils showing a

greater length to width ratio along all tooth positions with a single alveolus

approaching a ratio of 1.

There are fewer Quinkana specimens in the fossil collection, and so the data

is sparse. However, in comparison to the Crocodylus and Pallimnarchus

specimens the trend lines for the Quinkana specimens sit relatively flat and

consistently with a ratio longer than wide, which reflects the uniform

ziphodont nature of the teeth. More Quinkana specimens would need to be

measured to improve precision (Figure 40).

0.5

1

1.5

2

0 2 4 6 8 10 12

Alv

eo

li R

atio

Tooth Position

Quinkana sp.

QMF30582 QF Left QF Right

Linear (QMF30582) Linear (QF Left) Linear (QF Right)


61

5.5.4 Unknown species

Figure 41: Alveoli measurements of several unidentified jaw fossils with

various trends.

Four unknown jaw specimens were also measured to see if it is possible to

draw any conclusions as to which species they may belong, based on the

trend of the alveoli (Figure 41). Considering the previous, graphs it could be

inferred that specimens CF1 and QMF10141 belong to Pallimnarchus sp.,

CF2 could be a species of Crocodylus, and QMF52672 could be Quinkana

sp. Additional specimens would need to be added to confirm the accuracy of

using this method as a confirmation of species. Inferences from CF1 and

QMF10141 are particularly speculative.

6.0 Discussion:

Crocodilians first evolved approximately 250 million years ago, in the Early

Triassic Period (Brazaitis and Watanabe, 2011; Naish, 2001). Early

crocodilians are thought to have lived a primarily terrestrial lifestyle (Naish,

2001) with the more ‘modern’ aquatic lifestyle first appearing during the

Jurassic Period (Bellairs, 1969; Brazaitis and Watanabe, 2011). They have

been the dominant predator of the land-water interface since the Late

0.5

0.7

0.9

1.1

1.3

1.5

1.7

1.9

0 2 4 6 8 10

Alv

eo

li R

atio

Tooth Position

Unknown Species

CF1 CF2 QMF52672

QMF10141 Linear (CF1) Linear (CF2)

Linear (QMF52672) Linear (QMF10141)


62

Cretaceous Period, approximately 85 million years ago (Erickson et al.,

2012). From the Late Triassic Period through to the end of the Miocene

Epoch there were numerous species of crocodilians globally, some of which

possessed ziphodont dentition (Molnar, 1982). Australia, along with the

isolated South Pacific islands of New Caledonia, Vanuatu, and Fiji are the

only currently known regions in which ziphodont species survive well into

the Pleistocene and Holocene (Molnar, 2004).

Australia’s crocodilian fossil record begins during the early Late Cretaceous

with the small-sized species Isisfordia ducani (Salisbury et al., 2006),

recently considered to be from the family Susisuchidae, the sister taxon to

the Eusuchia (Turner and Pritchard, 2015). Crocodilian taxa are known from

the Paleogene and Neogene, representing almost exclusively a regionally

restricted subfamily, the Mekosuchinae (Willis, Molnar and Scanlon, 1993).

Two species of crocodilian have been identified from the Chinchilla Local

Fauna, Pallimnarchus pollens and Quinkana sp. (DeVis, 1886; Molnar,

1982; Louys and Price, 2015).

The majority of fossilised evidence for crocodilian species comes in the

form of isolated teeth and osteoderms (Mackness et al, 2010; Naish, 2001).

Teeth are the most numerous of these fossils to be found due to continuous

shedding throughout the crocodilians lifespan, therefore one individual

crocodile can provide multiple tooth generations for the fossil record in

comparison to a single skeleton. The shape and size of teeth are useful when

inferring palaeoecology and dietary preferences of species belonging to

these teeth (Brink et al., 2015; Dauphin and Williams, 2007).

The crocodilian tooth row has been described as having two main forms: the

anterior teeth, termed caniniform, and the posterior teeth, termed

molariform (Erickson et al, 2012). Caniniform teeth are generally larger and

more conically shaped, to easily grip prey items, whereas molariform teeth

are shorter and stouter, usually used for crushing the rigid components of

prey items like bone and shell (Enax et al, 2013).

According to previous studies, Pallimnarchus and Quinkana have both been

recovered from the Chinchilla Local Fauna (Louys and Price, 2015). These


63

two taxa could account for the two-different tooth types recorded at this

locality. The research undertaken here involved using different and new

methodologies to better determine the number of crocodilian species living

at a single locality during the Pliocene. To do this, a new method is

proposed using the largest available dataset, fossilised teeth. A total of 306

teeth were selected for the research based on the quality of preservation.

Teeth that were broken, cracked or otherwise damaged were excluded from

the original dataset. Initially the teeth were grouped based upon a visual

analysis. As the study progressed the teeth were CT Scanned and analysed

using three dimensional digital comparisons. Using the three-dimensional

models provided a way in which to compare teeth without being influenced

by biases from human error. The software Cloud Compare (Hassett and

Lewis-Bale, 2017) compares the mesh clouds of three-dimensional models

to provide a similarity index. These methods were used to determine the

crocodilian diversity within the Chinchilla Local Fauna, and if it matches

with previous studies presenting two taxa (DeVis, 1886; Louys and Price,

2015; Molnar, 1982).

Initial analyses are plotted with the inclusion of the two-modern species, C.

porosus and C. johnstoni (Figures 11 and 13). Crocodylus johnstoni had

very little overlap with the fossil specimens and C. porosus in Figure 13.

However, C. porosus had a large amount of overlap with the upper section

of the graph. This can be used to infer the difference between the modern

freshwater crocodile, C. johnstoni, when compared to C. porosus and the

fossil specimens. Teeth of C. johnstoni, though conical, tend to have a

different shape to those of C. porosus due to the different dietary

preferences. Figure 13 also shows how similar the teeth are between the

upper group of the fossil specimens and C. porosus. The similarity may be

linked to similar food sources and hunting preferences of the two-species

due to the overlap in tooth shape and size (Brink et al, 2015; Dauphin and

Williams, 2007). With the removal of the modern specimens, the two

groups within the fossil specimens become clearer (Figure 14 and 16).

I have focused primarily on the specimens that were used for the three-

dimensional analysis, which excluded some specimens from the initial


64

dataset due to inconsistencies within the teeth (Appendix 4). This reduced

set of teeth was also plotted on a PCA graph and the two groups are clearly

visible at the top and bottom of the graph (Figure 16). Given the two visible

groups, the data confirms previous studies commenting on two crocodilian

species living in the Chinchilla Local Fauna (DeVis, 1886; Louys and Price,

2015; Molnar, 1982). The key features that differentiate the two groups

from each other are with the size and shape of the teeth. Those that plot at

the top of the graph are generally larger and more conical, whereas those at

the bottom are more ziphodont not only at the base but across the crown of

the tooth.

The CT scanned teeth were further analysed using Non-Metric Multi-

Dimensional Scaling (MDS). This required the three-dimensional analysis

of the specimens using the output of a similarity index from Cloud

Compare. At the outset, it was difficult to determine how the teeth were

plotting across the MDS graph (Figure 17). However, as teeth were selected

to be placed on the graph, a definite trend was recognised. Using the three-

dimensional data, it is possible to determine the approximate positioning of

an isolated tooth along the tooth row, which is usually difficult to do (Figure

18). Furthermore, those teeth that plot towards the top section of the graph

are conical in shape with a conical base, and those towards the bottom

section are ziphodont, see Figure 19 for images of all teeth and their

corresponding base shape.

Given that one of the species currently known from the Chinchilla Local

Fauna is Pallimnarchus pollens, which has large conically based teeth, it

can be confirmed that the teeth plotting at the top of the graph belong to this

species. Based upon measurements taken of the alveoli of the modern

species C. porosus and C. johnstoni, the compression ratio increases

towards the posterior of the jaw (Figure 38, QMJ39230 and QMJ48127).

Given the lack of highly compressed molariform teeth specimens (Figure 28

and 29), it can be stated with certainty that neither C. porosus nor C.

johnstoni are present within the Chinchilla Local Fauna. C. johnstoni can

also be excluded due to the lack of overlap with the fossil specimens in the

initial dataset (Figure 13). C. porosus has also been excluded from the


65

Chinchilla Local Fauna due to the lack of fluting on the teeth, a distinctive

feature of C. porosus teeth not present on Pallimnarchus specimens.

The ratio of the Crown Base Length/Crown Base Width was calculated and

then used to determine if this influenced the composition of the two groups

seen in previous graphs and the positioning along the tooth row. Using these

ratios, a histogram was plotted and two groups were returned (Figure 31).

The teeth were then marked according to the groups shown on the histogram

and plotted accordingly on a scatterplot (Figure 32). The ratio can be used to

match alveoli to the tooth base and position isolated tooth specimens along

the tooth row where possible.

The ratio for the alveoli of available crocodilian specimens was also

collected. In Crocodylus spp. the anterior alveoli are the least compressed

with a ratio of nearly 1. However, the further towards the posterior end of

the jaw, the more compressed the alveoli become with the length being

almost double the width (Figure 38). This leads to a general upwards trend

in the alveolus ratio of Crocodylus species along the tooth row (anterior to

posterior). Both modern and fossilised specimens were included in the

Crocodylus alveoli analysis. For comparison, the alveoli shape and size for

specimens attributed to Pallimnarchus remain relatively consistent along the

tooth row (Figure 39). Aside from a few outliers, the majority of the

Pallimnarchus alveoli remain very close to having a ratio of 1, with minimal

compression towards the posterior end of the jaw. This is diagnostic of

Pallimnarchus and has been mentioned in previous studies (Willis and

Molnar, 1997a). The alveoli of Quinkana sp. are different again from

Crocodylus and Pallimnarchus. The compression of the teeth is much

greater with the majority of the alveoli having a ratio of 1.3 and higher

(Figure 40). This shows compression along the entire tooth row, which is

not seen in either of the other crocodilian species measured. The base and

roots of the teeth in the jaw would match the alveoli ratio, therefore these

ratios can be used as diagnostic features to determine which species the

isolated teeth may belong to if you now the general position of the tooth

within the tooth row. Several unknown jaw specimens were also measured

to determine the alveoli ratio and their taxonomic position (Figure 41).


66

The unknown specimens listed in Figure 41 were collected from Chinchilla

and surrounding areas on the Darling Downs, and are all from a similar age.

Two of the unknown specimens, QMF30582 and “CF2” have limited

locality data, however they were recovered from the Darling Downs,

Southeast Queensland. Due to the lack of information from the time of

collection and the age of the registration cards, it can be assumed that these

specimens were an early addition to the Queensland Museum Fossil

Collection and confirm the locality as being ‘Darling Downs’. Both

specimens are oxidised with an adhering matrix consisting of an iron-oxide

cement, coarse grained sand and gravel sediment. The only sites that have

been collected from the Darling Downs and form early Queensland Museum

collections that preserve such matrix come from the Chinchilla area, and in

particular from the Chinchilla Sands unit. None of the younger Pleistocene

sites from the Darling Downs preserve a sedimentary matrix as seen in these

two specimens. Therefore, with a fair degree of certainty, we can assign

these two specimens to the Pliocene-aged Chinchilla Sands as members of

the Pliocene crocodilian fauna from the Darling Downs.

Figure 42: Crocodylus porosus (left) and Crocodylus johnstoni (right)

dentary specimens from the Queensland Museum Vertebrate Collection.


67

Figure 43: From left to right, QMF17479 a fossilised partial dentary of

Crocodylus johnstoni, QMF30582 listed as a Quinkana sp. partial dentary

from the Darling Downs and CF2 an unknown ziphodont species of

crocodilian from the Darling Downs, probably a species of Quinkana.

Of particular note is specimen QMF30582. Details of the specimen list it as

a “Quinkana” dentary. It is clearly different from modern Crocodylus and

Pallimnarchus specimens (Figure 42 and 43). However, it also doesn’t fit

the morphology of known Quinkana species. Quinkana alveoli have a

consistently high compression ratio (Figure 40, whereas QMF30582 does

not. However, it does not possess the same alveolar ratios seen in

Pallimnarchus or Crocodylus. On comparison with the dentary element

from Chinchilla that more closely resembles known Quinkana species,

QMF30582 is much more robust. When looking from anterior to posterior,

the compression of the alveoli is also less pronounced reflecting a semi-

ziphodont morphology as opposed to a highly compressed ziphodont

dentition as seen in true Quinkana (Figure 41 and 44). Therefore, it is

proposed here that QMF30582 represents a new crocodilian taxon of semi-

ziphodont dentition, consistent with the semi-ziphodont teeth found at

Chinchilla, and different to the known species of Quinkana. CF2 is also the


68

first dentary element assignable to true Quinkana from the Chinchilla Local

Fauna.

The assessment of isolated teeth using two-dimensional and three-

dimensional morphometric and geometric comparisons, allow this study to

differentiate two distinct morphologies from the Chinchilla Local Fauna,

conical and semi-ziphodont. The distribution of these morphologies also

allowed the placement of these isolated teeth into their respective regions of

the jaw (e.g. caniniform vs molariform). Analysis of tooth row alveoli

enabled this study to match isolated teeth to the isolated, toothless jaws.

Two dominant morphologies were recovered pertaining to tooth rows with

conical based teeth (e.g. Pallimnarchus) and tooth rows with ziphodont-

based teeth (e.g. Quinkana). However, on comparison of the ziphodont-

based jaws it was clear that this morphology represents two distinct forms,

semi-ziphodont and ziphodont. The semi-ziphodont dentary (QMF30582)

differs from the ziphodont dentary (CF2) (Figure 43). The result of this

study suggests that there are three crocodilian taxa represented by three jaw

morphologies, yet potentially only two based solely on teeth alone. The

absence of the third tooth morphotype more typical of known Quinkana spp.

(i.e. highly compressed base) may be due to taphonomic bias in fossilisation

at Chinchilla with the larger, more robust conical and semi-ziphodont teeth

preserving more readily than the highly compressed true Quinkana teeth.

There is also the possibility that Quinkana have been a rare species during

the Pliocene or within the Chinchilla Local Fauna.

Crocodilian teeth, at first, appear to have a very similar shape and size

across the tooth row. However, when using three-dimensional data (both

morphometric and geometric), the subtle differences between each tooth

becomes apparent. With large enough samples, this data can differentiate

taxonomically or functionally informative groupings. This has aided the

present study to determine approximate position of isolated teeth along the

tooth row. The usefulness of three-dimensional datasets has been confirmed

with this research. The ultimate outcome of which has possibly identified

three crocodilian taxa from the Chinchilla Local Fauna, with a likely new

taxon needing full description. Similar bulk morphometric processing and


69

tooth row inference may be applicable to other vertebrate groups that also

appear to have homodont dentition, such as theropod dinosaurs and sharks.

Figure 44: Quinkana sp. tooth from Mt. Etna Cave National Park, central

Queensland.

6.1 Future Research

Future work should include a greater variety of modern species and

specimens, including ontogenetic series, to add to the dataset due to the vast

morphometric differences in the fossil specimens. Previously this would

have required collections to collect data from various osteological

specimens. However, based on this present study it is clear that modern

osteological collections do not always hold enough material available to

produce adequate datasets for comparison. This is primarily due to the lack

of individual wild-caught specimens being rendered into skeletons within

museum collections. One of the key innovations of this study was to

incorporate the use of CT data and extracted three-dimensional models from

these scans. The benefits of this are two-fold. Firstly, it allows full

geometric data to be collected simultaneously and with precise methods, as

demonstrated by the results provided for isolated teeth during this study.

Secondly, it provides a unique avenue to access geometric data on modern

species where skeletal remains are not available, especially with teeth

located within the alveoli.

The success of the three-dimensional data from this study demonstrates that

it is feasible to collect data from previously CT scanned specimens (i.e.

available online) that have been uploaded as supplementary data for other


70

crocodilian studies not necessarily focusing on dentition. Collecting and

rendering three-dimensional models from these datasets would provide a

much broader opportunity to assess morphological diversity in tooth shape,

position and alveolar shape and size. This would then have potentially

phylogenetic and ecomorphological value by being able to assign isolated

toothless jaw fragments to isolated teeth from similar fossiliferous

sedimentary units.

Using multiple extant species to add to the research would test if the

methodology works across all crocodilian species and how they compare to

each other. It would also be useful to add more molariform teeth and

specimens from other fossil crocodilian species from across Australia

allowing the present study to expand to assess morphological diversity in

crocodilians across time and space within the Australasian region.

Recently the right side of a Pallimnarchus dentary was found on the Darling

Downs (Figure 45). The specimen has many unerupted and in situ teeth

which can be rendered out in a CT Scan to add to the current dataset,

providing the unique opportunity to directly compare isolated teeth with

alveoli, ultimately testing the current studies results. The alveoli and teeth

will help determine if the pattern seen in this study from a Pliocene species

is subsequently found in a Pleistocene species of Pallimnarchus. This can

then be used to find possible patterns along the tooth row of other

crocodilian species. Another dentary and maxillary fragment of a species

from the Pleistocene of Winton, central Queensland, has recently been made

available for research. This dentary shows strikingly similar morphology in

both alveolar shape and dentary shape to the semi-ziphodont taxon proposed

here to be new (QMF30582). It clearly shows differences with the new

Pallimnarchus jaw and therefore allows for a more direct comparison of two

similarly sized crocodilians from the Plio-Pleistocene of Australia (Figure

46).


71

Figure 45: Three-dimensional photogrammetric model of a recently

discovered Pallimnarchus sp. dentary from the Darling Downs to be used

for future research.

Figure 46: Three -dimensional photogrammetric model of a recently

discovered crocodilian dentary from the Pleistocene of Winton to be used in

future research.


72

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87

8.0 Appendices:

Appendix 1: A summary of the raw data of all teeth measured by hand. M1

= Height; M1 ½ = Half height of tooth; M2 = Anterior-posterior half way up

the tooth; M3 = Labio-lingual half way up the tooth; M4 = Anterior-

posterior half way up the tooth; M5 = Labio-lingual half way up the tooth.

Tooth

Number

M1 M1 1/2 M2 M3 M4 M5

1.1 23.51 11.76 10.42 5.86 12.81 7.15

1.2 15.64 7.82 12.63 7.21 14.67 8.74

1.3 17.09 8.55 11.10 6.45 ? 8.01

1.4 16.95 8.48 12.31 7.17 13.01 8.13

1.5 16.03 8.01 12.15 6.64 13.24 8.28

1.6 24.12 12.06 11.00 6.43 12.97 8.44

1.7 24.06 12.03 10.86 6.34 12.59 8.35

1.8 9.71 4.85 9.11 5.50 10.35 ?

2.1 18.47 9.24 8.70 5.34 9.75 6.91

2.2 20.07 10.04 10.60 6.26 14.11 8.86

2.3 22.70 11.35 12.33 7.36 14.15 9.74

2.4 30.14 15.07 13.39 8.08 16.10 9.85

2.5 15.57 7.79 12.36 7.55 12.92 9.05

2.6 25.55 12.78 13.99 8.07 15.33 10.15

2.7 22.51 11.26 12.41 7.52 14.42 9.79

2.8 23.61 11.80 14.93 8.68 17.09 11.04

2.9 26.19 13.10 10.14 5.63 12.73 7.98

2.10 24.74 12.37 12.21 7.43 14.38 9.92

2.11 24.74 12.37 11.71 6.59 14.16 9.05

2.12 25.98 12.98 13.35 7.65 14.62 10.00

2.13 24.20 12.10 11.20 7.39 13.57 9.93

3.1 7.60 3.80 7.74 5.70 8.20 6.33

3.2 11.83 5.92 10.95 8.21 12.23 9.39

3.3 11.50 5.75 5.73 4.17 6.92 5.18

3.4 28.70 14.35 12.83 8.17 ? ?


88

3.5 26.28 13.14 11.26 7.22 14.08 10.29

3.6 29.96 14.98 12.40 7.36 15.34 10.45

3.7 25.54 12.77 12.91 9.91 14.84 10.39

3.8 30.60 15.30 12.85 7.87 15.83 11.23

3.9 30.22 15.11 12.96 7.81 ? 11.33

3.10 20.00 10.00 10.07 6.14 12.40 ?

3.11 38.46 19.23 13.48 8.67 ? ?

3.12 12.63 6.32 8.50 5.57 9.42 7.08

3.13 24.48 12.24 12.61 8.02 15.58 10.99

3.14 25.71 12.85 13.12 8.21 15.00 10.86

3.15 35.02 17.51 14.75 9.27 17.95 12.21

3.16 24.80 12.40 9.92 6.40 11.99 8.93

4.1 20.36 10.18 8.06 6.03 9.62 ?

4.2 9.95 4.97 8.03 6.29 9.59 7.14

4.3 18.52 9.26 7.22 5.66 8.98 7.15

4.4 15.83 7.92 6.77 5.21 8.28 6.57

4.5 29.12 14.56 13.59 8.55 17.76 13.08

4.6 27.98 13.99 15.03 9.29 17.93 12.05

4.7 24.61 12.31 12.20 8.11 14.98 10.74

4.8 21.41 10.70 12.06 7.21 13.85 9.83

4.9 31.65 15.83 12.01 7.80 14.15 10.56

4.10 20.38 10.19 11.38 7.45 14.87 10.26

4.11 26.57 13.28 14.46 9.25 17.55 12.19

4.12 24.68 12.34 10.44 6.93 ? 9.08

4.13 28.81 14.41 11.22 7.48 13.43 10.42

4.14 31.55 15.78 15.09 9.18 17.79 12.49

4.15 34.83 17.41 17.68 11.96 ? ?

4.16 23.99 12.00 10.21 6.83 12.39 9.28

4.17 28.79 14.40 13.96 8.49 16.90 11.45

4.18 29.71 14.85 12.50 8.26 16.36 11.19

4.19 17.12 8.56 11.92 7.53 12.57 9.70

5.1 26.77 13.38 11.79 10.42 ? ?

5.2 12.03 6.02 7.84 5.80 8.92 7.22


89

5.3 18.99 9.48 7.85 6.13 9.49 8.69

5.4 20.52 10.26 8.50 6.53 10.72 8.80

5.5 29.90 14.95 10.16 7.03 11.74 9.55

5.6 25.63 12.82 13.58 9.07 16.29 12.08

5.7 30.35 15.18 13.24 8.97 15.45 11.84

5.8 32.97 16.49 12.74 8.75 ? ?

5.9 38.07 19.04 15.44 9.68 16.66 13.16

5.10 29.30 14.65 15.73 10.33 19.24 13.73

5.11 30.65 15.33 17.18 10.22 20.09 13.94

5.12 28.00 14.00 14.31 9.76 16.89 12.29

5.13 30.87 15.44 12.33 8.77 15.40 11.45

5.14 24.60 12.30 13.43 7.98 16.06 10.56

5.15 28.22 14.11 14.61 9.21 16.44 12.26

5.16 28.90 14.45 11.73 7.32 13.29 9.83

5.17 32.79 16.39 16.59 10.01 17.79 13.03

5.18 34.39 17.20 13.63 8.80 16.53 12.00

5.19 38.96 19.48 15.53 9.93 ? ?

5.20 20.70 10.35 12.52 7.74 13.71 10.21

5.21 33.27 16.64 14.63 9.24 17.81 13.50

5.22 37.61 18.81 16.15 9.38 18.67 13.56

5.23 29.97 14.98 12.66 8.03 14.81 11.39

5.24 32.90 16.45 13.56 8.49 14.99 11.56

5.25 18.10 9.05 10.68 6.21 11.94 8.44

5.26 14.33 7.17 10.80 8.78 12.55 9.99

5.27 21.11 10.56 11.39 7.44 12.85 10.05

6.1 10.00 5.00 4.89 4.24 6.17 4.99

6.2 13.21 6.60 5.64 4.41 7.25 6.27

6.3 29.68 14.84 16.14 9.53 17.99 13.45

6.4 28.47 14.24 14.52 9.36 18.29 13.23

6.5 26.26 13.13 10.53 7.30 12.93 10.36

6.6 16.29 8.14 7.43 5.83 9.38 ?

6.7 30.97 15.48 13.72 8.57 14.99 11.74

6.8 12.18 6.09 7.04 5.52 9.09 7.33


90

6.9 34.84 17.42 14.03 9.47 18.20 14.05

6.10 29.44 14.72 14.34 8.74 16.00 11.88

6.11 29.42 14.71 ? 9.66 ? 12.84

6.12 30.81 15.41 12.96 8.93 17.15 12.89

6.13 31.19 15.59 14.12 8.50 16.11 12.19

6.14 29.32 14.66 14.08 9.38 17.29 ?

6.15 32.16 16.08 13.10 8.38 14.52 12.01

7.1 24.21 12.11 9.34 7.52 11.47 10.61

7.2 19.63 9.82 7.31 6.43 ? ?

7.3 16.34 8.17 8.74 6.27 9.34 9.32

7.4 20.51 10.26 8.17 6.12 10.27 9.31

7.5 34.26 17.13 16.09 11.03 20.97 16.24

7.6 12.30 6.15 10.57 8.77 11.66 9.63

7.7 28.28 14.14 11.54 7.74 13.84 10.86

7.8 32.50 16.25 13.76 9.45 17.39 13.68

7.9 29.98 14.99 13.85 8.40 15.80 12.26

7.10 31.15 15.57 12.90 9.11 15.10 12.09

7.11 14.50 7.25 10.96 8.80 12.45 10.24

7.12 34.62 17.31 13.87 8.76 16.73 13.31

7.13 17.62 8.81 11.62 9.29 14.51 11.24

7.14 16.57 8.29 10.08 7.23 12.01 9.65

7.15 29.41 14.71 12.96 8.34 14.61 11.36

7.16 12.99 6.50 8.07 6.07 9.07 7.54

7.17 32.38 16.19 12.66 8.00 ? 11.18

8.1 28.66 14.33 11.85 9.68 ? 13.36

8.2 23.66 11.83 10.32 8.02 12.80 11.93

8.3 18.10 9.05 12.34 10.27 13.86 12.65

8.4 11.88 5.94 7.20 6.00 8.77 7.46

8.5 10.98 5.49 9.38 9.19 11.17 9.75

8.6 13.53 6.77 11.63 9.59 13.81 12.24

8.7 24.95 12.48 10.24 7.82 12.75 11.07

8.8 25.73 12.87 13.89 9.28 16.57 12.42

8.9 11.09 5.55 7.13 6.10 8.77 7.35


91

8.10 35.73 17.86 15.66 10.67 18.53 14.72

8.11 38.76 19.38 15.29 10.40 17.96 14.88

8.12 39.12 19.61 14.27 11.34 17.39 ?

8.13 40.03 20.02 14.87 10.45 ? ?

8.14 25.01 12.51 10.23 6.95 12.03 9.85

8.15 40.07 20.04 16.46 10.61 ? ?

8.16 34.73 17.36 14.37 9.52 17.07 13.66

9.1 19.49 9.75 10.20 8.28 11.88 11.25

9.2 15.52 7.76 7.61 6.36 9.62 8.75

9.3 11.55 5.77 11.37 9.34 13.01 11.43

9.4 12.83 6.42 11.57 9.86 13.14 11.86

9.5 18.66 9.33 10.94 8.63 12.50 10.53

9.6 40.97 20.49 15.27 10.25 18.20 14.60

9.7 36.58 18.29 15.53 10.79 19.02 15.65

9.8 33.89 16.94 16.38 10.68 19.25 14.88

9.9 12.65 6.33 11.16 9.93 12.90 11.26

9.10 14.46 7.23 11.88 10.91 13.61 11.72

9.11 32.75 16.37 12.00 10.81 15.22 ?

9.12 30.97 15.49 12.92 9.56 15.60 13.40

9.13 21.70 10.85 14.08 11.59 15.86 14.28

9.14 10.40 5.20 9.29 8.30 11.11 ?

9.15 17.16 8.58 11.83 9.18 ? ?

9.16 36.94 18.47 14.17 10.09 ? 14.48

9.17 35.47 17.74 14.33 9.75 17.11 14.27

9.18 17.31 8.65 12.22 10.77 14.30 12.12

9.19 37.97 18.99 16.56 11.04 19.37 15.65

9.20 17.04 8.52 11.71 8.92 13.40 11.27

9.21 31.92 15.96 14.65 9.22 18.67 13.84

9.22 36.35 18.17 14.35 8.76 15.21 12.58

9.23 12.96 6.48 12.37 10.81 13.26 12.03

9.24 24.44 12.22 14.93 12.24 17.24 14.88

9.25 22.52 11.26 13.68 8.74 14.88 11.57

9.26 13.75 6.87 8.84 9.62 8.37 8.39


92

9.27 38.16 19.08 13.80 9.58 16.00 14.09

9.28 14.99 7.49 7.15 5.84 7.92 6.81

9.29 14.28 7.14 11.55 9.55 12.67 10.98

10.1 26.13 13.07 11.16 8.93 14.29 ?

10.2 18.39 9.20 9.23 8.30 10.19 10.44

10.3 17.48 8.74 ? 7.55 9.96 9.24

10.4 14.43 7.22 7.39 5.75 8.00 7.50

10.5 13.99 7.00 10.67 10.72 13.44 ?

10.6 22.41 11.21 12.53 10.60 15.01 13.22

10.7 14.21 7.11 7.13 5.90 9.33 8.40

10.8 35.66 17.83 14.54 10.07 16.70 14.82

10.9 19.81 9.91 13.12 10.98 14.66 ?

10.10 19.64 9.82 9.86 7.64 10.94 10.02

10.11 10.60 5.30 9.18 8.23 10.96 10.10

10.12 17.98 8.99 10.70 8.94 12.56 10.63

10.13 10.49 5.25 10.52 8.59 11.94 11.44

10.14 21.68 10.84 8.47 7.07 ? ?

10.15 15.25 7.62 10.08 7.67 11.27 9.65

10.16 9.49 4.72 9.19 8.92 12.31 11.74

10.17 12.51 6.26 9.52 9.08 11.76 10.79

10.18 28.16 14.08 ? 11.30 17.39 14.95

10.19 21.14 10.57 9.39 7.03 10.85 9.69

10.20 15.97 7.99 12.46 11.33 14.37 13.24

10.21 15.93 7.96 6.60 5.14 7.25 ?

10.22 34.12 17.06 14.22 11.25 17.14 15.11

10.23 20.52 10.26 9.03 6.87 10.17 9.21

10.24 29.89 14.95 11.29 8.90 14.75 13.47

10.25 31.56 15.78 15.37 10.97 16.47 14.20

10.26 25.03 12.52 10.47 8.02 13.22 11.69

10.27 15.05 7.53 12.27 11.10 15.07 14.02

10.28 18.15 9.08 8.05 6.45 9.58 8.49

10.29 18.38 9.19 8.65 7.40 9.95 9.05

10.30 17.30 8.65 12.72 10.93 16.15 13.96


93

10.31 12.92 6.46 11.17 9.43 14.16 12.19

10.32 19.86 9.93 12.32 9.98 14.75 12.29

10.33 18.98 9.49 12.35 10.02 13.96 12.35

10.34 17.14 8.57 7.23 5.82 8.41 ?

11.1 33.12 16.56 13.25 10.70 ? ?

11.2 29.22 14.61 12.79 9.85 15.99 14.49

11.3 21.10 10.55 10.37 8.35 11.45 11.86

11.4 16.66 8.33 8.06 6.50 9.21 8.60

11.5 15.67 7.83 7.04 5.63 7.84 7.07

11.6 20.38 10.19 12.13 9.73 14.00 12.47

11.7 21.22 10.61 8.95 6.82 11.57 10.60

11.8 22.43 11.21 10.89 8.82 12.62 11.66

11.9 13.11 6.56 10.92 9.38 12.99 11.81

11.10 33.08 16.54 12.26 10.03 15.94 14.56

11.11 33.78 16.89 14.07 11.01 15.84 15.03

11.12 18.77 9.38 12.94 10.14 14.45 13.24

11.13 26.46 13.23 10.88 8.47 14.28 13.26

11.14 16.25 8.12 11.78 10.42 13.82 12.10

11.15 11.60 5.80 8.49 7.34 9.93 ?

11.16 19.14 9.57 12.20 10.04 13.70 12.57

11.17 18.48 9.24 11.84 9.81 13.31 11.81

11.18 27.42 13.71 11.52 9.79 12.92 ?

11.19 22.20 11.10 12.81 11.42 14.93 13.96

11.20 18.08 9.04 8.77 7.36 9.74 9.24

11.21 17.65 8.83 9.78 7.70 10.48 9.75

11.22 18.50 9.25 12.31 11.31 15.58 14.20

11.23 16.02 8.01 9.16 7.15 9.79 9.09

11.24 21.79 10.90 10.37 8.47 12.03 11.06

11.25 29.66 14.83 11.43 9.37 13.33 12.58

12.1 29.02 14.51 10.82 8.56 13.23 12.45

12.2 18.29 9.15 13.60 12.31 16.35 15.50

12.3 14.20 7.10 6.13 4.48 6.91 5.86

12.4 11.93 5.97 ? 4.59 ? 6.20


94

12.5 16.25 8.13 7.95 6.14 8.54 8.00

12.6 13.46 6.73 10.55 9.73 12.62 11.86

12.7 31.06 15.53 12.69 9.35 15.79 15.10

12.8 22.86 11.42 10.35 8.61 11.28 11.05

12.9 17.36 8.68 10.45 8.94 12.25 11.87

12.10 33.53 16.76 12.55 10.74 15.97 14.86

12.11 38.54 19.27 14.29 12.15 18.21 16.78

12.12 24.73 12.37 9.21 7.53 ? ?

12.13 17.32 8.66 10.13 8.36 11.36 10.17

12.14 19.11 9.56 9.33 8.09 11.20 10.37

12.15 34.33 17.16 16.34 13.16 18.19 16.79

12.16 18.53 9.27 8.34 6.77 9.22 8.90

12.17 35.89 17.95 15.47 12.92 18.98 17.82

12.18 15.38 7.69 8.05 ? 8.76 8.40

12.19 19.05 9.52 10.34 8.70 12.99 11.82

12.20 24.85 12.42 12.23 10.35 15.08 14.17

12.21 11.22 5.61 10.32 9.05 11.21 10.01

13.1 29.93 14.96 12.11 10.90 16.44 16.29

13.2 18.62 9.31 7.25 5.90 8.76 8.21

13.3 15.08 7.54 7.77 6.14 9.07 8.24

13.4 31.53 15.77 16.88 14.50 21.53 ?

13.5 21.00 10.50 10.14 8.41 12.66 11.93

13.6 21.28 10.64 9.37 7.67 ? 9.76

13.7 32.42 16.21 15.21 12.38 18.11 17.28

13.8 30.71 15.36 13.32 11.32 16.49 15.95

13.9 14.07 7.03 11.75 10.32 13.60 ?

13.10 32.09 16.04 14.19 11.90 16.20 15.54

13.11 28.23 14.11 12.19 10.20 14.36 13.53

13.12 31.07 15.53 11.89 10.91 14.94 14.58

13.13 17.17 8.59 8.51 6.85 10.12 ?

13.14 29.85 14.93 13.20 12.14 16.40 16.77

13.15 35.98 17.99 14.44 10.41 18.52 16.58

13.16 34.49 17.25 12.43 10.08 17.17 15.76


95

13.17 17.07 8.54 8.05 6.33 10.04 9.00

13.18 26.82 13.41 12.52 10.97 15.12 14.97

13.19 14.35 7.18 10.94 9.65 13.22 12.52

13.20 20.61 10.31 9.97 8.35 ? 11.55

13.21 13.37 6.68 11.73 10.45 13.25 12.47

13.22 29.97 14.98 13.46 11.22 ? 15.87

13.23 27.96 13.98 11.76 10.94 13.75 13.86

13.24 28.93 14.46 13.14 11.47 ? 15.18

13.25 19.54 9.77 7.74 6.38 9.24 8.71

14.1 12.24 6.12 6.29 5.29 7.31 6.93

14.2 14.33 7.16 7.81 6.35 8.63 8.27

14.3 19.98 9.49 12.26 10.94 14.37 13.86

14.4 35.77 17.89 15.40 14.03 ? ?

14.5 31.50 10.75 12.59 10.53 14.62 14.39

14.6 31.23 15.61 12.60 9.92 ? 14.15

14.7 23.20 11.60 9.69 7.24 11.25 10.84

14.8 21.99 11.00 13.05 11.34 15.43 14.92

14.9 20.36 10.18 9.73 8.20 ? 11.03

14.10 16.04 8.02 8.50 6.85 8.98 8.60

14.11 19.58 9.79 9.99 7.83 11.24 10.67

14.12 20.05 10.02 9.11 7.73 ? 10.83

14.13 32.92 16.46 13.49 11.04 16.60 16.34

14.14 19.37 9.69 11.73 9.75 14.22 13.35

14.15 24.99 12.49 13.08 10.89 14.98 14.66

14.16 29.05 14.53 13.94 11.61 16.45 15.77

14.17 32.53 16.27 14.57 12.04 16.87 16.31

14.18 31.97 15.99 12.53 10.85 15.79 15.24

14.19 16.50 8.25 9.50 8.01 10.93 10.54

14.20 18.18 9.09 9.48 7.75 10.75 10.30

14.21 29.70 14.85 12.70 11.11 ? 15.25

14.22 31.43 15.71 14.28 11.80 ? 16.37

15.1 29.10 14.55 12.71 10.46 15.05 14.12

15.2 23.45 11.72 11.00 9.57 12.88 12.94


96

15.3 22.03 11.02 12.08 ? 13.16 12.82

15.4 18.16 9.08 7.50 6.56 9.84 9.23

15.5 27.70 13.85 12.44 10.76 15.25 15.35

15.6 20.44 10.22 9.45 8.43 11.02 11.00

15.7 16.07 13.04 12.09 10.25 13.95 13.55

15.8 16.52 8.26 8.85 7.84 10.43 9.74

15.9 18.74 9.37 9.34 7.84 10.39 10.52

15.10 22.68 11.34 9.72 8.93 11.75 11.63

15.11 19.82 9.91 8.73 7.46 9.55 9.64

15.12 19.97 9.99 10.56 8.74 11.82 11.53

15.13 21.22 10.61 13.83 12.55 16.59 16.37

15.14 22.55 11.28 11.01 9.93 13.86 14.05

15.15 15.63 7.81 7.07 5.61 8.75 8.50

15.16 34.40 17.20 13.75 12.47 17.82 17.46

15.17 33.06 16.53 13.35 11.93 17.64 17.15

15.18 37.50 18.75 14.84 13.23 ? ?

15.19 38.35 19.17 18.47 16.32 22.38 21.99

Appendix 2: Clean data set with no unknowns.

Tooth

Number M1 M2 M3 M4 M5

1.1 23.51 10.42 5.86 12.81 7.15

1.2 15.64 12.63 7.21 14.67 8.74

1.4 16.95 12.31 7.17 13.01 8.13

1.5 16.03 12.15 6.64 13.24 8.28

1.6 24.12 11.00 6.43 12.97 8.44

1.7 24.06 10.86 6.34 12.59 8.35

2.1 18.47 8.70 5.34 9.75 6.91

2.2 20.07 10.60 6.26 14.11 8.86

2.3 22.70 12.33 7.36 14.15 9.74

2.4 30.14 13.39 8.08 16.10 9.85

2.5 15.57 12.36 7.55 12.92 9.05


97

2.6 25.55 13.99 8.07 15.33 10.15

2.7 22.51 12.41 7.52 14.42 9.79

2.8 23.61 14.93 8.68 17.09 11.04

2.9 26.19 10.14 5.63 12.73 7.98

2.10 24.74 12.21 7.43 14.38 9.92

2.11 24.74 11.71 6.59 14.16 9.05

2.12 25.98 13.35 7.65 14.62 10.00

2.13 24.20 11.20 7.39 13.57 9.93

3.1 7.60 7.74 5.70 8.20 6.33

3.2 11.83 10.95 8.21 12.23 9.39

3.3 11.50 5.73 4.17 6.92 5.18

3.5 26.28 11.26 7.22 14.08 10.29

3.6 29.96 12.40 7.36 15.34 10.45

3.7 25.54 12.91 9.91 14.84 10.39

3.8 30.60 12.85 7.87 15.83 11.23

3.12 12.63 8.50 5.57 9.42 7.08

3.13 24.48 12.61 8.02 15.58 10.99

3.14 25.71 13.12 8.21 15.00 10.86

3.15 35.02 14.75 9.27 17.95 12.21

3.16 24.80 9.92 6.40 11.99 8.93

4.2 9.95 8.03 6.29 9.59 7.14

4.3 18.52 7.22 5.66 8.98 7.15

4.4 15.83 6.77 5.21 8.28 6.57

4.5 29.12 13.59 8.55 17.76 13.08

4.6 27.98 15.03 9.29 17.93 12.05

4.7 24.61 12.20 8.11 14.98 10.74

4.8 21.41 12.06 7.21 13.85 9.83

4.9 31.65 12.01 7.80 14.15 10.56

4.10 20.38 11.38 7.45 14.87 10.26

4.11 26.57 14.46 9.25 17.55 12.19

4.13 28.81 11.22 7.48 13.43 10.42

4.14 31.55 15.09 9.18 17.79 12.49

4.16 23.99 10.21 6.83 12.39 9.28


98

4.17 28.79 13.96 8.49 16.90 11.45

4.18 29.71 12.50 8.26 16.36 11.19

4.19 17.12 11.92 7.53 12.57 9.70

5.2 12.03 7.84 5.80 8.92 7.22

5.3 18.99 7.85 6.13 9.49 8.69

5.4 20.52 8.50 6.53 10.72 8.80

5.5 29.90 10.16 7.03 11.74 9.55

5.6 25.63 13.58 9.07 16.29 12.08

5.7 30.35 13.24 8.97 15.45 11.84

5.9 38.07 15.44 9.68 16.66 13.16

5.10 29.30 15.73 10.33 19.24 13.73

5.11 30.65 17.18 10.22 20.09 13.94

5.12 28.00 14.31 9.76 16.89 12.29

5.13 30.87 12.33 8.77 15.40 11.45

5.14 24.60 13.43 7.98 16.06 10.56

5.15 28.22 14.61 9.21 16.44 12.26

5.16 28.90 11.73 7.32 13.29 9.83

5.17 32.79 16.59 10.01 17.79 13.03

5.18 34.39 13.63 8.80 16.53 12.00

5.20 20.70 12.52 7.74 13.71 10.21

5.21 33.27 14.63 9.24 17.81 13.50

5.22 37.61 16.15 9.38 18.67 13.56

5.23 29.97 12.66 8.03 14.81 11.39

5.24 32.90 13.56 8.49 14.99 11.56

5.25 18.10 10.68 6.21 11.94 8.44

5.26 14.33 10.80 8.78 12.55 9.99

5.27 21.11 11.39 7.44 12.85 10.05

6.1 10.00 4.89 4.24 6.17 4.99

6.2 13.21 5.64 4.41 7.25 6.27

6.3 29.68 16.14 9.53 17.99 13.45

6.4 28.47 14.52 9.36 18.29 13.23

6.5 26.26 10.53 7.30 12.93 10.36

6.7 30.97 13.72 8.57 14.99 11.74


99

6.8 12.18 7.04 5.52 9.09 7.33

6.9 34.84 14.03 9.47 18.20 14.05

6.10 29.44 14.34 8.74 16.00 11.88

6.12 30.81 12.96 8.93 17.15 12.89

6.13 31.19 14.12 8.50 16.11 12.19

6.15 32.16 13.10 8.38 14.52 12.01

7.1 24.21 9.34 7.52 11.47 10.61

7.3 16.34 8.74 6.27 9.34 9.32

7.4 20.51 8.17 6.12 10.27 9.31

7.5 34.26 16.09 11.03 20.97 16.24

7.6 12.30 10.57 8.77 11.66 9.63

7.7 28.28 11.54 7.74 13.84 10.86

7.8 32.50 13.76 9.45 17.39 13.68

7.9 29.98 13.85 8.40 15.80 12.26

7.10 31.15 12.90 9.11 15.10 12.09

7.11 14.50 10.96 8.80 12.45 10.24

7.12 34.62 13.87 8.76 16.73 13.31

7.13 17.62 11.62 9.29 14.51 11.24

7.14 16.57 10.08 7.23 12.01 9.65

7.15 29.41 12.96 8.34 14.61 11.36

7.16 12.99 8.07 6.07 9.07 7.54

8.2 23.66 10.32 8.02 12.80 11.93

8.3 18.10 12.34 10.27 13.86 12.65

8.4 11.88 7.20 6.00 8.77 7.46

8.5 10.98 9.38 9.19 11.17 9.75

8.6 13.53 11.63 9.59 13.81 12.24

8.7 24.95 10.24 7.82 12.75 11.07

8.8 25.73 13.89 9.28 16.57 12.42

8.9 11.09 7.13 6.10 8.77 7.35

8.10 35.73 15.66 10.67 18.53 14.72

8.11 38.76 15.29 10.40 17.96 14.88

8.14 25.01 10.23 6.95 12.03 9.85

8.16 34.73 14.37 9.52 17.07 13.66


100

9.1 19.49 10.20 8.28 11.88 11.25

9.2 15.52 7.61 6.36 9.62 8.75

9.3 11.55 11.37 9.34 13.01 11.43

9.4 12.83 11.57 9.86 13.14 11.86

9.5 18.66 10.94 8.63 12.50 10.53

9.6 40.97 15.27 10.25 18.20 14.60

9.7 36.58 15.53 10.79 19.02 15.65

9.8 33.89 16.38 10.68 19.25 14.88

9.9 12.65 11.16 9.93 12.90 11.26

9.10 14.46 11.88 10.91 13.61 11.72

9.12 30.97 12.92 9.56 15.60 13.40

9.13 21.70 14.08 11.59 15.86 14.28

9.17 35.47 14.33 9.75 17.11 14.27

9.18 17.31 12.22 10.77 14.30 12.12

9.19 37.97 16.56 11.04 19.37 15.65

9.20 17.04 11.71 8.92 13.40 11.27

9.21 31.92 14.65 9.22 18.67 13.84

9.22 36.35 14.35 8.76 15.21 12.58

9.23 12.96 12.37 10.81 13.26 12.03

9.24 24.44 14.93 12.24 17.24 14.88

9.25 22.52 13.68 8.74 14.88 11.57

9.26 13.75 8.84 9.62 8.37 8.39

9.27 38.16 13.80 9.58 16.00 14.09

9.28 14.99 7.15 5.84 7.92 6.81

9.29 14.28 11.55 9.55 12.67 10.98

10.2 18.39 9.23 8.30 10.19 10.44

10.4 14.43 7.39 5.75 8.00 7.50

10.6 22.41 12.53 10.60 15.01 13.22

10.7 14.21 7.13 5.90 9.33 8.40

10.8 35.66 14.54 10.07 16.70 14.82

10.10 19.64 9.86 7.64 10.94 10.02

10.11 10.60 9.18 8.23 10.96 10.10

10.12 17.98 10.70 8.94 12.56 10.63


101

10.13 10.49 10.52 8.59 11.94 11.44

10.15 15.25 10.08 7.67 11.27 9.65

10.16 9.49 9.19 8.92 12.31 11.74

10.17 12.51 9.52 9.08 11.76 10.79

10.19 21.14 9.39 7.03 10.85 9.69

10.20 15.97 12.46 11.33 14.37 13.24

10.22 34.12 14.22 11.25 17.14 15.11

10.23 20.52 9.03 6.87 10.17 9.21

10.24 29.89 11.29 8.90 14.75 13.47

10.25 31.56 15.37 10.97 16.47 14.20

10.26 25.03 10.47 8.02 13.22 11.69

10.27 15.05 12.27 11.10 15.07 14.02

10.28 18.15 8.05 6.45 9.58 8.49

10.29 18.38 8.65 7.40 9.95 9.05

10.30 17.30 12.72 10.93 16.15 13.96

10.31 12.92 11.17 9.43 14.16 12.19

10.32 19.86 12.32 9.98 14.75 12.29

10.33 18.98 12.35 10.02 13.96 12.35

11.2 29.22 12.79 9.85 15.99 14.49

11.3 21.10 10.37 8.35 11.45 11.86

11.4 16.66 8.06 6.50 9.21 8.60

11.5 15.67 7.04 5.63 7.84 7.07

11.6 20.38 12.13 9.73 14.00 12.47

11.7 21.22 8.95 6.82 11.57 10.60

11.8 22.43 10.89 8.82 12.62 11.66

11.9 13.11 10.92 9.38 12.99 11.81

11.10 33.08 12.26 10.03 15.94 14.56

11.11 33.78 14.07 11.01 15.84 15.03

11.12 18.77 12.94 10.14 14.45 13.24

11.13 26.46 10.88 8.47 14.28 13.26

11.14 16.25 11.78 10.42 13.82 12.10

11.16 19.14 12.20 10.04 13.70 12.57

11.17 18.48 11.84 9.81 13.31 11.81


102

11.19 22.20 12.81 11.42 14.93 13.96

11.20 18.08 8.77 7.36 9.74 9.24

11.21 17.65 9.78 7.70 10.48 9.75

11.22 18.50 12.31 11.31 15.58 14.20

11.23 16.02 9.16 7.15 9.79 9.09

11.24 21.79 10.37 8.47 12.03 11.06

11.25 29.66 11.43 9.37 13.33 12.58

12.1 29.02 10.82 8.56 13.23 12.45

12.2 18.29 13.60 12.31 16.35 15.50

12.3 14.20 6.13 4.48 6.91 5.86

12.5 16.25 7.95 6.14 8.54 8.00

12.6 13.46 10.55 9.73 12.62 11.86

12.7 31.06 12.69 9.35 15.79 15.10

12.8 22.86 10.35 8.61 11.28 11.05

12.9 17.36 10.45 8.94 12.25 11.87

12.10 33.53 12.55 10.74 15.97 14.86

12.11 38.54 14.29 12.15 18.21 16.78

12.13 17.32 10.13 8.36 11.36 10.17

12.14 19.11 9.33 8.09 11.20 10.37

12.15 34.33 16.34 13.16 18.19 16.79

12.16 18.53 8.34 6.77 9.22 8.90

12.17 35.89 15.47 12.92 18.98 17.82

12.19 19.05 10.34 8.70 12.99 11.82

12.20 24.85 12.23 10.35 15.08 14.17

12.21 11.22 10.32 9.05 11.21 10.01

13.1 29.93 12.11 10.90 16.44 16.29

13.2 18.62 7.25 5.90 8.76 8.21

13.3 15.08 7.77 6.14 9.07 8.24

13.5 21.00 10.14 8.41 12.66 11.93

13.7 32.42 15.21 12.38 18.11 17.28

13.8 30.71 13.32 11.32 16.49 15.95

13.10 32.09 14.19 11.90 16.20 15.54

13.11 28.23 12.19 10.20 14.36 13.53


103

13.12 31.07 11.89 10.91 14.94 14.58

13.14 29.85 13.20 12.14 16.40 16.77

13.15 35.98 14.44 10.41 18.52 16.58

13.16 34.49 12.43 10.08 17.17 15.76

13.17 17.07 8.05 6.33 10.04 9.00

13.18 26.82 12.52 10.97 15.12 14.97

13.19 14.35 10.94 9.65 13.22 12.52

13.21 13.37 11.73 10.45 13.25 12.47

13.23 27.96 11.76 10.94 13.75 13.86

13.25 19.54 7.74 6.38 9.24 8.71

14.1 12.24 6.29 5.29 7.31 6.93

14.2 14.33 7.81 6.35 8.63 8.27

14.3 19.98 12.26 10.94 14.37 13.86

14.5 31.50 12.59 10.53 14.62 14.39

14.7 23.20 9.69 7.24 11.25 10.84

14.8 21.99 13.05 11.34 15.43 14.92

14.10 16.04 8.50 6.85 8.98 8.60

14.11 19.58 9.99 7.83 11.24 10.67

14.13 32.92 13.49 11.04 16.60 16.34

14.14 19.37 11.73 9.75 14.22 13.35

14.15 24.99 13.08 10.89 14.98 14.66

14.16 29.05 13.94 11.61 16.45 15.77

14.17 32.53 14.57 12.04 16.87 16.31

14.18 31.97 12.53 10.85 15.79 15.24

14.19 16.50 9.50 8.01 10.93 10.54

14.20 18.18 9.48 7.75 10.75 10.30

15.1 29.10 12.71 10.46 15.05 14.12

15.2 23.45 11.00 9.57 12.88 12.94

15.4 18.16 7.50 6.56 9.84 9.23

15.5 27.70 12.44 10.76 15.25 15.35

15.6 20.44 9.45 8.43 11.02 11.00

15.7 16.07 12.09 10.25 13.95 13.55

15.8 16.52 8.85 7.84 10.43 9.74


104

15.9 18.74 9.34 7.84 10.39 10.52

15.10 22.68 9.72 8.93 11.75 11.63

15.11 19.82 8.73 7.46 9.55 9.64

15.12 19.97 10.56 8.74 11.82 11.53

15.13 21.22 13.83 12.55 16.59 16.37

15.14 22.55 11.01 9.93 13.86 14.05

15.15 15.63 7.07 5.61 8.75 8.50

15.16 34.40 13.75 12.47 17.82 17.46

15.17 33.06 13.35 11.93 17.64 17.15

15.19 38.35 18.47 16.32 22.38 21.99

Appendix 3: Modern specimens Crocodylus porosus (CP) and Crocodylus

johnstoni (CJ) maxillae (T) and dentary (B) measurements without

unknowns. The numbers refer to the tooth, starting at the front of the mouth

and working towards the back.

Tooth


CPT1 8.88 3.84 4 4.89 5.09

CPT2 22.79 8.03 7.35 10.92 10.86

CPT3 26.79 9.57 8.59 12.76 12.59

CPT5 14.7 5.8 4.91 7.62 6.76

CPT6 13.53 6.31 4.83 7.99 7.04

CPT7 17.84 7.19 5.9 9.7 8.92

CPT8 21.02 9.18 7.91 11.74 10.47

CPT9 29.37 12.65 10.74 15.77 15.24

CPT10 15.51 7.05 6.61 10.09 8.8

CPT11 11.79 6.52 5.45 8.36 7.24

CPT12 11.2 6.41 4.95 8.31 7.16

CPT13 13.11 7.6 6.99 9.87 8.29

CPT14 11.74 7.4 6.54 9.53 7.76

CPT15 9.78 7.09 6.23 9.63 7.54


105

CPT16 7.68 6.78 4.72 7.98 6.34

CPT17 6.73 6.23 4.23 7.24 5.49

CPB1 25.76 9.12 7.37 12.16 10.73

CPB2 13.98 5.29 5.07 7.87 7.07

CPB3 15.71 5.86 4.91 8.23 7.34

CPB4 23.77 10.41 8.27 12.73 12.14

CPB5 12.88 5.61 4.72 7.7 6.9

CPB6 11.13 6.29 5.58 8.18 6.88

CPB7 11.75 5.71 4.64 8.11 6.92

CPB8 10.03 6.14 5.8 8.62 6.87

CPB9 12.18 6.53 5.66 8.85 7.13

CPB10 17.39 8.35 7.45 11.07 9.41

CPB11 16.12 9.11 7.29 11.26 9.53

CPB12 13.14 8.83 6.83 10.23 8.38

CPB13 10.8 6.81 5.78 9.34 7.26

CPB14 8.83 7.2 6.05 8.96 7.63

CPB15 6.68 5.62 4.67 7.06 5.58

CJT1 10.59 2.35 2.75 3.71 4

CJT2 9.07 3.11 2.59 4.15 3.96

CJT3 13.32 4.02 3.59 5.82 5.33

CJT4 12.4 4.79 4.75 7.03 6.69

CJT5 9.04 2.69 2.53 3.64 3.54

CJT6 11.24 3.44 2.79 5.29 4.01

CJT7 9.82 3.43 2.64 4.67 4.01

CJT8 12.45 3.47 2.77 5.42 4.53

CJT9 10.9 4.14 3.52 5.64 5.1

CJT10 11.83 5.69 3.72 6.96 5.35

CJT12 10.41 4.46 3.42 6.27 5.26

CJT13 11.64 4.42 3.78 6.9 5.69

CJT14 7.23 4.98 3.51 6.54 5.15


106

CJT15 8 5.25 4.18 6.96 5.53

CJT16 7.64 6.16 4.43 7.64 5.75

CJT17 8.03 4.68 3.44 6.17 4.62

CJT18 6.29 4.47 3.46 6.11 4.44

CJT19 5.3 3.92 2.48 4.86 3.38

CJB1 12.03 3.35 3.96 4.82 5.32

CJB2 11.49 2.65 2.68 4.44 4.24

CJB3 10.64 3.17 2.42 4.33 3.97

CJB4 13.82 4.91 3.82 6.52 5.14

CJB5 10.81 3.21 2.29 4.74 4.03

CJB6 9.72 3.33 2.16 4.33 3.52

CJB7 10.05 3.56 2.48 4.67 3.78

CJB8 10.61 3.67 2.76 4.78 3.92

CJB9 10.51 3.59 2.57 4.96 3.87

CJB10 10.57 4.38 3 5.77 4.44

CJB11 9.71 4.85 3.31 6.5 5.07

CJB12 8.93 5.02 3.3 6.1 4.81

CJB13 6.51 4.65 3.36 6.13 4.72

CJB14 7.61 4.83 3.83 6.88 4.61

CJB15 8.14 5.44 4.08 7.09 5.17

Appendix 4: CT Scanned data set, after removal of unrequired teeth from

the Clean Data set (Appendix 2).

Tooth


1.1 23.51 10.42 5.86 12.81 7.15

1.4 16.95 12.31 7.17 13.01 8.13

1.6 24.12 11.00 6.43 12.97 8.44

2.1 18.47 8.70 5.34 9.75 6.91

2.2 20.07 10.60 6.26 14.11 8.86


107

2.3 22.70 12.33 7.36 14.15 9.74

2.6 25.55 13.99 8.07 15.33 10.15

2.8 23.61 14.93 8.68 17.09 11.04

2.13 24.20 11.20 7.39 13.57 9.93

3.1 7.60 7.74 5.70 8.20 6.33

3.2 11.83 10.95 8.21 12.23 9.39

3.3 11.50 5.73 4.17 6.92 5.18

3.5 26.28 11.26 7.22 14.08 10.29

3.6 29.96 12.40 7.36 15.34 10.45

3.7 25.54 12.91 9.91 14.84 10.39

3.8 30.60 12.85 7.87 15.83 11.23

3.12 12.63 8.50 5.57 9.42 7.08

3.13 24.48 12.61 8.02 15.58 10.99

3.15 35.02 14.75 9.27 17.95 12.21

4.2 9.95 8.03 6.29 9.59 7.14

4.3 18.52 7.22 5.66 8.98 7.15

4.4 15.83 6.77 5.21 8.28 6.57

4.9 31.65 12.01 7.80 14.15 10.56

4.11 26.57 14.46 9.25 17.55 12.19

4.13 28.81 11.22 7.48 13.43 10.42

4.14 31.55 15.09 9.18 17.79 12.49

4.17 28.79 13.96 8.49 16.90 11.45

4.18 29.71 12.50 8.26 16.36 11.19

4.19 17.12 11.92 7.53 12.57 9.70

5.3 18.99 7.85 6.13 9.49 8.69

5.4 20.52 8.50 6.53 10.72 8.80

5.6 25.63 13.58 9.07 16.29 12.08

5.9 38.07 15.44 9.68 16.66 13.16

5.10 29.30 15.73 10.33 19.24 13.73

5.11 30.65 17.18 10.22 20.09 13.94

5.12 28.00 14.31 9.76 16.89 12.29

5.13 30.87 12.33 8.77 15.40 11.45

5.14 24.60 13.43 7.98 16.06 10.56


108

5.17 32.79 16.59 10.01 17.79 13.03

5.18 34.39 13.63 8.80 16.53 12.00

5.23 29.97 12.66 8.03 14.81 11.39

5.24 32.90 13.56 8.49 14.99 11.56

5.27 21.11 11.39 7.44 12.85 10.05

6.1 10.00 4.89 4.24 6.17 4.99

6.2 13.21 5.64 4.41 7.25 6.27

6.5 26.26 10.53 7.30 12.93 10.36

6.12 30.81 12.96 8.93 17.15 12.89

7.5 34.26 16.09 11.03 20.97 16.24

7.6 12.30 10.57 8.77 11.66 9.63

7.9 29.98 13.85 8.40 15.80 12.26

7.14 16.57 10.08 7.23 12.01 9.65

7.15 29.41 12.96 8.34 14.61 11.36

8.2 23.66 10.32 8.02 12.80 11.93

8.4 11.88 7.20 6.00 8.77 7.46

8.5 10.98 9.38 9.19 11.17 9.75

8.7 24.95 10.24 7.82 12.75 11.07

8.9 11.09 7.13 6.10 8.77 7.35

8.10 35.73 15.66 10.67 18.53 14.72

9.2 15.52 7.61 6.36 9.62 8.75

9.3 11.55 11.37 9.34 13.01 11.43

9.4 12.83 11.57 9.86 13.14 11.86

9.5 18.66 10.94 8.63 12.50 10.53

9.8 33.89 16.38 10.68 19.25 14.88

9.9 12.65 11.16 9.93 12.90 11.26

9.10 14.46 11.88 10.91 13.61 11.72

9.17 35.47 14.33 9.75 17.11 14.27

9.19 37.97 16.56 11.04 19.37 15.65

9.20 17.04 11.71 8.92 13.40 11.27

9.22 36.35 14.35 8.76 15.21 12.58

9.25 22.52 13.68 8.74 14.88 11.57

9.26 13.75 8.84 9.62 8.37 8.39


109

9.27 38.16 13.80 9.58 16.00 14.09

9.29 14.28 11.55 9.55 12.67 10.98

10.2 18.39 9.23 8.30 10.19 10.44

10.4 14.43 7.39 5.75 8.00 7.50

10.6 22.41 12.53 10.60 15.01 13.22

10.7 14.21 7.13 5.90 9.33 8.40

10.8 35.66 14.54 10.07 16.70 14.82

10.10 19.64 9.86 7.64 10.94 10.02

10.11 10.60 9.18 8.23 10.96 10.10

10.12 17.98 10.70 8.94 12.56 10.63

10.13 10.49 10.52 8.59 11.94 11.44

10.19 21.14 9.39 7.03 10.85 9.69

10.22 34.12 14.22 11.25 17.14 15.11

10.23 20.52 9.03 6.87 10.17 9.21

10.24 29.89 11.29 8.90 14.75 13.47

10.25 31.56 15.37 10.97 16.47 14.20

10.26 25.03 10.47 8.02 13.22 11.69

10.27 15.05 12.27 11.10 15.07 14.02

10.28 18.15 8.05 6.45 9.58 8.49

10.31 12.92 11.17 9.43 14.16 12.19

11.2 29.22 12.79 9.85 15.99 14.49

11.4 16.66 8.06 6.50 9.21 8.60

11.8 22.43 10.89 8.82 12.62 11.66

11.9 13.11 10.92 9.38 12.99 11.81

11.10 33.08 12.26 10.03 15.94 14.56

11.11 33.78 14.07 11.01 15.84 15.03

11.14 16.25 11.78 10.42 13.82 12.10

11.19 22.20 12.81 11.42 14.93 13.96

11.20 18.08 8.77 7.36 9.74 9.24

11.21 17.65 9.78 7.70 10.48 9.75

11.22 18.50 12.31 11.31 15.58 14.20

11.25 29.66 11.43 9.37 13.33 12.58

12.1 29.02 10.82 8.56 13.23 12.45


110

12.3 14.20 6.13 4.48 6.91 5.86

12.7 31.06 12.69 9.35 15.79 15.10

12.9 17.36 10.45 8.94 12.25 11.87

12.10 33.53 12.55 10.74 15.97 14.86

12.11 38.54 14.29 12.15 18.21 16.78

12.14 19.11 9.33 8.09 11.20 10.37

12.15 34.33 16.34 13.16 18.19 16.79

12.16 18.53 8.34 6.77 9.22 8.90

12.19 19.05 10.34 8.70 12.99 11.82

12.21 11.22 10.32 9.05 11.21 10.01

13.2 18.62 7.25 5.90 8.76 8.21

13.7 32.42 15.21 12.38 18.11 17.28

13.8 30.71 13.32 11.32 16.49 15.95

13.10 32.09 14.19 11.90 16.20 15.54

13.11 28.23 12.19 10.20 14.36 13.53

13.14 29.85 13.20 12.14 16.40 16.77

13.18 26.82 12.52 10.97 15.12 14.97

13.19 14.35 10.94 9.65 13.22 12.52

13.21 13.37 11.73 10.45 13.25 12.47

13.23 27.96 11.76 10.94 13.75 13.86

13.25 19.54 7.74 6.38 9.24 8.71

14.1 12.24 6.29 5.29 7.31 6.93

14.7 23.20 9.69 7.24 11.25 10.84

14.11 19.58 9.99 7.83 11.24 10.67

14.13 32.92 13.49 11.04 16.60 16.34

14.14 19.37 11.73 9.75 14.22 13.35

14.16 29.05 13.94 11.61 16.45 15.77

14.19 16.50 9.50 8.01 10.93 10.54

14.20 18.18 9.48 7.75 10.75 10.30

15.2 23.45 11.00 9.57 12.88 12.94

15.4 18.16 7.50 6.56 9.84 9.23

15.5 27.70 12.44 10.76 15.25 15.35

15.6 20.44 9.45 8.43 11.02 11.00


111

15.7 16.07 12.09 10.25 13.95 13.55

15.8 16.52 8.85 7.84 10.43 9.74

15.9 18.74 9.34 7.84 10.39 10.52

15.11 19.82 8.73 7.46 9.55 9.64

15.12 19.97 10.56 8.74 11.82 11.53

15.16 34.40 13.75 12.47 17.82 17.46

Appendix 5: Measurements taken of all caniniform teeth separated out from

the initial cleaned data set (Appendix 2).

Tooth


1.1 23.51 10.42 5.86 12.81 7.15

1.6 24.12 11.00 6.43 12.97 8.44

2.1 18.47 8.70 5.34 9.75 6.91

2.2 20.07 10.60 6.26 14.11 8.86

2.3 22.70 12.33 7.36 14.15 9.74

2.6 25.55 13.99 8.07 15.33 10.15

2.8 23.61 14.93 8.68 17.09 11.04

2.13 24.20 11.20 7.39 13.57 9.93

3.3 11.50 5.73 4.17 6.92 5.18

3.5 26.28 11.26 7.22 14.08 10.29

3.6 29.96 12.40 7.36 15.34 10.45

3.7 25.54 12.91 9.91 14.84 10.39

3.8 30.60 12.85 7.87 15.83 11.23

3.13 24.48 12.61 8.02 15.58 10.99

3.15 35.02 14.75 9.27 17.95 12.21

4.3 18.52 7.22 5.66 8.98 7.15

4.4 15.83 6.77 5.21 8.28 6.57

4.9 31.65 12.01 7.80 14.15 10.56

4.11 26.57 14.46 9.25 17.55 12.19

4.13 28.81 11.22 7.48 13.43 10.42

4.14 31.55 15.09 9.18 17.79 12.49


112

4.17 28.79 13.96 8.49 16.90 11.45

4.18 29.71 12.50 8.26 16.36 11.19

5.3 18.99 7.85 6.13 9.49 8.69

5.4 20.52 8.50 6.53 10.72 8.80

5.6 25.63 13.58 9.07 16.29 12.08

5.9 38.07 15.44 9.68 16.66 13.16

5.10 29.30 15.73 10.33 19.24 13.73

5.11 30.65 17.18 10.22 20.09 13.94

5.12 28.00 14.31 9.76 16.89 12.29

5.13 30.87 12.33 8.77 15.40 11.45

5.14 24.60 13.43 7.98 16.06 10.56

5.17 32.79 16.59 10.01 17.79 13.03

5.18 34.39 13.63 8.80 16.53 12.00

5.23 29.97 12.66 8.03 14.81 11.39

5.24 32.90 13.56 8.49 14.99 11.56

5.27 21.11 11.39 7.44 12.85 10.05

6.1 10.00 4.89 4.24 6.17 4.99

6.2 13.21 5.64 4.41 7.25 6.27

6.5 26.26 10.53 7.30 12.93 10.36

6.12 30.81 12.96 8.93 17.15 12.89

7.5 34.26 16.09 11.03 20.97 16.24

7.9 29.98 13.85 8.40 15.80 12.26

7.14 16.57 10.08 7.23 12.01 9.65

7.15 29.41 12.96 8.34 14.61 11.36

8.2 23.66 10.32 8.02 12.80 11.93

8.4 11.88 7.20 6.00 8.77 7.46

8.5 10.98 9.38 9.19 11.17 9.75

8.7 24.95 10.24 7.82 12.75 11.07

8.10 35.73 15.66 10.67 18.53 14.72

9.2 15.52 7.61 6.36 9.62 8.75

9.5 18.66 10.94 8.63 12.50 10.53

9.8 33.89 16.38 10.68 19.25 14.88

9.17 35.47 14.33 9.75 17.11 14.27


113

9.19 37.97 16.56 11.04 19.37 15.65

9.22 36.35 14.35 8.76 15.21 12.58

9.25 22.52 13.68 8.74 14.88 11.57

9.26 13.75 8.84 9.62 8.37 8.39

9.27 38.16 13.80 9.58 16.00 14.09

10.2 18.39 9.23 8.30 10.19 10.44

10.4 14.43 7.39 5.75 8.00 7.50

10.6 22.41 12.53 10.60 15.01 13.22

10.7 14.21 7.13 5.90 9.33 8.40

10.8 35.66 14.54 10.07 16.70 14.82

10.10 19.64 9.86 7.64 10.94 10.02

10.12 17.98 10.70 8.94 12.56 10.63

10.19 21.14 9.39 7.03 10.85 9.69

10.22 34.12 14.22 11.25 17.14 15.11

10.23 20.52 9.03 6.87 10.17 9.21

10.24 29.89 11.29 8.90 14.75 13.47

10.25 31.56 15.37 10.97 16.47 14.20

10.26 25.03 10.47 8.02 13.22 11.69

10.28 18.15 8.05 6.45 9.58 8.49

11.2 29.22 12.79 9.85 15.99 14.49

11.4 16.66 8.06 6.50 9.21 8.60

11.8 22.43 10.89 8.82 12.62 11.66

11.10 33.08 12.26 10.03 15.94 14.56

11.11 33.78 14.07 11.01 15.84 15.03

11.19 22.20 12.81 11.42 14.93 13.96

11.20 18.08 8.77 7.36 9.74 9.24

11.21 17.65 9.78 7.70 10.48 9.75

11.25 29.66 11.43 9.37 13.33 12.58

12.1 29.02 10.82 8.56 13.23 12.45

12.3 14.20 6.13 4.48 6.91 5.86

12.7 31.06 12.69 9.35 15.79 15.10

12.9 17.36 10.45 8.94 12.25 11.87

12.10 33.53 12.55 10.74 15.97 14.86


114

12.11 38.54 14.29 12.15 18.21 16.78

12.14 19.11 9.33 8.09 11.20 10.37

12.15 34.33 16.34 13.16 18.19 16.79

12.16 18.53 8.34 6.77 9.22 8.90

12.19 19.05 10.34 8.70 12.99 11.82

13.2 18.62 7.25 5.90 8.76 8.21

13.7 32.42 15.21 12.38 18.11 17.28

13.8 30.71 13.32 11.32 16.49 15.95

13.10 32.09 14.19 11.90 16.20 15.54

13.11 28.23 12.19 10.20 14.36 13.53

13.14 29.85 13.20 12.14 16.40 16.77

13.18 26.82 12.52 10.97 15.12 14.97

13.23 27.96 11.76 10.94 13.75 13.86

13.25 19.54 7.74 6.38 9.24 8.71

14.1 12.24 6.29 5.29 7.31 6.93

14.7 23.20 9.69 7.24 11.25 10.84

14.11 19.58 9.99 7.83 11.24 10.67

14.13 32.92 13.49 11.04 16.60 16.34

14.14 19.37 11.73 9.75 14.22 13.35

14.16 29.05 13.94 11.61 16.45 15.77

14.19 16.50 9.50 8.01 10.93 10.54

14.20 18.18 9.48 7.75 10.75 10.30

15.2 23.45 11.00 9.57 12.88 12.94

15.4 18.16 7.50 6.56 9.84 9.23

15.5 27.70 12.44 10.76 15.25 15.35

15.6 20.44 9.45 8.43 11.02 11.00

15.7 16.07 12.09 10.25 13.95 13.55

15.8 16.52 8.85 7.84 10.43 9.74

15.9 18.74 9.34 7.84 10.39 10.52

15.11 19.82 8.73 7.46 9.55 9.64

15.12 19.97 10.56 8.74 11.82 11.53

15.16 34.40 13.75 12.47 17.82 17.46


115

Appendix 6: Measurements taken of all molariform teeth separated out

from the initial cleaned data set (Appendix 2).

Tooth


1.4 16.95 12.31 7.17 13.01 8.13

3.1 7.60 7.74 5.70 8.20 6.33

3.2 11.83 10.95 8.21 12.23 9.39

3.12 12.63 8.50 5.57 9.42 7.08

4.2 9.95 8.03 6.29 9.59 7.14

4.19 17.12 11.92 7.53 12.57 9.70

7.6 12.30 10.57 8.77 11.66 9.63

8.9 11.09 7.13 6.10 8.77 7.35

9.3 11.55 11.37 9.34 13.01 11.43

9.4 12.83 11.57 9.86 13.14 11.86

9.9 12.65 11.16 9.93 12.90 11.26

9.10 14.46 11.88 10.91 13.61 11.72

9.20 17.04 11.71 8.92 13.40 11.27

9.29 14.28 11.55 9.55 12.67 10.98

10.11 10.60 9.18 8.23 10.96 10.10

10.13 10.49 10.52 8.59 11.94 11.44

10.27 15.05 12.27 11.10 15.07 14.02

10.31 12.92 11.17 9.43 14.16 12.19

11.9 13.11 10.92 9.38 12.99 11.81

11.14 16.25 11.78 10.42 13.82 12.10

11.22 18.50 12.31 11.31 15.58 14.20

12.21 11.22 10.32 9.05 11.21 10.01

13.19 14.35 10.94 9.65 13.22 12.52

13.21 13.37 11.73 10.45 13.25 12.47

9.0 Supplementary Figures:


117


118


119


120


121


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126


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pliocene crocodilians of chinchilla: identification using dental morphometrics · 2018. 6. 18. ·...

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