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UN' RSITY OF
ILLIi^iS LIBRARY

nIt?HB^NA"CHAMPA'GN
NATURAL HIST. SURVEY

FIELDIANA
Geology

Published by Field Museum of Natural History

Volume 31, No. 1 February 14, 1973

The Mammalian Fauna of Madura Cave,
Western Australia

Parti

Ernest L. Lundelius, Jr.

Professor of Geological Sciences, University of Texas at Austin

and Research Associate, Field Museum of Natural History

and

William D. Turnbull

Associate Curator, Fossil Mammals, Field Museum of Natural History

INTRODUCTION

This is the first of several planned reports of the second investiga-
tion of the stratified deposits of Madura Cave, one of the few caves in
the Eucla Basin known to contain such deposits. As such, it is im-
portant for the establishment of a sequence. Much material was
obtained during this investigation and the collections of the earlier
investigation have also been used in the preparation of the present
report.

The deposits of Madura Cave were investigated in 1955 by
Lundelius. A brief description of the cave, its sediments and con-
tained fossils, and their importance in the determination of the faunal
history of the region was published (Lundelius, 1963). This estab-
lished that the deposits were stratified and that extinct marsupial
remains (Sthenurus) present in the lower of the two stratigraphic
units then recognized demonstrated a Pleistocene age for that unit.
Thus it indicated a potential for gaining a longer and more complete
record of late Pleistocene faunal elements and changes if a more ex-
tensive study and excavation could be made.

Madura Cave is located in the Eucla Basin, which occupies a
large (nearly 170,000 sq. km. or 70,000 sq. miles) area in southern
Australia bordering on the Great Australian Bight (fig. 1). The
basin is vaguely crescent-shaped, bounded on the south for over 500

Library of Congress Catalog Card Number: 72-9756 %
Publication 1160 1

•> FIELDIANA: GEOLOGY, VOLUME 31

miles by the Southern Ocean from near Penong on Fowlers Bay in
the east to Israelite Bay in the west. Inland its border reaches north
from Penong to Lakes Pidinga and Ifould, then to Ooldea where it
makes a decided bend westward, in a great sinuous swing that takes
it past Lakes Jubilee and Gidgi. From this point it takes a south-
erly course to Balladonia and Israelite Bay. Thus about two-thirds
of the Eucla Basin lies within Western Australia, and one-third
within South Australia.

The basement complex of the basin is reported to be composed of
granitic or high grade metamorphic rocks (Ludbrook, 1958a, b).
The basin is a broad, shallow embayment that is filled with onlapped
Cretaceous marine conglomerates and shales and by Cenozoic mar-
ine limestones. The latter (the Eucla group for the most part, which
is widespread and over 900 ft. thick at Madura) have a youthful
karst development with numerous caves and dolines. Hence this
constitutes one of the world's largest karst regions (Jennings, 1963).
The Eucla group is comprised of the lower and thicker Wilson's
Bluff Limestone and the upper and thinner (100 ft. or less) Nullar-
bor Limestone (Singleton, 1954).

Physiographically it is an area of extremely low relief broken only
by the Hampton Scarp which extends from near Eucla on the east
(where it forms a sea cliff — the Bunda Cliff) for approximately 150
miles west near Eyre where it again runs along the Southern Ocean.
This scarp divides the region into the low-lying Roe, or Eyre, Plain
(about 100 ft. above sea level) to the south and the far larger Bunda
Plateau, or Hampton Tableland, (about 250 to 600 ft. or more above
sea level) to the north. The first geologic, biologic, and physio-
graphic reports of the region were made by Tate (1879). The large
central portion of the Hampton Tableland constitutes the true
Nullarbor Plain, although this term is sometimes loosely used for the
whole region of the Eucla Basin.

There has been speculation and some controversy as to the nature
of the Hampton Scarp. Frost (1958) and others before him (Wool-
nough, 1933; David & Browne, 1950, v. 1, p. 538; v. 2, p. 548) have
considered it to be a fault scarp. Ludbrook (1958a) concluded on the
basis of paleontological correlations and Pleistocene depositional
evidence that the Nullarbor Limestone was missing from the seaward

Fig. 1. Map of region of the Nullarbor Plain in Western Australia and South
Australia, with detail map of Madura Cave area showing location of the cave, the
Hampton scarp, and other caves nearby on the Hampton tableland.

4 FIELDIANA: GEOLOGY, VOLUME 31

side of the scarp as a result of marine erosion, being replaced uncon-
formably by thin shelly Pleistocene marine deposits. Frost's inter-
pretation was based on joint patterns and lithic correlations from the
Madura Bores (number 1 below the scarp and numbers 2 and 3
above). Prior fault scarp postulates were only based on physio-
graphic evidence. Jennings (1963) has thoroughly reviewed the
various arguments, concluding that, "Further stratigraphical evi-
dence may be necessary before this issue can be finally decided; one
thing seems certain, however, namely that the Hampton Range
[Scarp] has been subjected to marine erosion and is no longer a sim-
ple fault scarp. Air photographs show that the western part of the
Hampton Range takes the form of a series of shallow, smooth curves
in plan, best interpreted as the wavecut bays of a former sea cliff."

The Eucla Basin is the southern part of the central arid region of
Australia. Most of this area receives less than 10 in. of rain per year
and is subjected to high evaporation rates. The climate of most of
this area falls into the BSh category of the Koppen and the E B'd
of the Thornthwaite classifications of climate (Trewartha, 1954).

This area is interesting from the standpoint of paleo-biogeogra-
phy because it is situated between the more humid coastal areas of
southeastern and southwestern Australia, and its aridity is a barrier
to the free exchange of faunal elements of the two humid regions.
The floral zonation is directly controlled by climatic zonation. A
number of studies of speciation patterns of various groups of Recent
and fossil vertebrates in southwestern and southeastern Australia
have shown that these two regions have exchanged faunal elements
across this area at one or more times in the past (Serventy, 1951,
1953; Main et al., 1958; Gentilli, 1961; and others). The living
fauna is a desert adapted one. Most of these cited studies agree that
the Nullarbor Plain was the probable route of this exchange, and that
the exchange or exchanges took place when the climate was more
humid than it is today. Most also correlate the humid periods with
glacial stages of the Pleistocene. There is, however, disagreement on
the time of the most recent exchange.

The fossil record, which can provide the basis for the reconstruc-
tion of the faunal and climatic history, is poorly known for the Null-
arbor Plain. It is confined at present to several pond deposits near
Balladonia on the western edge and to cave deposits in many places.
The former contain remains of a number of extinct marsupials which
are widely distributed in Pleistocene deposits in Australia, (Glauert,

LUNDELIUS AND TURNBULL: MADURA CAVE 5

1912; Merrilees, 1968). The latter contain remains of both extant
and extinct forms (Lundelius, 1957, 1963). While the assemblages
mentioned above demonstrate a more humid climate in the past,
there is little or no data that permit the construction of a sequence of
faunas or the correlation of the known assemblages with those in
other places. Merrilees (1968) has recently reviewed various inter-
pretations of the evidence relating to the Pleistocene and post-
Plesitocene climates of the continent in comparison with those of the
rest of the world.

DESCRIPTION OF MADURA CAVE

Madura Cave is located on the Roe Plain, six miles south of the
settlement of Madura, 110 miles west of Eucla (fig. 1). It is one of
the few known caves on the Roe Plain and is the only cave south of
the Hampton Scarp investigated by us. The cave system consists of
a shallow doline, an oval depression whose long axis is oriented
NW-SE, with two tunnels extending outward from its margins (fig. 2,
and see Frost, 1958) . The oval depression quite clearly is the product
of a collapse of the roof of a cave. Frost believed the cave to be
formed in the Nullarbor Is., but if the Ludbrook (1958a, b) assess-
ments are correct, it is developed in the Wilson Bluff Is. One tunnel
extends southwestward from the doline's southern end, the other
northwestward from its northern end. The southern tunnel is open
for 160 ft. It is 40 ft. wide and 7 ft. high at the opening. Its floor
consists of loose gray sand and is at the same level as the bottom of
the central depression. No excavations were carried out in this
tunnel.

The northern tunnel is much larger. It extends northwestward
for 275 ft. where it divides into two, one branch trending southwest-
ward, the other north-northwest. Each of these subdivide again into
smaller tunnels which gradually shrink until they cannot be entered.
The main part of this tunnel is about 20 ft. wide with little variation
(fig. 2) . A small tunnel leaves the main tunnel to the southeast 140
ft. from the entrance.

The floor of the main tunnel is 8-10 ft. lower than the surface of
the depression. It has a gentle gradient toward the back of the cave.
The floor has a small meandering channel cut into it to a depth of one
to two feet. This channel sends a small distributary branch into the
small southeast trending passage at the halfway point of the main
tunnel. The main channel continues on to the branching of the main

FIELDIANA: GEOLOGY, VOLUME 31

.REAR TRENCH (#5)

MAP OF

MADURA CAVE

010 \\ — APPROX. POSITION TRENCH # 2 (ELL-1955)

APPROX.

POSITION TRENCH #1 (ELL-1955)

ROCK HOLE = O
REFERENCE POINTS ==©

MAGNETIC

Fig. 2. Map of cave.

tunnel where it follows the larger southwest-trending branch. This
channel clearly carries water from the surface depression during
heavy rains.

STRATIGRAPHY

In 1955 Lundelius dug two test trenches in the main tunnel of
Madura Cave, numbered Trenches 1 and 2. In 1964 we added three
more trenches (fig. 2), numbers 3-5. These are precisely located with
respect to the cave entrance and to one another, as well as to various

LUNDELIUS AND TURNBULL: MADURA CAVE

SECTION C-C MADURA CAVE

ENTRANCE PIT SECTION PARALLEL TO LONG AXIS OF CAVE— #3

1 GRAY-BROWN SILT

(Tx 1145) 15.600 ± 250 YBP
2 RED CLAY AND SILT

580 YBP

jjy — (Tx 1142) 22.400 +

LIMIT OF y-Z^r^^If^^^^S^Z r^zQ^^-p-^Z:

EXCAVATION

4 WHITE SANDY SILT

5 RED CLAY

Fig. 3. Section C-C, Entrance Trench, 3.

other landmarks within the cave. The exact locations of Trenches 1
and 2 were not recorded and we were unable to relocate them pre-
cisely in 1964. Trench 1 lay about 80 ft. in from the entrance, near
Trench 4, and Trench 2 lay about E of Instrument Station 10 (fig. 2),
near Trench 5. Trench No. 3, the Entrance Pit, is located 40 ft. from
the entrance in approximately the center of the cave. Trench 4, the
Middle Trench, is 110 ft. from the entrance and extends out from the
northeast wall to the midline of the cave. Trench 5, the Rear
Trench, is located in the northern branch and is oriented parallel to
the axis of the branch.

The sequence in the northeast wall of Trench 3 (Sec. C-C; fig. 3)
from top to bottom is as follows:

1. Loose, gray-brown silt with many limestone fragments and
boulders up to 4 in. in diameter and abundant small bones and or-
ganic materials. This unit forms the floor of the main passage of the
cave. Its surface is irregular because of the incised drainage channel
which is filled with limestone boulders. This unit is 2 ft. thick.

2. Loose, red clayey silt with lenses of gravel and bones and
numerous limestone boulders. This unit is 2 ft. thick at this place.

8 FIELDIANA: GEOLOGY, VOLUME 31

Two C-14 dates1 based on bone are available from this unit. One
(Tx 1145) from the upper one foot is 15,600 ± 250 years B.P. The
other (Tx 1142) from the lower one foot is 22,400 ± 580 years B.P.

The bottom of this unit forms the roof of an extensive void or
opening, a "tunnel" 12 to 18 in. high. This tunnel which is formed
completely in the cave fill has an arched roof and a floor which slopes
gently toward the present entrance. The floor of the "tunnel" is
covered with limestone cobbles in a matrix of gray-brown clay and
silt. This grades downward rapidly into the more reddish-colored
material typical of unit 3.

3. Very loose, brownish-red silt with many limestone cobbles
and some bones. This unit is 23 in. thick.

4. Loose, almost white, sandy silt with a few cobbles and some
very fragile bones. Many of the cobbles are composed of limestone
fragments cemented together. This unit is 7 in. thick.

5. Friable, red clay with limestone cobbles and sparse bones.
There were ten inches exposed.

The sequence in the southeast wall of Trench 4, the main trench,
(Sec. A-A'; fig. 4) from top to bottom is as follows:

1. Loose gray-brown silt with numerous limestone cobbles and
boulders. The contact of this unit with unit two is irregular and is
marked by a 1-2 in. zone of relatively clean gravel. The top has ap-
parently been eroded off by the present drainage. As a result of these
irregularities, the thickness varies from 42 in. near the wall to 12 in.
in the center of the cave. A C-14 date (Tx 1146) of 7,470 ± 120 B.P.
was obtained from bone from the top one foot of this unit.

2. Loose, red to orange-red clay and silt with numerous cobbles
and boulders and abundant bones. Thickness is 24 in. Two C-14
dates based on bone are available for this unit. One (Tx 1140) from
the upper 6 in. is 18,990 ± 220 years. The other (Tx 1141) from 6 to
12 in. below the top of this unit is 20,000 ± 430 years.

3. Loose to tight, yellow orange sand with few cobbles. Bones
are not abundant. It is composed mostly of fine limestone fragments
and dust. It is tightly cemented in place and incorporates cobbles
and pebbles of the underlying unit where the cemented areas are in
contact with it. This unit dips slightly toward the center of the cave

1 All C-14 dates reported here were obtained from bone samples by the method
developed by Haynes (1968). They are based on the calcium carbonate contained
in the structure of the apatite crystallites of the bone.

LUNDELIUS AND TURNBULL: MADURA CAVE

ROCK HOLE

YELLOW -ORANGE GRITTY SAND

'•>*t"i RED-GRAY SAND AND GRAVEL
TIGHT SILT AND CLAY SEAMS

RED-ORANGE SILTY CLAY

(Tx1143) 37,880 ± 3,520 YBP

Fig. 4. Section A-A', Middle Trench, 4.

and the thickness varies from 6 to 8 in. near the wall to 0 in. near the
middle of the cave.

4. Loose, dark red clayey silt with less rock than overlying unit.
Thickness varies from 6 in. near the wall to 3 in. near the center of
cave.

5. Loose, reddish gray sand and gravel with a minor amount of
limestone dust and abundant bones. This unit has a "speckled"
appearance because of the presence of well-rounded particles of dark
carbonaceous limestone. The thickness varies from 6 in. near the
wall to 16 in. toward the center of the cave. A C-14 date (Tx 1144)
of 22,220 ± 570 years B.P. was obtained on bone from units 4 and 5,
most of it from near the contact of these units or at the interface.

6. Soft, white limestone powder with thin seams of orange clay.
Bones absent. Thickness varies from 27 in. near the wall to 15 in.
near the center of the cave.

7. Tight, reddish-orange, silty to sandy clay with few large rocks.
Fifteen inches were exposed. A C-14 date (Tx 1143) of 37,880 ±
3,520 years B.P. was obtained from bone from the lower one foot of
this unit.

tr.

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-I

0
0.

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0)

1—

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UJ

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rr
I-

LUNDELIUS AND TURNBULL: MADURA CAVE 11

In the east wall of Trench 5 (Sec. B-B'; fig. 5), parallel to the long
axis of the passageway, the following sequence is exposed :

1. Loose, brownish-gray sandy silt with limestone cobbles and
boulders. This is lenticular and appears to be the filling of a channel
which was cut 30 in. into the underlying sediments. To the north, it
forms the surface of the cave floor, thickens to approximately 2 ft.
and acquires thin layers of rock dust which thin and disappear south-
ward.

2. Loose, light orange to reddish-orange sand and gravel with a
minor amount of silt and many limestone fragments. It is about 1 ft.
thick. To the north it ends against large limestone boulders.

3. Loose, reddish-brown silt with no cobbles. This is a channel
fill which has been cut through by a subsequent channel of unit 1,
leaving this unit preserved only on the sides of the channel it fills.

4. Alternating layers of brown silt and white rock dust. The
strata in the upper part are up to 6 in. thick with lenses of coarser
material and bones. The lower layers are 1-2 in. thick. The rela-
tionship of unit 5 to this one may indicate that unit 4 is compound.
Unit 5 appears to be emplaced into the lower part of unit. 4. No
evidence of any change was seen in unit 4, aside from the change in
the thickness of the layers of silt and rock dust.

To the north this unit becomes coarser and ends among large
limestone boulders. The upper part overlaps these boulders and a
few of the individual layers extend into the silts characteristic of unit
5, suggesting that the upper parts of units 4 and 5 interdigitate in this
area. The bedding in the parts of both units 4 and 5 that underlie the
boulders is disturbed in a way that suggests that the boulders fell
from the roof of the cave into the sediments.

5. Loose, reddish-brown silt with few rocks. It is similar to unit
3. This unit rests largely on large limestone boulders.

6. Large limestone boulders in a matrix of reddish-orange clayey
silt. The base of the unit was not reached in the excavations.

Multiple channelings and other disturbances of the sedimentary
regime, so clearly seen in the walls of Trench 5, indicate a very com-
plex genesis of the sequence of beds in the rear of the cave. This ap-
pears to be in contrast to the simpler-looking sequences in Trenches
3 and 4. These are somewhat deceptive because the evidences of
disturbances in deposition (cavity, lenses, and channels), being near

Fig. 5. Section B-B', Rear Trench, 5.

12 FIELDIANA: GEOLOGY, VOLUME 31

horizontal and low-angled features, are less apparent. All such evi-
dences of a complex depositional history argue for extreme caution
when attempting to correlate strata between trenches. Hence no
correlation will be made for Trench 5 beyond noting that the brown
silt (unit 1, the uppermost channel) is similar to that of the top unit
in Trenches 1, 3, and 4: presumably unit 1 in each of these trenches
is approximately equivalent.

It is possible to make some limited correlations of stratigraphic
units between Trenches 3 and 4 on the basis of the C-14 dates. It is
probable that the uppermost brown silt exposed in both is the same
age. It is similar in both trenches and is very different from the
underlying material. The C-14 date of 7,470 + 120 years B.P. was
obtained from bone adjacent to Trench 4. No C-14 date on this unit
from Trench 3 is available. The red clay and silt, designated unit 2
in Trenches 3 and 4 is similar in both places and is probably equiva-
lent to the red soil reported for Trench 1 (Lundelius, 1963). In gen-
eral, the C-14 dates support a correlation based on lithology.

The discrepancies concern the top and bottom of the red clay and
silt, designated unit 2 in both trenches, and units 4-5 of Trench 4.
The difference in the age of the top of unit 2 is readily explicable in
terms of the erosion surface that separates the upper brown silt from
the underlying red clay and silt. The discrepancies in the ages of the
unit 2 lower boundaries are interpreted as the result of different ages
for the onset of deposition of unit 2 material in the two areas of the
cave. In Trench 4 we have no C-14 dates for the lowermost foot of
unit 2, but the date for units 4-5 indicates that the bottom of unit 2
in this trench is younger than the bottom of this unit in Trench 3.
Different materials are being deposited in different parts of the cave
today. Light-colored, fine-grained, granular material similar to that
of unit 6 of Trench 4 is accumulating, apparently by spalation from
the walls, along the sides of the cave in some cases. This is different
from the brown silt and clay that is being deposited by runoff from
the surface.

Correlations with the units exposed in the other trenches seem im-
possible at present with the exception of the upper brown silt and the
uppermost 6 in. of the underlying red clay in Trench 1 reported by
Lundelius (1963) . They are probably the same as units 1 and units 2
in Trenches 3 and 4. No correlations can be made for units below
unit 2 of Trenches 3 and 4.

The fact that all of the deposits in the northern tunnel are
topographically lower than the material in the central doline raises

LUNDELIUS AND TURNBULL: MADURA CAVE 13

the possibility that material in the northern tunnel may be in part re-
worked. This is not believed to be a serious problem because stra-
tigraphy and C-14 date chronology are consistent. Reworking may
have happened with an occasional specimen, however.

An analysis of the salts in the dry screen concentrates has been
carried out for us by Mr. Gordon F. U. Baker (personal communica-
tion, 1965). The brown silt of unit 1 contained .26 per cent chlorides
and 2 per cent sulfates; the concentrates from unit 2 contained .05
per cent chlorides. Although these results were obtained from dry
screen concentrates and may not be completely accurate indicators
of the salt content of the two materials, they are highly suggestive
of significant climatic differences at the times of deposition of the
two units. This will be discussed in more detail in a later publica-
tion on environmental interpretation in our series.

METHODS

Material was removed from Trenches 3-5 in approximately 6 in.
intervals within each stratigraphic unit, with one exception (unit 1)
for which a sample of the top 1 ft. was taken adjacent to Trench 4.
All matrix was dry sieved in the field and all concentrate larger than
1/50 in. was kept. The better specimens were picked in the field and
treated with shellac to minimize breakage and loss of teeth.

The map of the cave (fig. 2) was made with a Brunton compass
and tape and an open-sight alidade.

Tooth measurements were made as shown in Figure 6 using either
micrometer calipers graduated to .01 mm. or a microscope ocular
micrometer graduated into 100 units, which, with the optics usually
used, permitted direct reading to either .038 mm. or .051 mm. Inter-
polation permitted accurate readings to half of these values per unit.
Other measurements were made with vernier calipers graduated to
.1 mm. and capable of interpolation to about .01 with the vernier
scale.

The dental nomenclature in general follows the familiar Cope-Os-
borne terminology with additions taken from Hershkovitz's (1971)
summary of terms used by a great number of authors. The first use
of a term is followed by the Hershkovitz numerical designation given
in parenthesis.

Because the dental morphology of many of the taxa of Australian
mammals is poorly known, we have figured dentitions of Recent
species in cases in which this aids the identification and description of

LUNDELIUS AND TURNBULL: MADURA CAVE 15

the fossil material. The drawings were made by Dr. Tibor Parenyi
from camera lucida drawings of the specimens. They are accurate as
to scale, dimensions, and proportions, and are shaded to show details
of surface texture and appearance, and to de-emphasize distracting
cracks, breaks, or other artifacts.

The bivariate scatter diagrams are plotted to the same scale for
each group of organisms to facilitate comparisons (see figs. 12, 13, for
examples) .

ABBREVIATIONS

Abbreviations used are as follows: FMNH or FM — Field Museum
of Natural History, Recent Mammal Collection; PM — Field Mu-
seum fossil mammal collection; MVZ — Museum of Vertebrate Zool-
ogy, University of California, Berkeley; TMM or UT — Texas
Memorial Museum, University of Texas at Austin1; WAM — Western
Australian Museum; L — length; W — width; AW — anterior width;
PW — posterior width; Tr. L — trigonid length; Tal.L — talonid length.
Other abbreviations are either in common use or are defined where
used in the text.

SYSTEMATICS

Class Mammalia

Subclass Theria

Infraclass Eutheria (Sensu VandeBroek, 1961, 1964)

Cohort Marsupiata (Sensu Turnbull, 1971; = Metatheria)

Order Marsupicarnivora (Ride, 1964)

Dasyuridae

Phascogalinae

Genus and species indet.

The fauna contains this indeterminate taxon, a pigmy antechinus,
which is close to Planigale ingrami and Antechinus maculatus in size
and most of its dental morphology, but which differs from both in
several ways that prevent an assignment to either genus.

1 Fossil vertebrates at Austin were formerly in the collections of the Bureau
of Economic Geology and (in earlier reports) bore the prefix BEG. They have been
transferred to the Texas Memorial Museum and will henceforth bear the prefix
TMM. An integral part of every TMM catalogue number is the five-digit
locality designation. That for Madura Cave is 41106-. In addition, recent mam-
mals in the UT or TMM collection have an M prefix designation.

16 FIELDIANA: GEOLOGY, VOLUME 31

Material. —

Entrance Pit (Trench 3), Lower red unit, under collapse (unit 3)

TMM 41106-660, right mandible with MT.
Middle Pit (Trench 4), unit 2, level 2

TMM 44106-661, -662, edentulous left mandibles with alveoli for

TMM 41106-663, right mandible with M^ and alveoli for rest

of cheek teeth and C.
WAM 72.3.6, right mandible with MT_^ and alveoli for other cheek

teeth and C.

TMM 41106-667, -668, edentulous mandibular fragments with
alveoli of last tooth or two.

TMM 41106-741, left M^ (fig. 8)

TMM 41106-742, left MT

TMM 41106-743, right M^ or 7

TMM 41106-744, right ramus fragment with MT

TMM 41106-745-6, two edentulous right ramus fragments

PM 25596, right ramus with M7 and alveoli for M^ and MT

PM 25597, right ramus with M7_T (fig. 9B)

PM 25710, right ramus fragment with M7

PM 25713, right ramus fragment with MT or M^

WAM 72.3.7, left mandible with M^ and alveoli for PT and other
molars

PM 26189, edentulous right ramus with alveoli for M^_T

PM 26193, left MT

PM 26194, -5, a right ramus fragment with alveoli for four teeth

and a left with alveoli for three, respectively.
PM 26268, left MT
PM 26269, right M^ or *
PM 26270-1, two edentulous right ramus fragments.

Units 4-5

WAM 72.3.3, right maxillary with M- and alveoli for other molars
TMM 41106-696, left ramus with PT-M7 and alveoli for other

cheek teeth (fig. 9A)
TMM 41106-699, left mandible with M^_T and alveoli for rest of

cheek teeth through C,

SMINTHOPSIS
CRASSICAUDATA

AND
ANTECHINOMYS
SPENCERI

PLANISALE C.F.
INGRAMI

MADURA
SPECIES
INDET.

ANTECHINUS
MACULATUS

Fig. 7. Diagram of condition of crowding of premolars of certain phascogales
as reflected in their alveolar patterns. Shown are the Madura Planigale-\\ke form
and several other small phascogales which are compared to it.

18 FIELDIANA: GEOLOGY, VOLUME 31

TMM 41106-740, left M^ (fig. 8)

WAM 72.3.1, edentulous right max.

TMM 41106-748, edentulous left max.

TMM 41106-749, -750, edentulous right mandible fragments

TMM 41106-751, -752, 753, edentulous left mandible fragments

TMM 41106-754, right MT

PM 25621, left maxillary fragment with P*-* and alveoli for C,
P^ and M1 (fig. 8)

PM 26190, left maxillary with M^ and alveoli for M^ and MA (fig.

8)
PM 26191, right ramus fragment with MT and alveoli for P7_T and

M¥.7
PM 26192, right ramus fragment with Mg and part of M7 and

alveoli for MT

PM 26216, left M-l

PM 26272, edentulous left max. with alveoli for C-M-

PM 26273, -274, -275, -276, edentulous left ramus fragments

WAM 72.3.4 and WAM 72.3.5, edentulous right ramus fragments

WAM 72.3.2, right M1

Unit 7, level 2
TMM 41106-659, right mandible with PT, MT

Descriptions. — This is the smallest and one of the rarest dasyurids
represented in the Madura collection. Only ten fragments were re-
covered that preserve any part of the maxillary bone or upper denti-
tion and, fortunately, one of them (PM 25621) shows the condition
of the root pattern of the upper premolars. The pattern corresponds
to that of A. maculatus, with P-* being crowded out of alignment,
rather than with that of Planigale c.f. ingrami in which the pre-
molars are all aligned (fig. 7A) .

The Pa and PA are each elliptical in outline with a single prominent
central cusp that is rounded anteriorly and ridged posteriorly (and
worn in PM 25621, fig. 8B). The posterior ridge runs to a prominent
cingular cuspule and a weak cingulum surrounds each tooth.

The M-l's (TMM 41106-740 and PxM 26216) are triangular in out-
line, with very small paracones. In both, the metacone is much the

* PL or P2. The first of the premolar teeth, whatever its true homology is.

Fig. 8. The Madura Planigale-\ike form. The right upper dentition in buccal
(A) and occlusal (B) views, based upon four of the fragmentary specimens that
contribute the most to knowledge of the maxillary and its dentition. Shown, left
to right are PM 25621 and PM 26190, the maxillary fragments with Pa-* and Ma
and TMM 41106-740 and -741, isolated M± and M2.

20 FIELDIANA: GEOLOGY, VOLUME 31

largest primary cusp, and the protocone is low and intermediate in
size (fig. 8) . The paracone-stylocone crest (stylocrista) portion of the
eocrista is very short, as in most phascogales, due to the very narrow
anterior part of the stylar shelf. The posterior half of the stylar shelf
is high and posterolaterally expanded. There is a well-formed ante-
rior cingulum which ties the protocone to the parastyle at the poste-
rior edge of that cuspule. No distinct conule is present, but this
anterior cingulum is inflected near the base of the paracone in a
manner suggestive of a vestigial protoconule. The posterior cingu-
lum also ties to the protocone, but shortly after leaving that cusp it
becomes very weak and continues in that manner past the base of the
metacone almost to the posterolateral corner of the tooth. In this it
is like both A. maculatus and P. ingrami (figs. 10, 11) and differs from
most Sminthopsis crassicaudata and S. murina in which it rarely
reaches as far as the posterior side of the metacone.

The M* (TMM 41106-741) is a deeply worn tooth that is closer in
its overall shape to that of the M- than to that of the M- (fig. 8) . It
is intermediate in the degree of development of the parastylar shelf.

The M^ in both specimens (TMM 41106-680 and PM 26190) is
worn and the relative development of the secondary cusps cannot
be determined with certainty. The tooth has the normal dasyurid
triangular shape but the protocone is smaller relative to the re-
mainder of the cusps than is the case in other dasyurids and it lacks
the posterior bulge seen in many dasyurids (fig. 8). The paracone is
smaller and lower than the metacone. The mesostyle (stylar cusp
C of Bensley, 1903) is the only stylar cusp preserved. It is about
the same size relative to the whole tooth as in other small dasyurids
(Planigale and Antechinus), but is less pronounced than in Sminthop-
sis and Antechinomys spenceri (AMNH 15012). The labial (ecto-
loph) margin is notably straighter with the pre-mesostyle notch
shallower than in Planigale cf. ingrami (AMNH 160308) or Smin-
thopsis crassicaudata (UT M-839). In this it is more like A. macu-
latus. There is a small procingulum in the region of the antero-
external root that extends lingually from the parastyle down to the
base of the tooth where it is weakly joined to the anterior crest from
the protocone.

The lower jaws and dentition are represented by more and bet-
ter, though also fragmentary, materials. Specimen TMM 41106-696
is the most complete of the lot (fig. 9A). In no case are incisor or
canine teeth preserved and, of the premolars, only the last (PT) is

4-1106 -696

PM 25597

Fig. 9. The lower dentition of the Madura Planigale-like form, seen in oc-
clusal and lingual views. A. Left mandibular fragment with P? through Ms, TMM
41106-696. B. Right mandibular fragment, PM 25597, with M3_T.

FIELDIANA: GEOLOGY, VOLUME 31

*J®

Fig. 10. Antechinus maculatus, FM 64350.
of right maxillary and its dentition.

A, B. Buccal and occlusal views

known in any of the specimens. In all specimens in which the evi-
dence of premolar condition is preserved by teeth or alveoli (TMM
41106-663, PM 26188, TMM 41106-666, TMM 41106-696, TMM
41106-699, and PM 26191 and TMM 41106-659) there is crowding
in this region. But unlike the condition of crowding in Recent Plani-
gale ingrami, in which PT is consistently crowded out of line and is
reduced considerably, here it is the anterior-most premolar that is
consistently out of line while the PT is usually aligned or just slightly
out of line in some specimens. Figure 7B diagrams these conditions
for P. ingrami, A. maculatus, the Madura specimens discussed here,
and other small phascogales. The depth of the horizontal ramus is

Q
23

FIELDIANA: GEOLOGY, VOLUME 31

tN-f»

TPi^f

Fig. 11. Planigale ingrami, AMNH 160308.
of the right maxillary and upper dentition.

A, B. Buccal and occlusal views

variable, but never very deep. The inferior dental canal is usually
divided so that the anterior foramen is at the level of P^ (or this may
be further divided as in PM 26191, with reduced openings in the
region of P^_7 and P^_T) and the posterior foramen is beneath MT_^,
often under the posterior root of MT.

The PT has a sub-elliptical outline in crown view. It is made up
of a primary cusp that lies somewhat anterior to the center of the
tooth. A ridge leads anteriorly from its apex to its base where it

26 FIELDIANA: GEOLOGY, VOLUME 31

joins the cingulum. Posterior to the apex there is a similar crest
that descends to the broadened cingular shelf with its cuspule. Wear
rapidly produces a flat surface on this crest, a surface that broadens
with increased attrition. The cingulum completely surrounds the
tooth at its base. It is weak anteriorly, and pronounced posteriorly,
especially postero-labially.

The MT is a triangular tooth with the protoconid and metaconid
about equal in height and placed very close together. The para-
conid is very much smaller and forms the anterior end of the tooth.
The talonid is basined. It is bordered labially by the hypoconid,
which is about one-half the height of the protoconid. It is bordered
lingually by a low, rounded ridge which joins the metaconid and the
hypoconulid. The posterior margin of the talonid is concave and
broadly "V"-shaped. The hypoconulid is a low cusp in TMM 41106-
666 located at the posterolingual corner of the talonid. It is the
posterior-most cusp of the tooth. The hypoconid and the hypo-
conulid are joined in TMM 41106-666 by a ridge that forms the
posterior margin of the talonid basin. The hypoconulid in TMM
41106-660 is a slightly elongate blunt cusp. It is not joined to the
hypoconid. The talonid basin is thus open posteriorly. There is no
entoconid in either TMM 41106-666 or TMM 41106-660. A post-
cingulum extends from the base of the hypoconid up the posterior
face of the tooth and merges with the hypoconulid.

The Mg and M^ have the normal dasyurid structure. The trigo-
nid is well developed. There is a deep, narrow cleft developed in
the paracristid (I') ridge that connects the paraconid and proto-
conid. The talonid is basined and is wider than long. As in the MT
there is no entoconid. The talonid is proportionately shorter than
in most other dasyurids. Its posterior edge is straight and parallel
to the posterior edge of the trigonid except at the posterolingual
corner where it turns abruptly posteriorly to join the hypoconulid.
The anterior cingulum extends upward from the antero-labial side
of the base of the protoconid but does not reach the anterior edge of
the paraconid. It ends abruptly where the postero-lingual corner of
the preceeding tooth is in contact with the paraconid. The posterior
cingulum extends upward along the posterior face of the talonid and
joins the hypoconulid.

The only MT is in PM 25597. It has a trigonid that is nearly the
same as that of M7 in its proportions (fig. 9B), being only slightly
smaller and more compressed laterally. The protoconid is much the
largest cusp in both height and bulk. It is followed by the meta-

LUNDELIUS AND TURNBULL: MADURA CAVE 27

conid which is rather columnar and stands nearly two-thirds as high
as the protoconid. The paraconid has a broad base but it barely
reaches to two-thirds the height of the metaconid in its worn state.
Since this cusp shows more wear than the others, it probably was
nearly as high as the metaconid in the unworn tooth. Both crests,
paracristid (I') and the combined epicristid-centrocristid (II' and
la'"), have a carnassial notch and groove. The posterior part of
the centrocristid (lb'") joins this main crest of the epicristid at its
base, medial to the groove. The talonid is laterally compressed,
progressively toward the rear. It has a well-developed hypoconid.
Antero-lingual to the hypoconid there is an elongated shallow basin
delimited by hypoconid, centrocristid, metaconid, and a low cingular
crest, apparently equivalent to the combined postmetacristid, disto-
cristid, and entocristid (I", VI, and V in the Hershkovitz symbolism)
There is no entoconid or hypoconulid. The anterior cingulum is
well developed with a good parastylid and another more labial
stylid, the two surrounding the hypoconulid of the M^.

Discussion. — Pigmy antechinuses are the smallest of the dasy-
urids and are among the smallest-sized mammals. They are not
well-represented in collections and are poorly known as to their
dental morphology, their ecology, and their taxonomy.

A comparison of the Madura Cave specimens with those of Re-
cent species shows many similarities, such as size and the tendency
to crowd the premolars. Comparison also reveals a number of dif-
ferences that indicate that the Madura Cave form is different from
the described Recent species. In general, the cheek teeth of the
Madura Cave form are slightly larger than their counterparts in
Recent Planigale ingrami, especially PT and M^_T (figs. 12, 13; tables
1, 2). Comparison with MT_^ of Antechinus maculatus is very close
(fig. 12). However, PT is much longer in the Madura Cave form than
it is in either Planigale ingrami or Antechinus maculatus. The MT
is also larger than that of Antechinus maculatus but the difference is
not as great as with Planigale ingrami.

It is clear that the Madura Cave material cannot be referred to
any of the described species of pygmy antechinuses. In addition,
it is clear, as stated by Ride (1970), that the generic grouping of the
species of pygmy antechinuses is still in doubt. For these reasons
we are not assigning the Madura Cave material to a genus or species.

General faunal and environmental significance of the presence
of this taxon in the Madura Cave deposits will be deferred until the
discussion in the final section of our series on the Madura Cave. It

D #•

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ex.

LUNDELIUS AND TURNBULL: MADURA CAVE

3.6 3.8 4.0 4.2 4.4 4.6 4.8 s.O

Length M 1 _ 4

Fig. 13. Bivariate graph comparing Planigale ingrami (circle), Antechinus
maculatus (square), and the Madura Cave Planigale-like species (star). Length of
lower molar tooth row is plotted against length of P? in each case. Individually
each of the lower teeth of the Madura form is, or tends to be, slightly larger than its
counterpart in the other species. This results in a cumulative difference seen in
these plots so as to give complete separation of the taxa. Wavy lines in the
P. ingrami plot show specimens for which only the MT-4 length value was avail-
able. The dashed line of the Madura species plot is an estimate of the probable
limits of the range of the MT.? length values. It is based upon the cumulative
engths of the smallest of each of the molars (MT + M2 etc.) for the shortest limit,
and the largest of each for the longest limit. The star is at the position of the mean.
In each case a 93 per cent factor was applied to accomodate for the overlapping of
the teeth in the series. This factor was chosen because it applied to both of the
other taxa shown, and hence gave the best indication of what could be expected in
the Madura form. The only P^'s available are both the same length, so there is no
way of estimating the range of variability from the sample in that case.

should be noted that Madura Cave is over 1,000 miles from the clos-
est known occurrence of any living pigmy antechinus as follows:
Antechinus maculatus in the extreme southeast corner of South
Australia; Planigale ingrami in north and central Queensland and
north of both Western Australia and the Northern Territory; Plani-

Fig. 12. Bivariate graph comparing proportions of the lower cheek teeth of
Planigale ingrami, Antechinus maculatus, and the Madura Cave Planigale-like form.
Width measurements used are anterior widths of molars and maximum widths of
premolars.

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32 FIELDIANA: GEOLOGY, VOLUME 31

TABLE 2b. Measurements of lower teeth of Planigale ingrami for
comparison with those of Antechinus maculalus and the Madura form.

Species and

Planigale :

ingrami

Trench,
Unit and Lc

ivel

FMNH AMNH

AMNH

Specimen numbers

66973

160066

160307-8

160311-2

Mean

Side

Distance from

front of F

'xtO

front of MT

.38

.08

.19

.22

L. MT_4

4.03

3.89

3.75

4.03

3.96

4.22

3.98

PT or 2

.55

.57

.53

.55

.36

.42

.34

.38

.46

.40

.70

.72

.68

.61

.69

.42

.46

.38

.49

.46

.49

.45

.44

.30

.48

.43

.32

.27

.34

.42

.34

.95

.99

.95

1.01

.99

1.03

.99

AW.

.51

.57

.46

.55

.59

—

.54

PW.

.57

.59

.51

.61

.58

1.18

1.14

1.10

1.14

AW.

.68

.65

.68

.72

.69

PW.

.62

.68

.61

.65

.68

.65

1.16

1.10

1.05

1.18

1.14

1.16

1.13

AW.

.68

.70

.72

.80

.76

.73

PW.

.57

.53

.63

.57

.68

.59

.98

.95

.93

1.05

.99

1.10

1.00

AW.

.58

.53

.61

.57

.68

.58

PW.

.27

.17

.23

.25

.19

.23

.22

gale tenuirostris in south-central Queensland and northeast New
South Wales.

ACKNOWLEDGEMENTS

The following people generously provided space, facilities, and
information: Prof. H. Waring, Dr. A. R. Main, Dr. J. Shield, of the
University of Western Australia; Dr. W. D. L. Ride, Dr. D. Mer-
rilees, of the Western Australian Museum; Profs. A. J. Marshall, J.
Warren, Dr. A. K. Lee, of Monash University; Dr. J. Jennings of
the Australian National University, Canberra; Mr. J. Calaby, of
CSIRO, Canberra; Mr. K. Quartermaine and Mr. H. Wheeler, of
the School of Mines, Kalgoorlie, W. A., and Mr. G. F. U. Baker,
Research Officer, Main Roads Department, Western Australia.

LUNDELIUS AND TURNBULL: MADURA CAVE 33

Prof. Warren, Dr. and Mrs. Lee, and Mr. L. Van der Velde pro-
vided assistance in the field, as did our wives.

The following persons provided technical assistance in the prepa-
ration and cataloguing of the material and manuscript preparation:
Mrs. M. Littlejohn, Miss Ethel Butler, Miss Susan Deutsch, Mr. E.
Krish, Mr. J. Joy, Mr. and Mrs. D. Witter, Mr. and Mrs. M. Col-
lins, Mr. B. Davidson, Mrs. K. Cooper, Mr. F. Corral, Dr. T. Per-
enyi, Mrs. W. Reinders, Mrs. R. Keppler, and Mrs. W. Krueger.

We are especially grateful for the assistance and encouragement
of our wives in every stage of the work.

The work was made possible by the financial support of Grants
GB 975, GB 3729, GB 7662 from the National Science Foundation;
Field Museum of Natural History; and from the Geology Founda-
tion of the University of Texas. The writing was done while Lun-
delius was on a research leave sponsored by the University Research
Institute of the University of Texas.

The Radiocarbon Laboratory of the University of Texas ran the
C-14 dates. We thank Mr. S. Valastro, Director of the laboratory,
for this important service.

REFERENCES

Bensley, B. A.

1903. On the evolution of the Australian Marsupialia; with remarks on the
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David, T. W. E. and W. R. Browne

1950. The geology of the Commonwealth of Australia. Edward Arnold & Co.,
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Frost, M.J.

1958. Jointing associated with the Hampton Fault near Madura, W. A. Jour.
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Gentilli, J.

1961. Quaternary climates of the Australian region. Ann. N. Y. Acad. Sci.,
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1912. Fossil remains from Balladonia in the Eucla Division. Rec. W. Austral.
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Haynes, C. V.

1968. Radiocarbon: Analysis of Inorganic Carbon of Fossil Bone and Enamel.
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34 FIELDIANA: GEOLOGY, VOLUME 31

Hershkovitz, P.

1971. Basic crown patterns and cusp homologies of mammalian teeth, pp. 95-
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Jennings, J. N.

1963. Some geomorphological problems of the Nullarbor Plain. Trans. Roy.
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Ludbrook, N. H.

1958a. The stratigraphic sequence in the western portion of the Eucla Basin.

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1879. The natural history of the country around the head of the Great Austral-
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LUNDELIUS AND TURNBULL: MADURA CAVE 35

Turnbull, W. D.

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VandeBroek, G.

1961. The comparative anatomy of the teeth of lower and nonspecialized mam-
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