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FIELDIANA gedi.oqi ubbab*

Geology

Published by Field Museum of Natural History
New Series, No. 14

THE MAMMALIAN FAUNA OF
MADURA CAVE, WESTERN AUSTRALIA
PART VI: MACROPODIDAE: POTOROINAE

ERNEST L. LUNDELIUS, JR.
WILLIAM D. TURNBULL

April 30, 1984
Publication 1354

THE MAMMALIAN FAUNA OF

MADURA CAVE, WESTERN AUSTRALIA

PART VI: MACROPODIDAE: POTOROINAE

TRENCH GOES ON FOR CA.20

MAP OF
MADURA CAVE

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DARK RED CLAV SILT

YELLOW - ORANGE GRITTY SAND

(TxH43) 37,880 ± 3. 520 YBP

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TIGHT SILT AND CLAV SEAM3

RED-ORANGE SILTY CLAV

y (Tx 1145) 15.600 t 250 YBP

LIMIT
EXCAVATION

4 WHITE SANDY SILT

5 RED CLAY

Left, bottom, Map of Madura Cave showing the locations of the trenches. Large arrows
point to cross sections of three of the five trenches in the northern tunnel; left, top,
Trench 5, Section B-B'; right, top, Trench 4, Section A-A'; and right, bottom, Trench 3,
Section C-C. (All illustrations are taken from Madura Cave, Part I.)

FIELDIANA
Geology

Published by Field Museum of Natural History

New Series, No. 14

THE MAMMALIAN FAUNA OF
MADURA CAVE, WESTERN AUSTRALIA
PART VI: MACROPODIDAE: POTOROINAE

ERNEST L. LUNDELIUS, JR.

Professor of Geological Sciences
University of Texas at Austin

Research Associate

Department of Geology

Field Museum of Natural History

WILLIAM D. TURNBULL

Curator, Fossil Mammals

Department of Geology

Field Museum of Natural History

Committee on Evolutionary Biology
University of Chicago

Research Associate
Texas Memorial Museum

Accepted for publication July 27, 1983
April 30, 1984
Publication 1354

© 1984 Field Museum of Natural History

Library of Congress Catalog Card No.: 72-97564

ISSN: 0096-2651

PRINTED IN THE UNITED STATES OF AMERICA

CONTENTS

List of Illustrations viii

List of Tables ix

Abstract 1

Introduction 1

Potorous Desmarest, 1804 2

Potorous platyops (Gould), 1844 2

Material 2

Comparative Material 2

Description 2

Discussion 12

Potorous lncertae Sedis 13

Material 13

Description 14

Caloprymnus Thomas, 1888 14

Caloprymnus campestris (Gould), 1843 14

Material 14

Comparative Material 20

Description 20

Discussion 29

Caloprymnus lncertae Sedis 30

Material 30

Bettongia 30

Bettongia lesueur (Quoy & Gaimard, 1824) 31

Material 31

Comparative Material 36

Description 36

Discussion 46

Bettongia penicillata 48

Material 48

Comparative Material 48

Description 49

Discussion 58

Bettongia Species Indet 58

Material 58

Conclusions 60

Acknowledgments 61

Literature Cited 61

LIST OF ILLUSTRATIONS

Map of Madura Cave and cross sections of trenches Frontispiece

1. Potorous platyops comparative materials 4, 5

2. Potorous platyops from Madura Cave 6,7

3. Potorous platyops from Madura Cave 8

4. Caloprymnus campestris from Madura Cave 21

5. Caloprymnus campestris from Madura Cave 23

6. Bivariate graphs of dP4s of Bettongia from Madura Cave 34, 35

7. Bettongia lesueur from Madura Cave, surface 38, 39

8. Bettongia lesueur from Madura Cave, Units 1 and 2 43

9. Bettongia lesueur from Madura Cave, surface and Unit 1 44

10. Bettongia lesueur from Madura Cave, surface and Unit 1 45

11. Bettongia penicillata, Recent 50, 51

12. Bettongia penicillata, Recent 52

13. Bettongia penicillata from Madura Cave, surface and Unit 1 53

14. Bettongia penicillata from Madura Cave, Unit 1 55

15. Bivariate graphs of P3s and P3s of Bettongia penicillata from Madura Cave and

Nannup Cave 57

LIST OF TABLES

1. Measurements of upper dentitions of Potorous platyops from Madura Cave 3

2. Measurements of lower teeth of Potorous platyops from Madura Cave 9

3. Measurements of upper dentitions of various samples of fossil and Recent

Potorous platyops 10

4. Measurements of lower dentitions of fossil and Recent samples of Potorous

platyops 11

5. Position of anterior end of masseteric canal in specimens of Caloprymnus

campestris of various dental ages 24

6. Measurements of upper dentitions of Caloprymnus campestris from Unit 1 of

Madura Cave 25

7. Statistical data on upper dentitions of Caloprymnus campestris from Units 2

to 4-5 of Madura Cave 26

8. Statistical data on upper dentitions of Caloprymnus campestris from Unit 7 of

Madura Cave 27

9. Measurements of lower dentitions of Caloprymnus campestris from Unit 1 of

Madura Cave 28

10. Statistical data on lower dentitions of Caloprymnus campestris from Units 2

to 4-5 of Madura Cave 29

11. Statistical data on lower dentitions of Caloprymnus campestris from Unit 7 of

Madura Cave 30

12. Dental and cranial characteristics used to distinguish the Madura Cave

Bettongia lesueur from Bettongia penicillata 32

13. Comparison of cranial, mandibular, and dental measurements of Bettongia

lesueur from Madura Cave with those of Bettongia penicillata anhydra

Finlayson 37

14. Statistical data on upper dentitions of Bettongia lesueur from Madura Cave 37

15. Statistical data on lower dentitions of Bettongia lesueur from Madura Cave 40

16. Statistical data on upper dentitions of Bettongia lesueur from Lake Victoria 40

17. Statistical data on lower dentitions of Bettongia lesueur from Lake Victoria 41

18. Statistical data on upper dentitions of Bettongia lesueur from Nannup Cave 41

19. Statistical data on lower dentitions of Bettongia lesueur from Nannup Cave 42

20. Statistical data on upper dentitions of Bettongia penicillata from Madura Cave

compared with those of other fossil and modern samples 54

21. Statistical data on lower dentitions of fossil and living samples of Bettongia

penicillata 56

ABSTRACT

The potoroines represented in the Madura Cave deposits are Potorous platyops,
Caloprymnus campestris, Bettongia lesueur, and Bettongia penicillata.

Potorous platyops, found in Units 1 through 4-5, is not significantly different
from Pleistocene, Holocene, or modern specimens from other parts of Australia.

Caloprymnus campestris, which occurs in all units, shows no significant mor-
phological or size change through the sequence and is smaller than the samples
from the Pleistocene of Lake Menindee and from the known Recent material,
with no overlap in size.

Bettongia is represented by B. lesueur, which is present throughout the strati-
graphic sequence, and B. penicillata, which is absent only from Unit 7. The
Madura Cave B. lesueur is smaller in size than other Pleistocene and modern
samples, but B. penicillata shows no differences.

Today, four species of potoroines are known only from an area near Albany,
Western Australia. The presence of Potorous platyops suggests that more effective
moisture existed in the vicinity of Madura Cave when these sediments were
deposited than at present. The association of now allopatric species such as
Potorous platyops and Caloprymnus campestris indicates a more equable climate in
the past.

INTRODUCTION

This section of the Madura Cave report covers the rat-kangaroos, Potoroinae,
and continues the systematic treatment of the marsupials (Lundelius & Turn-
bull, 1973, 1975, 1978, 1981, 1982). Measurements, abbreviations, and dental
terminology are either those in standard use or those outlined in Parts I through
V. Measurements in all tables and graphs are given in millimeters. Headings for
the statistical tables are as follows: N, sample size; OR, observed range; M,
mean; SE, standard error; SD, standard deviation; CV, coefficient of variation.
Abbreviations for institutions are: BMNH, British Museum (Natural History);
FMNH, Field Museum of Natural History (Recent mammal); PM, Field Museum
of Natural History (fossil mammal); SAM, South Australian Museum; TMM,
Texas Memorial Museum; WAM, Western Australian Museum.

2 FIELDIANA: GEOLOGY

Potorous Desmarest, 1804

Potorous platyops (Gould), 1844

Hypsiprymnus platyops Gould, 1844

Potorous platyops Thomas, 1888

Potorous morgani Finlayson, 1938

Potoroops platyops (Tate), 1948

Material

Trench 4, Unit 1

TMM 41106-576, right M, (fig. 3B)

TMM 41106-600, left M1 or M2 (fig. 2C)

TMM 41106-609, right M3 (fig. 3F)

TMM 41106-619, right P4 fragment (fig. 2B)
Trench 4, Unit 2, Level 1

PM 34418, left M1 or M2
Trench 4, Unit 2, Level 2

PM 34419, right maxillary fragment with M3 (fig. 2E)

PM 34420, right M, (fig. 3C)

PM 34421, left M1 or M2

PM 34422, left M2 (fig. 3E)
Trench 4, Units 4-5

PM 34423, left M1 or M2 (fig. 2D)

TMM 41106-647, juvenile ramus with I, P„, alveoli of P3, dP4, M, (fig. 3A)

PM 34424, right M,

PM 34425, right M, (fig. 3D)

WAM 75.1.130, left M1 or M2
Trench 5, Unit 4

PM 34426, right M1 or M2
Trench 5, Unit 6

WAM 75.1.131, left M' or M2

TMM 41106-632, right dP4 (fig. 2A)

Comparative Material

Potorous platyops

BMNH 46.4.25.11, P. platyops type

BMNH 53.10.22.19

SAM PI 68, P. morgani cotype
Hastings' Cave

PM 6313 (fig. 1D,E), 6314-6318, 6695, 7073-7078, 36844-36848, 36852, 36869-36871
Wedge's Cave

PM 5598-5602, 36843
Webb's Cave

PM 4355 (fig. 1A-C)

Description

Upper Dentition.— The dP4 (TMM 41106-632; fig. 2A) is rooted, but unworn.
Its nearly equally spaced cusps and primary lophs are low and sharp. The tooth
is rectangular in outline with the anterior ridge of the paracone extending
forward as a short blade. As compared with PM 6313 (fig. 1D,E) and PM 6315,
the Madura Cave specimen is more elongate with more pronounced constric-
tions both labially and lingually between the protoloph and metaloph. A low

LUNDELIUS & TURNBULL: MADURA CAVE J

Table 1. Measurements of upper dentitions of Potorous platyops from Madura Cave.

Anterior

Posterior

Teeth

Specimen No.

Length

width

width

dP4

TMM

41106-632

2.85

2.13

2.20

P4

TMM

41106-619

> 3.5

> 1.7

1.62

M1 or M2

TMM

41106-600

3.42

2.96

2.74

PM

34423

3.34

3.04

2.89

PM

34424

3.45

2.96

2.81

PM

34421

3.08

3.00

2.81

PM

34426

3.42

2.04

2.70

PM

34418

3.12

2.74

WAM

75.1.130

3.04

2.74

2.66

WAM

75.1.131

3.34

2.81

2.74

M3

PM

34419

3.12

2.74

2.25

angular ridge extends posteriorly from the apex of the protocone parallel to
the labial side of the tooth. This ridge almost reaches the median valley, then
swings abruptly lingually and disappears. A similar low ridge runs anteriorly
from the apex of the metacone along the labial edge of the tooth and disappears
near the valley without turning lingually. Low, blunt, and more massive ridges
connect the protocone and hypocone, meeting at a nearly right-angled V where
they cross the bottom of the median valley. The protoloph forms a broadly
open saddle, with the paracone making the greater contribution. The metaloph
is nearly straight and horizontal. It is weakly notched near the hypocone. The
metacone causes the labial half of the metaloph to bulge posteriorly. The post-
cingulum is a heavy ridge, especially near the hypocone. It is U-shaped in
posterior view and it and the metaloph delimit a shallow, crescentic posterior
cingular basin.

The P4 (TMM 41106-619) lacks the anterior end. It is unworn, lacks roots, and
probably was unerupted (fig. 2B). The long axis of the tooth is concave labially.
This appears to be a variable character since PM 4355 (fig. 1A-C) from Webb's
Cave (about 55 miles [89 km] east of Madura Cave) is nearly straight, while PM
6313 (fig. 1D-E) and PM 6315 from Hastings' Cave show considerable difference
in their curvature. Of all the specimens here compared, PM 6315 shows the
most curvature. The anterior cusp is higher than the others. The two specimens
from the Nullarbor Plain differ from the two Hastings' Cave specimens in
having less pronounced vertical ridges on the labial side of the two anterior
cusps.

It is difficult to distinguish between isolated M's and M^. The M1 in most of
the available comparative material has a more pronounced anterolabial projec-
tion on the procingulum than does the M2. However, the Madura Cave speci-
mens do not fall into two distinct groups on the basis of this character or on
the basis of size. Both M1 and M2 probably are represented. There is much more
similarity between M1 and M2 in the Madura Cave sample than in the Hastings'
Cave and Wedge's Cave samples.

The M1 or M2 (as represented by TMM 41106-600; fig. 2C) is slightly longer
than wide. The posterior part of the tooth is narrower than the anterior part,
both in the spacing of the two posterior cusps with respect to the anterior ones
and in the overall width of the tooth. Its protocone and hypocone are relatively

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Fig. 2. A-E, Potorous platyops from Madura Cave. A, TMM 41 106-632. Right dP4 shown
in labial (top), crown, and lingual views. B, TMM 41106-619. Right P4 shown in labial
(left), crown, and lingual views. C, TMM 41106-600. Left M1 or M2 shown in labial (top),
crown, lingual, posterior (left), and anterior views. D, PM 34423. Left M1 or M2 shown
in labial (top), crown, and lingual views. E, PM 34419. Right M' in maxillary fragment
shown in labial (top), crown, and lingual views. F,G, lncertae sedis, probably P. platyops,
Madura cave. F, TMM 41106-682. Right premaxillary with I'2, alveolus for I\ shown in
labial (left), crown, and lingual views. G, PM 34510. Left premaxillary with I1, alveoli for
I23, and C, shown in crown and labial views.

LUNDELIUS & TURNBULL: MADURA CAVE 9

Table 2. Measurements of lower dentitions of Potorous platyops from Madura Cave.

Anterior

Posterior

Teeth

Specimen No.

Length

width

width

P.

TMM 41106-647

3.80

1.34

1.14

M,

PM 34420

3.01

2.13

2.20

PM 34425

2.96

1.98

2.05

TMM 41106-576

3.34

2.37

3.43

M2

PM 34422

3.27

2.74

2.58

M5

TMM 41106-609

3.08

2.37

2.09

high, blunt cusps. The metaloph is nearly straight, but the protoloph is slightly
bowed and lies slightly oblique to the long axis of the tooth, joining the pro-
tocone at its anterior margin. In profile the crests of the lophs are gently con-
cave between the cusps. A low but continuous crest connects the protocone to
the hypocone. Crests from the paracone and metacone nearly meet at the bases
of these cusps. These crests and the lophs define a nearly square, almost fea-
tureless central basin. Pro- and postcingula are both present. The procingulum
is connected to the protocone by a weak crest and to the paracone by a some-
what stronger crest, forming a broad asymmetrical basin. Similarly developed
distinct crests extend downward and posteriorly from the apices of the hypo-
cone and metacone, meeting about halfway to the base of the crown to form a
symmetrical posterior cingular basin (fig. 2C). The other three M's or M2s (PM
34423, PM 34424, WAM 75.1.131) show minor variations in the condition of
the notch that separates the crests from the paracone and metacone. PM 34423
(fig. 2D) and PM 34424 have a narrow U-shaped notch, while WAM 75.1.131
has a narrow cleft. Wear has made it impossible to determine the condition of
this feature in TMM 41106-600. Specimen PM 34423 differs from the other three
M's or M2s in possessing a protoloph that in profile has an open V shape rather
than a U shape. This character may be affected by wear. The posterior cingular
basins of PM 34423 and WAM 75.1.131 are narrower than those of TMM 41 106-
600 and PM 34424.

The M3 (PM 34419) differs from the M's and M2s in having the posterior part
of the tooth distinctly narrower than the anterior part. It has a square, feature-
less central basin, a broad, asymmetrical procingular basin, and a well-defined
oval posterior cingular basin (fig. 2E).

No M4s have been recognized.

Loicer Dentition.— The mandible (TMM 41106-647; fig. 3A) is shallow. The
lower incisor is narrow and procumbent, as is normal in Potorous, and is curved
inward and upward throughout its length. It shows slight wear at the tip and
on the ventromedial edge. The ventrolateral surface of the tooth is convex and
enamel covered throughout its length. In cross section the dorsolateral surface
is generally convex in its dorsal half, with a narrow groove on its dorsal edge.
A broad trough extends the length of the tooth on the ventromedial edge.
Enamel is present only on the distal end of the medial convexity. There do not
appear to be any differences between the Madura tooth, the Western Australian
specimens of P. platyops, and the Kangaroo Island P. morgani which cannot be
accounted for by individual variation.

All the cheek teeth except the unerupted P4 have been lost from the man-
dible. The crown of the P4 (fig. 3A) is well formed but the tooth lacks roots.

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12 FIELDIANA: GEOLOGY

The long axis of the tooth is straight, in contrast to the situation in P4. The
tooth is compressed and divided on both sides by two broad grooves into three
well-defined cusps. The posterior cusp has an indistinct groove that forms a
low notch on the crest. The posterior end of the tooth is very flat. The length
of the P4 is 3.80 mm, which is just below the size range of P4s of a sample of P.
platyops from Hastings' Cave, Western Australia (table 3). The type (BMNH
46.4.25.11, female) and one additional Recent specimen from Western Australia
(BMNH 53.10.22.19, male) measure 3.8 and 4.1 mm, respectively. The length of
the P4 of one of the cotypes of Potorous morgani is 3.9 mm (Finlayson, 1938, p.
134).* An examination of Tables 2 and 4 shows that the lengths of the P4s of
all available samples of P. platyops, both fossil and Recent, overlap the sample
from Hastings' Cave.

The M, (as represented by TMM 41106-576; fig. 3B) is slightly narrower across
the protolophid than across the metalophid (the protolophid and metalophid
are more nearly equal in PM 34420). The central basin is large, longer on the
lingual than the labial side, with no significant irregularities. Labial and lingual
ridges connect the protoconid and hypoconid and the metaconid and entoconid
along the margins of the tooth. In TMM 41106-576 these ridges are not fused
where they meet; in PM 34420 and PM 34425 the ridges are fused, with no sign
of a cleft. The pro- and postcingula are distinct and are connected with the
apices of their associated cusps. The anterior and posterior cingular basins are
asymmetrical with their centers labial to the midline of the tooth. In TMM
41106-576 the posterior cingular basin is subdivided by a low bulge on the floor
of the basin; this feature is lacking in the other two M,s, and PM 34420 has a
more symmetrical posterior cingular basin.

The only M2 (PM 34422; fig. 3E) is rectangular with the metalophid slightly
narrower than the protolophid. The ridges from the major cusps join along the
sides of the central basin without fusing, as in TMM 41106-576. The posterior
cingular basin is smaller than that in M1 and is more lingually situated.

The only M3 (TMM 41106-609; fig. 3F) is moderately worn. It is slightly wider
across the protolophid than across the hypolophid, its central basin is nearly
square, and there is a trace of a transverse valley across the middle, as in MP
Wear has reduced the ridges labial and lingual to the central basin, but the
clefts are still in evidence where the ridges meet but remain unfused. The
procingulum is expanded anteriorly in its middle, so that the anterior cingular
basin is more oval than in the M! but lacks the prominent anterolabial cuspule
seen in PM 7074 from Hastings' Cave. Wear has truncated the postcingulum to
the point where the basin is nearly obliterated. Only a small, deep pit remains
almost directly behind the saddle of the hypolophid. The thick enamel tips of
the protoconid and hypoconid have both been breached by wear so that small
circles of dentine are exposed.

Discussion

The dentition of Potorous platyops differs from that of other species of Potorous
primarily in its smaller size. The P4 of P. platyops differs from that of P. tridac-

* The measurement is listed as P4. This, as well as the references to M1 through M4, is
surely a typographical error, as the paragraph on p. 134 refers to the mandible, and the
length of P4 in the table on p. 139 is 5.0 mm.

LUNDELIUS & TURNBULL: MADURA CAVE 13

tylus (including P. gilberti) in having a shallower notch between the second and
third cusps and the absence of a flared base. Three specimens of Potorous apicalis
from Tasmania (FM 98902, FM 57805, FM 98789) differ from P. platyops in
having three or four small cusps between the two large end ones rather than
the two characteristic of P. platyops.

This is the fifth reported occurrence of P. platyops in cave deposits on the
Nullarbor Plain. The species was reported earlier from Webb's Cave (55 miles
[89 km] east of Madura) on Mundrabilla Station (Lundelius, 1963) and more
recently (Butler & Merrilees, 1971) from caves near Madura, Eucla, and Koon-
alda. The material from the older deposits of Madura Cave shows that the
species occupied the Nullarbor Plain during the latter part of the Pleistocene.

Lundelius (1963) suggested that it is likely that P. platyops from Western
Australia, P. morgani from Kangaroo Island, and the specimen from Webb's Cave
are representatives of a single species, P. platyops. The extensive overlap in
dental dimensions of the various samples of P. platyops (including the cotypes
of P. morgani) supports this view. Finlayson (1938) listed six characters as criteria
which separated the type of P. platyops from P. morgani. Four of these are minor
differences in proportions which Finlayson himself pointed out might break
down if larger series of specimens were examined. The other two, the size and
shape of the first upper incisor and the degree of specialization of the lower
incisor, seem to us to be equally subject to geographic variation, although little
material from Madura Cave bears on this.

Potorous platyops formerly was widespread in coastal southern Australia. In
addition to the Madura Cave Pleistocene and Holocene fossils reported here,
and to the Webb's Cave and Kangaroo Island occurrences discussed above, P.
platyops is also known from Pleistocene deposits in Victoria Cave, southeastern
Southern Australia (Smith, 1971), and from the following Holocene deposits:
Dongara-Hill River area on the west coast of Western Australia (Lundelius, 1957,
1960), Bremer Bay on the south coast of Western Australia (Butler & Merrilees,
1971), several caves on the Nullarbor Plain (Lundelius, 1957, 1963; Butler &
Merrilees, 1971), and from aboriginal sites on the lower Murray River (Wake-
field, 1964). Several of these occurrences indicate a late Holocene age for the
specimens: the Webb's Cave specimen was collected from the surface (Lunde-
lius, 1963); the Wedge's Cave material from the west coast is known to be
younger than 3750 BP; the Bremer Bay material has Cu dates indicating an age
between 1190 and 620 BP; the material from caves near Madura, Eucla, and
Koonalda all appears to be of late Holocene age (Butler & Merrilees, 1971); and
the species is represented in the lower Murray River sites with C14 dates of
1800 BP. Evidently P. platyops, like Cercartetus concinnus (Lundelius & Turnbull,
1982), persisted until quite recently, perhaps as relict populations in locally
favorable areas. The restriction and fragmentation of the mainland population
probably resulted from the post-Pleistocene trend toward aridity.

Potorous Incertae Sedis

Material

Trench 3, Unit 2, Level ?

TMM 41106-682, premaxillary fragment with I1 2, alveolus of I3 (fig. 2F)

14 FIELDIANA: GEOLOGY

Trench 4, Unit 2, Level 1

PM 34510, premaxillary fragment with I1, alveoli for I23, C (fig. 2G)

PM 36877, 36878, two I's
Trench 5, Unit 6

PM 38673, I"

Description

These specimens probably belong to Potorous platyops. The available compar-
ative specimens have no upper incisors preserved; hence, the uncertainty. The
Ps thought to belong to Potorous are uniformly curved, nearly prismatic teeth
with only a slight taper throughout their length. If not too worn, they may
have a faint groove on their posterolabial edges. The I's lack enamel on their
posterior side except at the extreme labial edge where it folds around from the
labial side. The I2 has an oval crown in occlusal view. The occlusal surface is
flat. The crown is expanded forward so that more of it lies anterior to the root
than posterior to it. The alveolus for the canine lies largely within the premax-
illary, and the anterior edge of the incisive foramen is located opposite the
alveolus for P.

Caloprymnus Thomas, 1888

Caloprymnus campestris (Gould), 1843
Bettongia campestris Gould, 1843

Material

Trench 3, Unit 2, Level 1

TMM 41106-160, left ramus with P3, dP4, M,.2, alveoli for M3.4 (fig. 5A)

TMM 41106-161, right ramus with P3, dP4, M,_3, alveoli for M4 (fig. 5B)

TMM 41106-162, right ramus with shattered roots of all teeth

TMM 41106-163, left ramus with I, P3_4 removed, M,_3 (fig. 5C)

TMM 41106-164, left ramus fragment with P4, M, (fig. 5D)

TMM 41106-165, right ramus fragment with worn M2

TMM 41106-166, left ramus with P4, M2_4 (fig. 5E)

TMM 41106-167, left maxillary fragment with P3, dP4, P4 in crypt, M12, alveoli for M3

(fig. 4C)
TMM 41106-304, right maxillary fragment with M12
TMM 41106-305, right maxillary fragment with M1-3
TMM 41106-306, left maxillary fragment with M12
TMM 41106-307, left maxillary fragment with worn M12 (fig. 4F)
TMM 41106-308, right maxillary fragment with P4 broken, M1
TMM 41106-309, right maxillary fragment with worn M3 4 (fig. 4G)
TMM 41106-310, right maxillary fragment with P4-M' (fig. 4D)
TMM 41106-313, left M2 or M3
TMM 41106-2829, left M2, M1, or M3
TMM 41106-2830, -5099, two right M3s or M2s
TMM 41106-5100, M2 or M, or M3
TMM 41106-5101, right M2
TMM 41106-5102, right M,
TMM 41106-5103, left M2, M1, or M3
TMM 41106-5104, right M1 or M2
TMM 41106-5105, left M' or M2
TMM 41106-5106, right M2 or M3
TMM 41106-5107, right M' or M3
TMM 41106-5108, left dP4

LUNDELIUS & TURNBULL: MADURA CAVE 15

TMM 41106-5120, left P4

TMM 41 106-51 22, left P4

TMM 41106-5123, left P4

TMM 41106-5124, posterior % right P4

TMM 41106-5125, -5126, two right dP4s

TMM 41106-5127, badly worn lower molar

TMM 41106-5128, right M2 or M'

TMM 41106-5129, right dP4

WAM 75.1.145, left M1 or M2

PM 34427, right maxillary fragment with M1'2

PM 34428, left maxillary fragment with P3, dP4

PM 34429, left maxillary fragment with M1"2

PM 34430, right M, or M2

PM 34431, 34432, two left dP4s

PM 34433, right maxillary fragment with P4 in crypt

PM 34435, left ramus with all teeth shattered

PM 34461, right ramus fragment with I,, P4

PM 34462, left ramus fragment with P4-M3

PM 34463, edentulous right ramus

PM 34464, left ramus with M3

PM 34465, right ramus fragment with I,, P3

PM 34466, right ramus fragment with I,, M2_3, crypt for P4

PM 34467, right ramus fragment with M2_3

PM 34468, right ramus fragment with M2„3 or M3_4

PM 34470, edentulous maxillary fragment with alveoli for dP4-M', crypt for P4
Trench 3, Unit 2, Level 2

PM 34436, right maxillary fragment with alveoli for P\ dP4, crypt for P4
Trench 3, Unit 2, Level 4

WAM 75.1.132, left M,

WAM 75.1.133, left M2 or M3

PM 34437, 34438, two right P3s

PM 34439, posterior % right P3

PM 34440, right P3

PM 34441, 34442, two right dP4s

PM 34443, right P4

PM 34444, left P4

PM 34445, right dP4

PM 34446, right M,

PM 34447, left M,

PM 34448, right M1

PM 34449, right M2

PM 34450, very worn left M2

PM 34451, right M'

PM 34452, left M,

PM 34453, right M2

PM 34454, right M,

PM 34455, left M2 or M3

PM 34456, right M2 or M3

PM 34457, 34458, two left M's or M^

PM 34459, right M2, M', or M3

PM 34460, right M2 or M,
Trench 3, Unit 3, Level ?

TMM 41106-34, maxillary with P3, P4 removed from crypt, M1-2 (fig. 4B)

TMM 41106-35, left dP4

TMM 41106-36, left ramus fragment with M,, alveolus for M2, part of crypt for P4

TMM 41106-311, right ramus fragment with M3.4

TMM 41106-312, left ramus fragment with M2 broken, M3

TMM 41106-344, left ramus with I, P3, dP4, P4 in crypt, M,_2

TMM 41106-345, left ramus with dP4, M,_2, unerupted P4 and M3.„

TMM 41106-346, left ramus with P3, dP4

TMM 41106-348, edentulous left ramus fragment

16 FIELDIANA: GEOLOGY

TMM 41106-349, left ramus fragment with dP4/ M2.3

TMM 41106-351, right ramus with M,

TMM 41106-352, right ramus with I, dP4, M,

TMM 41106-353, right ramus with M2

TMM 41106-354, left ramus with dP4-M,

TMM 41106-357, right maxillary with P3, dP4, M1"2

TMM 41106-358, right maxillary with M2"3 (fig. 4E)

TMM 41106-359, left maxillary with P3, dP4, M1"2

WAM 75.1.134, left ramus with I, broken, alveoli for P3, dP4, M,_2

WAM 75.1.146, right ramus fragment with dP4, P4 in crypt, alveoli for I, P3, M,_2

WAM 75.1.147, right ramus fragment with P4 exposed in crypt, alveoli for P3-M,

PM 34471, 34472, two left dP4s

PM 34473, right M2 or M3

PM 34474, right M2

PM 34475, edentulous right ramus fragment

PM 34476, juvenile edentulous left ramus

PM 34477, left maxillary fragment with very worn M12

PM 34478, right ramus fragment with I,, P3

PM 34479, edentulous left ramus

PM 34480, edentulous right ramus

PM 34481, right ramus with very worn M3.4

PM 34482, left ramus with I,, probably P4 in crypt

PM 34483, juvenile edentulous left ramus

PM 34484, maxillary fragment with P4 (inverted in crypt)
Trench 4, Unit 1, top 1 foot

TMM 41 106-497, left M2

TMM 41106-540, left maxillary fragment with M12, part of M3

TMM 41106-541, left ramus with P4-M4

TMM 41106-570, left dP4

TMM 41106-577, left M1

TMM 41106-5131, left dP4

TMM 41106-5132, right M1

TMM 41106-5133, right M2

TMM 41106-5134, right dP4

TMM 41106-5135, anterior % left M1

TMM 41106-5136, broken left P4

TMM 41106-5137, left M1

TMM 41106-5138, right P4

TMM 41106-5139, left P3

TMM 41106-5140, left P3

PM 34485, right ramus fragment with P4-M,

PM 34486, left ramus fragment with I„ P3, dP4

PM 34487, left ramus fragment with I,

PM 34488, right maxillary fragment with broken M2"3

PM 34489, left P3
Trench 4, Unit 2, Level 1

TMM 41106-314, right maxillary with P3, dP4, M1, part of M2

WAM 75.1.135, left maxillary fragment with Mu2

WAM 75.1.136, right ramus fragment with M2_3

PM 34490, left P3

PM 34491, worn right P4

PM 34492, right M1

PM 34493, right M2

PM 34494, right M2

PM 34495, right M,

PM 34496, left M,

PM 34497, right M2

PM 34498, right M3

PM 34499, left M,

PM 34500, right M'

PM 34501, left M2

LUNDELIUS & TURNBULL: MADURA CAVE 17

PM 34502, left M1

PM 34503, left M2

PM 34504, left M', M2, or M3

PM 34505, right M2

PM 34506, anterior Vi left M2 or M3

PM 34507, right M2

PM 34508, left M3

PM 34509, edentulous left ramus

PM 34511, right maxillary fragment with M2

PM 36873, posterior Vi lower molar

PM 36874, right ramus fragment with M2.3

PM 36875, left maxillary fragment with dP\ P4, and M2 in crypts, alveoli for P\ M' (fig.
4A)

PM 36876, right maxillary fragment with M'2
Trench 4, Unit 2, Level 2

TMM 41106-12, left maxillary with P3, crypt for P4

PM 36880, X right M2

PM 36881, left M3 or M2

PM 36882, left M2

PM 36883, left M3

PM 36884, 36885, two left M's

PM 36886, right M2 or M3

PM 36887, left M3

PM 36888, left ramus fragment with dP4

PM 36889, edentulous right ramus fragment

PM 36890, edentulous left ramus fragment

PM 36891, edentulous left maxillary fragment

PM 36892, left maxillary fragment with M1

PM 36893, left maxillary fragment with broken P4, M1

PM 36895, left dP4

PM 36896, right dP4

PM 36897, left P4

PM 36898, posterior % right P4

PM 36899, posterior Vi left P4

PM 36900, left M,

PM 36901, left P3

PM 36902, left P3 (uncertain since tooth is broader than rest of sample, blade is not
compressed, and has labial groove)

PM 36903, left M, (has peculiar uninterrupted protoconid ridge anteriorly)

PM 38712, left M2
Trench 4, Unit 2, Level 3

TMM 41106-2, right ramus fragment with M,_2
Trench 4, Unit 4

TMM 41106-63, right maxillary with P3, dP\ M1
Trench 4, Units 4-5

WAM 75.1.149, 150, two right P4s

WAM 78.3.1, 3, two left dP4s

WAM 78.3.2, right dP4

PM 36904, right P4

PM 36905, right P4

PM 36906-36909, four left dP4s

PM 36910-36912, three right dP4s

PM 36913, right P4

PM 36914, left P4

PM 36915-36918, four right P4s

PM 36919, right P4

PM 36920, left P4

PM 36921, left dP4

PM 36922, 36923, two right M,s

PM 36924, right M1 or M2

PM 36925-36929, five right dP4s

18 FIELDIANA: GEOLOGY

PM 36930, 36931, two left dP4s

PM 36932-36934, three left P3s

PM 36935-36938, four right P3s

PM 36939-36942, four left P3s

PM 36943-36948, six right P3s

PM 36949-36951, three right M,s

PM 36952, right M3

PM 36954-36956, three right M's or M2s

PM 36957, left M2

PM 36958, left M,

PM 36959-36962, four right M3s

PM 36963-36965, three right M's

PM 36966, left M3

PM 36967, right M2

PM 36968, 36969, two left I,s

PM 36970, molar fragment

PM 36971, left M,

PM 36972, left I,

PM 36973, right dP4

PM 36974, 36975, two molar fragments

PM 36976, left I,

PM 36977, right M,

PM 36978, right dP4

PM 36979, 36980, two molar fragments
Trench 4, Unit 7, Level 1

WAM 78.3.4, right M2

WAM 78.3.5, left M2

WAM 78.3.6, left M'

WAM 78.3.7, right M2

WAM 78.3.8, right M2 or M3

PM 36989, 36990, two left P4s

PM 36991, 36992, two right P4s

PM 36993-36996, four I,s

PM 36997, 36998, two right M3s

PM 36999, 37000, two right M2s

PM 37001, 37002, two left M,s

PM 37703, left M3

PM 37004, 37005, two left M's

PM 37006, left M2 or M3

PM 37007, right M,

PM 37008, 37009, two left M2s

PM 37010, 37011, two right M2s or M3s

PM 37012, left dP4

PM 37013, left P3

PM 37014, left M,

PM 37015, left M2 or M3

PM 37016, right ramus fragment with M3, alveoli for M2, M4
Trench 4, Unit 7, Level 2

PM 37017, left maxillary fragment with M'

PM 37018, right P4

PM 37019, 37020, two ramus fragments

PM 37021, left ramus fragment with broken molar, probably M2

PM 37022, 37023, two left P3s

PM 37024-37026, three right P3s

PM 37027-37029, three left P3s

PM 37030, 37031, two right P3s

PM 37032, 37033, two right P3s

PM 37034-37037, four broken left P4s

PM 37038, 37039, two broken right P4s

PM 37040, left P3

LUNDELIUS & TURNBULL: MADURA CAVE 19

PM 37041, left P3

PM 37042, 37043, two left dP4s

PM 37044, left P4

PM 37045, right P4

PM 37046, 37047, two right dP4s

PM 37048, left dP4

PM 37049-37057, nine I,s

PM 37058-37060, three right M2s

PM 38617, 38618, two right MjS

PM 38619-38623, five left M's

PM 38624, left M2

PM 38625-38627, three left M2s or M3s

PM 38628-38631, four right M,s

PM 38632-38635, four right M3s

PM 38636, left M,

PM 38637-38639, three right M's

PM 38640, right M,

PM 38641, right M3 or M2

PM 38642, left M2

PM 38643, 38644, two left M,s

PM 38646, 38647, two left M4s

PM 38648, right M,

PM 38649, left M,

PM 38650, broken tooth
Trench 5, Unit 6

TMM 41106-589, left P3

TMM 41106-593, left P4

TMM 41106-598, right P4

TMM 41106-630-633, three left dP4s

TMM 41106-634, left lower molar fragment

TMM 41106-635, right dP4

TMM 41106-637, left M, or M2

TMM 41106-638, left dP4

TMM 41106-639, left lower molar

TMM 41106-642, left dP4

TMM 41106-645, right M, or M2

TMM 41106-646, right dP4

TMM 41106-5028, left P4

TMM 41106-5109, right P4

TMM 41106-5111-5115, five right P3s

TMM 41106-5116, right P3

TMM 41106-5118, -5119, two left P3s

PM 38651, 38652, two left M2s

PM 38653, broken M, or M2

PM 38654, right M3 or M2

PM 38655-38658, four left M's

PM 38659, broken left M2

PM 38660-38663, four left M3s or M^

PM 38664-38666, three right M's

PM 38667, 38668, two right M^

PM 38669, left M2 or M,

PM 38670, right M2 or M,

PM 38671, left M, or M2

PM 38672, left M3

PM 38678-38685, eight right lower incisors

PM 38686, right M2
Trench ?, Unit ?; probably Trench 3, Unit 3

WAM 75.1.148, right ramus fragment with P„ dP4
Trench ?, Unit ?, Level ?

TMM 41106-5141, right M, or M2

20 FIELDIANA: GEOLOGY

Comparative Material

Webb's Cave

PM 36894, right ramus with P4, broken M,_„

Description

Skull. — Several maxillary fragments show some morphological characters
which permit comparison with the skulls of various rat-kangaroos. Since most
of these fragments are from juvenile individuals, we have used juveniles for
comparison where they were available. The anterior opening of the postorbital
canal lies approximately over the midpoint of P3, usually as a simple opening.
In one specimen (TMM 41106-34) this foramen opens into a broad, deep depres-
sion on the side of the maxillary. Within the central part of this depression
there is a flat arching strut of bone that forms a tunnel about one-fourth the
diameter of the infraorbital foramen, large enough to pass a sizable branch(es)
of the infraorbital nerve or artery (fig. 4B). This character apparently is variable
in potoroines; a specimen of Bettongia lesueur from Koomooloobooka Cave (TMM
41230-108) shows this "bifurcated" condition only on one side of the skull. The
posterior opening of the infraorbital canal lies close to the sphenopalatine fo-
ramen, as it does in Bettongia; it differs in this character from Potorous tridactylus
(PM 6706 and TMM M-2549). Behind and slightly lateral to the posterior open-
ing of the infraorbital canal are two or three small nutrient foramina which
extend downward into the alveoli of the teeth. One branch of the infraorbital
canal extends downward and anteriorly and emerges as a small foramen on the
palate just anterior to the P3. At least one lacrimal foramen is present. The canal
leading downward and anteriorly from it can be seen on the inside of the
maxillary (fig. 4B). This foramen probably is equivalent to the inferior lacrimal
foramen described by Stirton (1963) for Protemnodon, which is present in most,
if not all, macropodids.

Upper Dentition. — No incisors have been found among the Madura Cave ma-
terials which can definitely be associated with skull fragments identified as
Caloprymnus. One edentulous premaxillary (PM 34434; fig. 4H) has been ten-
tatively referred to this genus on the basis of alveolar dimensions and positions,
but this specimen also might belong to Lagorchestes, whose premaxillary is very
similar to that of Caloprymnus.

The P3 is a laterally compressed, serrate blade, with three strong grooves
followed by one weak groove dividing the crown labially and lingually into

Opposite:

Fig. 4. Caloprymnus catnpestris from Madura Cave. A, PM 36875. Left maxillary of a
juvenile with dP4, P\ and M2 in their crypts, alveoli of P3 and M1, shown in ventral and
labial views. B, TMM 41106-34. Left maxillary of a juvenile with P3, P4 removed from
crypt, M12, shown in lingual (left), ventral, and labial views. C, TMM 41106-167. Left
maxillary of a juvenile with P\ dP4, M'~2, alveoli for M\ shown in labial (left), ventral,
and lingual views. D, TMM 41106-310. Right maxillary fragment of an adult with P4, M1,
shown in labial and crown views. E, TMM 41106-358. Right maxillary fragment of an
adult with M2-3, shown in crown and lingual views. F, TMM 41106-307. Left maxillary
fragment of an adult with M13, shown in crown and lingual views. G, TMM 41106-309.
Right maxillary fragment of an adult with M34, shown in crown and lingual views. H,
PM 34434. Incertae sedis, probably C. catnpestris, possibly Lagorchestes. Edentulous right
premaxillary shown in labial and ventral views.

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22 FIELDIANA: GEOLOGY

three well-defined anterior cusps and two less well-defined posterior cusps.
The combined length of the posterior pair of cusps is two to two and one-half
times that of either of the preceding cusps. The two posterior cusps are strongly
compressed laterally and are aligned with the long axis of the tooth. The crest
of the blade is almost straight. The tooth has three roots, one anterior and two
posterior. The posterior roots lie side by side and diverge upward. A small
fourth root is fused to the lingual side of the anterior root in two specimens
(TMM 41106-167, 41106-359). The lingual and labial edges of the tooth have
cingular shelves which are widest over the posterior roots. The shelves bear
two to four (most often three) broad, low cusps which are located opposite the
ridges on the sides of the blade, but are not always joined to them. Both sides
of the tooth usually have the same number of cingular cusps.

The dP4 is a rectangular tooth with four major equal-sized cusps (fig. 4A,C).
A laterally compressed parastyle, indistinctly set off from the paracone, forms
a short blade which is aligned with the blade of P3. Crests extending posteriorly
from the protocone and anteriorly from the hypocone join near the midline of
the tooth to form a V-shaped lingual boundary for the central basin. Very weak
crests extend from the apices of the paracone and metacone toward the center
of the tooth but they do not join, thus leaving the labial side of the basin open.
The posterior crests of the metacone and hypocone join to form a narrow pos-
terior cingular basin. The protocone lacks an anterior crest. The anterior cin-
gular area consists of a concave surface which slopes rootward from the para-
style to a point about halfway across the anterior face of the protocone. There
is a very weak procingular ridge.

The protoloph is straight, with its lowest point close to the protocone. The
metaloph has a slight sigmoid curve, with its lowest point close to the hypo-
cone. As in Recent material from South Australia (Finlayson, 1932), the para-
cone and metacone contribute more to the lophs than do the protocone and
hypocone. The posterior face of the metaloph has a broad convexity extending
from the midpoint of the crest to the posterior cingular basin. The anterior face
of the metaloph and both faces of the protoloph are flat.

A spur running anterolabially from the hypocone into the central valley was
described by Finlayson (1932) in Recent specimens of Caloprymnus campestris
from South Australia and by Tedford (1967) in a tentatively referred specimen
from Lake Menindee, New South Wales. The only sign of this spur on the
Madura Cave specimens is a faint bulge in that position on some specimens.

The P4 is a straight, serrate, laterally compressed blade with no arch to the
edge (fig. 4B,D). Six grooves divide the crown into seven laterally compressed
cusps. The anterior six cusps are approximately the same size. The posterior
cusp is approximately twice as long as any other cusp and has a broad ridge on
its lingual face which extends to the valley which separates this cusp from the
posterointernal cuspule. The posterointernal cuspule is conical when unworn
and is about half the height of the posterior cusp of the blade. A lingual cin-
gulum is present which bears three to five low, rounded cuspules which are
located opposite the grooves in the blade and are joined to the cusps of the
blade by weak ridges. The anterior cusp of the blade has a well-defined anterior
ridge.

The general pattern of M1, M2, and M3 (fig. 4A-G) is similar to that of dP4.
There is no parastyle, but anterior crests of the protocone and paracone join to
form a prominent ridged procingulum and an anterior cingular basin. The
posterior crest of the protocone and the anterior crest of the hypocone are

3 #

•J

Fig. 5. Caloprymnus campestris from Madura Cave. A, TMM 41106-160. Left mandible
of a juvenile with P3, dP4, M,.2, alveolus for M3, crypt for M4, shown in dorsal and lingual
views. B, TMM 41106-161. Right mandible of a juvenile with P3, dP4, M,_3, alveoli for M4,
shown in dorsal and labial views. C, TMM 41106-163. Left mandible of a juvenile with
I, P3, P4 (all removed), M,_2, M3 in crypt, crypt for M4, shown in labial (left), crown, and
lingual views. D, TMM 41106-164. Left ramus fragment of an adult with broken incisor
and very worn P4 and M„ shown in labial (left), crown, and lingual views. E, TMM
41106-166. Left ramus fragment of an adult with P4, M2_4, alveolus for M,, shown in labial
(left), crown, and lingual views.

23

24

FIELDIANA: GEOLOGY

Table 5. Position of anterior end of masseteric canal in specimens of Caloprymnus
campestris of various dental ages.

Position of anterior

Oental

end of masseteric

stage

Specimen No.

Eruption stage

canal

1

TMM 41106-354

dP4 in place, M, erupting

Posterior to M,

2

TMM 41106-351

M, in place, M2 erupting,
M3 unerupted

Anterior end of M2

3

TMM 41106-346

P3, dP4, M,* in place, M2
erupting*

Posterior end of M,

3

TMM 41106-349

dP4-M,* in place, M2

Posterior end of M,

erupting

alveolus

3

TMM 41106-352

dP4, M, in place, M2
erupting*

Posterior end of M,

4

TMM 41106-353

P3/* dP4,* M„* M2

Midpoint of M, alveo-

erupting

lus

5

TMM 41106-2

M„ M2 in place, M3
erupting?

Protolophid of M,

5

TMM 41106-344

P3-M2 in place, M3 erupt-
ing*

Hypolophid of M,

5 +

TMM 41106-345

P3-M2 in place, M3 un-
erupted

Posterior end of M,

6

PM 36894 (Webb's Cave)

P4-M4 in place

Anterior end of M,

Based on condition of alveoli.

directed labially and meet in the central valley in either a V or a U. The pos-
terior crest of the paracone and the anterior crest of the metacone are weakly
developed. They extend from the apices of the cusps toward the central valley
and then turn lingually in varying ways. In some specimens the crests almost
reach the central valley and then turn sharply lingually, while in other speci-
mens the crests swing lingually in a smooth curve about halfway to the bottom
of the central valley. In some specimens the posterior crest of the paracone
bifurcates and one ridge is directed lingually, while the other, which usually
is more rounded, remains near the labial edge of the cusp, disappearing near
the bottom of the median valley. The labial crests usually do not join in the
central valley as do the lingual crests. The posterior crests of the metacone and
hypocone join to form a well-developed postcingulum with a deep posterior
cingular basin.

The protoloph and metaloph are slightly wavy. They join the paracone and
metacone at their apices, but join the protocone and hypocone on their labial
sides below their apices. The protoloph joins the side of the protocone slightly
anterior to its apex; the metaloph joins the side of the hypocone slightly pos-
terior to its apex. The protoloph has a low rounded bulge on its anterior face.
The metaloph has a similar, but larger, bulge on its posterior face; this posterior
bulge may be ridged. The faces of both metaloph and protoloph which border
on the central valley are flat.

The anterolabially directed spur from the hypocone mentioned by Finlayson
(1932) and Tedford (1967) is variable in its expression. An enamel column may
be present between the protocone and hypocone of M1. The cusps and ridges
of all of these molars are higher and sharper than those of Bettongia.

The only M4 in the Madura Cave collection (TMM 41106-309; fig. 4G) is so
heavily worn that few features can be discerned. It is markedly smaller than
M3, and the posterior end is reduced so that the tooth has a triangular shape.

LUNDELIUS & TURNBULL: MADURA CAVE

25

Table 6. Measurements of upper dentitions of Caloprymnus campestris from Unit 1 of
Madura Cave.

TMM TM TM

PM 34488 41106-570 41106-5140 4106-5138

3.30
1.25
1.22
1.71

dP<

M

M'

M

TMM

41106-540

L

AW

PW

x. W

L

AW

PW

L

AW

PW

L

3.04

AW

2.96

PW

2.93

L

3.46

AW

3.27

PW

3.27

L

AW

PW

...

2.89
2.20
2.43

3.50
2.89
2.43

5.85
1.78
1.98

Mandible. — The mandible is short and deep, as in Bettongia and unlike Poto-
rous. No specimen from the Madura Cave collection has a complete ascending
ramus, but the basal part is preserved in two specimens (TMM 41 106-160, 41 106-
345; fig. 5A). The anterior edge of the ascending ramus is at a 45° angle to the
tooth row, steeper than the angle found in either Bettongia or Potorous. The
masseteric fossa is deep and opens anteriorly into a tunnel-like masseteric canal,
which has been shown by Abbie (1939) to be confluent with the inferior dental
canal and also serves as the area of insertion for the deep masseter. This canal
extends to the posterior end of the alveolus for the lower incisor, which lies at
the level of the posterior end of M, in juvenile individuals and at the level of
the anterior end of M, in adult specimens (table 5). Other genera of potoroines
show some variation in the forward extent of this canal. In Aepyprymnus and
Potorous it extends to the level of M2, and in Bettongia, to the middle of M,
(Abbie, 1939). Thus, Caloprymnus shows as specialized a state of this character
as any of the other potoroines.

The mental foramen is high on the labial side of the mandible just ahead of
P4 or P3. All mandibles have a shallow pit on the labial side just below M,. In
some specimens this pit is elongated, somewhat like the labial groove in Pro-
temnodon (Stirton, 1963, p. 123). There is an equidimensional concave-upward
pterygoid fossa. The mandibular foramen is located at the anterior end of this
fossa directly under the posterior end of the tooth row. The symphysis is not
ankylosed. The symphyseal areas are prominent and roughly oval with an ir-
regular surface. The posterior end of the symphysis lies under dP4 in juvenile
individuals and under P4 in adult individuals. There is a large pit immediately
posterior to the symphysis.

Lower Dentition. — The lower incisor is long, gently tapered, and weakly curved,
with a suboval cross section except for a flattening on the ventral edge of the

26 FIELDIANA: GEOLOGY

Table 7. Statistical data on upper dentitions of Caloprymnus campestris from Units 2 to
4-5 of Madura Cave.

I"

dP<

M1

M2

M3

M'

N

Mean ± SE

SD

CV (%)

OR

L

24

3.61

± .049

.24

6.65

3.09-3.95

AW

24

1.31

± .022

.11

8.40

1.14-1.52

PW

24

1.28

± .033

.16

12.50

1.14-1.90

Max. W

24

1.85

± .039

.19

10.27

1.43-2.20

L

21

2.93

± .035

.16

5.46

2.58-3.19

AW

21

2.14

± .031

.14

6.54

1.97-2.43

PW

21

2.43

± .037

.17

7.00

2.05-2.66

L

6

5.53

5.32-5.78

AW

6

1.64

1.44-2.05

PW

7

2.05

1.86-2.28

L

16

3.26

± .035

.14

4.29

3.04-3.50

AW

17

2.90

± .046

.19

6.55

2.66-3.30

PW

16

2.84

± .025

.10

3.52

2.74-3.12

L

16

3.43

± .045

.18

5.25

3.12-3.72

AW

16

3.17

± .040

.16

5.05

2.96-3.50

PW

14

2.97

± .035

.13

4.38

2.81-3.34

L

4

3.28

3.19-3.42

AW

4

3.12

2.96-3.19

PW

3

2.63

...

2.48-2.66

L

1

2.36

AW

1

1.78

...

PW

medial surface (fig. 5C). It shows somewhat greater lateral compression distally
than proximally. The ventral and lateral surfaces of the crown are covered with
enamel, except for the most proximal quarter of the lateral surface. The medial
surface has a fairly even enamel band along its ventral edge which gradually
thins to nothing in the most proximal quarter of the crown. The rest of the
medial surface and all of the dorsal surface except for its extreme lateral edge
lack enamel. Most of the lower incisors from the lower units in the cave have
enamel extending slightly higher on the medial side.

The P3 is a serrate, bladelike tooth with three grooves which divide it into
four laterally compressed cusps. The anterior and posterior cusps are larger
than the two central cusps. The posterior cusp, which is the largest, is weakly
subdivided on both sides and across the ridge by a shallow groove. This groove
usually is removed by wear. The posterior half of this cusp is turned lingually,
and considerably overhangs the posterior roots of the tooth. The labial side of
the tooth has a prominent bulge centered over the posterolabial root. This bulge
shows subdued extensions of the grooves that divide the crown. The lingual
side of the tooth has a low, rounded cingulum which extends from the anterior
part of the posterior cusp to the posterior part of the anterior cusp. This cin-
gulum bears two low, blunt cusps which lie opposite the two central cusps of
the blade. The labially expanded base of the crown shows some variation at its
posterior end. In TMM 41106-160 and 41106-161 its edge is a smooth arc. In
TMM 41106-344 the edge is sharply constricted at its posterior end. TMM 41106-
163 is less sharply constricted. Like P3, P3 has one anterior root and two side-
by-side posterior roots (fig. 5A,B).

The dP4 is a subtriangular tooth with one anterior root and two posterior

LUNDELIUS & TURNBULL: MADURA CAVE 27

Table 8. Statistical data on upper dentitions of Caloprymnus campestris from Unit 7 of
Madura Cave.

P>

dP<

N

Mean ± SE

SD

CV(%)

OR

L

10

3.55 ± .060

.19

5.35

3.27-3.84

AW

10

1.23 ± .002

.08

6.50

1.14-1.37

PW

10

1.33 ± .002

.08

6.02

1.22-1.44

Max. W

10

1.75 ± .051

.16

9.14

1.44-1.82

L

8

3.02 ± .049

.14

4.64

2.81-3.19

AW

8

2.28 ± .078

.22

9.65

1.90-2.66

PW

8

2.45 ± .078

.22

8.98

2.20-2.88

L

3

5.23

4.98-5.55

AW

2

1.45

...

2.37-1.52

PW

2

1.64

...

1.60-1.67

L

1

3.50

...

AW

1

3.04

...

...

...

PW

1

2.92

...

...

...

M1

roots. The anterior end of the tooth is laterally compressed all the way to the
base of the crown. The blade thus formed has its anterior half oriented parallel
to the long axis of the tooth and its posterior half turned lingually at an angle
of about 45° to the long axis of the tooth. The change in orientation between
the two halves is abrupt. In some specimens the blade is weakly divided into
two cusps by a shallow groove. At the point of inflection there is a ridge on
the labial side of the blade which extends rootward and posteriorly to join the
anterior crest of the hypoconid, forming the labial boundary of the central basin
well in from the labial edge of the tooth. The lingual border of the central
basin is formed by the merging of the bases of the lingual cusps. In many
specimens this border is enhanced by the joining of crests extended from the
lingual cusps. The hypoconid and entoconid are subequal in size when unworn
and are joined by a sharp ridge. The crest of the entoconid makes the greatest
contribution to this ridge. Crests extend posteriorly from the apices of the
hypoconid and entoconid to surround a broad posterior basin.

The P4 is a relatively low-crowned, serrate, bladelike tooth with two roots
(fig. 5C-E). In profile its cutting edge is almost straight rather than being slight-
ly arched as it is in Bettongia. In crown view the crest bends lingually in a
gentle arc which is most pronounced in the posterior half of the blade. Five
well-developed pairs of grooves divide the tooth into six cusps. The anterior
cusp may be larger than the next four cusps. The posterior cusp is two to three
times the size of any of the four preceding cusps. It is strongly compressed
laterally and is inflected lingually at an angle of about 40° to the long axis of
the tooth. The lingual surface of the tooth has a cingular bulge which is variably
subdivided by extensions of the grooves on the blade. The labial surface of the
tooth is almost flat, but may have a similar, though smaller, cingular bulge.

The M,, M2, and M3 are rectangular teeth with four nearly equal-sized cusps
(fig. 5A-E). The anterior moiety of M, is slightly narrower than the posterior
moiety (tables 9, 10). The protolophid and the hypolophid are well developed.
These lophids are sharp when unworn and are much higher than those of
Bettongia. The lophids join the lingual conids at their apices and slope down
from there to the midline of the tooth. As pointed out by Finlayson (1932) for
the Recent population from South Australia, the metaconid and entoconid con-
tribute more to the lophids than do the protoconid and hypoconid. The hy-

28

FIELDIANA: GEOLOGY

Table 9. Measurements of lower dentitions of Caloprymnus campestris from Unit 1 of
Madura Cave.

TMM

TMM

TMM

TMM

PM

PM

PM

41106-

41106-

41106-

41106-541

34485

34486

34489

5131

5134

5139

Mean

p3

L

3.27

3.04

3.16

AW

1.14

1.22

1.18

PW

1.37

1.06

.99

1.14

Max. W

1.75

1.82

1.90

1.82

dP4

L

2.81

2.81

2.96

2.86

AW

1.98

1.75

1.56

1.76

PW

2.05

2.32

2.05

2.14

P4

L

4.94

4.94

AW

1.75

1.75

PW

1.67

1.52

1.60

M,

L

2.81

2.96

2.89

AW

2.13

PW

2.51

M2

L
AW
PW

3.08

2.74
2.74

M3

L
AW
PW

3.12
2.96

2.74

M4

L
AW
PW

2.74
2.81
2.05

M,_3

L

8.97

M,_4

L

11.67

polophid is straight. The protolophid is slightly convex anteriorly. In unworn
teeth prominent crests extend downward and lingually from the apices of the
protoconid and hypoconid toward the midline of the tooth and join in the
central valley to form the labial boundary of the central basin. The metaconid
and entoconid have weakly developed crests which extend down the cusps to
meet at the midline of the tooth. These crests form the lingual boundary of the
central basin. The procingulum of Ml is variable. In some specimens it is only
half the width of the tooth, lacks a well-defined basin, and is joined by a ridge
only to the metaconid. In most specimens it is larger, occupying up to three-
quarters of the width of the tooth, has a definite basin, and is joined by ridges
to both the metaconid and the protoconid. A few specimens have procingular
basins, but lack the ridge from the protoconid. The M2, M3, and M4 all have
procingula which are connected to the protoconid by prominent crests and
which have well-defined basins. In all specimens which are not extremely worn,
the procingulum is joined to the base of the anterolabial corner of the proto-
conid. The posterior crests of the hypoconid and entoconid of Mj_3 swing down-
ward and toward the midline of the tooth, joining to form a well-defined post-
cingulum with a well-developed basin.

The M4 is markedly narrower posteriorly than the other molars (fig. 5E). The

L

14

3.05

+

.040

.15

4.92

2.85-3.34

AW

14

1.26

±

.027

.10

7.94

1.06-1.44

PW

14

1.27

t

.032

.12

9.45

1.06-1.44

Max. W

14

1.83

f

.021

.08

4.37

1.67-1.98

L

19

2.74

±

.028

.12

4.38

2.51-2.96

AW

19

1.51

4

.044

.19

12.58

1.37-1.90

PW

18

2.00

±

.028

.12

6.00

1.82-2.36

L

11

4.80

±

.108

.36

7.50

4.26-5.40

AW

9

1.47

f

.070

.21

14.74

1.22-1.75

PW

11

1.57

+

.060

.20

12.74

1.22-1.82

L

12

3.17

±

.052

.18

5.68

2.89-3.60

AW

12

2.29

±

.029

.10

4.37

2.12-2.47

PW

13

2.64

±

.078

.27

10.22

2.36-3.43

L

15

3.48

±

.036

.14

4.02

3.27-3.72

AW

13

2.78

*

.028

.10

3.60

2.62-2.96

PW

13

2.95

fc

.036

.13

4.41

2.74-3.12

LUNDELIUS & TURNBULL: MADURA CAVE 29

Table 10. Statistical data on lower dentitions of Caloprymnus campestris from Units 2
to 4-5 of Madura Cave.

N Mean ± SE SD CV (%) OR

dP4
P<

M,
M,

hypoconid and entoconid are well defined. The postcingulum, formed by crests
from the hypoconid and entoconid, has a basin which is located partly between
the hypoconid and entoconid rather than entirely posterior to them as it is in
the other molars.

Discussion

A comparison of measurements of teeth from various stratigraphic units shows
few differences in size (tables 6-11). The lengths of the P4 and M2 of two
specimens from Unit 1 are below the lower limit of those teeth of the sample
from Units 2 to 4-5, but larger samples would be needed to demonstrate sig-
nificant differences. Comparisons of samples from Units 2 to 4-5 and Unit 7
reveal minor differences, but f tests give nonsignificant results (P > .05) in
every case.

The Madura Cave specimens are smaller than the Pleistocene specimens from
Lake Menindee and the Recent specimens from northeastern South Australia.
A comparison of the dental measures given by Tedford (1967, p. 30) and Fin-
layson (1932, p. 166) shows that there is no overlap in any measurement be-
tween those samples and the sample from Madura Cave.

Caloprymnus campestris is known as a living animal only from a small area in
northeastern South Australia and southwestern Queensland (Finlayson, 1932).
This area has sand ridges, claypans, and gibber plains. According to Finlayson,
Caloprymnus does not occur on the sand ridges, but seems to be restricted to the
flat areas.

Caloprymnus is known from late Pleistocene and Holocene deposits at Lake
Menindee, western New South Wales (Tedford, 1967) and various caves on the
Nullarbor Plain (Lundelius, 1963; Archer, 1972, 1974). The time of its disap-
pearance from the Nullarbor Plain is not known precisely, but was certainly
Holocene. Its presence in surficial deposits in Webb's Cave (Lundelius, 1957,
1963) has been taken to indicate a very recent occurrence as a living animal in
this area, but C14 dates of 1500 ± 70 BP on charcoal (TX 1148) and 5030 ± 90

L

8

3.14

2.96-3.50

AW

8

1.20

1.06-1.44

PW

8

1.28

1.14-1.52

Max. W

8

1.79

1.67-1.98

L

3

2.79

2.66-2.98

AW

3

1.37

1.29-1.44

PW

3

1.95

1.94-1.98

30 FIELDIANA: GEOLOGY

Table 11. Statistical data on lower dentitions of Caloprymnus campestris from Unit 7 of
Madura Cave.

N Mean OR

dP4

BP on bone (TX 1147) from a depth of 1 to 3 inches (2 to 7 cm) in this cave
throw doubt on this conclusion. In addition, on two mummified specimens of
Thylacinus cynocephalus from two other Nullarbor caves, C14 dates are 3280 ± 90
BP for one and 4650 ±104 and 4550 ± 153 BP for the other (Partridge, 1967;
Lowry & Merrilees, 1969). These dates indicate that there has been little or no
deposition in some of these caves for significant periods of time. This, together
with the dry conditions in many of these caves, which ensure good preserva-
tion, demonstrates that caution should be used in drawing conclusions about
the age of material based only on its appearance or on its position on the
surface. It is possible that Caloprymnus survived to the time of European settle-
ment in locally favorable areas as the report of its presence on the Bunda
Plateau by Tate (1879) suggests, but as yet there is no supporting evidence from
the cave deposits.

Like Dasyuroides byrnei, Caloprymnus disappeared from the Nullarbor Plain
some time during the Holocene when the climate was becoming drier, but still
survives in a very arid area in central Australia. The reasons for the disappear-
ance of these two animals from the Nullarbor Plain are not known. Finlayson
(1958) has suggested that Caloprymnus campestris is competitively inferior to
Bettongia lesueur in arid areas because of the latter's burrowing habit. He points
out that, today, C. campestris survives only in areas in which B. lesueur does not
occur. If Finlayson's hypothesis is correct, then we need to account for the
exception to it given by the Madura Cave and Lake Menindee records. This
will be considered in the conclusion.

Caloprymnus Incertae Sedis

Material

Trench 3, Unit 2, Level 1

PM 34434, edentulous right premaxillary, possibly Lagorchestes (fig. 4H)
Trench 3, Unit 2, Level ?

PM 38718, broken molar
Trench 5, Unit 6

PM 38674-38677, four upper incisors

Bettongia

Bettongia Gray, 1837
Bettongiops Matschie, 1916

The genus Bettongia is represented in the Madura Cave deposits by two species,
B. lesueur and B. penicillata. Specimens with diagnostic characters such as sec-

LUNDELIUS & TURNBULL: MADURA CAVE 31

torial premolars and auditory bullae can be confidently assigned to species.
Table 12 lists the diagnostic features. However, the majority of the Bettongia
material from Madura Cave consists of isolated teeth that usually cannot be
identified to species; many of these specimens are listed as Bettongia sp. These
materials show no characters that would rule out their assignment to either B.
lesueur or B. penicillata or that suggest the presence of any other species of
Bettongia.

The M1 and M2 of B. penicillata frequently have a mesostyle (Finlayson, 1958;
Tedford, 1967; Marshall, 1973). All of the Madura Cave specimens which can
be assigned to B. penicillata on other grounds have mesostyles on dP\ M1, and
M2. Therefore, isolated upper molariform teeth with mesostyles have been re-
ferred to B. penicillata. Because some specimens of B. penicillata have been re-
ported to lack a mesostyle on M1 and M2, isolated upper molariform teeth which
lack a mesostyle cannot be assigned to either species. The situation is further
complicated by the fact that, on rare occasions, a specimen of B. lesueur may
have a trace of mesostyle on dP4, M1, or M2 (an example is PM 4786, positively
identified as B. lesueur on the basis of the form and groove count of its P3s and
P*s).

It is difficult to identify individual dP*s to species on the basis of standard
dental measurements. Considering only teeth in jaws, which can be identified
on the basis of other features, the dP4s of B. penicillata are smaller on the average
than those of B. lesueur, but the two size distributions overlap. A plot of length
versus anterior width for all known dP4s from Madura Cave (fig. 6A) illustrates
the extent of this overlap. In Figure 6B the points for isolated teeth have been
added to the plot from Figure 6A. Small x's indicate those isolated teeth which
we considered to belong to B. penicillata because they are small and their crowns
are simple. Small circles indicate those isolated teeth which we considered to
belong to B. lesueur because they are larger and more bulbous and have more
complex crowns. Seven teeth which did not fall into either group are indicated
by dots with specimen numbers. Most of the isolated teeth fall within or near
the plots of the known teeth. The fact that it is only tooth proportions, rather
than other consistent morphologic features, which seem to divide the two groups
makes the identifications less certain. Figure 6C shows a plot of length versus
posterior width for the same specimens. Overall, the results of this plot are the
same as those of Figure 6B, although the two differ in detail.

Bettongia lesueur (Quoy & Gaimard, 1824)

Hypsiprymnus lesueur Quoy and Gaimard, 1824

Hypsiprymnus graii Gould, 1840

IBettongia graii Gray, 1841

Perameles harveyi Waterhouse, 1842

Bettongia lesueuri Thomas, 1888
Bettongiops lesueuri Matschie, 1916

Material

Surface pickup

TMM 41106-20, adult skull with left and right I', P4-M\ and right ramus with I, P4-M,

(figs. 7, 9A-C)
TMM 41106-26, left ramus with I, P3, dP4, P4 in crypt, M„ M2, and M3 in crypt (fig.
10A-C)

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A

3.0

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2.2

# b. penicillata
Ob. lesueur

/ ° )

0 6

o

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2.8

3.2

3.6

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LENGTH

□ INDIVIDUAL TOOTH SCORED AS
B. LESUEUR

X INDIVIDUAL TOOTH NOT SCOREABLE
ON MORPHOLOGIC CRITERIA

2.8

3.2

3.6

4.0

LENGTH

34

LUNDELIUS & TURNBULL: MADURA CAVE

35

LENGTH

Fig. 6. Bivariate graphs of dP4s of Bettongia from Madura Cave. A, Plot of teeth of
known specific identity. B, Plot of individual teeth added to A to show the degree of
correspondence of morphological and quantitative criteria for specific assignment. The
dashed line outlines the probable area of Bettongia lesueur. Note that, although there is
an overlap zone, only one specimen is unassignable on the basis of both morphological
and quantitative criteria. C, Plot of dP*s of known specific identity and of individual
teeth. The dashed lines outline the probable areas of Bettongia penicillata and Bettongia
lesueur. Note the area of overlap.

Trench 1, Unit 1, top 1 foot

PM 4786, right maxillary with P\ dP4, P4 removed from crypt, M1 2, crypt for M3 (fig.

8A,B)
PM 4787, 4788, two right maxillaries with P3, dP4, P4 exposed in crypt, M1, crypt for M2

(fig. 8C,D: PM 4787)
PM 4790, right ramus with I, P„ dP4, P4, and M2.j in crypts
PM 4791, right ramus with I, P4 in crypt, M2, alveoli for P„ dP4, and M,
PM 4793, right ramus with I, P3, dP4, P4 in crypt
PM 4795, left ramus with I, dP4, P4 in crypt, M,, M2 removed from crypt, alveolus for

P, (fig. 9D-F)
PM 4796, left ramus with I, P3, dP4, P4 in crypt, M2, M, in crypt, alveolus for M, (fig.

10D-F)
PM 4797, left ramus with I, P4 and M2 in crypt, alveoli for P„ dP4, M,
Trench 3, Unit 2, Level ?

TMM 41106-5, right ramus with broken I, P4, M,.3, alveoli for M4

TMM 41106-117, left maxillary with P4-M2 (fig. 8E-G)

TMM 41106-118, right ramus fragment with broken I, P4, alveolus for M,

36 FIELDIANA: GEOLOGY

Trench 3, Unit 3, Level ?

TMM 41106-33, left maxillary with P3, dP4, M1

PM 38727, left maxillary with broken P4 in crypt, dP4, M1
Trench 4, Unit 1, Level 1-2

TMM 41106-496, right P3
Trench 4, Unit 7, Level 1

PM 38740, 38741, two left P3s
Trench 4, Unit 7, Level 2

PM 38645, right P3

PM 38772, left P3 or right P3
Trench 5, Unit 6

TMM 41106-585, left P4

TMM 41106-587, right P3?

TMM 41106-588, right P3 or left P3

TMM 41106-5147, left P4

Comparative Material

Koomooloobooka Cave, southwestern South Australia

TMM 41230-108
Cave N-31, southwestern South Australia

TMM 41235-31
Nannup Cave, southwestern Western Australia

PM 4998, 5141, 5144, 5146, 5150-5152, 5154, 5161-5163, 5166, 5168, 5170, 5171, 5174,
5178, 5180, 5181, 5183, 5188, 5980, 5983, 25173, 25179

Description

Skull— The single well-preserved skull (TMM 41106-20; figs. 7A-D, 9A-C)
has all the characteristic features of B. lesueur noted by Finlayson (1958), Mar-
shall (1973), Tedford (1967), and Wakefield (1967). Some of these characters are
summarized in Table 12. The principal characters listed by Wakefield (1967)
are: (a) relatively short and narrow rostrum; (b) great inflation of auditory bul-
lae, with bulla length approximately 25% and bulla depth below the glenoid
fossa approximately 23% of the basal skull length (the values for the Madura
Cave skull are 28% and 26%); (c) large premolars, with P4 length approximately
14% of the basal skull length (12% in the Madura Cave skull); and (d) rapid
decrease in size from M1 to M4.

Cranial, mandibular, and dental measurements of the Madura Cave material
are compared in Tables 13-19 with the measurements of fossil samples from
Nannup Cave, southwestern Australia, and Lake Victoria, New South Wales,
and with Recent specimens from Victoria and southwestern Australia. The Ma-
dura Cave material also has been compared with measurements given by Ted-
ford (1967, tables 5-8) for a fossil sample from Lake Menindee and Recent
samples from central Australia, South Australia, southwestern Australia, and
Bernier Island. Except for the dimensions of the bulla, the dimensions of the
Madura Cave specimen are below the size ranges of all of these samples except
for the Recent one from central Australia. The dimensions of the bulla of the
Madura Cave skull are the same as or greater than the means of these dimen-
sions for all samples except for the Recent one from South Australia, and they
are within the observed ranges of that sample. The Madura Cave skull is almost
as small as one from the western Northern Territory described by Finlayson
(1957) as B. penicillata anhydra and later referred by Wakefield (1967) to B. lesueur.

Table 13. Comparison of cranial, mandibular, and dental measurements of Bettongia
lesueur from Madura Cave with those of Bettongia penicillata anhydra Finlayson.

Measurement

Condylobasal length

Basal length

Greatest length

Zygomatic breadth

Nasals, length

Nasals, greatest breadth

Interorbital constriction

Palate, length

Palate, inside breadth across M2

Diastema, P-P4

Incisive foramen, length

Rostrum, depth

Bulla, length

Bulla, maximum breadth

P\ length

P4, posterior width

M'-\ length

P4, length

P4, maximum width

M,_3, length

Depth of ramus to M2

Breadth of ascending process

* Data from Finlayson, 1957. t Labeled "maximum length" by Finlayson. It is not known
how this measurement was taken as the specimen is reported to lack the occiput,
t Estimate.

Madura Cave

B. penicillata

TMM 41106-20

.in/n 1//.1'

59.3

55.5

63.4

62.lt

39.5

37.4

23.2

23.7

10.6

9.5

13.4

12.2

31.34:

32.5

10.3

9.4

5.15

8.0

2.46

2.4

11.6

11.6

16.3

12.5

10.4

7.30

7.5

2.43

3.0

11.66

10.5

6.08

6.4

2.05

2.8

11.16

10.2

7.95

8.7

13.2

13.3

Table 14. Statistical data on upper dentitions of Bettongia lesueur from Madura Cave.

P3
dP4

M

M'

M

M-

M1

L

W

L
AW
PW

L
AW
PW

L
AW
PW

L
AW
PW

L
AW
PW

L
AW
PW

OR

4.41-5.47
1.90-2.96

3.27-4.10
2.66-3.12
2.69-3.57

7.30-8.73
2.20-2.74
2.43-2.89

3.80-4.56
3.50-4.33
3.80-4.71

4.18-4.86
4.26-4.64
3.95-4.41

Mean ± SE

4.90 ± .155
2.49 ± .132

3.68
2.87
3.27

7.84
2.51
2.63

4.02 ± .110
3.90 ± .113
4.04 ± .136

4.46
4.49
4.23

3.80
3.80
3.19

11.66

SD

41
35

CV (%)

8.37
14.06

7.21
7.69
8.91

37

38

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c

a.

u

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3 ""

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2P -
ID

39

Table 15.

Statisti*

:al da

P3

L

N
6

W

6

dP4

L

5

AW

5

PW

5

P<

L

3

AW

3

PW

3

M,

L

4

AW

4

PW

4

M2

L

8

AW

8

PW

8

M3

L

3

AW

3

PW

3

M4

L
AW
PW

Statistical data on lower dentitions of Bettongia lesueur from Madura Cave.

OR

3.80-4.94
2.28-2.51

3.50-3.72
1.98-2.20
2.54-2.93

6.08-6.84
2.20-2.51
2.05-2.66

3.65-4.10
3.04-3.34
3.27-3.72

3.88-4.56
3.57-4.56
3.57-3.95

3.57-3.65
3.57-3.80
2.96-3.42

M,

Mean ± SE

4.23
2.39

3.60
2.09

2.77

6.49
2.33
2.36

3.88
3.22
3.53

4.14 ± .085
3.90 ± .102
3.70 ± .046

3.62
3.72

3.24

11.16

SD

CV <%)

5.80
7.44
3.51

Table 16. Statistical data on upper dentitions of Bettongia lesueur from Lake Victoria
(data from Marshall, 1973).

dP<

M1

M2

M3

M<

M'

N

OR

Mean ± SE

L

5

4.8-5.3

5.02 ± .079

W

6

2.4-2.7

2.52 ± .054

L

10

3.3-3.9

3.60 ± .061

Max. W

10

2.8-3.4

3.15 ± .065

L

18

7.7-9.0

8.21 ± .097

Max. W

18

2.7-3.5

3.01 ± .062

L

49

3.7-4.6

4.16 ± .031

AW

45

3.4-4.5

4.93 ± .039

PW

47

3.6-4.6

4.03 ± .034

L

48

4.0-4.8

4.43 ± .029

AW

48

3.7-4.8

4.30 ± .034

PW

49

3.7-4.5

4.07 ± .028

L

35

3.6-4.5

4.00 ± .035

AW

34

3.1-4.2

3.81 ± .042

PW

34

2.8-3.6

3.25 ± .039

L

19

2.5-3.0

2.73 ± .044

AW

19

2.3-3.1

2.66 ± .055

PW

L

31

11.6-13.3

12.58 ± .093

SD

CV (%)

179

3.57

133

5.28

194

5.39

207

6.57

411

5.01

265

8.80

217

5.22

266

6.77

233

5.78

202

5.56

233

5.42

195

4.79

206

5.15

242

6.35

227

6.98

191

7.0

241

9.06

.518

4.12

40

Table 17. Statistical data on lower dentitions of Bettongia lesueur from Lake Victoria
(data from Marshall, 1973).

N

OR

Mean ± SE

SD

CV(%)

p3

L

36

3.7-4.7

4.28

±

.042

.253

5.91

W

39

2.2-2.7

2.44

+

.025

.154

6.31

dp4

L

41

2.9-3.8

3.37

±

.030

.195

5.79

Max. W

40

2.6-3.1

2.77

±

.021

.135

4.87

p.

L

48

6.1-7.6

6.80

±

.050

.346

5.09

Max. W

53

2.2-3.0

2.49

♦

.023

.165

6.63

M,

L

98

3.5-4.2

3.92

±

.019

.192

4.90

AW

89

3.0-3.8

3.33

±

.020

.192

5.77

PW

91

3.2-4.2

3.68

±

.021

.200

5.43

M2

L

87

3.8-4.8

4.42

±

.023

.214

4.84

AW

86

3.6-4.5

4.06

±

.024

.220

5.42

PW

84

3.3-4.5

3.95

±

.027

.245

6.20

M3

L

68

3.8-4.6

4.09

±

.030

.147

3.59

AW

64

3.6-4.5

3.94

±

.023

.185

4.70

PW

62

3.2-4.3

3.53

±

.029

.231

6.54

M4

L

38

2.8-3.6

3.17

t

.032

.200

6.31

AW

35

2.6-3.7

2.98

±

.040

.238

7.99

PW

29

2.0-2.8

2.34

±

.043

.229

9.79

M..3

L

59

11.2-13.2

12.50

±

.064

.493

3.94

Table 18. Statistical data on upper dentitions of Bettongia lesueur from Nannup Cave.

CV(%)

P3
dP4

M1

M

M'

M4

M1

L

W

L
AW
PW

L
AW
PW

L
AW
PW

L
AW
PW

L
AW
PW

L
AW
PW

N

2
2

2
2
2

6
6
7

13
13
13

12
12
12

10
10
10

3
3
3

10
3

OR

4.86-5.14
2.74-2.81

2.66-3.12
2.96-3.19

7.75-8.66
2.58-2.96
2.74-3.19

3.95-4.71
3.80-4.26
4.03-4.60

4.10-4.82
4.10-4.56
3.80-4.41

3.42-4.18
3.27-4.03
2.81-3.34

2.51-3.12
2.36-2.81
1.14-2.13

11.85-13.52
14.71-15.38

Mean ± SE

5.00
2.78

3.72
2.89
3.08

8.31
2.76
2.91

4.23
4.02
4.35

4.47
4.38
4.18

3.38
3.64
3.05

2.85
2.51
1.50

12.48
14.96

SD

.050
.039
.053

.055
.038
.046

.076
.070
.057

± .175

53

4.26
3.48
4.37

4.25
2.97
3.83

6.27
6.04
5.90

4.25

41

42

FIELDIANA: GEOLOGY

Table 19. Statistical data on lower dentitions of Bettongia lesueur from Nannup Cave.

N

dP,

P.

M,

M2
M3
M4

M,_3

M,_4

L
W

L
AW
PW

L
AW

PW

L
AW
PW

L
AW
PW

L
AW

PW

L
AW
PW

L
L

4

4
4

13
13
13

13

13
13

9
9
9

1
1
1

9
1

OR

4.41-4.94
2.58-2.89

3.24-3.80
2.20-2.51
2.66-3.04

6.38-6.84
2.51-2.66
2.28-2.54

3.72-4.26
3.27-3.57
3.57-4.03

4.03-4.86
3.95-4.33
3.88-4.26

3.80-4.33
3.80-4.18
3.27-3.50

11.32-12.62

Mean ± SE

4.78

2.77

3.46

2.32
2.88

6.69
2.59
2.39

3.96 ± .053
3.42 ± .025
3.82 ± .031

4.39 ± .067
4.14 ± .039
4.02 ± .036

3.95
4.01
3.39

3.04
2.89
1.98

12.10
15.26

SD

CV (%)

.19
.09
.11

.24
.14
.13

Upper Dentition. — The P3 is a serrate, bladelike tooth which is aligned with
the molar row (fig. 8A-D). It has six grooves on the labial side (table 12). This
agrees with the number of five to six reported by Marshall (1973, p. 106) for
the P3s of a Pleistocene sample from Lake Victoria and by Finlayson (1958, p.
256) for the P3s of a modern population from central Australia. The anterior
and posterior crown heights are approximately the same. In crown view the
tooth has an even oval shape with prominent cingula.

The dP4 is roughly square and molariform in structure, with four approxi-
mately equal-sized principal cusps (fig. 8A-D). The parastyle is large and lat-
erally compressed and joins with the paracone to form an anterior blade that
is aligned with the main ridge of the P3. The dP4 usually lacks the mesostyle
which is frequently present on the dP4 of B. penicillata, but PM 4786 (fig. 8A,D)
has a small mesostyle.

The P4 is elongate, bladelike, and usually straight, and like the P3, is aligned
with the molar row (figs. 7, 8). It has nine to ten grooves on the labial side
(table 12). This agrees with the number of eight to ten reported by Marshall
(1973, p. 103) and by Finlayson (1958, p. 266). Tate (1948, p. 266) reported nine
grooves for B. lesueur. Bettongia penicillata is reported by Finlayson (1958) to
have seven grooves.

The upper molars are square with four principal cusps. Their size decreases
markedly from M2 to M4; the relative sizes are M2 > M1 > M3 > M4. This is said
by Wakefield (1967) and Tate (1948) to be characteristic of this species.

The protoloph and metaloph are straight, with bulbous expansions on the
labial segments. The posterior crest of the protocone and the anterior crest of

Fig. 8. Bettongia lesueur from Madura Cave, Units 1 (post-Pleistocene) and 2 (late Pleis-
tocene). A,B/ PM 4786, right maxillary with P3, dP4, M12 in place, removed P\ and crypt
for M3, shown in labial and ventral views. The P4 is also shown in lingual view. C,D, PM
4787, right maxillary fragment with P\ dP4, exposed P\ M1, and crypt for M2, shown in
labial and ventral views. E-G, TMM 41106-117, left maxillary fragment with P-M2, shown
in lingual, ventral, and labial views.

43

Fig. 9. Bettongia lesueur from Madura Cave, surface (Recent) and Unit 1 (post-Pleisto-
cene). A-C, TMM 41106-20. Right ramus with I, P4-M3, and alveoli for M4, shown in
lingual (A), dorsal, and labial views. D-F, PM 4795. Left ramus of a juvenile with I, dP4,
P4 in crypt, M,, and M2 removed from crypts, shown in labial (D), dorsal, and lingual
views. The M2 is shown in anterior (left), crown, and posterior views.

44

H

1cm

Fig. 10. Bettongia lesueur from Madura Cave, surface (Recent) and Unit 1 (post-Pleis-
tocene). A-C, TMM 41106-26. Left ramus with I, P3, P4 unexposed in crypt, M,, M2 erupt-
ing, M3 in crypt, shown in labial (A), dorsal, and lingual views. D-F, PM 4796. Left ramus
with I, P3/ dP4, P4 in crypt, M, missing, M2 erupting, M, in crypt, shown in labial (D),
dorsal, and lingual views.

45

46 FIELDIANA: GEOLOGY

the hypocone are thick. They turn into the central basin, where they may or
may not join. The inturned crest from the hypocone usually is larger than the
one from the protocone. The crests of the paracone and metacone are sharp.
They also turn into the central basin, but they do not join. There usually is no
mesostyle (Tedford, 1967; Marshall, 1973), but PM 4786 (fig. 8A,B) has a small
mesostyle on dP4 and a trace of one on M1.

Lower Dentition. — The P3 is a straight, serrate, bladelike tooth which is aligned
with the molar row (fig. 10). In crown view it is somewhat oval, with cingular
shelves. One-fourth of our sample has five grooves on P3; the rest of the spec-
imens have six grooves. This agrees with the five to six grooves reported by
Marshall (1973, p. 105) and by Finlayson (1958, p. 256). Bettongia penicillata has
five grooves on P3 (Finlayson, 1958). The anterior and posterior crown heights
are the same.

The dP4 (figs. 9D-F, 10) is triangular in crown view because it lacks a pro-
toconid. The metaconid, which is the highest cusp, is laterally compressed and
has an anterior bladelike process which is a functional continuation of the P3.
The entoconid is higher than the hypoconid and is joined to it by a hypolophid.
The hypoconid is joined to the metaconid by a ridge. A postcingulum borders
a small cingular basin.

The P4 is straight, serrate, and bladelike and is aligned with the molar row
(figs. 9, 10). The posterior cusp is laterally compressed and about three times as
large as the other cusps on the blade. Crown height is uniform along the entire
length of the tooth. The blade has seven to nine grooves, with nine most
common. This is slightly less than the eight to ten grooves reported by Marshall
(1973, p. 106) and the nine reported by Finlayson (1958, p. 256). Bettongia pen-
icillata has seven grooves (Finlayson, 1958).

The lower molars (figs. 9, 10) are rectangular and slightly longer than wide.
The M4 may be triangular because of reduction of the hypolophid. The proto-
lophid is shorter than the hypolophid in Mu equal in size in M2, and longer
in M3. The lingual segments of both lophids are swollen. Crests run from the
protoconid and hypoconid into the central basin. The metaconid and entoconid
are joined by sharp crests at the labial edges of the teeth. The pro- and postcin-
gula are large and outline oval cingular basins.

Discussion

A comparison of dental dimensions of the Madura Cave sample (tables 13-
15) with the dental dimensions of fossil samples from Lake Menindee, New
South Wales (Tedford, 1967, p. 35), Lake Victoria (Marshall, 1973; tables 16, 17),
and Nannup Cave, southwestern Australia (tables 18, 19) and of Recent samples
from central Australia and South Australia (Finlayson, 1958, pp. 255, 256, 258,
264; summarized by Tedford, 1967) shows only minor differences. The Madura
Cave sample has slightly lower mean values for most measurements than those
of the other samples, although the ranges overlap considerably. The exceptions
to this rule are the widths of dP4 and M2 and the length of dP4, which have
larger mean values than do those of any of the other samples. The Madura Cave
sample is closest in size to the samples from Lake Menindee and Lake Victoria.

Only M1 and M2 have large enough sample sizes to make calculation of coef-
ficients of variation worthwhile. The coefficients are slightly higher than the

LUNDELIUS & TURNBULL: MADURA CAVE 47

corresponding values for the other samples (tables 13-19), probably because of
the small size of the Madura Cave sample.

Size comparisons of the Madura Cave sample with other samples of fossil
and recent B. lesueur are somewhat uncertain because of the small sample size
and wide age range of the Madura Cave material. The Madura Cave sample has
smaller average values for most measurements of the upper dentition than do
samples from Lake Menindee, Lake Victoria, and Nannup Cave (tables 14, 16,
18; Tedford, 1967, table 7). The samples from Lake Menindee, Lake Victoria,
and Nannup Cave are similar in size, but for a number of measurements, the
Lake Menindee sample has the highest mean values. The Madura Cave sample
also shows smaller mean values than the modern sample from central Australia
in the six measures available for comparison (tables 13, 14; Tedford, 1967, ta-
ble 5).

A decrease in size of modern examples of B. lesueur from the humid area of
southwestern Australia through southern South Australia to arid central Aus-
tralia has been reported by Finlayson (1958) and Tedford (1967). This size cline
does not seem to exist in the fossil samples. Pleistocene specimens from Lake
Victoria and Lake Menindee, both now in the arid zone, are as large as or larger
than Pleistocene specimens from Nannup Cave in the humid area of south-
western Western Australia. The Pleistocene specimens from Madura Cave, also
now in the arid zone, are as small as or smaller than modern specimens from
arid South Australia in spite of other indications that, at that time, the climate
in the vicinity of Madura Cave was more humid than it is today.

Bettongia lesueur was distributed through a wide range of environments at the
time of European settlement (Finlayson, 1958, p. 240). In the more humid parts
of its range, it was sympatric with B. penicillata. Bettongia lesueur is regarded by
Finlayson (1958, pp. 297, 298) as being competitively superior to both B. peni-
cillata and Caloprymnus campestris in arid areas. This advantage probably is due
in part to its burrowing habit, which the other two species lack (Marshall, 1973).
The much greater abundance of B. lesueur than of B. penicillata in the deposits
at Lake Menindee (101 B. penicillata to five B. lesueur; Tedford, 1967) and Lake
Victoria (119 B. penicillata to two B. lesueur; Marshall, 1973) has been attributed
by Marshall (1973) to this difference in adaptation to arid conditions. Caloprym-
nus is rare in the Lake Menindee collections (two specimens) and not recorded
from Lake Victoria. In contrast, the number of Madura Cave specimens defi-
nitely assignable to B. lesueur (20) is smaller than the number assignable to B.
penicillata (38), and Caloprymnus campestris is much more abundant than either
species of Bettongia.

Any attempt to account for these differences in representation of the two
species of Bettongia in the various samples must consider the fact that the bur-
rowing habit of B. lesueur must greatly increase the probability of remains of
this species being preserved in open sites such as Lake Menindee and Lake
Victoria. The numerous articulated skeletons of B. lesueur found at both of these
sites and the clumped distribution of bones of B. lesueur and of Lasiorhinus sp.,
another communal burrower, at Lake Menindee suggests that many of these
animals died in their burrows (Tedford, 1967, p. 23; Marshall, 1973). In contrast,
the principal agents for introduction of bones into the Madura Cave deposits
were avian and mammalian predators, which would bias the sample differently,
but which might more accurately reflect the relative abundance of the living
populations in the area.

48 FIELDIANA: GEOLOGY

Bettongia penicillata

Bettongia penicillata Gray, 1837

Hypsiprimnus ogilbyi Waterhouse, 1841

Bettongia gouldi Waterhouse, 1846

Material

Surface pickup

TMM 41106-27, left maxillary with P3, dP4, P4 removed from crypt, M1, M2-3 in crypt,
edentulous premaxillary (fig. 13A-C)
Trench 1, Unit 1, top 1 foot

PM 4789, left maxillary with P3 in crypt, dP4, P4 removed from its crypt, M1, crypt for
M2 (fig. 13G-I)

PM 4792, right ramus with I, P3, dP4, P4 in crypt, M, (fig. 14D-F)

PM 4794, left ramus with I, P3, dP4, P4 in crypt, M„ alveolus for M2 (fig. 14A-C)
Trench 3, Unit 2, Level ?

TMM 41106-6, left maxillary fragment with worn M1-2 (fig. 13D-F)

TMM 41106-62, left P4

TMM 41106-603, left M2

TMM 41106-5142, right P4 fragment
Trench 3, Unit 2, Level 4

PM 38688, left P4
Trench 3, Unit 2, Level ? or Trench 4, Unit 1

PM 38690, right M1 or M2
Trench 4, Unit 1

TMM 41106-533, right P3

TMM 41106-586, -605, two left M's or M2s

TMM 41106-614, right P3

TMM 41106-622, right P3

TMM 41106-623, right P3

TMM 41106-685, left P3

TMM 41106-686, right P3

TMM 41106-5144, broken upper molar
Trench 4, Unit 2, Level 1

WAM 78.3.10, left M' or M2
Trench 4, Unit 2, Level 2

PM 38691, 38692, two left P3s
Trench 4, Units 4-5

TMM 41106-18, left dP4

TMM 41106-648, left ramus fragment with P4-M, (fig. 14G-I)

WAM 78.3.12, left P3

PM 38694, left P4

PM 38695, right P3

PM 38696, right P4

PM 38697, left M1

PM 38698, right M2

PM 38701, left dP4

PM 38703, right M2 or M3

PM 38705, left M' or M2

PM 38736, right P3 or left P3

PM 38737, right P3
Trench ?, Unit ?, Level ?

PM 38715, right dP4

Comparative Material

Recent, southwestern Western Australia
FMNH 35325, 35330

LUNDELIUS & TURNBULL: MADURA CAVE 49

Nannup Cave, southwestern Western Australia

PM 5142, 5143, 5149, 5156, 5159, 5167, 5169, 5172, 5173, 5175-5177, 5222, 5232, 5973
5974, 5975-5977, 5979, 5981, 5982
Weebubbie Cave, southeastern Western Australia (north of Eucla)

TMM 41107-67
Cave N-31, southwestern South Australia

TMM 41235-97

Description

Skull.— Other than maxillary fragments with teeth, the only skull material is
the left side of the facial region of a juvenile skull (TMM 41106-27; fig. 13A-
C). Because it is a juvenile, comparisons of cranial characters must be made with
caution. The tooth row turns sharply linguad at the level of M2, suggesting that
the tooth rows converged strongly posteriorly. This is a diagnostic character of
B. penicillata (Finlayson, 1958, p. 284).

Upper Dentition. — The P3 is a broadly oval tooth with a laterally compressed
serrate ridge (fig. 13A-C). The base is bulbous and is widest about at the mid-
point. Cingula are absent or weak. The anterior end of the tooth usually is
turned outward with respect to the molar row. There are five grooves on the
labial surface. The crown is highest at the anterior end. PM 4789 does not fit
the foregoing description. It has a more elongate P3 with a crest which is slight-
ly curved labially at its anterior end. There is a prominent notch in the labial
cingulum and the lingual cingulum is prominent, with three cuspules which
are placed opposite the three posterior lingual ridges. Although these charac-
ters are reminiscent of B. lesueur, PM 4789 has been referred to B. penicillata
because it has seven grooves on the P4 and mesostyles on dP4 and M1.

The dP4 is molariform. The paracone and parastyle are joined to form a lat-
erally compressed, anteriorly projecting blade which is a functional continua-
tion of the P3 (fig. 13A-F). Crests from the paracone and metacone and from
the protocone and hypocone close the central basin on the labial and lingual
sides. Unlike M13, dP4 does not have a spur from the hypocone extending into
the central basin. A mesostyle is present in PM 4789 but not in TMM 41106-27
(although this specimen has a mesostyle on M1). Finlayson (1958) found the
mesostyle to be present only rarely on the dP4s of a sample from central Aus-
tralia.

The P4 is elongate, laterally compressed, and serrate (fig. 13A-F). The long
axis of the tooth is turned outward anteriorly with respect to the molar row.
The crest may be straight or curved labially at the anterior end. Seven grooves
are present on the labial side of the blade. The posterointernal cusp is com-
pressed anteroposteriorly, and is two to three times wider than, but lower than,
the posterior cusp of the blade. The posterior cusp of the blade usually is turned
outward. The tooth is constricted at the middle. The crown is highest at the
anterior end.

Of the upper molars (fig. 13), only M's and M^ can be positively assigned to
B. penicillata. The width of M1 and M2 is equal to or slightly greater than the
length. The protoloph and metaloph are nearly straight and have less promi-
nent bulges on their labial segments than do the lophs of B. lesueur. The lophs
and ridges are lower and the central basin is more open than in B. lesueur. A
mesostyle is present on M1. It varies from single and minute in TMM 41106-27
to complex and prominent in PM 4789.

50

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Fig. 13. Bettongia penicillata from Madura Cave, surface (Recent) and Unit 1 (post-
Pleistocene). A-C, TMM 41106-27. Edentulous premaxillary and left maxillary with P3,
dP", P4 removed from crypt, M1, M2, M3 in crypt, shown in lingual (A), ventral, and labial
views (each with inset of P4). D-F, TMM 41106-6. Left maxillary fragment with M\ M2,
shown in lingual (D), ventral, and labial views. G-I, PM 4789. Left maxillary fragment
with partly erupted P\ dP4, P4 removed from crypt, M', crypt for M2, shown in lingual
(G), ventral, and labial views (each with inset of P4).

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Fig. 14. Bettottgia penicillata from Madura Cave, Unit 1 (post-Pleistocene). A-C, PM
4794. Left ramus with I, P3, dP4/ P4 in crypt, M,, alveoli for M2, M3 in crypt, shown in
labial (A), dorsal, and lingual views. D-F, PM 4792. Right ramus with I, P3, dP4, P4 in
crypt, M,, shown in lingual (D), dorsal, and labial views. G-I, TMM 41 106-648. Left ramus
fragment with P4, M,, shown in labial (G), dorsal, and lingual views.

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LENGTH

LENGTH

Fig. 15. Bivariate graphs of PJs and P3s of Bettongia penicillata from Madura Cave and
Nannup Cave. A, Plot of P's and P3s from Madura Cave, showing probable assignment.
B, Plot of length vs. maximum width of P3s and one P3 of Bettongia penicillata from Nannup
Cave. Note the apparent reversal of position of the scatters of the two teeth. Symbols as
in A.

57

58 FIELDIANA: GEOLOGY

Lower Dentition. — The dP4 is triangular and lacks a protoconid (fig. 14A-F).
The metaconid is the highest cusp; it is laterally compressed and projects for-
ward to the posterior end of the P3. The entoconid is higher than the hypoconid
and is joined to it by a hypolophid. The metaconid and entoconid are connected
by a ridge. The posterior cingulum is small.

The P4 is a simple, laterally compressed blade with a straight crest (fig. 14).
The posterointernal cusp is either absent or reduced to a bulge on the lingual
side of the posterior ridge. The long axis of the tooth is turned outward ante-
riorly with respect to the molar row. There are seven grooves on the labial side
of the blade. The anterior part of the tooth is widest. As with P\ crown height
is greatest at the anterior end of the tooth.

Lower molars which are associated with sectorial premolars and which thus
can be positively identified as B. penicillata are rare in the Madura Cave mate-
rials. Only TMM 41106-648, PM 4792, and PM 4794 have M, in place (fig. 14).
The protolophid of the M, is shorter than the hypolophid. The ridge which
joins the protoconid and hypoconid is less prominent than it is in B. lesueur.
The lophids of M, are less bulbous than those of B. lesueur, and thus the central
basin is more open.

Discussion

The P3 and P3 of B. penicillata are so similar that differentiation of isolated
teeth is difficult. The length /width ratios of the four specimens which are still
in a maxillary or ramus differ between P3 (.56, .57) and P3 (.65, .69). The length/
width ratios of the isolated teeth tend to cluster around those of known posi-
tion, but there is no clear separation between the two groups. A bivariate scatter
diagram of length versus width does suggest a separation into two groups
which correspond to the groups formed on the basis of length /width ratios
(fig. 7A). This is the tenuous basis for the identifications of isolated P3s in this
paper. The plots of points representing each tooth are so close to each other
that a larger sample size might produce a wide zone of overlap.

A scatter diagram of length versus width for a sample of P3s of B. penicillata
from a late Pleistocene fauna from Nannup Cave in southwestern Australia (fig.
7B) yields a very different distribution. Even so, it appears that the Nannup
Cave P3s might be separable on this basis. The difference between the two
samples indicates that criteria for distinguishing between isolated P3s and P3s
will have to be developed independently for each sample.

Bettongia Species Indet.

Material

Trench 3, Unit 2, Level ?

TMM 41106-116, right ramus fragment with M2^3, alveoli for P4, M,, and M4
TMM 41106-602, right M2

TMM 41106-681, right M3 (possibly Potorous platyops)
PM 38687, right dP4 (probably B. lesueur)

PM 38716, right ramus fragment with unerupted M2 in crypt, proximal tip of alveolus
for I, alveoli for posterior roots of dP4, posterior end of crypt for P4, alveoli for M,,
small crypt for M3
PM 38717, left dP4

LUNDELIUS & TURNBULL: MADURA CAVE 59

Trench 3, Unit 2, Level 2

PM 38774, right M4
Trench 3, Unit 2, Level 4

PM 38689, left M,
Trench 4, Unit 1, Level ?

TMM 41106-564, left dP4

TMM 41106-565, right dP«

TMM 41106-567, -569, two right M,s

TMM 41106-571, broken right P*

TMM 41106-572, broken upper molar

TMM 41106-574, right M2

TMM 41106-578, left M2 or M,

TMM 41106-582, broken right upper molar or dP4

TMM 41106-591, left lower molar

TMM 41106-592, right M,

TMM 41 106-595, left M3

TMM 41106-596, left M2

TMM 41106-597, -599, two right M2s

TMM 41106-601, right M,

TMM 41106-604, right I, fragment

TMM 41106-606, right dP4

TMM 41106-607, left M2 or M3

TMM 41106-608, left dP4 (probably B. penicillata)

TMM 41106-610, right M,

TMM 41106-615, posterior half right P3 (probably B. penicillata)

TMM 41106-616, left M,

TMM 41106-617, left M2

TMM 41106-618, posterior half left P3 (probably B. penicillata)

TMM 41106-620, right M,

TMM 41106-621, left M2 or M3

TMM 41106-2828, left P,

TMM 41106-5143, upper molar (probably B. lesueur)

TMM 41106-5145, left dP4 (probably B. penicillata)

PM 38719, right M2 or M3

PM 38720, left upper molar

PM 38752, anterior half ?upper premolar

PM 38753-38756, four right I's

PM 38757-38760, four Ps

PM 38761-38768, eight Fs
Trench 4, Unit 2, Level 1

WAM 78.3.9, right M,

WAM 78.3.11, right dP4 (probably B. penicillata)

WAM 78.3.13, right M3

WAM 78.3.14, dP4 (possibly not Bettongia)

WAM 78.3.15, M3 or M2

WAM 78.3.16, right M2

PM 38709, broken left dP

PM 38724, right M, or M2

PM 38725, right M,

PM 38726, right M2

PM 38728, left dP4 (probably B. lesueur)

PM 38729, left dP4 or M'

PM 38730, broken right molar

PM 38769, edentulous maxillary fragment
Trench 4, Unit 2, Level 2

PM 38710, 38711, two left dP4s

PM 38731, right M2

PM 38732, right M3 or M2

PM 38733, left M2 or M'

PM 38734, left M3

PM 38735, left M,

60 FIELDIANA: GEOLOGY

Trench 4, Unit 2, Level ?

PM 38693, ramus fragment with dP4-M, (probably B. penicillata)

PM 38770, right M,

PM 38771, left maxillary with shattered tooth roots
Trench 4, Units 4 and 5

WAM 78.3.17, right M,

WAM 78.3.18, left M1

WAM 78.3.19, right M3

PM 36983-36985, three left I's

PM 36986, ?F (possibly other potoroine or phalangerid)

PM 36987, I2 (possibly other potoroine or phalangerid)

PM 36988, left P (possibly other potoroine)

PM 38699, left M2 or M3

PM 38700, broken right M2

PM 38702, right M, or M2

PM 38704, right M2

PM 38706, left M2

PM 38738, broken molar

PM 38739, right M2
Trench 4, Unit 7, Level 2

PM 38707, right M2

PM 38708, right M2 or M,

PM 38742, left M2

PM 38743, right M3

PM 38773, left upper molar
Trench 5, Unit 6

TMM 41106-629, left M4

TMM 41106-636, left dP4

TMM 41106-643, right dP4

TMM 41106-5146, left dP4

PM 38744, right I,

PM 38745, left M2 or M3

PM 38746, right M1 or M2

PM 38747, left M2

PM 38748, right M,

PM 38749, right M1 or M2

PM 38750, left M2

PM 38751, left M2 or M3
Trench, Unit, and Level data lost

PM 38713, broken right P4

PM 38714, left M2

CONCLUSIONS

The Madura Cave deposits contain four species of Potoroinae, Potorous pla-
tyops, Caloprymnus campestris, Bettongia lesueur, and Bettongia penicillata. Potorous
platyops and Caloprymnus campestris are not known to have occupied the Nullar-
bor Plain since the beginning of European settlement, although there is an
early report of Caloprymnus campestris by Tate (1879) with no supporting evi-
dence. The only part of Australia where four species of potoroines have been
reported as living animals is a small area near Albany in southwestern Austra-
lia. The species reported there are Potorous tridactylus, P. platyops, Bettongia le-
sueur, and B. penicillata. Potorous platyops was known as a living animal from two
localities in southwestern Western Australia, one near the town of Goomalling
(Calaby, 1954), and the other near the Pallinup River, 110 km northeast of
Albany (Ride, 1970). Both localities are in areas of higher average annual rain-
fall (20 to 30 inches [50 to 75 cm]) than the Madura Cave area, which has an

LUNDELIUS & TURNBULL: MADURA CAVE 61

average annual rainfall of approximately 10 inches (25 cm). Caloprymnus cam-
pestris has been reported from an extremely arid area in central Australia (Fin-
layson, 1932). The increased diversity of potoroines and the presence of now
allopatric species of that group are similar to the situation seen in the Dasyu-
ridae from this site (Lundelius & Turnbull, 1978). This supports the interpre-
tation, based on the dasyurid assemblage, of a climate with more available
moisture and more equability than the present climate in this area. The pres-
ence of all four potoroines in Unit 1 (CM date of 7470 ± 120 BP from the top 1
foot) demonstrates that the change that resulted in the loss of Caloprymnus and
Potorous took place some time during the last 7,500 years. This is consistent
with the conclusions regarding the changing climatic conditions which were
made in the earlier parts of this study based on Dasyuridae, Peramelidae, and
Phalangeridae.

ACKNOWLEDGMENTS

In addition to those individuals mentioned in earlier parts of this faunal
report, we wish to thank Dr. Larry Marshall of the University of Arizona for
allowing us to use his unpublished data on Bettongia penicillata and B. lesueur
from Lake Victoria, Mr. Zbigniew Jastrzebski for illustrations, and Ms. Elizabeth
Moore and Ms. Melissa Winans for typing and editing.

Financial support was provided by Field Museum of Natural History, the
Geology Foundation of the University of Texas at Austin, and by grants GB
975, GB 3729, and GB 7662 from the National Science Foundation.

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