FIELDIANA
Geology
Published by Field Museum of Natural History
Volume 31. No. 2 August 29. 1975
The Mammalian Fauna of Madura Cave,
Western Australia
Part II
Ernbst L. Lundbuus. Jr.
Propbssor or gbolooical Sciences, University op Texas at Austin
AND
RESEARCH A8S0CUTB. FIELD MUSEUM OP NATURAL HISTORY
AND
WiLUAM D. TURNBULL
CURATOR. Fossil Mammals, Field MusEim op Natural History
AND
Research AssoaATK Texas Memorial Museum
INTRODUCTION
This work constitutes Part II of a series of reports on the
Madura Cave fauna. It continues the systematic section, Order
Marsupicamivora, begun in Part I (Lundelius and Tumbull, 1973).
Included are accounts of the Madura Cave representatives of the
genera Sminthopsis and Antechinomys.
The lack of detailed description and illustration of the
dentitions of most living taxa of Marsupicamivora severely hampers
paleontological study. To remedy this situation, we have continued
the procedure initiated in Part I of describing and illustrating the
dentitions of certain of the living species, as well as those of the
Madura fossils. The description of the remainder of the fauna,
analysis of its composition, and the paleoenvironmental and
zoogeographic implications will be dealt with in subsequent parts of
this volume.
Section I dealt with the regional setting of the cave, the
stratigraphy of the deposits, the C-14 dates, and the small
phascogaline allied to Planigale or Antechinus maculatus. Thus far.
Library of Congress CtUalog Ctud Number: 72-97564
Publication 1209 87
38 FIELDIANA: GEOLOGY, VOLUME 31
none of the four taxa described (Sminthopsis crassicaudata, S.
murina, and Antechinomys spenceri treated below, and the
Planigale-\ike form of Part I) show any significant evolutionary
change throughout the sequence. Small sample size of all taxa
except A. spenceri rule out the chances of our being able to detect
small changes if they do exist.
Abbreviations, measurements, and tooth element terminology
are either those in standard usage or they follow the form and
procedures given in the methods section of Part I, unless otherwise
noted.
SYSTEMATICS
Class Mammalia
Subclass Theria
Infraclass Eutheria (Sensu VandeBroek, 1961, 1964)
Cohort Marsupiata (Sensu Turnbull, 1971; = Metatheria)
Order Marsupicarnivora (Ride, 1964)
Dasyuridae
Phascogalinae
The Madura Cave fauna contains several other phascogales in
addition to the minute, indeterminate form of pygmy antechinus
near Planigale ingrami and Antechinus maculatus treated in Part
I. These are {1) a complex of three small species (treated here)
currently put into the genera Sminthopsis and Antechinomys,
which are so similar in size, proportions, and dental morphology
that separation of their skeletal and dental remains is exceedingly
difficult, and (2) six species belonging to the genera Antechinus,
Phascogale, Par antechinus, Dasycercus, and Dasyuroides.
Generic separation of Sminthopsis and Antechinomys on the
basis of cranial features rests largely upon differences in degree of
development of the auditory bulla (larger in Antechinomys) and the
following characteristics of the mandibles (fig. 1). In Sminthopsis
the masseteric fossa is broad and flares upwards, the condyle is
relatively low (only slightly raised above the occlusal area of the
dental battery), and the angular process, while elongated and
arched, is relatively less developed, so that its tip and the tip of the
coronoid process are about equidistant from the condyle. In
Antechinomys the ascending ramus has a more horizontal
inclination (about 60° to occlusal plane in contrast to about 70° in
Sminthopsis), the masseteric fossa is narrow, and its front and back
LUNDELIUS AND TURNBULL: MADURA CAVE
39
SnuiiKoptii Mwrmo
AM»cKino«r« )onie*r
Anrvchinomyi ftp*nc«
Fig. 1. Outline drawings of medial views of mandibles of two of the living species
of Sminthopsis (above), and of the two species of Antechinomys (below). Inserts show
the lateral views of angle and ascending ramus of each. Compare: (/) breadth of the
ascending ramus, broad in Sminthopsis, narrow in Antechinomys (see the oblique
arrows and spread angle of the fossae); (2) height of the condyle above the occlusal
area of the cheek teeth (proportionally higher in Antechinomys than in Sminthopsis;
posterior- most vertical arrows); and {3) ratio of A = distance from condyle to tip of
coronoid process, to B " distance from condyle to tip of angle. (A/B is > '/i and
approaches 1/1 in Sminthopsis while in Antechinomys A/B < '/4 and approaches '4).
The extent of expansion of the angular process is emphasized by the other set of
vertical arrows.
edges are nearly parallel; the condyle is high absolutely and relative
to the tooth row; and the angle is delicately built, highly arched,
and attenuated to accommodate the large bulla of this genus, and
thus its tip is much farther removed from the condyle than is the
tip of the coronoid process. On dental criteria alone, generic
separation of Sminthopsis and Antechinomys is more obscure.
Within the genus Sminthopsis, two groups' can be recognized
on the basis of the lower molars. In one of these, here designated
'These groups merely comprise species that have in common one condition of
entoconid form and development. They are not meant to be formal taxonomic groups
for they are presently baaed upon only this one character, but this is not to deny that
they may have taxonomic implications.
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LUNDEUUS AND TURNBULL: MADURA CAVE 41
group A (comprised of S. crassicaudata, S. macrura, S. rufigenis, S.
stalkeri, and S. larapinta), each of the lower molar teeth excepting
M4 possesses a well-developed, usually high columnar entoconid as
a characteristic feature. In the other, group B (comprised of S.
murina, S. hirtipes, S. psammophilla, S. leucopus, S. longicaudata,
and S. granuUpes), the lower molars either lack entoconids entirely,
or, if present, they are small to minute and low, usually
anteroposteriorly enlongated. According to Archer (pers. comm.),
Sminthopsis murina shows greater variation in the development of
the entoconids than we have observed, but, in general, its
entoconids are not as large as those of the species within group A.
In both groups the crest from the hypoconid (post-metacristid or
I") extends far lingually then swings sharply posteriorly to join the
hypoconulid. In group A it runs onto the side, or at least the base,
of the entoconid before it turns toward the hypoconulid, and with
wear becomes joined to the entoconid. In group B there is no
entoconid-postmetacristid fusion, even when entoconids are
relatively well formed; instead there is usually a low subsidiary
diagonal ridge that runs into the center of the talonid basin from
the point of flexure of the postmetacristid toward the hypoconulid,
and this ridge may even become somewhat accentuated by wear.
Unfortunately for the worker confined to teeth, these lower molar
features of group B are all found also in the lower molars of species
of Antechinomys.
For the upper dentition there is no parallel dichotomy based
upon such a clearcut discrete feature as entoconid development of
the lower molars; instead, we have only subtle proportion
differences without clear-cut separations.
At the species level, both within the genus Antechinomys and
within the two species groups of Sminthopsis, size and dental
profjortions of the lower teeth serve to distinguish some species
(figs. 2, 3, 4, tables 1-15). For Antechinomys, dental proportions of
the living A. spenceri and those of A. laniger differ from one
another only slightly (fig. 4). The Madura fossil representatives of
this genus also differ slightly from both of the living species, but
they show a closer affinity to A. spenceri than to A. laniger. For
species within group A, S. crassicaudata is the smallest, and
thereby separable. For the sp>ecies within group B, not only are size
and dental proportions close, but the lower teeth are very much like
those of Antechinomys.
SMINTHOPSIS SR
OUP A
Fig. 3. A. Bivariate graphs showing length (abscissa) X anterior width (ordinate;
maximum width for the premolars) dimensions of the lower cheek teeth for a series of
samples of species in Sminthopsis group A. The Madura Cave sample of Sminthopsis
crassicaudata is shown at the bottom and a Recent sample is just above it. Other
related species are shown for comparison. Note both extent of variation for each
tooth and the patterns of the series of plots for each taxon as one proceeds from P2 to
M4 along the series. A key to the symbols is given in Figure 3B. Specimens measured
are listed in Appendix 2.
42
Pig. 3. B. Comparable graphs to those shown in A represent a series of species of
Sminthopsis group B. The Madura Cave sample of Sminthopsis murina is at the
bottom and the Recent samples of two of the subspecies of that species are
immediately above it. As in A, some other related species are also shown for
comparison. Specimens measured are listed in Appendix 3.
43
44
FIELDIANA: GEOLOGY, VOLUME 31
ANTECHINOMVS
Fig. 4. Bivariate graphs comparable to those of Figure 3 for samples of the two
Recent species of Antechinomys together with that of the Madura Cave sample
(bottom). With the Madura sample the points are not included as there are too many
for clarity — instead the clouds are tightly drawn so as to delimit each set of plots
and the N for each is given along with the tooth symbol. For key see Figure 3B.
Specimens measured are listed in Appendix 4.
The lower teeth of Sminthopsis murina and Antechinomys
spenceri are so similar that were it not for association with discrete
osteological differences, such as the shape of the masseteric fossa,
their separation would be almost impossible. Some scatter diagrams
of measurements of lower molars do show separate groups that
correspond with the groups based on the shape of the masseteric
fossa in those specimens in which it is known. This permits the
assignment of most of the ramal fragments with teeth (but which
lack the diagnostic masseteric fossa or any part of the ascending
ramus) and possibly even isolated teeth, so that only a small
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Pig. 5. Bivariate graphs of length (abscissa) x width (ordinate) for each of the
lower premolars of the Madura Cave specimens in the Antechinomys-Sminthopsis
murina group. Teeth in jaws with a broad fossa are shown as filled .squares, those in
jaws with a narrow fossa are shown as filled circles. Teeth in partial jaws thought to
be of broad fossa tyf>e are each shown as a dot with a b subscript, those in partial
jaws thought to be narrow fossa type are each shown as a dot with an n subscript.
Very worn teeth are indicated by a dot with a « subscript. Isolated teeth or thoae in
jaws which do not preserve enough of the angle and ascending ramus for an
assessment of the form of the fossa are shown as a dot. Specimens included are listed
in Appendix 5.
For comparison, the Recent specimens are also included: open square -
Sminthopsis murina murina; square with vertical line - Sminthopais murina
fuUginoaa; open circle - Antechinomys laniger; circle with verticle line -
Antechinomys apenceri. The Recent specimens are listed in Appendices 2-4.
45
46
FIELDIANA: GEOLOGY, VOLUME 31
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Fig. 6. Bivariate graphs (length x anterior width) for each of the lower molar
teeth in the Madura Cave Antechinomys-Sminthopsis murina samples. Symbols as in
Figure 5. Specimens included are listed in Appendix 6.
proportion of such remains cannot be assigned and must be carried
incertae sedis. All such assignments depend upon the closeness of
fit of the plots of the unknown specimens to those of the modern
species and the "known" series of the fossils (see the series of
bivariate scatter diagrams on figs. 5, 6, 7). The procedure is as
follows:
Scatter diagrams of various pairs of cheek teeth measurements
were made of all teeth remaining in mandibles that, because they
lack the crassicaudata-type of entoconid, are thought to be
assignable to either Antechinomys spenceri or Sminthopsis murina
(figs. 5, 6, 7). In each case those associated with a broad masseteric
fossa are indicated by a black square. Those associated with a
LUNDEUUS AND TURNBULL: MADURA CAVE
47
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Fig. 7. Bivariate graphs of trigonid length x anterior width (— trigonid width)
for each of the lower molar teeth in the Madura Cave Antechinomys-Sminthopsis
murina samples. Symbols as in Figure 5. Specimens included are listed in Appendix 7.
narrow masseteric fossa are indicated by a black circle. All others
are plotted as points, with severely worn teeth further designated
by a "w." Teeth in fragmentary mandibles that preserve some
evidence of masseteric width and can, therefore, be at least
questionably assigned are further indicated by subscripts: n for
narrow, b for broad.
Scatter diagrams of length versus width of all premolars
associated with molars with the small or absent entoconids from
Madura Cave show a tendency to form two groups (fig. 5). When
48
FIELDIANA: GEOLOGY. VOLUME 31
Fig. 8. Diagram of manner of taking upper molar measurements and of detail of
the labial border of the ectoloph. A. Schematic outline of an upper molar showing
measures used. Abbreviations as follows: P-M = paracone-metacone length; Pr =
protocone length; A W = anterior width; P W = posterior width; PSW =
parastylar width (parastyle-paracone width); MSW = metastylar width (metastyle-
metacone width). The "thegosed" wear facet of the protocone of S. crassicaudata
(see text) is indicated by diagonal hatchure.
B C. The two basic forms of ectoloph margin of M2-3 In the Madura Cave fossils
the deeply notched margin (B) characterizes Sminthopsis, especially S.
crassicaudata, and the nearly straight margin (C) characterizes Antechinomys.
the specimens in which the ascending ramus is preserved are
considered, it is clear that although there is overlap, the premolars
of those which can be identified as Antechinomys on the basis of
narrow masseteric fossae are somewhat larger than those identified
as S. murina on the basis of broad masseteric fossae.
Scatter diagrams of the length versus anterior width of the
lower molars show much the same pattern (fig. 6), but the Ma's and
M4's of Antechinomys are proportionately wider than those of S.
murina.
LUNDEUUS AND TURNBULL: MADURA CAVE
49
Fig. 9. Three bivariate graphs in which various combinations of measurements of
M''8 (in mm.) belonging to animals in the Sminthopsis-Antechinomys assemblage
from the Madura Cave fauna are given. In each case, the features measured are
indicated along the abscissa and ordinate axes. Note that the two Sminthopsis
species usually overlap one another broadly, but that Antechinomys, being slightly
larger, only overlaps S. murina and the overlap is to a lesser degree. This sort of
pattern is repeated in a number of combinations of dental measures, and in a few
cases, as seen in Figure lOA, E, there is no zone of overlap between the two genera.
Large circles represent the Sminthopsis teeth, dots the Antechinomys teeth. Open
circles, all of which bear a tail indicative of the presence of a thegosed wear facet on
the posterolingual side of the protocone, represent S. crassicaudata. Filled circles,
which with one exception (531) lack the thegosed facet, represent S. murina. The dots
represent probable Antechinomys spenceri. A few specimens that occasionally, or
even repeatedly, are found to plot in the overlap zone between the genera are tagged
by their last three catalogue numbers as their assignment may be somewhat less
certain. Abbreviations as in Figure 8. Specimens plotted are listed in Appendix 8.
50
61
52 FIELDIANA: GEOLOGY, VOLUME 31
Scatter diagrcims of lengths of trigonid versus anterior wddth
(=trigonid width) of the molars show that the Mi's of
Antechinomys are slightly larger than those of S. murina (fig. 7).
The Ma's, Ma's and M4's of Antechinomys have sUghtly shorter and
wider trigonids than their counterparts in S. murina. In the Ms
there is no overlap of the clouds of points of the two species, and
the specimens that are not classifiable on the basis of the masseteric
fossa fall unequivocally in one or the other of the groups. The
scatter diagram of the M2 shows nearly as clear a separation of the
two groups, while those of Mi and M4 show considerable overlap.
Identification of upper dentitions and maxillary fragments
within this complex of species is even more difficult, for there
usually are no such clearcut, discrete structures as columnar
entoconids or broad versus narrow masseteric fossae upon which to
base identifications. With the Madura teeth, the combination of
dental measurements with several somewhat discrete and usually
distinguishable features (fig. 8) definitely helps separate the upper
teeth of Smmthopsis from those of Antechinomys. Features that
distinguish the genera are (1) the detail of the form of the buccal
edge of the ectoloph of the upper molars, especially M^, and (2) the
size and proportions of the molars (width, symmetry, relative length
of the eocrista).
Size and proportions overlap somewhat, but they are consistent
enough within the Madura sample that they afford a basis for
separating the two genera. In Sminthopsis, M^^ are narrower and
more symmetrical in respect to protocone position and form of
metastylar area than are the counterparts in Antechinomys where
greater breadth and an anteriorly protruding protocone condition
usually makes for fairly easy recognition. Within the available
Recent comparative materials, however, neither of these features is
as clearcut.
The most consistent proportional differences of both the
Recent and fossil samples in the upper molars between Sminthopsis
and Antechinomys are the lengths of the anterior and posterior
segments of the eocrista (parastyle-paracone width and metastyle-
metacone width) relative to other parts of the teeth. In the upper
molars of Antechinomys these two segments are longer relative to
other dimensions of the teeth than in Sminthopsis. This is clearly
shown in a number of scatter diagrams that involve these
characters (figs. 9, 10). The elongation of these parts of the eocrista
LUNDEUUS AND TURNBULL: MADURA CAVE 53
is correlated functionally with the relatively wider trigonids in the
lower molars of Antechinomys.
The form of the buccal edge of the ectoloph also appears to be
consistent for the Madura materials. Within the Madura sample
there are two basic forms of margin (fig. 8b, c) that correspond to
the two genera as here differentiated. These do not hold for the
Recent materials of these same taxa in which the condition \s
variable or may even be reversed. This disconcerting apparent
inconsistency may be explained by the nature of the sampling of
both the Recent and fossil materials. The variable Recent samples
of each species are composed of specimens drawn from discrete
remote local populations, while those with consistent patterns are
from single local populations in which one or the other variant of
this morphologic feature appears to have been selected for. The
Madura fossils represent a single local fauna, and it appears that
the taxa considered here have maintained their local morphologic
identities throughout the 30,000-year time span represented by the
specimens.
The species assignment of the upper molars of the Sminthopsis
materials is a much more difficult problem. As is shown in Figure 2,
Recent specimens of Sminthopsis crassicaudata are smaller than
other species of Sminthopsis. Presumably the same relationship
holds in the Madura fossils. Also the presence or absence of a
peculiar sort of facet' in some ways similar to a thegosis-type of
wear facet (Every and Kiihne, 1970) on the posterolingual side of
the protocone of M'^ suggests another means of separating the two
species. This wear facet, as in thegosis facets, has parallel
microstriae which result from contact with the well-developed
entoconid of the opposing lower molar. Such "thegosed" facets are
present on the Recent specimens of S. crassicaudata if the
individual is old enough to show wear elsewhere in the dentition.
Unfortunately, some attrition may also be caused by the food,
causing the facet to be variably formed. Older specimens of S.
murina and Antechinomys frequently show indistinct facets formed
'This facet forms on the gently oirved side of the protocone where at first it does
not relate to the sharply curving surface of a crest, although with time and enough
wear it usually comes to do so. Hence, for much of its existence, there is no distinct
cutting edge involved with such a facet, and even though the thegoeis type of
microstriae do form as a result of tooth- to- tooth contact, the tooth sharpening
aspects of true thegosis are missing until wear brings the facet and crest into
proximity. To call attention to this distinction we refer to such facets as being
"thegoeed."
64 FIELDIANA: GEOLOGY, VOLUME 31
by food attrition, but they lack the "thegosed" microstriae that
result from enamel contact.
The smallest Madura specimens have "thegosed" facets, but
there is extensive overlap in size with those that lack them (figs. 9,
10, tables). This is similar to the relationship between the Recent
specimens of S. crassicaudata and S. murina. On this basis, those
specimens with a "thegosed" facet on the posterolingual surface of
the protocone are assigned to S. crassicaudata, and those that lack
it are tentatively assigned to S. murina with the realization that
some S. crassicaudata may be included.
Sminthopsis Thomas, 1888
Sminthopsis crassicaudata (Gould, 1844)
This small phascogale, like the pygmy antechinus, is also a
scarce element of the fauna. There are 12 specimens representing
the lower jaws and mandibular dentition, and seven maxillaries and
upper dentition £is follows.
Material. —
Mandibles and Lower Dentitions
Trench 1, Unit 1, Top 1'
PM 25525, right ramus with P2, P4, Mi, M4 and alveoU for other
cheek teeth
WAM 72:3.8, right ramus with P2-M2 and alveoli for other cheek
teeth (fig. 12)
PM 25528, right ramus with P2, P4, M2-4 and alveoH for other
cheek teeth
PM 25529, right ramus with P2, Mi, and M4 and alveoh for other
cheek teeth
PM 25530, right ramus with M3-4 and alveoH for other cheek teeth
PM 25534, left ramus with P2-M1, M4 and alveoU for other cheek
teeth
PM 25535, left ramus with C, P2, P4-M4 and alveoli for other
cheek teeth (fig. 13)
PM 25537, left ramus with P2-M4 and alveoli for other cheek teeth
Trench 2, Unit 2?
PM 25268, right ramus with Ms and alveoli for other cheek teeth
LUNDEUUS AND TURNBULL: MADURA CAVE 66
Trench 3, Unit 2
TMM 41106-521, right ramus with P3-M2 and alveoli for other
cheek teeth
WAM 72.3.9, right ramus with M2-4 and alveoli for other cheek
teeth
Trench 4, Unit 1
TMM 41106-755, right M3 (or M2)
Maxillaries and Upper Dentitions
Trench 1, Unit 1, Top 1'
PM 25639, left maxillary fragment with M^^
Trench 3, Unit 2
PM 26139, right maxillary h-agment with M^-*
Trench 4, Unit 2, Level 1
PM 25744, right maxillary with M'^ and all premolar alveoli (fig.
14A, B)
Trench 4, Unit 2, Level 2
WAM 72.3.10, right maxillary fragment with M^^ alveoli of other
molars, and P^"*
PM 25604, left maxillary fragment with P*-M^ and alveoli of
canine and other premolars (fig. 14C, D)
Trench 4, Units 4-5
PM 25620, right maxillary fragment with M^*^
Trench 4, Unit 7, Level 2
PM 25630, left maxillary fragment with M^, alveoli for other
molars, and P*
Descriptions. — Lower molars of the animals assigned to this
species are readily recognizable because of their distinctive, high,
pillar-like entoconids, as can be seen in examples of Recent (fig.
lie, D) and fossil specimens (figs. 12A, B and 13A, B). Compare
these also with Sminthopsis murina and Antechinomys spenceri
(figs. 15C, D; 18C, D). This distinctive morphologic feature of the
main lower molars (it is variable on M4) may become quite reduced
by wear. Even on a very worn tooth, wear facet evidence indicative
56
FIELDIANA: GEOLOGY, VOLUME 31
Fig. 11. Dentition of Sminthopsis crassicaudata, Recent specimen, FMNH 34723,
shown enlarged approximately S^A times. Right upi>er dentition in (A) lateral and (B)
occlusal views.
of the original presence of this well-formed cusp is apparent. Hence,
if a specimen preserves one of the lower molars with an entoconid of
such relatively large proportions, it can be sissigned to a member of
Sminthopsis group A, and within that group size and proportions
permit one to separate the smallest S. crassicaudata from some of
its sister species (S. rufigenis and S. larapinta). From others, where
proportions are nearly identical to those of S. crassicaudata {S.
macrura and S. stalkeri), zoogeographic considerations are all we
have, and they thus assume prime importance.
As noted earUer, the presence of a relatively wide masseteric
fossa helps to distinguish Sminthopsis from Antechinomys, but it
does not afford the means of separating the various Sminthopsis
d-s
67
58 LUNDELIUS AND TURNBULL: MADURA CAVE
species from one another — or even the species group with certainty,
although Sminthopsis crassicaudata does have a slightly higher
condyle than S. murina in the few specimens available to us (fig. 1 ).
Where lower molars are involved, S. crassicaudata is readily
identified by the entoconids, in spite of the fact that dental
dimensions and proportions are extremely close both to the other
taxon of Sminthopsis in the fauna and to Antechinomys.
The mandible has the usual phascogaline form. There is an
elongate, gently-tapered, and downwardly-arched horizontal ramus
which bears three incisors, a short, somewhat premolariform canine,
three small premolars (Pi or Pa, Pa, and P4) and four molars. There
is a vertical, elongated ligamentary symphysis which runs from the
tip of the jaw back to the level of the last premolar. The position of
the mental foramen is variable on the lateral surface of the ramus.
It lies beneath Mi or even M2, with subsidiary ones occasionally
under the premolars. The ascending ramus has a broad, upward-
flaring masseteric fossa and an upward- and backward-curving
coronoid process that is somewhat hooked at its tip. It also has an
articular process which, together with the posteroventral edge of
the masseteric fossa, continues the arc of the curve of the ventral
edge of the horizontal ramus in a bony strut that runs to and
supports a flattened articular condyle. An inflected, delicate,
attenuated, and arcuate angular process protrudes ventromedially
from the arched lower edge of the jaw. The mandibular foramen
opens posteriorly on the medial side just above the junction of the
angle of the horizontal and ascending rami.
The Madura Cave material referred to S. crassicaudata
resembles Recent specimens in the following ways: (i) the
masseteric fossa is relatively broad, ranging from 24°-40°, usually
about 30°-35°; (2) the condyle is in a relatively low position; (5) and
as a result of 1 and 2, the mandibular condyle is fairly evenly
positioned between tips of angular and coronoid processes (fig. 1);
{4) it has the distinctive high, columnar entoconids on M1-3 (often
with a trace on M4); and (5) details of dental proportions
correspond as shown in the graphs (fig. 3A).
No specimen preserves any of the incisor teeth, nor aie there
any complete jaws, but WAM 72.3.8 and PM 25535 between them
are representative (figs. 12, 13).
The lower canine is relatively low and elongated in crown view
and its tip stands only slightly above the tips of the premolars;
FIELDIANA: GEOLOGY. VOLUME 31 59
hence it is somewhat premolariform. The three premolars are oval
in occlusal outline. They are in line and are nearly the same size,
the middle one, Pa, being slightly larger than the others. In each,
the primary cusp is stout and high and is located over the anterior
part of the tooth. Each is ridged anteroposteriorly, especially behind
the main cusp. There is a developed posterior cingulum with a
cuspule that is joined to the axial crest and a smaller anterior
cingulum with a small central cuspule.
The lower molars have trigonids that are larger and stand
higher than their talonids. The protoconid is the largest trigonid
cusp in each, followed in order by metaconid and paraconid in M1.2.
In M34, paraconid and metaconid are subequal. In M13, the
hypoconid is a stout, well-formed cusp about equal in bulk to
paraconid and metaconid, but it arises from a lower base level. The
distinctive pillar-like, attenuated entoconids already have been
mentioned. Those of M2 and M3 are nearly as tall as the paraconid
of the following tooth when unworn and are almost round in cross-
section. That of Ml is smaller, and the cusp is very small or absent
in M4. The entoconid and hypoconid, in most specimens with some
wear, appear to be joined by a postcristid in Mi. 3, and the
hypoconulids are much lower but distinct and posteriorly projecting
and joined to the posterior cingulum. In unworn teeth, the
postcristid runs to the base or side of the entoconid where it turns
sharply toward the hypoconulid. It may be weakly separated from
the entoconid by a tiny elongate valley, which is readily obliterated
by a slight amount of wear. In M4 the talonid is nearly as long as
the trigonid, but its cusp and crest development is variable. Each
lower molar has a well-developed, steeply-inclined anterior
cingulum. In M24, between its highest point and the nearby
parastylid, there is a notch that receives the preceding hypoconulid
in an interlocking manner. In Mi the cingulum runs to the
paraconid, the parastylid being absent. Well-developed posterior
cingula are present in Mi 3.
The most complete maxillaries in the collection (PM 25744 and
PM 25604, fig. 14) between them preserve P*-M*. In PM 25604, the
maxillary bone is nearly complete, lacking only bits from its
posterior edge. Also, the area medial to the lacrymal and orbit and
extending to the region of the junction of the frontal and nasal
bone is missing. It appears that the snout was more vaulted and
slightly larger than in our Recent materials, and it had a decidedly
larger infraorbital foramen that opened above the anterior part of
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62
FIELDIANA: GEOLOGY, VOLUME 31
Fig. 14. Sminthopsis cf. crassicaudata, PM 25744, from Madura Cave. A, B. R.
maxillary with M '"' and alveoli of the premolars, shown in occlusal (A ) and labial
(buccal )(B) views.
M^ In this last regard, it more closely resembles FM 60116, a
captive specimen derived from South Australian stock. The
premaxillary suture is a shallow, grooved structure with its medial
flange protruding more anteriorly than the lateral one. The
posterior limit of the left anterior palatal foramen is preserved; it
lies close to the midline, directly opposite the center of the anterior
premolar. The anterolateral edge of the main palatal vacuity Ues
near the medial root of M\ slightly behind the position of its
counterpart in TMM M-839 and more like that in FM 34723,
modern specimens from Western Australia. The rest of the palate is
missing behind this point except for the part directly supporting
molars 1-3. The alveolus of the canine is oval with its longitudinal
axis being about equal to the combined length of the next three
alveoli (those for P' or P^ and anterior root of P^). All three
premolars are double-rooted, the P" being the largest of them.
The P" has the usual form; it consists of a tall, single central
cusp surrounded at its base by a cingulum that is continuous except
LUNDELIUS AND TURNBULL: MADURA CAVE
<^^
Fig. 14. Smintfiopsis sp. from Madura Cave, PM 25604, left maxillary with P*-M'
and the alveoli of the other premolars and canine shown in (C) labial and (D)
occlusal views. Both approximately X 8'/i
for a gap near the back of the anterior root. It is more massive than
in the Recent TMM M-839 and FM specimens from Western
Australia, and it has a more triangular occlusal outline because of
the expansion of the crown lingually beneath the posterior root.
There is an axial crest that is weak and blunt anterior to the cusp
and which bears a heavy-wear facet behind the cusp from its tip to
base and out onto the posterior cingular cusplet.
The molars are very much like those of the Recent FM and
TMM specimens in most details, the few differences being a slightly
more massive build to each tooth. These subtle differences can be
seen most readily by comparison of the "lengths" of the M''s,
measured from the anterior edge of the parastyle to the back side of
the tooth at the metacone, and by comparison of the lengths and
areas of the stylar shelves in the M^'s. All three are basically
64 FIELDIANA: GEOLOGY, VOLUME 31
triangular in crown view outline, and the anterior and buccal edges
of each are nearly at right angles to one another. The protocones
are low and ridged from the tip. The anterior ridge runs buccad to a
low protoconule in M^ and to the protoconular region in the others
where the conule is weak or absent. In M^ this crest then continues
onward weakly along the anterior base of the paracone, then
abruptly expands to become stout as it joins the longitudinally-
ridged parastyle. In M*-^ this ridge is interrupted along the bsise of
the paracone; the break is short on M^, larger on M^. The posterior
crest of the protocone runs back to a weak but definite metaconule
in each tooth, then rapidly fades away at the posterolingual side of
the base of the metacone. In descending order of height, the
primary cusps are metacone > paracone > protocone, and while the
protocone has a more massive appearance in crown view, it is about
equal in bulk to the taller metacone. The paracone is much the
smallest of the three cusps. The stylar shelf is expanded buccad in
each tooth, except for the anterior (paracone) region in M', and the
eocrista is strongly W-shaped. The ectoloph has a weak stylocone
that is connected by one ridge to the paracone and by another to
the parastyle. There is a notch in the buccal border of the ectoloph,
and a valley runs from this to the central flexure of the W of the
eocrista. This notch is deep and is bordered anteriorly and
posteriorly by strongly curved shoulders. The anterior shoulder is
made up of the stylocone and the posterior one by a \aige stylar
cusp (cusp C or D of Bensley, 1903) that sits immediately behind
the notch. This cusp is largest in M\ smallest in M^, and in all
three is ridged so that weak crests run from the tip. One of these
crests goes back along the buccal edge of the tooth to the
metastyle, the other runs a short distance lingually toward the base
of the cusp. The metastyle is drawn out posteriorly and buccally,
especially on M ^ ^. Wear is greater on PM 25744 than on PM 25604,
but it preserves M* (as does PM 25639), which consists of a
prominent eocrista that runs from a fused parastyle-stylocone to a
stout paracone and on to a reduced but distinct metacone. Lingual
to the metacone, and reduced almost to a cingular vestige is a
small, ridged protocone. Like the other molars, the M* is three-
rooted, but the posterior of the buccal pair of roots is extremely
reduced, and it is shifted lingually to lie close against the lingual
root.
Discussion. — Several characters of the Madura specimens,
such as the large infraorbital foramen and the deep notch between
LUNDEUUS AND TURNBULL: MADURA CAVE 65
the stylocone and cusp D, are more similar to those of Recent
specimens from South Australia than to material from Western
Australia. However, the Recent samples available to us are too
small to permit one to draw conclusions from this.
This species is widely distributed in AustraUa today but is not
known to occur on the NuUarbor Plain, although Lundehus (1957,
1963) recorded it in surficial deposits from various caves in this
region. Its presence in every stratigraphic level in Madura Cave
indicates that it has been an element of the fauna of this region
through the late Pleistocene, as well as part of the Holocene.
Sminthopsis murina (Waterhouse, 1838)
This taxon, like S. crassicaudata and the indeterminate pygmy
antechinus described earlier, is scarce in the Madura fauna.
Materials. —
Mandibles and Lower Dentitions
Trench 1, Top 1'
PM 25532, left mandible with P2-M4 (fig. 16C, D)
Trench 2, Top 1'
PM 25269, left ramus fragment with M*
Trench 3, Unit 2
WAM 72.3.11, right ramus fragment with Pa^, Mjm (fig. 16A, B)
PM 25757, right ramus fragment with M3.4
WAM 72.3.12, left ramus with P4-M2
PM 25791, right ramus with P4-M3
PM 25793, right ramus with Pa, M2-4
Trench 4, Unit 1, Level 1
TMM 41106-479, right ramus with P4-M1 and alveoli of all other
teeth
PM 25636, left ramus fragment with M4?
Trench 4, Unit 1, Top 1'
PM 26306, ramus juvenile with P4
Trench 4, Unit 2, Level 1
PM 25715, right mandible with M4
PM 25737, left ramus with M*4
66
FIELDIANA: GEOLOGY, VOLUME 31
Fig. 15. Dentition of Sminthopsis murina, Recent juvenile specimen with the
deciduous premolars functional, TMM M-840. The dP" is somewhat molariform, the
P* and dP4are not.
R. upper dentition is shown in lateral (A) and occlusal (B) views. The R.
mandible with its full dentition is shown in lingual view (D), and the lower teeth are
shown in occlusal view (C). In A, M^ is shown partially erupted, as it is in the
specimen, while in B it is shown as fully erupted so as to expose its crown surface to
view. Approximately x SVs.
Trench 4, Unit 2, Level 2
PM 25600, left ramus fragment with M4
PM 25606, left ramus with P3
PM 25711, right ramus fragment with M4
WAM 72.3.13, left ramus fragment with M4
67
68
FIELDIANA: GEOLOGY, VOLUME 31
r
ramus
Fig. 16. Sminthopsis murina from Madura Cave. A, B. WAM 72.3.11, R.
with P2-4, M3-4 and alveoli of C and M1-2, in occlusal (A) and lingual (B) views.
Trench 4, Unit 2, Level 3
PM 25635, left ramus with M4
Trench 4, Unit 7, Level 2
PM 25629, left ramus with M3-4
Maxillaries and Upper Dentitions
Trench 1, Top 1'
PM 25531, left maxillary with P^ or P^- M^, alveoli for M\ and
canine (fig. 17C, D)
Trench 3, Unit 2
WAM 72.3.14, right maxillary with M^ =• and part of M*
PM 25797, left maxillary with M^"* and alveoli of canine and
premolars
PM 26140, right maxillary with M^^
PM 26148, right maxillary with M^^
/^
6e
70 FIELDIANA: GEOLOGY, VOLUME 31
Trench 4, Unit 1, Level 1
TMM 41106-480, right maxillary with M'^
WAM 72.3.15, left M^
Trench 4, Unit 2, Level 1
PM 25745, right maxillary with M^ -» (fig. 17E, F)
Trench 4, Units 4-5
TMM 41106-761, left M^
Descriptions. — The horizontal ramus of the mandible is
slender and tapers anteriorly. The ventral margin is gently convex
from the base of the angular process to the anterior end and does
not show any tendency for the anterior portion to be flattened, as
in Antechinomys.
A variable number (up to three) of mental foramina are
present. They are located under the premolars and Mi, with that
beneath Mi usually being the largest and most consistent.
The symphysial joint is ligamental. It extends from the
anterior end of the horizontal ramus to a position under the
anterior end of the P4.
The posterior part of the mandible has the characteristic
structure of Sminthopsis. The anterior margin of the ascending
ramus rises steeply, the masseteric fossa is broad, and the condyle is
located relatively low with respect to the tooth row.
The articular surface of the condyle is slightly rounded
anteroposteriorly but not laterally. This is in contrast to the flat
condition in Antechinomys. It also differs from that of
Antechinomys in its more oval shape.
The angular process is inflected at about 45° to the vertical
plane of the ramus. None of the Madura Cave species preserves a
complete angular process, so its size cannot be determined.
No incisors or canines are preserved in the mandibles. The
alveoli of the incisors and the canine indicate that these teeth were
procumbent as in Recent specimens. The alveoli of the incisors are
elliptical in cross-section and show the same kind of crowding seen
in the Recent specimens. The alveolus of the canine is usually
elongate or elliptical, but the lateral wall may be reduced so that it
may be triangular.
LUNDEUUS AND TURNBULL: MADURA CAVE 71
The lower premolars are all double -rooted and have a single
major cusp located over the anterior root. The Pa is the largest, and
the P2 is the smallest of the premolars. They are roughly eUiptical
in cross -section. The overlap of ?< and Mi is variable but never
great. The cusps of each are crested anteroposteriorly, especially
posteriorly. The posterior crest descends rapidly and joins a
posterior cingular cuspule at the posterior edge of the tooth. This
crest is clearly an important functional part of the tooth as it shows
a considerable amount of wear in all the specimens we have. The
posterior cingulum and its posterior cuspule form a talonid-like
structure at the back of the tooth.
The molars show the usual tribosphenic form seen in the
Phascogalinae (figs. 15C-D, 16). In crown view, M1.2 are always
tapered: narrower anteriorly. In M.) the two moieties are about
equal, and in M* the taper is sharp posteriorly.
The molars all have high trigonids that are dominated by the
protoconid. The paraconid is weakest and lowest on Mi. In all
molars except M* the metaconid is the second largest trigonid cusp,
and the paraconids show a progressive increase in size from Mi to
M4 so that in M4 the paraconid and the metaconid are subequal.
Crests connect the protoconid to the paraconid [paracristid (I')] and
to the metaconid [epi-centrocristid (IF and I'")] across V-shaped
valleys that initially have a weak cleft (carnassial notch) and
groove at the bottom. Wear may quickly erase this feature, but it
usually persists longer on the paracristid than in the epi-
centrocristid except in Mi, in which the paracristid is poorly
developed.
Viewed from above, the epi-centrocristids are oriented 5" off
from the transverse axis of the teeth (and jaw ramus) in M24, with
the metaconids slightly posterior to the protoconid. The angle
formed by the paraconid, protoconid, and metaconid is about 40*.
This is different from Antechinomys in which the same angle in M2.3
is slightly smaller (M2, 35-40°; M3, 30-35°). This is reflected in the
differences between those two taxa in the scatter diagrams of
trigonid length versus trigonid width (anterior width) in which the
trigonids of Sminthopsis murina are narrower relative to their
length than in Antechinomys (fig. 7). This is particularly true of Ma
in which there is virtually no overlap in the clouds of points of the
two taxa (fig. 6).
The scatter diagrams (figs. 6, 7) and Tables 6, 10, 12, 13 show
that the sample of fossil S. murina is on the average slightly smaller
72
Fig 17. C, D, E, F. Sminthopsis murina from Madura Cave PM 25531, L.
maxillary with all three premolars, M'-^, and the alveoli for C and M*, shown in labial
and (D) occlusal views. PM 25745, R. maxillary fragment with M2-< shown in (E)
labial and (F) occlusal views. All approximately x 8V^.
73
74 FIELDIANA: GEOLOGY, VOLUME 31
in most dental dimensions than fossil Antechinomys. The exceptions
are the trigonid lengths of M2-3 in which the means of the S. murina
sample are sUghtly larger than those of Antechinomys.
The Ml is much more tapered anteriorly than the other molars
with the metaconid close against the protoconid. The short epi-
centrocristid is oriented at an angle of 70° to the long axis of the
tooth row. The antero-extemal cingulum is variable in size, ranging
from a large feature that gives an angled appearance to the antero-
extemal corner of the tooth to virtual absence.
In unworn teeth a valley runs diagonally across the floor of the
trigonid from the bottom of the V in the paracristid to the gap
between paraconid and metaconid. With progressive wear the center
of the trigonid becomes smoothly beisined. Each trigonid is situated
over the anterior root.
The talonids are broad, basined, lower than the trigonids, and
are located over the posterior root of the tooth. They are dominated
by the hypoconids which are so formed that their anteromedially-
and posteromedially- directed crests form V's in an en echelon
arrangement behind and below those of the protoconids.
Entoconids are variably developed; small to absent. When
present they lie at the extreme medial edge of the tooth and are
usually elongated antero-posteriorly. They may be connected by a
weak crest to the metaconid and also to the hypoconulid.
The hypoconulids are distinct and well formed. They project
backward so as to interlock between the parastylid and the medial
end of the anterior cingulum of the next tooth. In old individuals
with extremely worn teeth, they become reduced. The posterior
crest from the hypoconid extends nearly to the medial edge of the
tooth before turning sharply posteriorly to join the hypoconulid. At
the point of inflection, it joins a variably-developed, short, diagonal
crest which trends inward toward the center of the talonid basin.
With wear that portion of the talonid basin behind this ridge loses
its enamel over a trianguljir-shaped area and the ridge becomes
accentuated.
Stout anterior and posterior cingula commence low on the
labial side of the front and back edges of each molar tooth (except
the rear of M4). They rise as they cross the tooth toward the medial
corners.
LUNDELIUS AND TURNBULL: MADURA CAVE 76
The talonid of M« is narrow and elongate, even more so than
Antechinomys, with a single, dominant posterior cusp, the
hypoconid. In many specimens it is connected to the base of the
metaconid (and epicristid) by a low cristid obliqua {\"'h) that
extends from its apex obliquely forward across the talonid.
The maxillary and upper molars of Sminthopsis murina (figs.
15A-B, 17C-F) are so similar to those of S. crassicaudata that they
are separable only with extreme difficulty and some uncertainty.
Separations are based largely on dental measurements and the
absence of "thegosed" wear facets on the posteromedial sides of the
protocones.
The following description is based on PM 25797 except as
otherwise noted. A small part of the premaxillary suture is
preserved, but there is no sign of a flange to the suture.
The infraorbital foramen is preserved in PM 26140 and PM
26148, as well as PM 25797. It is variable in size, being appreciably
larger in PM 26148 than in the other two specimens. It is located
over the anterior half of M' and occupies the middle third of the
space between the alveolar margin and the suture with the lacrimal.
The posterior margin of the anterior palatal foramen is opposite
the posterior root of the anterior-most premolar. The anterior
margin of the posterior palatal vacuity is opposite the center of P*,
and its lateral limits can be discerned posteriorly to opposite the
posterior half of M^ behind which the defmitive edge is broken
away.
The alveoli of the premolars indicate that these teeth were
shorter than any of the molars except M*.
The molars of PM 25797 show little wear and are proportioned
like those of TMM M-840. The morphology of the upper molars is
very much like those of S. crassicaudata. In one detail the Madura
Cave specimens seem to differ from the Recent S. murina. The
posterior crest of the protocone is sharper and more pronounced in
the fossil specimens. The stylar shelves are sUghtly more expanded
buccally than in the modern specimen, which gives the teeth a more
laterally stretched-out appearance.
In the little-worn specimen, PM 25797, M* and M' have a weak
but definite stylar cusp between the large central stylar cusp and
the metastyle. A faint trace of this can be seen in TMM M-840 and
in one of the two specimens (MVZ 133183) from eastern Australia.
This cusp is generally absent from S. crassicaudata but is
76 FIELDIANA: GEOLOGY, VOLUME 31
sometimes present, though weakly developed. In the M"* the basin of
the stylar shelf is larger; the tooth is slightly larger than in S.
crassicaudata.
Specimen PM 25531, one of the most complete of the
Sminthopsis specimens, is, unfortunately, not unequivocally
referrable to species on the basis of the criteria used here. It has
extensive (but non -striated) facets on the posteroUngual faces of the
protocones, features which do not support one assignment over the
other. The parastyle to paracone distance suggests S. crassicaudata
affinities. However, the overall size appears to ally it with S.
murina, as does the broader stylar shelf and the generally
broadened aspect of the molars. Overall, its affinities appear to be
closest to S. murina, and we tentatively assign it to that species.
It is the only specimen with a complete premolar series
preserved. The premolars increase in size from front to back. They
are double-rooted teeth, and each has a single main cusp centered
over the area between the anterior and posterior roots. The cusp is
rounded anteriorly and crested and somewhat worn posteriorly.
Anterior and posterior cingula are present, the latter being
prominent.
The palate is nearly complete. The anterior palatal foramen
extends posteriorly to a point opposite the region between the first
two premolars. The large posterior palatal vacuity does not extend
as far forward as in PM 25797, but it appears to go only to a point
opposite the region between P* and M^ In the Recent specimens
available these vacuities reach only to the level of the medial root
ofM'.
Discussion. — The distribution map in Marlow (1962) shows
that the species occurs in the wetter coastal areas in both Western
Australia and the eastern part of the continent. It has been
reported from the Nullarbor from Doldea (Troughton, 1964) and
from cave N-59 (Archer, 1972).
Remains of S. murina occur in units 1, 2, and 7 of the Madura
Cave sequence. Like S. crassicaudata, it was part of the fauna of
the Nullarbor Plain during the late Pleistocene and early Holocene.
Its absence in unit 4-5 is probably an accident of sampling.
Antechinomys Krefft, 1867
Antechinomys spenceri Thomeis, 1906
LUNDELIUS AND TURNBULL: MADURA CAVE 77
This is the most abundantly represented species of phascogaie
in the Madura fauna. There are 127 specimens in all, not counting
many isolated teeth.
Materials: — Specimens with an asterisk (*) are those which
could not be identified by the masseteric fossa criterion but which
plotted within the range of points for positively -known A. spenceri
(narrow fossa) and are therefore included tentatively.
Mandibles and Lower Dentitions
Trench 1, Top 1'
PM 25527, complete right ramus with M2.4 and alveoli of other
teeth
PM 25533, left ramus with P2, P4-M4, and alveoU of other teeth;
lacks only condyle and end of angular process
PM 25536, nearly complete left ramus with P2-M2 and alveoli of
other teeth; lacks only end of angular process
Trench 1, Unit 1, Top 30"
PM 26166, right ramus with P2 and alveoh of all cheek teeth;
lacks condyle, tip of coronoid process, and much of angular
process
Trench 3, Unit 2
TMM 41106-120, right ramus with P4-M1
TMM 41106-121, right ramus with P2 and alveoh of rest of teeth
WAM 72.3.16, left ramus with P2.3 and M2.4
TMM 41106-123, left ramus with P2-M4
TMM 41106-124, left ramus with M3.4
TMM 41106-125, right ramus with M2.4
TMM 41106-126, right ramus with P3-M4
TMM 41106-128, left ramus with P4-M4
TMM 41106-129, right ramus with P2, P4, and M2.4
TMM 41106-130, left ramus with P2 and M.
TMM 41106-131, left ramus with P23
PM 25750, right mandible edentulous
PM 25754. left ramus with P4-M4
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PM 25755, right ramus with M2.4
PM 25756, right ramus with M2.3
PM 25759, right ramus with M3-4
PM 25760, left ramus with P2-3
PM 25761, right ramus with M4
PM 25762, right ramus with P2-M4
PM 25763, left ramus with P2.3 and Mi^
PM 25764, left ramus with M1.2
PM 25765, left ramus with M3-4
PM 25766, right ramus with M2-4
PM 25767, right ramus with P2 and M1.2
PM 25768, right ramus with M3.4
PM 25769, right ramus with M1.4
PM 25770, right ramus with P3-4
PM 25772, left ramus with M3-4
PM 25774, left ramus, edentulous
PM 25775, left ramus with P2 and P4-M4
PM 25776, left ramus with Mi-4
PM 25777, left ramus with M1.4
PM 25778, left ramus with M2^
PM 25779, left ramus with P2 and Mi
PM 25780, left ramus with P3.4
PM 25781, left ramus with M3 and angle
WAM 72.3.17, left ramus edentulous, with angle
PM 25785, left ramus with P2-M1
PM 25786, left ramus with M2-4
PM 25788, right ramus, edentulous
PM 25790, right ramus with M3 and angle
WAM 72.3.18, right ramus with P4-M1
Trench 3, Unit 2, Level 2
PM 26161, left ramus with M4
LUNDEUUS AND TURNBULL: MADURA CAVE 81
Trench 3, Unit 2, Level 4
WAM 72.3.19, left ramus with M1.2
PM 26153, right ramus with Mm
PM 26156, left ramus with Mi-a
PM 26157, right ramus with Mi^
PM 26158, right ramus with Ms^
PM 26159, right ramus with Pa-Ms
PM 26160, right ramus with Mm
Trench 4, Unit 1
TMM 41106-520, left ramus with Pa^ and alveoli of rest of teeth
(fig. 19A, B)
Trench 4, Unit 2, Level 1
TMM 41106-325, left ramus with P4-M4
PM 25716*, right ramus with P2 and Mi-4
PM 25717, right ramus with P4-M4
PM 25718, left ramus with P2 and P4-M4
PM 25719, left ramus with P2 and M1.4
WAM 72.3.20, left ramus with P2-M1 and M3.4
WAM 74.9.1, right ramus edentulous
PM 25723, right ramus edentulous
PM 25724, right ramus edentulous
PM 25725, right ramus edentulous
PM 25726, right ramus edentulous
PM 25727, left ramus with Ma and angle
PM 25728*, left ramus fragment with Mi
PM 25729*, left ramus with P2-3
PM 25730*, right ramus fragment with Mi
PM 25732*, left ramus with Mi
PM 25733*, left ramus with P4
PM 25734, right ramus with P4-M4
PM 25735, right ramus with Ft-M*
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LUNDEUUS AND TURNBULL: MADURA CAVE 83
WAM 74.9.2, right ramus with Mi 2
PM 25747, right ramus with M3
PM 26305*, left ramus with M.s
Trench 4, Unit 2, Level 2
PM 25598, right ramus with Mi <
PM 25599', left ramus with M34
PM 25603*, left ramus fragment with Mi
PM 25605, left ramus with P4-M1 and M4
Trench 4, Unit 7, Level 2
PM 25628*, right ramus with Ma
Maxillaries and Upper Dentitions
Trench 3, Unit 2
WAM 74.9.3, left maxillary with M'-»
PM 25749, right maxillary with M'^
PM 25758, right maxillary with P^^
PM 25771, left maxillary with M'"
PM 25773, right maxillary with M'
PM 25796, right maxillary with M^'
PM 25798, right maxillary with P-M* (fig. 21 A, B)
PM 25799, left maxillary with P, M'" (fig. 20A, B)
PM 26141, left maxillary with M' *
PM 26142, right maxillary with M'^
PM 26143, right maxillary with M^^
PM 26144, right maxillary with M^
PM 26145, right maxillary with M^^
WAM 74.9.4, right maxillary with P'^*
PM 26147, left maxillary with M^^
PM 26149, right maxillary with M^^
PM 26150, left maxillary with M^^
PM 26151, left maxillary with M'
PM 26154, left maxillary with M'
//
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84
LUNDEUUS AND TURNBULL: MADURA CAVE 85
PM 26162, left maxillary with M»
PM 29910, left maxillary with broken M» and alveoli of M« and
M*
TMM 41106-139, right maxillary with M'''
Trench 4, Unit 1
PM 25642, right maxillary with M'»
WAM 74.9.5, right maxillary with P*-M=»
Trench 4, Unit 1, Level 1
TMM 41106-556, left maxillary with M'*
Trench 4, Unit 2, Level 1
PM 25731, left maxillary fragment with M'
PM 25738, right maxillary with M'"
WAM 74.9.6, left maxillary with M', M^-*
PM 25740, left maxillary with M' '
PM 25741, right maxillary with M' ^
PM 25742, left maxillary with M'^
PM 25743, right maxillary with M*""
PM 25746, right maxillary with P^-M>
TMM 41106-824, right maxillary with M'-^
Trench 4, Unit 2, Level 2
PM 25607, left maxillary with P^-M*
Trench 4, Units 4-5
TMM 41106-701, left maxillary fragment with M«
TMM 41106-703, right maxillary fragment with M'
PM 25613, left maxillary with P'" (figs. 20C, D)
PM 25614, right maxillary fragment with M'
PM 25619, left maxillary fragment with M^
Oppoaite:
Fig 20. Antechinomys apenceri from Madura Cave shown approximately x 8V1
A, B. PM 25799, L maxillary with P», M'-« shown in labial (A) and occlusal (B)
views, C, D. PM 25613, L maxillary with all premolars, shown in labial (C) and
occlusal (D) views.
86 FIELDIANA: GEOLOGY, VOLUME 31
WAM 74.9,7, right maxillary fragment with M^
PM 29458, left maxillary fragment with M^
PM 29459, right maxillary fragment with M^
PM 29460, right maxillary fragment with M^
Trench 5, Unit 5
PM 25632, right mandible edentulous
We have available for comparison eight modem specimens of
Antechinomys (A. laniger MVZ 133197 through 133202; A.
spencen MVZ 134276, AMNH 15012, fig. 18). When their dental
measures are added to the scatter diagrams (figs. 4-7), they show
that, in general, the modern representatives of these species are
slightly smaller than the Madura fossil A. spenceri.
This is the case for P2 and Pa, where one P2 and one Pa of the
modern A. spenceri specimens, and one Pa of the A. laniger
specimens, fall just within the clouds of points for the Madura
fossils assigned to A. spenceri. All other P23 of the modern A.
spenceri and A. laniger are still smaller, falling just outside the
clouds, within the area of the fossil Sminthopsis murina teeth (fig.
5). The P4's of the fossils are slightly broader than those of either of
the modern representatives, but the modern A. spenceri tooth is
slightly longer than any of the fossils, and the modern A. laniger
teeth are at the lower end of the range for both dimensions.
The clouds of points representing the lower molars of the
Recent A. laniger sample overlap the short, narrow end of the
clouds of the Madura Cave Antechinomys sample. Both A. spenceri
specimens fall well within the cloud for M2 but adjacent to it for
Ma. For Ml and M4, in each case one specimen plots within the
cloud and one outside, on the narrow side. The M4 that falls outside
is also the longest tooth of the lot.
Description. — The mandible of the Madura Cave
Antechinomys has the general form seen in most of the small
dasyurids. The horizontal ramus is slender and tapers anteriorly
(fig. 19). The ventral margin is gently convex, but there is a
tendency for the portion anterior to Pa to be flattened, giving a
slightly concave appearance to this section of the mandible. The
mental foramen lies under the Mi. The symphysial joint is
ligamental. It extends from the anterior end of the horizontal
ramus to a position between Pa and P4.
LUNDEUUS AND TURNBULL: MADURA CAVE
87
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,rt^**
Fig. 21. Antechinomys spenceri from Madura Cave shown approximately X 8'/i.
PM 25798, R. maxillary with P*-M* in labial (A) and occlusal (B) views.
The posterior part of the mandible has a number of features
that distinguish Antechinomys from Sminthopsis. The anterior
border of the ascending ramus is more inclined posteriorly than in
the species of Sminthopsis we have examined. In most specimens of
Antechinomys it is slightly concave anterodorsally. The coronoid
process is turned strongly posteriorly in most specimens in which it
is preserved. The articular process of the condyle is flat and slightly
inclined forward and medially. It is almost rectangular (the lateral
end is slightly wider than the medial), with the long axis being
transverse to the plane of the ramus. It is located higher above the
tooth row than in Sminthopsis. This high position of the condyle
and the more posterior inclination of the anterior edge of the
ascending ramus results in a masseteric fossa that is narrower than
in Sminthopsis. Its front and back edges are more nearly parallel
rather than flaring upward as they do in Sminthopsis.
The angular process of Antechinomys is inflected at about 45**
and is very long and greatly arched to extend under the large
auditory bulla. This, and the relatively high position of the condyle,
result in the condyle being located closer to the end of the coronoid
process rather than being equidistant between the coronoid and
angular processes, as in Sminthopsis.
88 FIELDIANA: GE0LCX3Y, VOLUME 31
The mandibular foramen is small and is located close to the
midline of the ascending ramus at or below the level of the teeth.
None of the fossil jaw rami preserve the lower incisors or canine
teeth. Alveoli of the incisors are weakly oval (higher than broad),
and crowded. That of the canine is elongate and oval, the long
dimension being about the combined length of the P2 alveoli lying
immediately behind it.
The premolars are small, elUptical in crown view, with one
major anteriorly-placed cusp each. This main cusp in unworn teeth
turns slightly lingually near its tip. The Pa is the largest of the
premolars. P2 and P4 are approximately the same size with the P4
tending to be somewhat wider.
The long axis of the P4 is oriented parallel to that of the other
teeth in most specimens, but its posterior end lies labial to the
anterior end of the Mi. The degree of overlap is variable.
All of the premolars have well -developed posterior cingula
which in Ps and P4 have small cuspules. The Pa and P4 also have
smaller anterior cingula which may or may not be connected to the
posterior ones labially or lingually.
The molars all have high trigonids that are dominated by the
protoconid. The paraconid is weakest and lowest on Mi. In all
molars except M4 the metaconid is the second largest trigonid cusp,
and the paraconid shows a progressive increase in size from Mi to
M4 so that in M4 paraconid and metaconid are subequal. Crests
connect protoconid to paraconid [paracristid (!')] and to metaconid
[epicristid (IF and L'")] across V-shaped valleys that initially have
a weak cleft (carnassial notch) and groove at the bottom. Wear may
quickly erase this feature, but it usually persists longer on the
paracristid than in the epicristid, except in Mi in which the
paracristid is poorly developed.
Viewed from above, the epicristid is oriented transverse to the
long axis of the tooth and jaw ramus in M2-4, and the paracristid Ues
at about a 35-40" angle from this (with the angle centered on the
protoconid). In Mi the corresponding angle is much wider (60-70"),
and the epicristid, instead of having a transverse orientation as in
M2-4, runs diagonally posteromedially at about 15-25° from the
transverse axis. The paracristid runs diagonally anteromedially at
about 30-45° from the long axis. In unworn teeth, a valley Ues
diagonally across the floor of the trigonid from the bottom of the V
LUNDELIUS AND TURNBULL: MADURA CAVE g9
in the paracristid to the gap between paraconid and metaconid.
With progressive wear, the center of the trigonid becomes smoothly
basined. Each trigonid is situated over the anterior root.
The talonids are broad and low, lower than the trigonids, and
located above the posterior root in each case. They are dominated
by the hypoconids, which are so formed that their anteromedially-
and posteromedially-directed crests form V*s in an en echelon
arrangement behind and below those of the protoconids. The
talonids are basined initially and remain so until extreme wear
causes their posterior crests {post-cristids) to be breached, thus
opening the basins to the interdental spaces. Entoconid
development varies from small to absent. When present, it lies at
the extreme medial edge of the tooth and is usually elongated. It is
connected by a weak crest to the metaconid and usually also to the
hypoconulid. The hypoconulids are distinct and well formed. Each
projects backward so as to interlock between the parastylid and
medial end of the anterior cingulum of the next tooth. In old
individuals with extremely worn teeth, they become reduced. The
p>osterior crest from the hypoconid runs nearly to the medial edge of
the tooth before turning sharply to join the hypoconulid. At the
point of inflection it joins a variably-developed, short, diagonal crest
which trends inward toward the center of the talonid basin.
Stout anterior and posterior cingula commence low on the
labial side of the front and back edges of each tooth (except rear of
M4). They rise as they cross the tooth towards the medial corners.
There is some variation in development of the anterior cingulum
and the parastylid, but the usual pattern is as described above in
the discussion of the hypoconulid.
The talonid of M4 is narrow and elongate with a single
dominant posterior cusp, the hypoconid, at the back. The hypoconid
is connected to the base of the metaconid (and epicristid) by a low
crest, cristid obliqua, that runs from its apex obliquely forward
across the talonid. Traces of hypoconulid and entoconid may or
may not be present, and they are readily removed by wear, as can
be the cristid obliqua. In crown view Mi and M2 are always tapered;
narrower in front, wider behind. In Ma the two moieties are about
equal, and in M4 the taper is sharp in the posterior direction.
Maxillaries and Upper Dentitions
The description is based on PM 25613, WAM 74.9.5 and .6, PM
25741, PM 25771, PM 25798, PM 25799, and TMM 41106-824,
90 FIELDIANA: GEOLOGY, VOLUME 31
except as otherwise noted. No specimen has the entire maxillary
preserved, but taken together the materials allow the
reconstruction of all of the maxillary except for part of the palate
and that portion dorsal to the infraorbital foramen (figs. 20, 21).
The size of the infraorbital foramen is fairly constant, but its
shape varies from an elongate double opening in PM 25771 and
TMM 41106-824 to a single opening in WAM 74.9.5. Most of the
other specimens show an intermediate condition, and two of them,
PM 25741 and PM 25613, have a tiny groove that runs anteriorly
from the main opening for a short distance, then leads into a small
foramen which runs forward within the bone. In WAM 74.9.6, where
a break goes through the area of the P^ it can be seen that this
canal branches. One branch is directed ventrally into the palatal
part of the bone. The other follows along the base of the roots of
the teeth. In PM 25771 this dorsal branch of the canal can be
followed all the way to the alveolus of the canine. These canals are
for the alveolar nerves and blood vessels. In the Recent specimens
available to us, the infraorbital foramen is variable but tends to be
doubled.
Specimen PM 25613 preserves two sutures, that for the
premaxillary, which has a tongue and groove structure, and a
portion of that for the lacrimal. Specimen PM 25771 shows that the
posterior margin of the anterior palatine foramen lies opposite the
middle of the first premolar. The large palatal vacuity can be traced
from the posterior root of P'' at least as far posteriorly as the
posterior edge of M^, where breakage makes its limit uncertain.
No specimen has the canine preserved, and PM 25613 is the
only specimen with three premolars preserved. There is considerable
variation in the size of P^ and F\ All the premolars are more
massive, especially basally, than the premolars of Sminthopsis.
Each has a central cusp that is rounded in front and crested behind.
A small cingulum is present on the front part of each tooth, from
the lingual side of the anterior root around the front edge to the
labial side of the tooth where it becomes very weak between the
roots. From this point it rapidly enlarges to form a prominent
posterior cusplet. The cingulum disappears on the lingual side of
the posterior root.
The premolars increase in size from front to back with P^ and
P3 being subequal. In specimen PM 25613 all three teeth show
heavy wear along their posterior edges from the tips of the main
cusps to the backs of the teeth.
LUNDEUUS AND TURNBULL: MADURA CAVE 91
There are four specimens with complete molar series, PM
25771, PM 25793, PM 25799, and TMM 41106-824. As stated above,
the molars are similar to those of Sminthopsis, especially S. murina,
but they differ in their width measures primarily in the area of the
stylar shelf. This is most readily seen in those scatter diagrams of
M^ that involve the anterior and pK)sterior segments of the eocrista
(parastyle-paracone width and metastyle-metacone width) and the
anterior and posterior widths of the teeth (figs. 9A, C; lOB, D-F).
The notch on the ectoloph between the stylocone and cusp C
(Bensley, 1903) is generally bordered by a straight or slightly -curved
anterior shoulder (fig. 21B). Specimens PM 25799 and TMM 41106-
824 are exceptional in having the strongly-curved anterior shoulders
characteristic of Sminthopsis. The posterolingual face of the
protocone may or may not have a wear facet. In those specimens in
which it does occur, it is not thegosed. The angle formed by the
ectoloph and the anterior edge of the tooth tends to be appreciably
greater than 90° in Antechinomys, in contrast to the situation in
Sminthopsis in which it is very close to 90°.
Discussion. — Antechinomys spenceri is widely distributed in
the arid regions of Western Austraha, South Australia, and the
Northern Territory. This includes at least part of the Nullarbor
Plain with a record from Rawlinna (Troughton, 1962). Its
continuous presence in the Nullarbor Plains area through the late
Pleistocene and post- Pleistocene indicates the presence of open
areas throughout this period of time. The minor morphological
differences noted here between the fossil material and the Recent
specimens suggest that there have been small evolutionary changes
over this period of time.
INCERTAE SEDIS
Eight specimens, each displaying distinct morphologic features,
had to be included here because neither tooth measurements nor
other features allowed their unequivocal assignment.
Trench 4, Unit 2, Level 2
PM 25601, left ramus fragment with Mi 2. Probably Antechinomys
spenceri since most dental measures plot within the range of
the other specimens of that taxon, but the plot for Ma of
trigonid length versus trigonid width is intermediate between
those of Sminthopsis murina and A. spenceri.
92 FIELDIANA: GEOLOGY, VOLUME 31
PM 25706, left ramus with M2.3. Probably A. spenceri, but both
trigonid length versus trigonid width plots (M2 and Ma) are
intermediate between those of A. spenceri and S. murina, while
the other measures give plots that fall within those of A.
spenceri.
PM 25709, left ramus fragment with M2. Probably S. murina as
the plot of length versus anterior width indicates, but that for
trigonid length versus trigonid width is intermediate between
those for S. murina and A. spenceri.
TMM-41106-690, isolated lower molar. Probably S. murina since
most measures plot with those of S. murina, but they are
frequently quite far off to one side (long trigonid).
Trench 4, Units 4-5
PM 25617, right maxillary fragment with P"
and M* I Probably Sminthopsis
PM 25618, right maxillary fragment with P^ ( or Antechinomys, but
PM 25622, left maxillary fragment with M^
}
possible Antechinus.
Trench 4, Unit 7
PM 25626, right ramus fragment with Mi. Some of the bivariate
plots are intermediate between S. murina and. A. spenceri.
Included among the sieving concentrates are hundreds of
isolated phascogaline teeth and edentulous ramal and maxillary
fragments. Those which we believe to belong to the three taxa
covered by this section of the Madura Cave fauna are listed by
trench and level:
Trench 1, Top 30"
PM 26167
Trench 3, Unit 2
PM 25782, PM 25789, PM 25794
Trench 3, Unit 2, Level 4
TMM-41 106-152
Trench 3, Unit 3
TMM-41 106-53, PM 26133
LUNDEUUS AND TURNBULL: MADURA CAVE 93
Trench 4, Unit 1, Top 1'
PM 25637-8, PM 25640-1, PM 26280, TMM-41 106-555, TMM-
41106-756, TMM-41 106-758-9
Trench 4, Unit 2, Level 1
PM 25720
Trench 4, Unit 2, Level 2
PM 25712, PM 26286 through PM 26302, TMM-41 106-757, TMM-
41106-762, TMM-41106-779, TMM-41106-823, and one
uncatalogued bulk lot
Trench 4, Unit 2, Level 3
PM 25634, TMM-41 106-780 through 784
Trench 4, Units 4-5
PM 25544, PM 25608, PM 25623, PM 26281 through 26285,
TMM-41 106- 19 (a bulk lot with over 100 isolated teeth), TMM-
41 106-763A and B through 778, TMM-41 106-788-9, TMM-
41 106-790-822, two uncatalogued edentulous rami, and an
uncatalogued bulk lot with approximately 50 teeth and 25
jaws.
Trench 4, Unit 7
PM 25625 and an uncatalogued bulk lot with dozens of isolated
teeth and six edentulous jaw fragments
Trench 5, Unit 5
1 uncatalogued ramus fragment
ACKNOWLEDGEMENTS
In addition to those people listed on pages 32 and 33 of Part I
of this faunal report, we thank Professor R. T. Prider and Dr. B.
Logan of the Department of Geology, University of Western
Australia for use of facilities and the following people for assistance
in preparing this part of the paper: Dr. M. Archer of the
Queensland Museum supplied data on the condition of the
entoconid in several species of Sminthopsis; Mr. J. Calaby of
Commonwealth Scientific and Industrial Research Organization,
Canberra provided much general information about dasyurids; Dr.
P. Aitkin of the South Australian Museum, Mr. R. Green of Queen
Victoria Museum, Launceston, Tasmania, Mr. H. Van Deusen of
94 FIELDIANA: GEOLOGY, VOLUME 31
the American Museum of Natural History, Dr. W. Z. Lidicker of the
Museum of Vertebrate Zoology, Berkeley, and Drs. P. and G.
Crowcroft and Mr. T. Strickler of the Brookfield Zoo, Chicago
loaned material in their care.
We are especially appreciative of the fine drawings of the fossils
and the Recent comparative specimens. These drawings were made
by Dr. Tibor Perenyi, Scientific Illustrator, Field Museum.
Financial support was provided by Grants GB 975, GB 3729,
GB 7662 from the National Science Foundation; Field Museum of
Natural History and the Geology Foundation of the University of
Texas at Austin.
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LUNDEUUS AND TURNBULL: MADURA CAVE 95
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Belgie, Brussels. Pp. 215-320 (part 1); pis. 1-44 (part 2).
1964. R^cherches sur I'origine des mammif^res. Ann. Soc. Roy. Zool. Belg., 94, pp.
117-160.
Waterhouse. G. R.
1838. Catalogue of the Manunalia preserved in the museum of the Zoological
Society, 2nd ed. Richard & John E. Taylor, London, 68 pp.
r
APPENDIX 1
Specimen List for Figure 2
Sminthopsis murina Wedges Cave N-25; PM 4235 through 4237, PM 4240-1, PM
4243-4, PM 4246 through 4260, PM 4262-3. and PM 4265.
S. murina Hastings Cave N - 41; PM 16559. PM 16562. PM 16565. PM 16568-9. PM
16576 through 16582 (PM 16581 Ls shown in fig. 17 A-B).
S. "hirtipes" ( -S. murina) W.A.M. Recent N- 2; M 1547, M 1577.
S. murina Madura Cave fossils N>-5; only one specimen, PM 25532, gave both
measurements while in four others, WAM 72.3.11-12, PM 25791. and TMM 41106-
479, P4 was measured but only a minimal estimate of the molar series was possible,
and, finally, three others afforded only the latter estimate, PM 25635, PM 25791,
and PM 25793.
S. murina W.A.M. and U.S.A. Recent N-34 (30 WAM. 3 MVZ, 1 TMM); M 4, M
81, M 286, M 662. M 709. M 981. M 1005-6. M 1177, M 1231, M 1314, M 1333. M
1564. M 1674. M 1714. M 1758, M 1835-6, M 1841, M 1945, M 1960. M 1969. M 2059,
M 2193. M 2263. M 2345, M 2401, M 2453, M 2534, M 7384; MVZ 133183-4, MVZ
133342; and TMM M-840.
S. rufigenis lumholtzi AMNH N-9; 154479 through 154487 (2 juveniles lacked
erupted P4'8).
S. larapinta MVZ Recent N-5; 133173, 133188, 133190 through 133192.
S. granuUpes W.A.M. N-3; M 1397, M 1968, M 2333.
S. crassicaudata Murrafelellevan Cave N-10; PM 4128, PM 4130-1, PM 4133
through 4138. and PM 4157.
S. crassicaudata W.A.M. Recent N - 33; M 549, M 564. M 670. M 690. M 799. M 911-
2, M 1011, M 1015, M 1031, M 1055-6. M 1140, M 1157, M 1174, M 1278, M 1387, M
1565, M 1676, M 1735. M 1764. M1830, M 1965. M 2275. M 2277, M 2283, M 2286, M
2290, M 2413, M 2413*, M 2452, M 2560, and M 2624.
S. crassicaudata Recent USA N-7; FMNH 34722 through 34724, 60116, 104788,
TMM M-839, and AMNH 160032.
S. crassicaudata Madura Cave fossils N-6; PM 25525, PM 25534-5, and PM 25637
each gave both measures, while WAM 72.3.8, PM 25528-25529 each gave one
measure only.
Antechinomys laniger MVZ N-5; MVZ 133197, MVZ 133199 through 133202.
A. spenceri Murraelellevan Cave N-16; PM 4140 through 4147, PM 4150 through
4156. and PM 4158.
A. spenceri W.A.M. and other N-7; M 1546. M 2230, and M 2368, BMNH 5.5.3.3,
BMNH 15.2.22.6, AMNH 15012, MVZ 134276.
A. cf. spenceri Madura Cave foasils N-22; (17 specimens gave both measures, 13
gave one or the other as follows): Both TMM 41106-123, 41106-126, 41106-128-129.
41106-325, 41106-520, and PM 25533, PM 25536, PM 25605, PM 25717-26718, PM
25721, PM 25724, PM 25735. PM 26754. PM 26762. and PM 26775; P4 only PM
25770, PM 26780, PM 26786, WAM 72.3.18, and PM 26169; with Mi.4 only PM
26698, PM 25716, PM 26719, PM 26763, PM 26769, PM 26776-26777, and PM 26160.
'EHfferent data — probably recording error in specimen number
97
APPENDIX 2
Specimen List for Figure 3A
Sminthopsis rufigenis lumholtzi AMNH 154479 through 154487.
S. larapinta MVZ - University of California 133173, 133188, 133190 through 133192.
Plotted with a flag is AMNH 108934 labelled Sminthopsis macrura but which plots
with Sminthopsis larapinta.
S. stalkeri MVZ - University of California 134305.
S. macrura MVZ - University of California 134244.
S. crassicaudata Recent sample TMM M-839, AMNH 160032, 196685, FMNH 34722
through 34724, 60116, and 104788.
S. crassicaudata (Madura Cave) PM 25525, WAM 72.3.8, PM 25528 through 25530,
PM 25534-25535, and PM 25537 and TMM 41106-755, all from Unit 1; and 41106-
521 and WAM 72.3.9 from Unit 2.
APPENDIX 3
Specimen List for Figure 3B
Sminthopsis psammophila South Australia Museum M 7971.
S leucopus R. H. Green private collection. Registered #19/1 (Queen Victoria
Museum).
S. hirtipesl MVZ 133187.
S. munna Hastings Cave fossil PM 16559, PM 16562, PM 16565, and PM 16568
through 16582.
S. murina fuliginosa MVZ 133342, TMM M-840.
S. murina murina, MVZ 133183-133184.
S. murina (Madura Cave) TMM 41106-479, PM 25269, PM 25532, PM 25600, PM
25606, PM 25629, PM 25635-25636, PM 25711, PM 25714-25715, PM 25737, WAM
72.3.11, PM 25757, WAM 72.3.12, PM 25791, and PM 25793.
APPENDIX 4
Specimen List for Figure 4
Antechinomys spenceri AMNH 15012, MVZ, U. of California 134276.
A. laniger MVZ 133197 through 133202.
A. spenceri Madura TMM 41106-120-121, WAM 72.3.16, TMM 41106-123 through -
126, TMM 41106-128 through -131, TMM 41106-325, PM 25527, PM 25533, PM
25536, PM 25598-25599, PM 25603, PM 25605, PM 25628, PM 25716 through 25719,
WAM 72.3.20, PM 25727 through 25730, PM 25732 through 25735, WAM 74.9.2, PM
25747, PM 25754 through 25756, PM 25759 through 25770, PM 25772, PM 25775
through 25781, WAM 72.3.17, PM 25785 through 25786, PM 25790, WAM 72.3.18
and 19. PM 26153, PM 26156 through PM 26161, PM 26166, and PM 26305.
98
APPENDIX 5
Specimen List for Figure 5
Sminthopsis murina TMM 41106-479. PM 25532. PM 25606. WAM 72 3 11 and 12
PM 25791. and PM 25793.
Antechinomys apenceri TMM 41106-120-121. WAM 72.3.16. TMM 41106-126. TMM
41106-128 through -131. TMM 41106-325. TMM 41106-520. PM 25533. PM 25536.
PM 25605. PM 25716 through 25719, WAM 72.3.20. PM 25733 through 25735. PM
25754. PM 25760. PM 25762 through 25763. PM 25767. PM 25770. PM 25775, PM
25779 through 25780, PM 25785. WAM 72.3.18. PM 26159, and PM 26166.
APPENDIX 6
Specimen List for Figure 6
Sminthopsis murina WAM 72.3.12, PM 25791. and PM 25793.
Antechinomys spencen TMM 41106-120-121, 123, WAM 72.3.16. TMM 41106-125-126,
TMM 41106-128 through -130. TMM 41106-325. PM 25598. PM 25603, PM 25605.
PM 25716 through 25719. WAM 72.3.20, PM 25728, PM 25730, PM 25732, PM 25734
through 25735. WAM 74.9.2. PM 25754 through 25756, PM 25762 through 25764,
PM 25766 through 25767, PM 25769, PM 25775 through 25779, PM 25785 through
25786, WAM 72.3.18 and 19, PM 26153, PM 26156 through 26160, and PM 26305.
APPENDIX 7
Specimen List for Figure 7
Sminthopsis murina PM 25600. PM 25635. PM 25711. WAM 72.3.13, PM 25715, PM
25737. WAM 72.3.11. PM 25757, WAM 72.3.12. PM 25791, and PM 25793.
Antechinomys spencen TMM 41106-120-121, 123, WAM 72.ai6 TMM 41106-125-126.
TMM 41106-128 through -130. TMM 41106-325. PM 2559a PM 25603. PM 25605.
PM 25716 through 25719. WAM 72.3.20. PM 25728, PM 25730, PM 25732, PM 25734
through 25735, WAM 79.9.2, PM 25754 through 25756. PM 25762 through 25764.
PM 25766 through 25767. PM 25769. PM 25775 through 25779, PM 25785 through
25786. WAM 7Zai8and 19. PM 26153, PM 26156 through 26160, and PM 26305.
APPENDIX 8
Specimen List for Figures 9, 10
Sminthopsis crassicaudata WAM 72.3.10, PM 25604, PM 25620, PM 25630, and PM
25744.
S. munna PM 25531, PM 25745, WAM 72.3.14, PM 25797, PM 26140, PM 26148,
TMM 41106-480, and TMM 41106-761.
ArUechinomys spencen TMM 41106-139, TMM 41106-556, TMM 41106-701, TMM
41106-824, PM 25607, PM 25642-3, PM 25738, PM 25740 through 25743, PM 25748-
9, PM 25771, PM 25796, PM 25798-9, PM 26141 through 26143, PM 26145, PM
26147, and PM 26149-50.
APPENDIX 9
Statistical Tables
TABLE 1. Statistical data on upper dentitions of Sminthopsis crassicaudata
from Madura Cave.
N
M^
m2
m3
L
2
AW
2
PW
2
P8W
2
msw
2
PSW + MSW
2
L
4
AW
4
PW
4
PSW
4
MSW
4
PSW + MSW
4
L
5
AW
5
FW
4
PSW
5
MSW
5
PSW + MSW
5
Observed
Mean
Range
1.67-1.75
1.71
1.25-1.37
1.31
1.82
1.82
.61-70
.66
1.29-1.41
1.35
1.99-2.02
2.00
1.62-1.67
1.64
1.52-1.67
1.58
1.98-2.13
2.04
.89-99
.94
1.46-1.56
1.50
2.37-2.55
2.44
1.44-1.65
1.52
1.71-1.84
1.77
1.92-2.13
2.05
1.10-1.23
1.18
1.44-1.56
1.50
2.54-2.79
2.68
100
TABLE 2. Statistical data on lower dentitions ot Sminthopsis crassicaudata
from Madura Cave units 1 and 2.
N
Observed
Range
Umn
LM
1-4
5.76-6.02
5.91
Distance - ant
end P4 to ant.
end M]^
P2
L
W
P3
L
W
P4
L
W
Ml
L
AW
PW
Mft
L
^
AW
PW
Mg
L
AW
PW
M4
L
AW
PW
.68-79
.73
.91-96
.95
.48-56
.53
1.12-1.17
1.15
.61-66
.64
.94-1.12
1.08
.51-56
.53
1.44-1.52
1.48
.73-76
.75
.79-.81
.80
1.47-1.67
1.58
.81-96
.90
.89-.99
.96
1.52-1.72
1.64
1.01-1.08
1.03
.86-99
.94
1.47-1.62
1.67
.91-99
.95
.30-.46
.38
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TABLE 5. Statistical data on upper dentitions of Sminthopsis murina from
Madura Cave.
N Observed Mean
Range
M^ L 6 1.61-1.79 1.70
1.10-1.31 1.20
1.79-1.94 1.87
.53-.66 .62
1.31-1.48 1.41
1.96-2.09 2.02
M^ L 8 1.56-1.71 1.63
1.51-1.63 1.56
2.05-2.17 2.09
.91-1.03 .97
1.50-1.63 1.57
2.47-2.63 2.54
M** L 5 1.29-1.52 1.43
1.65-1.82 1.77
1.86-2.09 1.98
1.18-1.27 1.22
1.29-1.56 1.45
2.54-2.75 2.67
M^ L 1 .72
1.71
1.25
L
6
AW
6
PW
6
PSW
6
MSW
6
PSW + MSW
6
L
8
AW
8
PW
8
PSW
8
MSW
8
PSW + MSW
8
L
5
AW
5
PW
5
PSW
5
MSW
5
PSW + MSW
5
L
1
AW
1
FW
1
106
TABLE 6. Statistical data on lower dentitions of Sminthopsis murina from
Madura Cave.
N Observed Mean ±- Standard Standard Coefficient of
Range Error Deviation Variation (%)
P2 L 2 .91-.94 .93 - -
W 2 .51--.56 .54 — —
P3 L 3 1.12-1.22 1.17 - —
W 3 .56-.58 .57 - —
P4 L 2 .92-.97 .95 — —
W 2 .48-.56 .52 - —
Ml L 1 1.52 - —
AW 1 .81 - —
PW 1 .89
M2 L 1 1.67 — —
AW 1 1.04 - —
PW 1 .96 - —
M3 L 4 1.57-1.72 1.67 — —
AW 4 1.06-1.12 1.09 — —
PW 5 .86-. 96 .89 - —
M4 L 10 1.52-1.77 1.63 -±.02 .07 4.59
AW 11 .86-1.06 .98 ±.02 .06 6.34
PW 9 .30-.46 .35 ±.02 .05 1.66
106
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TABLE 9. Statistical data on upper dentitions of Antechinomys spenceri
from unit 1, Madura Cave.
N Observed Mean
Range
m1 L 3 1.79-1.83 1.80
1.25-1.34 1.28
2.01-2.09 2.04
.66-.68 .67
1.56-1.60 1.59
2.22-2.28 2.26
m2 L 3 1.63-1.68 1.65
1.63-1.71 1.68
2.17-2.38 2.28
.95-1.06 1.00
1.71-1.82 1.76
2.70-2.88 2.76
M^ L 2 1.51-1.52 1.52
1.82-1.85 1.84
2.12-2.20 2.16
1.25-1.27 1.26
1.61-1.71 1.66
2.88-2.96 2.92
M^ L 1 .76
1.68
L
3
AW
3
FW
3
PSW
3
MSW
3
PSW + MSW
3
L
3
AW
3
PW
3
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3
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3
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3
L
2
AW
2
PW
2
PSW
2
MSW
2
PSW -•• MSW
2
L
1
AW
1
110
TABLE 10. Statistical data on lower dentitions of Antechinomys spenceri
from unit 1, Madura Cave.
N Observed Mean
Range
3 .89-1.00 .94
2 1.06-1.12 1.09
2 .56 .56
2 1.32-1.37 1.35
2 .61-.66 .64
3 1.01-1.12 1.06
3 .58— .61 .60
3 1.52-1.57 1.54
3 .74-.89 .80
3 .89-1.01 .94
4 1.52-1.75 1.60
4 1.02-1.12 1.08
4 1.04-1.12 1.09
3 1.49-1.67 1.56
3 1.12-1.22 1.17
3 1.01-1.09 1.05
3 1.52-1.67 1.57
3 .99-1.01 1.00
3 .41-.51 .46
DUt.
ant. end
P4 to ant. end
Ml
P2
L
W
P3
L
W
P4
L
W
Ml
L
AW
PW
M2
L
AW
PW
M3
L
AW
PW
M4
L
AW
PW
111
TABLE 11. Statistical data on upper dentitions of Anfechmomys spencer/
from unit 2 Madura Cave.
N Observed Mean ±- Standard Standard Coefficient of
Range Error Deviation Variation (%)
m1 L 11 1.71-1.92 1.80 :L. 02 .06 3.34
AW 11 1.24-1.44 1.33 t. 02 .06 4.27
PW 11 1.90-2.15 2.05 ±.02 .08 3.88
PSW 11 .61— .80 .69 ±..02 .06 9.13
MSW 11 1.48-1.67 1.57 ±.02 .06 3.77
PSW+
MSW 11 2.13-2.39 2.26 ± .02 .08 3.41
m2 L 12 1.58-1.76 1.67 ±.02 .07 4.31
AW 12 1.63-1.90 1.74 ±.02 .08 4.93
PW 12 2.20-2.55 2.33 ± .03 .11 4.62
PSW 12 .95-1.10 1.03 ±.01 .04 3.91
MSW 12 1.71-1.90 1.78 ±.02 .06 3.64
PSW+
MSW 12 2.70-2.92 2.81 ± .02 .08 2.74
M^ L 12 1.41-1.60 1.51 ±.02 .06 3.96
AW 12 1.78-2.05 1.92 ±.02 .08 4.33
PW 12 2.13-2.32 2.24 ±.01 .05 2.34
PSW 12 1.14-1.37 1.28 ±.02 .07 5.25
IVfSW 12 1.54-1.84 1.67 ±.02 .07 4.47
PSW+
MSW 12 2.82-3.10 2.95 ± .02 .09 2.96
M^ L 4 .78— .95 .84 - —
AW 4 1.71-1.90 1.84 - -
112
TABLE 12. Statistical data on lov/er deniiiions of Antechinomys spenceri
from unit 2, Madura Cave.
N Observed Mean — Standard Standard Coefficient of
Range Error Deviation Variation (%)
L M1.4 7 6.01-6.48 6.23 — —
Dist. ant. end
of Pa to ant. 5 .93-1.01 .96 — —
end M
P2
L 6 1.06-1.17 1.13
W 6 .53-. 58 .56
P3 L 4 1.22-1.42 1.35 — —
W 4 .66-.71 .68 — —
P4 L 6 1.09-1.17 1.14 — —
W 6 .56 -.66 .61 — —
Ml L 11 1.59-1.77 1.67 ±.02 .06 8.80
AW 10 .81-.92 .87:L.01 .04 4.88
PW 10 .91-1.04 .98 ±.01 .04 4.64
M2 L 11 1.57-1.77 1.67 +.02 .07 4.42
AW 11 1.06-1.18 1.13 ±.01 .05 4.12
PW 11 1.04-1.17 1.10 ±.01 .04 8.87
M3 L 12 1.57-1.77 1.65 ±.02 .06 8.58
AW 14 1.14-1.24 1.20 ±.01 .03 2.88
PW 13 .96-1.17 1.08 ±.02 .06 6.92
M4 L 13 1.52-1.77 1.65 ±.02 .07 4.42
AW 13 .96-1.12 1.04 ±.01 .05 4.68
PW 13 .33--.51 .43 ±.01 .05 12.31
113
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