UNIVtRSlTv C^ ILLINOIS I aRARY AT URBAMA-CHAMPAIGN L IOLOGY ^ CO CO h 3 FIELDIANA Geology Published by Field Museum of Natural History Volume 38 THE MAMMALIAN FAUNA OF MADURA CAVE, WESTERN AUSTRALIA PART III ERNEST L. LUNDELIUS, JR. and WILLIAM D. TURNBULL cn % % S* i « (J/ 2 ■ ■ -¥~ ■o c 0. 1 5 n3 0) u = « 1 X Q s 5 Ifl ■ j » ': 0 ?, * " z s Z r a a> z > 5 j * . « (D a. \ 0 i », a 3 TJ 10 h .-<- -I 11 5 « \ z " \ > ioU « » u <" — c a- ■- c 5£ K +- 00 (ft z 0 K (0 < a SJ * 2° O 3 £ ■o — 2 2 u-q; \ ["•< 0. \ o i V _ 1 J r Z tt> / > s cJ / ° ' 1 3 M 0! 3 T3 \ ^z ° • 2 01 5 otf li >o 2 N« o >0 (A n a 1 3 r> > 1 « / X 5^ 5" < LL< N (0 U r-i Z E i xo5 « ! o 5 10 Madur 18889- eucog m q. -: zz - <; < "-< £ 14 1 BMNH and TMM N* R tapoatafa Wedges Cave Nr3 Madura Cave N=!x2 BMNH and|FM N = 5 D. byrnei ••:-//: 9.9 10.6 B ■+■ 11.0 11.2 1 1.4 1 1.6 LENGTH M ,_4 11.8 12.0 12.2 12.4 Fig. 5. Bivariate graphs of length of lower molar series vs . length P4 for various taxa of dasyurids showing relative size of the P»'s. This graph is an extension of Figure 13 of Part I, and Figure 2 of Part II. Shown in A are: Antechinus stuartii adustus Antechinus flavipes Phascogale calura Pseudantechinus mimulus and Ps. macdonnellensis Parantechinus apicalis Dasycercus cristicauda Shown in B are: Dasyuroides byrnei Phascogale tapoatafa 15 16 FIELDIANA: GEOLOGY, VOLUME 38 ->.'~\~<'\.--- \^J -/ Fig. 6. Phascogale calura FM 36052. A, B, Rostrum with right upper dentition shown in labial and crown views. WAM 74.9.13, right ramus fragment with M2or3, alveolus for M.3 or 4 WAM 74.9.14, right ramus, edentulous, with alveoli for P4-M4 WAM 74.9.15, left ramus fragment with M3-4 Unit 7, level 2 TMM 41106-733, right ramus fragment with M2-.3, alveoli for C, P3-4, Mi, M4 (fig. 7A, B) Trench 5 Unit 3 PM 29431, right M2or3 ■ I JS bo 2 Q CO ci 17 18 fieldiana: geology, volume 38 Upper Dentitions Trench 3 Unit 2 WAM 74.9.16, right maxillary fragment with M3; alveoli for M2, M4 Trench 4 Unit 1, level 1 TMM 41106-739, left maxillary fragment with M34; alveoli for M1 2 (fig. 8D, E, F) Unit 2, level 1 PM 25580, left maxillary fragment with M2; alveoli for other molars, P4 (fig. 8G, H, I) Unit 2, level 2 TMM 41106-738, left maxillary fragment with M1; alveoli for P4, M2 (fig. 8A, B, C) Description. — Mandibles: The jaws are slightly more mas- sive than in Antechinus flavipes, and they each have a masseteric fossa with prominent surrounding ridges and an upward- and backward-flaring configuration, more like that of Sminthopsis crassicaudata than that of Antechinomys spenceri. The condyle lies slightly above the dental occlusal plane; when viewed from the side it is seen as being nearer to the tip of the coronoid pro- cess than to the tip of the inflected angular process. A large men- tal foramen opens either beneath the posterior root of Mi or be- tween Mi and M2 . A smaller anterior one may be single or divided into two openings located beneath P2 or P3, or, if split, with one foramen under the front of Mi . Occasionally, a smaller additional one is found behind the large posterior foramen. Lower Dentitions: The lower premolars are best seen in PM 4798-4800, PM 25266, and TMM 41106-132. They are double- rooted, broad at the base, with well-developed lateral and pos- terior cingula. There are small cingular cuspules on the posterior ends of the premolars, and their principal cusps are relatively low. A crest extends from the principal cusp of each premolar to its posterior cingular cuspule. This ridge is located slightly labial to the midline in each tooth, and a slight furrow is present lingual to the ridge. The furrow is best developed in the Pi. There is a notch at the posterior end of the furrow in P2 and P3 (best de- veloped in P2) into which the anterior end of the following tooth c is 0 CO is • 00 > £ e is 3 B - ■* es d s a c tc o fl •~ c c > c S is "3 o c X cs CO CO CO .2 CO •8 9 .^ E ~o at Sj h > *> as J3 M"P *c a OS „ CO w 3 t> cC X 3 o ~r s is 1 CO Ei 3 E X es C < £ CO V > es BS „ o Q a es X 9 •* T3 5 s E E Q cfc W -3 es e? C 3 -n a, cs fa ° c 19 20 FIELDIANA: GEOLOGY, VOLUME 38 fits. Wear progressively truncates the crest and produces a flat- tened or broadly convex surface on the posterior half of P2 and P3 . The P3 is the largest premolar, followed by P2 and Pi. The P4 is usually oriented at a noticeable angle (10°- 40°) to the long axis of the tooth row. Its posterior end lies labial to the anterior end of the Mi. The lower molars have the normal dasyurid structure. Seven specimens have complete, or nearly complete, lower series (TMM 41106-132, PM 4798-4800, PM 25548, and PM 25556). In each, the protoconid is the largest cusp, followed by the metaconid and paraconid. The talonid is basined. The hypoconid is joined to the hypoconulid by the postcristid. The labial border of the talonid is formed by the cristid obliqua. This ridge joins the protolophid at the center of the protoconid in Mi. The junction is located pro- gressively farther lingually in the posterior molars until, in M4, it is located at the center of the protolophid. The entoconid is usu- ally a prominent cusp, is higher than the hypoconulid, and is lo- cated on the lingual border of the talonid slightly closer to the hypoconulid than to the metaconid. It is oval to almost circular in cross-section. The entoconid is small on Mi and variably de- veloped on M3. The labial side of each lower molar tooth has a well-developed cingulum that joins the pre- and post-cingula. Usually there is some cingular development in the valley be- tween the paraconid and metaconid, but its extent is varied. Another specimen, PM 25542, shows some variations. It has a somewhat larger-than-average M3-4, and the M3 has a better de- veloped entoconid — even more developed than in the Recent specimen. This specimen preserves the alveoli of C-M2. These show that the P2 and Pi were both set at an angle to the tooth row, since the anterior alveolus of P2 is labial to that of the canine and the posterior alveolus of Pi is anterolabial to the an- terior alveolus of the Mi . Opposite: Fig. 8. Phascogale calura from Madura Cave. A-C, TMM 41106-738, a left maxillary fragment with M1 and alveoli of P4 and M2 shown in labial, crown, and lingual views. D-F, TMM 41106-739, a left maxillary fragment with M3-4 and alveoli of M1-2 shown in labial, crown, and lingual views. G-I, PM 25580, a left maxillary fragment with M2 and alveoli of P4 and the other molars shown in labial, crown, and dorsal views. The black fiber marks the path of a canal for a branch of the infraorbital artery. I ( J 21 22 FIELDIANA: GEOLOGY, VOLUME 38 Upper dentitions: The largest maxillary fragments are PM 25580 and TMM 41106-739 and between them they preserve the alveoli of P4 and all of the upper molars. The former also pre- serves part of the infraorbital foramen and palate, as does TMM 41106-738, the fragment with M1. Specimen PM 25580 shows a divergent canal branching off from the infraorbital foramen (fig. 81) which extends forward and medially into the interior of the palatal portion of the maxillary bone. The P4 was double-rooted with the roots aligned with the edge of bone. In crown view the molars are all triangular in outline and have the usual dasyurid morphology with a well-developed stylar shelf and strong eocrista. The protocone is low and crescentic in all of the molars. In M13 the metacone is taller and larger than the other cusps and the paracone is the smallest. In M4, as would be expected, the paracone is near normal size while the metacone is very reduced. In M1 the stylocone is quite distinct, as is the parastyle, while in the other molars they are fused. A stylar cus- pule (C2) is present in M1 3, situated just anterior to the metastyle and behind stylar cusp C of Bensley (1903). The anterior ridge from the protocone runs to a protoconule which is developed in M1. There is a metaconule in M1 3 and, in addition, either a low ridge or a cuspule between the metaconule and the base of the metacone. A posterior cingulum runs well labiad in M13, extend- ing farthest on M1. Discussion. — The lower teeth of Phascogale calura (tables 5-9) are readily distinguishable from those of P. tapoatafa (tables 10-12) by their substantially smaller size. P. calura is somewhat larger than Antechinus flavipes leucogaster, its premolars are slightly broader and more massive, and the entoconids tend to be relatively better developed. The mandibles and lower dentitions of P. calura from Madura Cave cannot be distinguished from those from Hasting's Cave and Wedge's Cave, Western Australia, and comparisons with a Recent specimen (FM 36052, fig. 6) show a close similarity. Most of the lower dentition of Phascogale calura , both fossil and Recent, is so similar in size and proportions to that of a Re- cent specimen of Antechinus flavipes rufogaster that they could be easily confused. There are differences that appear to be valid. A comparison of the Recent specimens shows the lower canine of P. calura to be larger and more massive than that of A. f. rufogaster; the Ii of P. calura is larger than that of A. f. rufogaster. Speci- mens PM 4798-4800 from Madura Cave all have alveoli for large Ii's. LUNDELIUS & TURNBULL: MADURA CAVE 23 The P2 of Phascogale calura from Madura Cave is longer than that of Recent specimens of Antechinus flavipes rufogaster and A. f. leucogaster (fig. 3A, B; tables 1, 5, 6). The principal cusp of the P4 of A. f. rufogaster when viewed from the labial side is rela- tively higher, and is slightly more extensive anteroposteriorly than that of the Recent P. calura. The Madura Cave fossils con- form to the P. calura pattern. Also, the labial cingulum of the P4 dips lower over the posterior root in both fossil and Recent speci- mens than it does in A . f rufogaster and the paraconid of the Mi is larger in both the Recent and Madura Cave specimens of P. calura than it is in A. f. rufogaster. This taxon is represented by samples from three statigraphic levels within trench 4 (units 1, 2, and 4-5) and equivalent units from trenches 1, 2, 3, and 5 that are large enough to allow a check on possible size change through time. An examination of Tables 5-7 shows that there was no significant change, although mean values of some measures show a decrease from older to younger units. Comparison of the upper dentitions with the Recent specimen (FM 36052) shows that the M1 of the fossil has a slightly de- veloped stylar cusp behind cusp D which is lacking in the Recent specimen. There are no other differences in morphology or size in the upper dentition worthy of mention (tables 8, 9). This species has not been recorded living from the Nullarbor Plain, but the presence of its remains throughout the entire known sequence in Madura Cave and surface deposits in other caves on the Nullarbor Plain (Lundelius, 1963) indicates that it was an element of the Pleistocene and sub- Recent fauna that has only recently disappeared from this region. Its current distribu- tion is in more humid parts of southwestern Australia, in the Adelaide area, and in isolated refugia near Alice Springs and the junction of the Murray and Darling Rivers (Jones, 1923; Marlow, 1962). According to Ride (1970), it is an inhabitant of sclerophyll forest and woodland. This distribution pattern plus its recent dis- appearance from the Nullarbor Plain suggests that the species was very widespread until fairly recently and that the environ- ment has continued to deteriorate since the end of the Pleis- tocene. ?Phascogale calura Material. — Trench 4 24 FIELDIANA: GEOLOGY, VOLUME 38 Unit 2, level 1 PM 25581, left ramus fragment with P4, Mi; alveoli for C, P2-3, anterior root M2 Description. — This specimen has a P4 that is oval, with the principal cusp located in the anterior half of the tooth and a well-developed cingulum all around the tooth. The long axis is oriented 45° off the axis of the tooth row, and the Mi overlaps the posterior half of the Pi lingually. The Mi has a trigonid with the protoconid as the largest cusp, followed by the metaconid and paraconid. The talonid has a small entoconid which has been eliminated by wear. Discussion. — This specimen resembles the Madura Cave Phascogale calura in every character except the larger overall size of the Mi and the greater development of the anterior part of the trigonid. The larger size is primarily in the anteroposterior length. It is probable that this specimen is a larger example of Phascogale calura, but we are calling attention to it on the chance that it represents another taxon. Phascogale tapoatafa (Meyer), 1793. Tables 10-12; Figures 5B, 9, 10A-B, 15 Material. — Lower Dentitions Trench 4 Unit 1, level 1 PM 29437, left Ms PM 29438, left ramus fragment with M4 Unit 2, level 2 PM 30580, left ramus fragment with broken Mi; M2; alveoli for C, P2-4, M3 Units 4-5 PM 29901, right ramus fragment, edentulous, alveoli for M2-4 PM 29902, right ramus, edentulous, alveoli for C-M4 PM 29903, left ramus fragment with P4, partial Mi; alveoli for I1-P3, M2 PM 29904, right ramus with C-M4, alveoli for I1-2 (fig. 10A, B) PM 29905, right ramus with M3-4, alveolus for M2 LUNDELIUS & TURNBULL: MADURA CAVE 25 WAM 74.9.17, right ramus fragment with Mi 2, anterior part M3, alveoli for C-P4 Description. — Phascogale tapoatafa, the largest of the phas- cogalines, is not well represented in the Madura Cave material. Only nine specimens with teeth have been recovered, and none from the 1954 excavation (Lundelius, 1963). We have had only one Recent specimen (TMM M-838; fig. 9) and four fossil speci- mens (PM 30438-PM 30441) from Western Australia for comparison. P. tapoatafa is easily distinguished from its congener, P. cal- ura, by its much greater size (tables 5-12). It usually can be dis- tinguished from the comparably-sized dasy urines, Dasycercus and Dasyuroides , by the presence of double-rooted Pi's. The P4 of P. tapoatafa is approximately the size of the P3, while the P4 is much smaller than the P3 In Dasycercus the I^'s are usually much re- duced or absent, but occasionally the P4's approach the size of those of P. tapoatafa. In Dasyuroides the P4 is absent or repre- sented by a small vestige of a tooth, while the P4 is variable — most often double-rooted and somewhat smaller than the P2 or P3, but sometimes reduced to a single-rooted peg. Our one Recent specimen of Dasyuroides (FMNH 104786) has a single-rooted RP4; however, the LP4 is not present. Mandibles: The mandibular ramus has no features that are strongly divergent from those of other comparably-sized phas- cogalines. There is some variation in the depth of the horizontal ramus in the Madura Cave specimens which is related to the dif- ferent ages of the individuals at death. The anterior border of the ascending ramus is more horizontally inclined than that ofSmin- thopsis, Dasycercus, or Dasyuroides, but is less inclined than in Antechinomys . The masseteric fossa is deep and is bordered both dorsally and ventrally by well defined ridges. The lower of these ridges is the larger and is flared to a distinct, laterally projecting shelf between the base of the angular process and the condyle. The coronoid process in lateral view is broad anteroposteriorly at its apex and has a small, slightly postero ventrally directed point. The condyle is located slightly above the tooth row, closer to the coronoid than to the angular process. Its articular surface is oval and convex upward as in the one Recent specimen that is available for comparison. The dental canal opens immediately above the base of the angular process. There are two or three 26 FIELDIANA: GEOLOGY, VOLUME 38 Fig. 9. Phascogale tapoatafa. TMM M 838. A, B, Rostrum with right upper dentition shown in labial and crown views. mental foramina. The posterior one(s) open beneath the Mi in all the Madura Cave specimens and in the Recent specimen. The an- terior one opens beneath either the P2 or P3 in the fossils. In the Recent specimen the anterior foramina opens beneath P3 on the right side, and beneath P4 on the left. Lower Dentitions: None of the fossil P. tapoatafa from Mad- ura Cave preserve the lower incisors, but two specimens, PM 29903 and PM 29904, have alveoli for the incisors. The alveoli provide some information about these teeth. The alveoli show that there were three lower incisors, that h was much larger than the other two, and that all three were procumbent. This is also the situation in the Recent specimen. The canine (PM 29904) is large, procumbent, and somewhat incisiform, and it is long and broad at the base. The lingually T3 C CO H 3 bo C Q d OS 6 fa 27 28 FIELDIANA: GEOLOGY, VOLUME 38 ^ Fig. 10. A, B,Phascogale tapoatafa from Madura Cave, PM 29904. Right ramus with canine through Mi shown in lingual and crown views. C, D, Dasyuroides byrnei, PM 30437 from Madura Cave. Right maxillary fragment with P4 alveoli of P2"3 and M1 shown in labial and crown views. Originally identified as Whascogale tapoatafa, assignment revised on basis of Figure 15E. recurved main cusp has a longitudinal ridge that climbs its an- terior edge to the apex and runs down the posterior side of the cusp to the center of the posterior cingulum; the posterior half of this ridge is very weak. A strong cingulum extends from the an- terolingual corner posteriorly and ventrally around the posterior end of the tooth to the posterolabial corner. At the posterior end of the tooth there is a small cingular cuspule. Wear takes place on the posterolingual side of the main cusp. The canine shows no structural differences from the Recent specimen. LUNDELIUS & TURNBULL: MADURA CAVE 29 There are three premolars, all of which are double-rooted. The P2 and P3 are subequal, similar in morphology, and are nearly twice the length of the P4 . The first two are elongate with their long axes in line and parallel to the long axis of the tooth row. They are bluntly pointed anteriorly, and the squared posterior edge of each has a weak groove near its midpoint into which the adjoining tooth fits. The P3 may even overlap P2 slightly. The principal cusp of each is located in the anterior one-third of the tooth. The posterior edge is a ridge that extends backward across a talonid-like area to the posterolabial corner of the tooth where it joins a small cingular cusp. The posterior occlusal surfaces of both P2 and P3 are flattened. This feature becomes emphasized with wear. The first two premolars each have a cingulum that is large on the sides and posterior end and is weaker anteriorly. The P4 is oval in shape and about half the length of the anterior pre- molars. It is oriented en echelon to the other teeth, with more than half of its length located labial to the anterior end of Mi . It has one main cusp, which is located anterior to the midpoint of the tooth and which is joined to cingular cusps anteriorly and posteriorly by ridges. A cingulum encircles the tooth. The pre- molars of the known P. tapoatafa fossil specimens show no differ- ences in structure from those of the Recent specimen; however, they are larger in the fossil (tables 10-12) and are more crowded in the jaw. The lower molars have the usual dasyurid structure of a high trigonid and a low talonid. The Mi is distinctive, is narrower an- teriorly than the other molars, and retains a functional paraconid. The paraconid is well formed but is much smaller rela- tive to the protoconid and metaconid than it is in the posterior molars; the paraconid is located almost directly anterior to the protoconid. This condition pertains in the three Madura Cave specimens (PM 29904, WAM 74.9.17, PM 30580), as well as in the one Recent specimen available (TMM M-838) and in the three specimens from Wedge's Cave (PM 30439, 30440, 30441). This re- sults in an anteroposterior orientation of the paracristid (D. De- spite this reduction, there is a distinct, well-developed (though small) carnassial notch in the paracristid. The protoconid is taller and about twice the size of the metaconid. They are joined by a short but well-formed epicristid (II', III') with a distinct carnas- sial notch. The epicristid is oriented approximately 45° to the long axis of the tooth. In crown view the talonid is strongly basined and broad, almost as broad as in M2-3. The hypoconid is by far the 30 FIELDIANA: GEOLOGY, VOLUME 38 largest cusp of the talonid. It is joined to the base of the pro- toconid by the cristid obliqua, which has a small notch close to the protoconid. The hypoconulid is the smallest and the post- eriormost cusp of the talonid. It is joined to the hypoconid by the postmetacristid (F). It protrudes backward into a notch in the front edge of the adjacent tooth. The entoconid is larger than the hypoconulid, stands nearly as high as the paraconid, and forms the internal border of the talonid basin. It is laterally compressed at its base but becomes conical toward the apex. A ridge, the en- tocristid (V), joins it to the metaconid. The entocristid, like the cristid obliqua, has a small but distinct notch close to the epicris- tid. The entoconid and hypoconulid are separated by an open groove. The floor of the talonid basin is smooth and featureless. Well-developed cingula extend posteriorly on both sides of the tooth from the paraconid. The labial cingulum is part of the main cingulum of the tooth. The lingual cingulum disappears abruptly anterior to the base of the metaconid. This produces a distinct lingual bulge anterior to the metaconid. The M2 and M3 are similar in size and morphology. The trigonids of M2-4 are almost identical. In each the protoconid is the largest cusp, followed by the metaconid and the paraconid. The protoconid is joined to the paraconid by the paracristid and to the metaconid by the epicristid. Both of these cristids have prominent carnassial notches as in the other dasyurids. All three teeth have blade-like parastylids that project forward and lie linguad of the hypoconulid of the preceding tooth. The paraconid and metaconid are separated by an open valley. The talonids of M2-3 are deeply basined and much alike and, except for their slightly larger size relative to the entire tooth, resemble the talonid of Mi The talonid of M4 is reduced in Phascogale tapoatafa as in all dasyurids. The Madura Cave sample shows some variation in the size of this structure. All three cusps of the talonid are present on the two M4S that are available, although they are much reduced in size. The talonids of both teeth have distinct basins. All the lower molars have cingula on their labial, anterior and posterior borders. The cingulum is weak where it skirts the pro- toconids and hypoconids and absent on the posterior side of M4. The anterior cingulum is prominent on M2-4 and ends at a notch that receives the hypoconulid of the preoeding tooth. The posterior cin- gulum is prominent on M1-3 and joins the hypoconulid. LUNDELIUS & TURNBULL: MADURA CAVE 31 Discussion . — The lower dentition of Phascogale tapoatafa can be readily distinguished from other similar-sized forms (Dasycer- cus and Dasyuroides) on the basis of the presence of a P4, the structure of the trigonid of Mi. and the well-developed basin of the M4 Neither of these other genera has the well-developed paraconid and expanded anterior end of the Mi or the broadly basined talonid on M4 . In both Dasycercus and Dasyuroides , the anterior ends of Mi are highly compressed laterally, and the paraconids are reduced. The metaconids are relatively much smaller than the protoconids and are located more posteriorly than in P. tapoatafa. The talonids of the Ma's in Dasycercus and Dasyuroides are much reduced, and if they are basined, they are much narrower. P. tapoatafa can also be distinguished from Dasycercus by its larger size and the large size of the entoconids of the lower molars. The entoconids in Dasycercus are low and in cross-section are narrowly elliptical. Unfortunately, these last two characters will not reliably separate P. tapoatafa from Dasyuroides . Phascogale tapoatafa is known to occur as a living animal in the coastal area of southern Queensland, New South Wales, and Victoria; in the area of Adelaide, South Australia; in extreme southwestern Australia; in the Kimberly District; and from northeast of Kalgoorlie, Western Australia eastward to the bor- der (Marlow, 1962; Ride, 1970). This would place it immediately north of the Nullarbor Plain, if not on it. It appears that this species has long been a resident of the Nullarbor Plain. Its ab- sence from stratigraphic units other than Units 1, 2, and 4-5 is probably the result of sampling accidents. Subfamily Dasyurinae Gill, 1872 cf. Parantechinus Tate, 1947 cf. Parantechinus apicalis (Gray), 1842. Tables 13-14; Figures 5A, 11, 15. Material. — Trench 1 Top first foot (possibly Unit 1) PM 25330, right ramus fragment with Mi, anterior half M2, M4; alveoli for li 3, C, P24, posterior alveolus for M2, alveolus for M3 (fig. 11) Description. — We have no Recent comparative materials of 32 FIELDIANA: GEOLOGY, VOLUME 38 this taxon. Our comparisons are based on a few Holocene fossils (PM 7268-7270) collected from Hasting's Cave, Western Australia, in 1954 and compared directly with Recent specimens in the collections of the Western Australian Museum at that time. The horizontal ramus is shallow and slender and has the nor- mal dasyurid form, with a convex ventral margin. The ascending ramus is incomplete but appears to have had a wide masseteric fossa with the anterior margin steeply inclined with respect to the tooth row. There are three mental foramina — a posterior one located under the Mi and two anterior ones located under P3 and P2, respectively. The alveoli for the P2 and P3 are oriented slightly en echelon to the tooth row, and the alveolus for the anterior root of P2 opens into that of the canine. Between the posterior alveolus for the P3 and the Mi is a small circular alveolus 0.5 mm. in diameter for a single-rooted P4. The Mi is deeply worn, and part of the anterior end is broken. As a result, little can be said about the trigonid. The metaconid is in the same position relative to the protoconid as in the Mi of specimens of Parantechinus apicalis from Hasting's Cave, West- ern Australia. The talonid has a well-developed hypoconid and a posteriorly projecting hypoconulid, as in the Hasting's Cave specimens. The presence or absence of an entoconid cannot be de- termined because of the worn condition of the tooth. The postcin- gulum is large and extends around the base of the hypoconid to the base of the protoconid. The development of this cingulum is variable in the Hasting's Cave sample, but the degree of de- velopment of the Madura Cave specimen is matched by specimen PM 7268. The M2 is so incomplete that little can be learned from it. A broad anterior cingulum that does not reach the parastylid is pre- sent. The Mi has a well-developed trigonid. The protoconid is the largest cusp. The metaconid and paraconid are subequal in size, but breakage of the former renders a precise determination of rel- ative size impossible. The metaconid appears to have been higher than the paraconid, which is more massive. The paracristid and protocristid have well-developed carnassial notches. The talonid is narrow and basined with a small but distinct hypoconid with a weak cristid obliqua. No entoconid is discernable. The anterior cingulum is broad but does not reach the blade-like parastylid. Discussion. — In all these characters except size (tables 13, LUNDELIUS & TURNBULL: MADURA CAVE 33 Jwl3\ . t Fig. 11. Parantechinus apicalis from Madura Cave. PM 25330 right ramus with Ml, M2 Broken, M4 and alveoli for canine, premolars, posterior root of M2 and M3, shown in (A) lingual and (B) crown views. 14; fig. 11) this specimen resembles the Recent sample and the sample of Parantechinus apicalis from Hasting's Cave. In tooth size this specimen is larger than the Hasting's Cave specimens and is much more comparable to Dasycercus cristicauda, but the depth of the horizontal ramus is less than in D. cristicauda and is similar to that of P. apicalis. The trigonids of Mi and Ma are nar- rower and less massive than those of D. cristicauda (fig. 15A, B, C). These features, plus the presence of the P4, lead to the identi- fication as P. apicalis. Dasycercus Peters, 1875 Dasycercus cristicauda (Krefft), 1867. Tables 15-18; Figures 5A, 12-15. Material. — Lower Dentitions Trench 1 Top first foot (possibly Unit 1) PM 6070, right ramus with C-M4, alveoli for incisors PM 6071, right ramus with C-M2, M4; alveoli for incisors, M3 PM 6072, left ramus with P2-M4, alveoli for incisors, C PM 6073, left ramus with C-M2, alveoli for incisors, M34 PM 6074, right ramus with C-M4, alveoli for incisors PM 6075, left ramus with C, P3, M3-4; alveoli for incisors,P2, Ml-2 PM 25331, left ramus with C, P2, alveoli for other teeth 34 FIELDIANA: GEOLOGY, VOLUME 38 Trench 2 2x/2 ft. below surface (possibly Unit 2) PM 6233, left ramus with M1-4; alveoli for incisors, C, P2-3 PM 25264, left ramus, edentulous, with ascending ramus, angular process PM 25265, right ramus fragment with M4 Trench 3 Unit 2 TMM 41106-127, right ramus, edentulous PM 25586, left ramus, edentulous PM 25588, right ramus fragment with M2-4 WAM 74.9.19, left ramus fragment with M2-3; alveoli for two incisors C-Mi Unit 3 PM 29383, right ramus fragment with M2-3; alveoli for C, P2-3, Mi, M4 PM 29390, left ramus fragment with M3; alveoli for M2, M4 PM 29391, right ramus fragment with M3, broken Mi Trench 4 Unit 1, level 1 TMM 41106-517, right ramus with Mi; alveoli for P2-3, M2-4 PM 29911, left ramus fragment with M2, part M3 PM 29912, left ramus fragment with Mb (or M2) Unit 2 PM 30443, left ramus fragment with P2-3, broken Mi ; alveoli for one incisor, C Unit 2, level 1 WAM 74.9.20, right ramus fragment with M1-3 PM 29430, right ramus fragment with M2; alveoli for P2-3, Mi, M3 Unit 2, level 2 TMM 41106-16, left ramus fragment with P3, M1-2 PM 30444, left ramus fragment with M3, alveolus for M2 (or M2, alveolus for Mi) PM 30445, left ramus fragment with M2; alveoli for M3-4 Units 4-5 PM 25553, right ramus fragment with broken M3; alveoli for one incisor, C, P2-3, rest of molars LUNDELIUS & TURNBULL: MADURA CAVE 35 PM 29426, left ramus fragment with Mi; alveoli for M2-3 PM 29446, right ramus with Mi-..; alveoli for P23 (fig. 13C,D) PM 29447, left ramus fragment with Mi 3, broken Mi, al- veolus for P3 PM 29448. right ramus fragment with M3; alveoli for M2, M» PM 29449, left ramus fragment with M2; alveoli for R2-M1 , M3 PM 29450, right ramus fragment with M2-3; alveoli for C, P2-3, Mi PM 29451, left ramus fragment with M3; alveoli for M1-2, M4 PM 29452, left ramus fragment with M3; alveoli for M2, M4 PM 29453, left ramus fragment with M3; alveoli for M2, M4 PM 29454, right ramus fragment with M2; alveoli for P2-3, Mi, M3 PM 29455, right ramus fragment with Mi 2, alveolus for P3 WAM 74.9.21, left ramus fragment with Mi, alveoli for P2-3,M2 PM 30446, right M4 PM 30447, left Mi (or Ms) broken Unit 5 WAM 74.9.22, right ramus fragment with M4, alveolus for M3 PM 29418, right ramus fragment with M2; alveoli for Mi, M3 PM 29419, left ramus fragment with Mi PM 29420, right ramus fragment with M2-4 PM 29421, right ramus fragment with Mi; alveoli for M2-3 PM 29422, left ramus fragment with M1-3; alveoli for C-P3, Mi WAM 74.9.23, right M2 Unit 7, level 1 PM 29415, right ramus fragment with Mi Trench 5 Unit 4 PM 29427, right ramus with M3; alveoli for P2-M2 Unit 5 PM 29428, right M3 PM 30448, right ramus fragment with Mi; alveoli for M2-3 Upper Dentitions Trench 1 Top first foot (possibly Unit 1) PM 6067, right maxillary with P2, M1 4; alveoli for C, P34 PM 6068, right maxillary with P2 \ M1 4, alveolus for C, (fig. 14A, B) 36 FIELDIANA: GEOLOGY, VOLUME 38 PM 6069, left maxillary with M1 4; alveoli for C-P3 Trench 3 Unit 2 PM 25589, left maxillary fragment with M3, roots for M4 Unit 2, level 2 PM 29368, left maxillary fragment with M2 4 WAM 74.9.24, left maxillary fragment with M24 Trench 4 Unit 2, level 1 PM 25591, left maxillary fragment with M34 PM 25592, left maxillary fragment with M23 Unit 2, level 2 WAM 74.9.25, right maxillary fragment with M2 Units 4-5 PM 29439, right maxillary fragment with M2'3; alveoli for C P2"3, M\ M4 PM 29440, left maxillary fragment with M3; alveoli for M2 M4 PM 29441, right maxillary fragment with M1; alveoli for C p23 M2 PM 29442, right maxillary fragment with M3; alveoli for M2 M4 PM 29443, left maxillary fragment with M3; alveoli for M2 M4 PM 29444, right maxillary fragment with M3; alveoli for M2 M4 WAM 74.9.26, right maxillary fragment with M3; alveoli for M2, M4 PM 29461, right M3 Trench 5 Unit 5 PM 29429, right M3 (or M2) Description. — Mandibles: The mandible is robust, and the horizontal ramus is relatively deep. The ascending ramus is broad with the condyle widely separated from the coronoid process. The anterior edge is inclined at a steep angle to the tooth row. The coronoid process is short, broad, and only slightly turned pos- teriorly. The central part of the ventral edge of the masseteric LUNDELIUS & TURNBULL: MADURA CAVE 37 fossa is expanded laterally to form a broad shelf. The condyle is high above the level of the tooth row. Its articular surface is ellip- tical in shape and is gently convex anteroposteriorly. Despite the robustness of the mandible, there are no rugosities for muscle at- tachments on the medial surface of the ascending ramus except that for the deep portion of the M. temporalis. On one specimen, PM 6072, there is a mylohyoid groove that extends anterior from just below the mandibular foramen. The mandibular symphysis is ligamentous and extends from the anterior end of the mandible posteriorly to the position of the anterior end of Mi. A prominent foramen is present near the center of the symphyseal surface. There are two mental foramina, an anterior one opening under P2 and a posterior one under Mi . Lower Dentitions: The lower dentition of Dasycercus is characteristic of the more advanced dasyurids (e.g., Dasyuroides, Dasyurus , Sarcophilus) in the absence of the P4. Although none of the Madura Cave specimens have the incisors preserved, the al- veoli of some specimens (PM 6072-6075, TMM 41106-407) show that three were present, as in the Recent population. The canine is prominent and stands almost vertically in the jaw. It is recurved posteriorly and slightly inwardly. There is a basal cingulum which is weak externally and strong internally. The cingulum rises on the tooth anteriorly and joins a low, lon- gitudinally oriented ridge on the anterior face of the tooth, best seen on an unworn specimen (PM 6073). In these characters the canines of Dasycercus differ from those of Phascogale tapoatafa in which they are strongly procumbent with a well-developed post- erior shelf. The P2 and P3 are both oval to subquadrate in shape. Each has a principal cusp located in the anterior half of the tooth, which is connected to anterior and posterior cingular cuspules by low crests. A cingulum encircles each tooth. The posterior border of P2 has a concavity into which the anterior end of P3 fits. The apex of the principal cusp of P3, when unworn, turns lingually. The Mi is narrow and tapers anteriorly. The trigonid is highly compressed laterally with a large, centrally-placed protoconid and with a much smaller metaconid located very close to the pro- toconid. The paraconid is reduced to a small cingular cuspule on the anterior end of the tooth. The talonid is basined in the un- worn state but becomes flat as wear reduces the bordering cusps. The hypoconid is approximately as tall as the metaconid when 38 FIELDIANA: GEOLOGY, VOLUME 38 \ FlG. 12. Dasycercus cristicauda AMNH 15009. A, B, Rostrum with right upper dentition shown in labial and occlusal views. unworn. The premetacristid from the hypoconid and the post- paracristid from the protoconid almost meet on the labial side of the talonid to form a narrow slit that is similar to the carnassial notches in the paracristid and epicristid. The lingual border of the talonid has a small entoconid whose shape in cross-section is that of an elongate ellipse. The hypoconulid is located at the posterior end of the tooth only slightly lingual to the midline of the tooth. The postmetacristid joining the hypoconid and hypoconulid is oriented more anteroposteriorly, especially the posterior part, than it is in M2 and Ma. A cingulum extends from the vestigial paraconid around the labial side of the tooth to the hypoconulid. 39 c ^ I o s 00 X 4J o s 'S _> CO 3 E 0 C« OD (h o -U c X he -a c 5 03 CO £ 03 -c CM 3 s 0 IV -4^ Q 9) ^ C u '8 OS l£9 o t- C J3 CO *j 'i S 00 & 3 £ E o 03 *4M >J -o c— GO 91 >-. J o +» en CM aj t> (h o co c S '2 c- 03 u n -a i < 03 cc 0) cc > CD oB CJ O a 33 a U s* 3 JS T3 03 3 s a £ OS c e» ,1- tt-1 >, a -a "^3 c 3 o O ^CJ •hi ■Si £ CO o "> CO a c c s- u * -a c cr: IS i— i 3 be 6 ti £ *-* c •8 40 LUNDELIUS & TURNBULL: MADURA CAVE 41 The M2 and M3 are similar in size and morphology. The trigonid has no features that distinguish it from those of other dasyurids of comparable size. The talonid is basined when un- worn. When worn it becomes shallower and flatter. It is bordered labially by the hypoconid which, in contrast to the condition in the Mi, is not as high as the metaconid. As in the Mi, the pre- metacristid and postparacristid meet and form a tight notch on the labial side of the talonid. The talonid is bordered lingually by the entoconid which is somewhat variable in size but usually is low and elongate. The hypoconulid is the posteriormost part of the tooth. A broad cingulum is present on the anterior and pos- terior ends of the tooth. It is not present around the bases of the protoconid and hypoconid and does not reach the parastylid be- cause of the notch for the hypoconulid of the next anterior tooth. The Mj has a trigonid that differs from those of the M2 and M3 only in its slightly smaller size (tables 15-16). The talonid is re- duced and may or may not be basined. In those specimens in which it is basined, a vestigial hypoconid can be identified, but no entoconid is discernable. Maxillaries and Upper Dentitions: The identified skull ma- terial of this taxon from Madura Cave is limited to maxillary fragments. These show that the infraorbital foramen is large and opens above the M1. A subsidiary canal leaves the main canal and extends into the parietal part of the maxillary. The infraorbital canal usually opens into a well-defined fossa on the lateral sur- face of the maxillary, but occasionally no fossa is developed. No incisors or canines are preserved. Alveoli for the canines are pre- served in some specimens which show that the size of the canine in the fossils was comparable to that of the Recent specimen and to a sample of Recent to sub-Recent specimens from Mur- raelelleven Cave. Two of the specimens, PM 6067-8, have the upper premolars preserved, and specimens PM 6069, PM 29441, and PM 29439 have the alveoli for P2 3. These show that P2 and P3 are double- rooted and about the same size as those of the Recent specimen. Two specimens, PM 6068, which has a P4, and PM 6069, which has an alveolus for P4, indicate that the tooth was single-rooted as in the Recent specimen. In PM 6067 the P4 which has been broken off or shed was smaller than that of PM 6068. The absence of the alveolus in the other two specimens is consistent with a sample of 29 specimens (18 left, 20 right) from post- Pleistocene deposits in Murraelellevan Cave in which the tooth is missing on 42 FIELDIANA: GEOLOGY, VOLUME 38 one or both sides in 30-33 per cent of the sample. When present, the tooth shows considerable variation but almost always is a single-rooted peg. One specimen (PM 4880) from Murraelellevan Cave has premolars that raise a question as to the identity of the peg-like tooth usually termed the P4. This is discussed in the sec- tion dealing with Dasyuroides byrnei . The M1 is triangular and asymmetrical, with the protocone located as far anterior as the paracone. The stylocone and paracone are apparently fused. The parastyle is an anteriorly pro- jecting process. It is connected to an anterior cingulum that ex- tends only halfway across the base of the paracone. The metacone and stylar cusp C are approximately equal in size and are close together but distinct. The protocone is triangular with convex sides. A distinct protoconule is located at the lingual edge of the base of the paracone. The posterior margin of the protocone ex- tends posteriorly around the base of the metacone in varying de- grees. There is no posterior cingulum. The M2 is triangular and slightly asymmetrical. The labial edges of the major stylar cusps are almost in line, resulting in a nearly straight edge for the stylar shelf. There is a moderate em- bayment in the labial edge of the stylar shelf between cusp C and the metastyle. The metacone is the tallest cusp, followed in order of decreasing size by cusp C, the stylocone, and the paracone. A short cingulum is present from the parastyle to the midpoint of the base of the paracone. The sides of the protocone are straight to convex. A well-defined protoconule is present at or im- mediately anterior to the lingual edge of the paracone. The pos- terior border of the protocone may extend onto the posterior side of the metacone. There is no posterior cingulum. A small cusp D is located on the edge of the stylar shelf just anterior to the meta- style. M3 is a triangular, symmetrical tooth with the protocone lo- cated at the apex of the triangle. Stylar cusp C is located lingual to a line that connects the stylocone and the metastyle. The metacone is the tallest cusp, followed closely by cusp C, then the stylocone, paracone, and metastyle. Small, ridge-like stylar cusps Bi and D are present. The protocone is triangular with straight to convex margins. A protoconule is present anterolingual to the base of the paracone, and a metaconule is present posterolingual to the base of the metacone. The edges of the protocone do not extend around the paracone or metacone. LUNDELIUS & TURNBULL: MADURA CAVE 43 W£^%*f°* Fig. 14. Dasycercus cristicauda from Madura Cave. A, B, PM 6068. Right maxillary with P2 - M4, alveoli for canine shown in labial and crown views. C, D, PM 29439. Right maxillary fragment with M2"3, alveoli for P3 - M\ M4 shown in labial and crown views. M4 is much reduced posteriorly. The stylocone and paracone — the largest cusps — are widely separated and are joined by a slightly sinuous paracrista which is oriented transverse to the tooth row. The metacone is small and poorly separated from the paracone. The stylar shelf is a narrow, posteriorly sloping surface with no cusps. The protocone is a small, lingually rounded cusp Length j 1.50 Length 2 0Q B M, y^*~^ S^ © \ / © © \ / x x x x i — D /^ / 1047868 / i Q/ /8 ( ° / o o • \ o • •/ O ^__^<-^/l5009 TMM-M-838 < .c ■o 2.00 Length 2 50 3 00 Fig. 15. Bivariate graphs comparing various dental measures of Parantechinus apicalis, Dasycercus cristicauda , Dasyuroides byrnei, and Phascogale tapoatafa. 44 I I Par antechi nus apicalis Madura Cave Parantechinus apicalis Hastings Cave Dasycercus Madura Cave (unit I) Dasycercus Madura Cave (units 2-7) Dasycercus Murraellelevan Cave AMNH 15009 Dasycercus cristacauda FMNH 104786 Dasyuroides byrnei Dasyuroides (unit 2-3) Madura Cave Dasyuroides (unit 4-5. 7) Madura Cave Phascogale tapoatafa Madura Cave Phascogale tapoatafa Wedge's Cave Phascogale tapoatafa Recent L e ng t h ®4880 >-A42* « 368 dp >• Dasyuroid Recent samples - Phase, tap. p4 0 dp4 o es byrnei p- x . dp-4 BMNH £ 15009 486 368 9++412 + 368 I 0 X» Length Fig. 15 45 46 FIELDIANA: GEOLOGY, VOLUME 38 located lingual, to the paracone. Its margins extend around the base of the paracone. An anterior cingulum is present from the stylocone to the midpoint of the paracrista. Discussion. — Two species of Dasycercus have been described: D. cristicauda from the Nullarbor Plain and central Australia and D. blythi from northwestern Australia. They are distin- guished on the basis of the lack of a P4 in the latter species; Ride (1970) indicates that they may be conspecific. More information on the nature of the zone of contact between the two populations is needed to establish the validity of the two species. The Madura Cave material is assigned to D. cristicauda on the basis of the presence of the P4. Criteria for distinguishing the lower dentition of Dasycercus cristicauda from that of Phascogale tapoatafa have been discussed in the section dealing with the latter taxon. It can be distin- guished from that of Dasyuroides byrnei on the basis of the slight- ly smaller size of most of the molar teeth (tables 15-16, 19-23; figs. 5, 15), the form of the Mi, and the size of the entoconids on Mi through M3. The Mi of Dasycercus cristicauda has a smaller paraconid and smaller anterior cingulum than Dasyuroides byrnei. The entoconids are smaller and more ridge-like on all lower teeth of Dasycercus cristicauda. The entoconids of the M2 and M3 of Dasyuroides byrnei are almost as high as the hypoconids and are almost circular in cross-section at their apices. The lower canine of Dasycercus cristicauda is straighter, with a less well-developed base than that of Dasyuroides byrnei. The upper dentition of Dasycercus cristicauda differs from that of Dasyuroides byrnei in its generally smaller size and less mas- sive form. This is usually reflected in the more robustly developed metacone and the consequent greater anteroposterior length of the M4 in the latter taxon (tables 17, 23; figs. 12A, B; 14A, B; 16A B; 17A, B). The P4 of Dasycercus cristicauda is a single-rooted tooth, while in Dasyuroides byrnei it is double-rooted and usually has a more complex crown. The presence of Dasycercus cristicauda in the Madura Cave deposits is not unexpected in view of its presence in the Recent fauna. This species is widely distributed in desert areas in the central part of Australia where it occupies sandridge, stony de- sert, and spinifex grassland situations (Ride, 1970). Its presence in the lower units of Madura Cave indicates that it inhabited more humid areas in the past. LUNDELIUS & TURNBULL: MADURA CAVE 47 An examination of Tables 15-17 and Figure 15A, B, C suggests that the species underwent a size increase between the time of deposition of unit 2 and unit 1. The differences between various dental measures of the two samples were tested for statis- tical significance by means of the Mann- Whitney U test (Siegel, 1956, pp. 116-126). The results are summarized in Table 18. The measures that show statistically significant differences at P < .05 are those of the molars, primarily the lengths. This trend toward larger size is contrary to that shown by most mammals through the Pleistocene-Recent transition, but it is not unknown. Kurten (1965) has shown that the red fox {Vulpes fulva) in the Pales- tinian cave sequence increased in size through this time. Such changes may reflect some improvement of a limiting environmental requirement. Dasyuroides Spencer, 1896 Dasyuroides byrnei Spencer, 1896. Tables 19-24; Figures 5B, 15-17. Material. — Lower Dentitions Trench 3 Unit 2 WAM 74.9.27, left ramus with broken M2, M34, alveoli for P23 TMM 41106-369, right ramus with M2-4; alveoli for other teeth TMM 41106-370, right ramus with P3, M2-3; alveoli for P2, Mi, M4 TMM 41106-371, right ramus fragment with M24 TMM 41106-372, left ramus fragment with M1-4; alveoli for P23 TMM 41106-373, right ramus with P3, Mi-4; alveoli for other teeth TMM 41106-374, right ramus with dP4 or P4, Mi-4; alveoli for incisors, C, P23, ?dP3. (fig. 17E, F.) WAM 74.9.28, left ramus fragment with P2, broken P3, Mi-4 TMM 41106-376, left ramus with P3-M4, alveoli for other teeth TMM 41106-377, right ramus fragment with M1-4; alveoli for P3, dP4, or P4 TMM 41106-378, right ramus with Mi-4; alveoli for C-P3 48 FIELDIANA: GEOLOGY, VOLUME 38 TMM 411Q6-379, left ramus with P3-M4; alveoli for other teeth TMM 41106-380, right ramus with P3, M2-4; alveoli for C, P2, Mi WAM 74.9.29, right ramus with P3-M4; alveoli for other teeth WAM 74.9.30, right ramus fragment with M1-3; alveoli for C, P23, M4 TMM 41106-383, right ramus with P2-3, M1-3; alveoli for other teeth TMM 41106-384, right ramus fragment with P3-M4, alveolus for P2 TMM 41106-385, left ramus fragment with M2; alveoli for P3, Mi, M34 TMM 41106-386, left ramus, edentulous, with alveoli for all teeth TMM 41106-387, left ramus fragment with M2-3; alveoli for incisors, C, P2-M1 TMM 41106-388, right ramus with Mi-4; alveoli for other teeth TMM 41106-389, left ramus fragment with M2-3; alveoli for C, P2-3, Mi, Mi TMM 41106-390, right ramus fragment with M2-4 TMM 41106-391, right ramus fragment with M2-4, alveoli for Mi TMM 41106-392, right ramus fragment with M2-4, alveoli for Mi TMM 41106-393, right ramus fragment with broken P3, Mi 3; alveoli for C, P2 TMM 41106-394, left ramus fragment with M3-4, alveoli for M2 TMM 41106-395, left ramus fragment with P3, M2-3; alveoli for C, P2, Mi TMM 41106-396, left ramus fragment with Ms, alveoli for Mi TMM 41106-397, left ramus fragment with M3-4 TMM 41106-398, right ramus fragment with Mi 3; alveoli for incisors, C, P2-3 PM 25583, left ramus fragment with P3, alveoli for Mi PM 28360, left ramus fragment with M2; alveoli for C, P2-3, Mi PM 28361, right ramus fragment with M4 PM 29356, right ramus fragment with M3; alveoli for Mi 2, Mi PM 29357, left ramus fragment with Mi; alveoli for C, P2-3 PM 29359, right ramus fragment with P3-M1; alveoli for in- cisors, C, P2, M2 LUNDELIUS & TURNBULL: MADURA CAVE 49 PM 29362, right ramus fragment with M2-4 PM 29363, right ramus fragment with Mi 3; alveoli for C, P2-3 PM 29364, left ramus fragment with M4 PM 29365, left ramus, edentulous, (juv.) PM 29366, left ramus fragment with M2-4 PM 29367, right ramus fragment with P3-M4; alveoli for C, P2 PM 30506, right ramus fragment with M2-4 Unit 2, level 2 TMM 41106-138, right ramus, edentulous, with alveoli for all teeth TMM 41106-401, left ramus with P2-3, Mi-4 TMM 41106-403, right ramus with M1-4; alveoli for incisors, C, P2-4 TMM 41106-404, left ramus fragment with M3-4, alveoli for M2 TMM 41106-406, right ramus with P3-M4; alveoli for C, Rz TMM 41106-407, left ramus with M1-3, broken Mi; alveoli for other teeth TMM 41106-409, right ramus fragment with M3; alveoli for Ml-2, Ml TMM 41106-410, left ramus fragment with M3; alveoli for M2, M4 PM 29370, right ramus fragment with M4 Unit 3 TMM 41106-48, right ramus with M2-4; alveoli for C-Mi TMM 41106-49, left ramus with M2-4; alveoli for C-Mi TMM 41106-50, right ramus with M2-4, alveoli for Mi PM 29384, right M2 or 3 PM 29385, left ramus fragment with M4, alveoli for M3 PM 29386, right ramus fragment with M3-4, alveoli for M2 PM 29387, right ramus fragment with M2; alveoli for Mi , M3-4 PM 29388, right ramus fragment with Mr, alveoli for M2-3 PM 29389, left ramus fragment with Mi PM 30563, right ramus fragment, edentulous, with alveoli for M3-4 Trench 4 Unit 1, level 1 PM 30507, left ramus fragment with M3, alveolus for M* PM 30508, left ramus fragment with broken Mi; alveoli for P3, M2-3 PM 30509, right ramus fragment with M2-3 50 FIELDIANA: GEOLOGY, VOLUME 38 Unit 2, level 1 PM 30459, right ramus with M1-4; alveoli for other teeth PM 30460, right ramus with M1-4; alveoli for C, P2-3 PM 30461, right ramus with P3-M4; alveoli for other teeth PM 30462, right ramus fragment with M2-4 PM 30463, right ramus with M3-4; alveoli for other teeth PM 30464, right ramus fragment with M3-4 PM 30465, right ramus fragment with M2-4 PM 30466, left ramus fragment with M4, alveolus for M3 PM 30467, left ramus with C, P2-3, M2-4; alveoli for incisors, Mi PM 30510, right ramus fragment with M2or3 PM 30511, right ramus fragment with M2-3; alveoli for in- cisors, C, P2-3, Mi WAM 74.9.31, right ramus fragment with M2-3; alveoli for Mi, M4 WAM 74.9.32, right ramus fragment with M3; alveoli for M1-2 Unit 2, level 2 PM 30482, left ramus with M2-4; alveoli for P3, Mi PM 30483, left ramus fragment with M3, broken Ma PM 30484, left ramus fragment with M3-4 PM 30485, left ramus fragment with M2or3, alveolus for M3or4 PM 30491, right ramus fragment with M3-4 WAM 74.9.33, right ramus with P2-3, M1-4; alveoli for incisors, C. (fig. 17G, H) WAM 74.9.34, right ramus fragment with M* PM 30494, right ramus with M2-4; alveoli for other teeth PM 30495, right ramus with P3-M4; alveoli for other teeth PM 30496, right ramus fragment with M2-3; alveoli for M4 PM 30497, left ramus with M2-4; alveoli for other teeth PM 30498, left ramus fragment with M3, broken M4 PM 30499, left ramus fragment with Mi PM 30500, left ramus fragment with M1-2, roots of Mb Units 4-5 PM 29457, left ramus fragment with Mi PM 30514, left ramus fragment with M3-4 PM 30515, left ramus fragment with M2or3 PM 30516, right ramus fragment with M4, alveolus for M3 PM 30517, right ramus with M2-4; alveoli for C-Mi including P4 or dP4 PM 30518, right ramus fragment with Mi; alveoli for incisors, C, P2-3, M2 LUNDELIUS & TURNBULL: MADURA CAVE 51 PM 30519, right ramus fragment with Mi Unit 5 PM 29423, right ramus fragment with Mr, alveoli for M2-3 PM 29424, left ramus fragment with M3; alveoli for M2, M4 Unit 7 (or possibly Unit 5) PM 30472, right ramus with P2-3; M1-3; alveoli for incisors, C, M4 PM 30473, right ramus fragment with M2-3; alveoli for P3, Mi, M4 PM 30474, right ramus fragment with M2-4 PM 30475, right ramus with M2-4; alveoli for C-Mi PM 30476, right ramus with M2-4; alveoli for C-Mi PM 30477, right ramus fragment with P3; alveoli for incisors, C, P2-M1-2 PM 30478, right ramus fragment with M3; alveoli for M1-2; Mi PM 30479, right ramus fragment with Mi PM 30480, right ramus fragment with M2-4 PM 30481, left ramus fragment with broken M2, M3-4; alveoli for Mi Unit 7, level 2 PM 29392, left ramus fragment with M3; alveoli for M2, Mi PM 29393, right ramus with M3-4; alveoli for P2-M2 PM 29394, right ramus fragment with M3 (or 2); alveoli for M2 and 4 (Or Ml and 3) PM 29395, left ramus fragment with M2-3; alveoli for P2-3, M PM 29396, right ramus fragment with Mi; alveoli for P3, M2 PM 29397, right ramus fragment with P2-3, M2; alveoli for C, Mi, M3 PM 29398, right ramus fragment with M3-4, alveoli for M2 PM 29399, left ramus fragment with M2-4, alveoli for Mi PM 29400, right ramus fragment with M2; alveoli for C, P2-3, M PM 29401, left ramus fragment with Mi; alveoli for M1-3 PM 29402, right ramus fragment with M3-4; alveoli for Mi 2 PM 29403, right ramus fragment with Mi; alveoli for C, P2-3, M2-4, also minute pits possibly for Pi or dPi Trench 5 Unit 5 or 6 PM 30501, left ramus fragment with Mi ; alveoli for C, Pw, M2 52 fieldiana: geology, volume 38 Upper Dentitions Trench 3 Unit 2 TMM 41106-364, right maxillary with P34, M1 4, alveoli for C, P2 (fig. 17A, B) TMM 41106-365, right maxillary with M14, alveoli for C, P24 TMM 41106-368, left maxillary with dP4, unerupted P4, M12; alveoli for C, P23, M3, (fig. 17C, D) PM 29348, right maxillary with P4, roots of dP4, M13; alveoli for C, P23, M4 PM 29349, right maxillary fragment with M23, alveolus for M4 PM 29350, right maxillary with M1"3, alveoli for C, P24, M4 PM 29351, right maxillary fragment with M2"4 PM 29353, right maxillary with M12; alveoli for P2 4, M3"4 WAM 74.9.35, right maxillary fragment with M1; alveoli for C, P24 WAM 74.9.36, left maxillary fragment with P2"3; alveoli for C, P4, M1 Unit 2, level 4 PM 26163, right maxillary fragment with P4; alveoli for C, P2-3, M1 Unit 3 TMM 41106-51, left maxillary fragment with M23, alveolus for M4 PM 29379, right maxillary with P4-M2; alveoli for C, P2"3, M34 PM 29380, left maxillary with P4-M4; alveoli for P23 PM 29381, right maxillary fragment with M23; alveoli for M1, M4 PM 29382, left maxillary fragment with M1; alveoli for M24 Unit 3, level 2 TMM 41106-323, left maxillary fragment with P4-M3, alveolus for M4 TMM 41106-324, right maxillary with M1 3; alveoli for C, P24, M4 TMM 41106-325, left maxillary with M1 4; alveoli for P24 TMM 41106-411, left maxillary with M12, broken M3; alveoli for C, P3 4, M4 TMM 41106-412, right maxillary with M12; alveoli for C, ?dP4, M3 TMM 41106-413, left maxillary fragment with P24, M1, al- veolus for C LUNDELIUS & TURNBULL: MADURA CAVE 53 WAM 74.9.37, left maxillary fragment with M2, broken M3, M4, alveolus for M1 PM 29371, left maxillary fragment with M3; alveoli for M2, M4 PM 29372, left maxillary fragment with M1, roots of dP4 (nearly resorbed), crypt for P4, alveolus for M2 PM 29373, left maxillary fragment with M1 PM 29374, left maxillary fragment with M12 PM 29375, left M3 PM 29376, right maxillary fragment with M2 or 3 PM 29377, left maxillary fragment with M3; alveoli for M2, M4 PM 29378, right maxillary fragment with M34 Trench 4 Unit 1, level 1 TMM 41106-511, right maxillary fragment with M14; alveoli for P2 4 Unit 2, level 1 WAM 74.9.38, left maxillary with P^M3; alveoli for P23, M4 PM 30469, left maxillary fragment with M1; alveoli for C-P4 PM 30470, right maxillary fragment with M1"4; alveoli for P3"4 PM 30471, right maxillary fragment with M12, alveolus for M3 PM 30486, right maxillary with P3"4, M14; alveoli for C, P2 PM 30487, right maxillary fragment with M1"3, alveolus for M4 PM 30488, right maxillary fragment with P34, M1, alveolus for P2 WAM 74.9.39, right maxillary fragment with M3; alveoli for M2, M4 WAM 74.9.40, right maxillary with P»-M2; alveoli for C, P23, M3 Unit 2, level 2 PM 29407, right maxillary with P^M4, alveolus for P3 PM 29408, right maxillary fragment with M2; alveoli for M3"4 PM 29409, left maxillary fragment with M2 3, alveolus for M1 PM 29410, right maxillary fragment with M2 PM 29411, right maxillary fragment with M2, alveolus for M1 PM 29412, right maxillary with M\ broken M2, M3"4; alveoli for C-P4 PM 30449, left maxillary fragment with M3; alveoli for M2, M4 PM 30450, right maxillary fragment with M3; alveoli for M12; M4 54 FIELDIANA: GEOLOGY, VOLUME 38 WAM 74.9.18, right maxillary fragment with P4; alveoli for C, P23 M1 PM 30452, right maxillary fragment with M3; alveoli for M2, M4 PM 30453, left maxillary fragment with broken M3; alveoli for M2, M4 PM 30454, left maxillary fragment with M1; alveoli for C, P2 4 PM 30455, right maxillary fragment with M1 2, alveolus for M3 PM 30456, right maxillary fragment with P4, M1 PM 30457, left maxillary fragment with M2; alveoli for M1, M3 PM 30458, right maxillary fragment with P4, M1; alveoli for C, P23 Units 4-5 PM 30504, left maxillary fragment with M2 or 3 PM 30505, right maxillary fragment with M3; alveoli for M1 2, M4 Unit 7, level 1 PM 29414, left maxillary fragment with M3; alveoli for M2, M4 WAM 74.9.41, left maxillary fragment with M2; alveoli for M3-4 Unit 7, level 2 PM 29404, left maxillary fragment with M3; alveoli for M2, M4 PM 29405, left maxillary fragment with M3; alveolus for M4 PM 29406, ?left M3 Trench 5 Unit 5 PM 30437, right maxillary fragment with P4; alveoli for P2 3, M1 (fig. 10C, D) Unit 5 or 6 PM 30502, right maxillary fragment with M3, alveolus for M4 PM 30503, left M2 or 3 Description . — Mandibles: The mandible of Dasyuroides byrnei is similar to that of Dasycercus cristicauda . The horizontal ramus is convex ventrally and has two mental foramina, one under the Mi and one under the posterior end of P2. The anterior border of the ascending ramus rises steeply above the tooth row; the mas- seteric fossa is wide. Lower Dentitions: No specimen has the lower incisors pre- served, but a number have alveoli indicating that three lower in- LUNDELIUS & TURNBULL: MADURA CAVE 55 cisors were present, as in the Recent specimen, and that the teeth were arranged in the same way with the root of the Ii extending under the roots of the two lateral incisors at depth. The canine is preserved in only one specimen of Dasyuroides byrnei (PM 30467). It is a prominent tooth, recurved posteriorly and lingually. Its base is expanded posteriorly. A lightly defined cingulum extends from the edges of this base anteriorly and up- ward on each side of the tooth and joins on the anterior edge. The lower part of the posterior edge has a small wear facet where it comes in contact with the upper canine. The two lower premolars, P2 and P3, are similar in size and shape and will be described together. Both are oval in occlusal view and each has one principal cusp located in the anterior half of the tooth. The principal cusp of each has anterior and posterior ridges that extend to cingular cuspules on each end of the tooth. The posterior cuspule is the larger in each premolar. Each is en- circled by a cingulum that is more developed posteriorly. The posterior border of P2 is convex rather than concave as in many phascogalines. The degree of crowding of the premolars is vari- able. In those specimens in which the premolars are arranged en echelon , the posterior end of each tooth is lingual to the anterior end of the following tooth. There is normally no P4 present, but three specimens (TMM 41106-373, -374, -377) have either small teeth or alveoli for small teeth that are probably the dPi and/or the Pi . The first specimen has a remnant of a tooth root between the P3 and Mi. TMM 41106-374 has one alveolus for P2 immediately behind that of the canine, and it is separated from that for the anterior root of the P3 by a small diastema with a pitted alveolar surface suggesting that the posterior part of P2 may have been broken away during life, and the alveolus occluded by bone. The alveoli for P3 appear to be normal sized for Dasyuroides but are located slightly more labially than usual. Lingual to the anterior alveolus of P3 are two tooth fragments which appear to be either the remnants of two small single-rooted teeth, or of a somewhat larger double-rooted tooth. The anterior of these two tooth stubs shares an enlarged alveolus with an unerupted tooth. Lingual to the posterior al- veolus of P3 is a double-rooted tooth with an expanded base and single conical cusp (dimensions: 1 = .95 mm., w = .65 mm.). TMM 41106-377 has a single rooted Pi .38 mm. in diameter. The Mi is a distinctive tooth. The trigonid is laterally com- 56 FIELDIANA: GEOLOGY, VOLUME 38 \ V * M\ %? Fig. 16. Dasyuroides byrnei FM 104786. A, B, Rostrum with right upper denti- tion shown in labial and crown views, I4 restored from left side. P4 not present on left side. pressed, especially its anterior part. The protoconid is the largest cusp, is located centrally in the anterior part of the tooth, and is elongated anteroposteriorly. The metaconid is much smaller and is located on the posterolingual flank of the protoconid. The para- conid is reduced to a small cingular cuspule on the anterior end of the tooth. The anterior cingulum disappears on the lingual side of the tooth over the anterior root of the tooth. Labially the cingu- lum is continuous from the anterior end of the tooth to the hypo- conulid. In occlusal view the anterolabial corner of the tooth is a right angle. The hypoconid is the largest cusp of the talonid. It is almost as tall as the metaconid and is more massive. The pre- metacristid (from the hypoconid) is separated from the postpara- cristid (from the metaconid) by a narrow notch. The hypoconid is connected to the hypoconulid at the posterior end of the tooth by a J3 £ Q 57 58 FIELDIANA: GEOLOGY, VOLUME 38 continuous postmetacristid that is oriented approximately 30° from the transverse axis of the tooth for most of its length. It turns abruptly posteriorly just as it joins the hypoconulid. The entoconid is somewhat variable in size but is a distinct and well- formed conical cusp on most specimens. It may or may not be joined to the metaconid by a low entocristid. It is joined by a low crest to the postmetacristid at the point where it turns posteriorly to join the hypoconulid. The talonid basin is oval with its long axis extending from the base of the protoconid to the entoconid. The margin formed by the metaconid is noticeably steeper than that formed by the hypoconid. The M2 and Mb are similar in size and morphology and will be described together. The trigonid shows no features that distin- guish it from other dasyurids of comparable size. The anterior and posterior cingula are very broad. In occlusal view, the an- terior cingulum makes a right angle as it joins the much weaker cingulum on the labial side of the protoconid. The cingulum is variably developed on the labial side of the tooth. In some speci- mens it is absent on the sides of the protoconid and hypoconid. The talonid basin is square and smooth. The hypoconid is the largest cusp of the talonid. A premetacristid extends anteriorly from the hypoconid and is separated by a narrow cleft from the postparacristid, which runs posteriorly from the protoconid. The entoconid is large in most specimens; in some specimens it stands almost as high as the hypoconid. In a few specimens the entoconid is relatively low and elliptical on one or two of the molars. In one specimen (PM 30467) the entoconids are low and elliptical on all the molars. In its typical form, the entoconid is broadly oval at its base and circular near its apex. The entoconid may be joined to the metaconid by a low entocristid but is separated from the hypoconulid by an open groove. A small cuspule is present on the postmetacristid of most specimens where it turns posteriorly to join the hypoconulid. This cuspule is removed early by wear. The M4 differs from the M2 and M3 only in the small size of its talonid. The talonid is highly variable in size, but is usually basined and has a hypoconid. A small entoconid is sometimes pre- sent. No distinctive hypoconulid is present on the M4 of any of the Madura Cave specimens, although most have a terminal cingular bulge in this position behind the hypoconid. Maxillaries and upper dentitions: Skull material of Dasyuroides byrnei is limited to maxillary fragments with teeth. LUNDELIUS & TURNBULL: MADURA CAVE 59 The infraorbital foramen is large and opens over M1 as in the Recent specimen. Anterior to the upper canine in specimen TMM 41106-413 is the posterior part of the pit into which the lower canine fits when the jaws are occluded. No incisors or canines are represented in the Madura Cave sample, but alveoli in a number of specimens show that size of the canine tooth in the fossils was comparable to that in the Recent specimen. The P2 and P3 are similar in shape, but the P2 is markedly smaller than the P3. They are each oval in shape with a principal cusp located just anterior to the midpoint of the tooth. Each is encircled by a cingulum that is weak, or sometimes absent, lingually. There may be other small interruptions in the cingu- lum that give an irregular appearance to the edge of the tooth in occlusal view. There are usually cingular cuspules, one at each end of the tooth. The ridge connecting the principal cusp with the posterior cingulum is usually the better defined of the two. One specimen (TMM 41106-368) has the dP4 in place and above it a developing P4 in the crypt (fig. 17C, D). The dP4 is single-rooted, peg-like oval in crown outline with its long axis oriented obliquely to the tooth row. Its crown consists of a pointed central cusp with tiny anterior, posterior, and labial cingular tubercles. In the Madura Cave sample the P4 is variable in size. In most specimens it is small, oval, or triangular in occlusal view. It usu- ally has a distinct central cusp and an encircling cingulum. It may have one or two roots. The larger P^'s (such as PM 30437) may have posterior and labial cingular cuspules. One specimen (PM 30486) has a single-rooted, peg-like tooth in the P4 position. This tooth is similar to the dP4 of TMM 41106-368 in size, but unlike that specimen there is no developing P4 to be found in the maxillary above it. The situation in the Recent materials appears to be similar. The one Recent specimen we have available for comparison (FM 104786) has a peg-like tooth in the P4 position on the right side, and no tooth on the left. Tate (1947, p. 153) gives length measures of 1.1 and 1.25 mm. for the P^s of two Recent specimens from central Australia. Our measurements of a small series in the BMNH are 1.09, 1.34, 1.32, and 0.97 mm. These measurements are within the range of size of the undoubted P^'s of the Madura Cave sample (fig. 15E). The resemblance of the small teeth in the Recent specimen and PM 30486 to the un- c aj X3 c Q c c 08 o a u "3 _c -a co c C SL e8 j^ C M =2 h a a i 'o 0) 03 o "5 a> CO C »" 3 "8 s "o 03 u 9 -a 5? _> c PL, 03 03 -C T3 BO 4-1 C *H '5 CO l"~l CO XI >» u c L 03 03 03 O £ "o 'B CJ 03 i j> 1 03 *a ap * be 3 o £ s 03 E £ "rf o CD § x CO 03 +-> CO ■^ '£ o CO I— 1 i-H 2 3 E a) Tf -c s +j s- s 'i p CO 3 be PQ E 03 s h CO < ~> CO rC 05 oj M •* > CC jS l> O ^ s 09 t~ < Si 3 cd £ T3 03 o i— i s £ ^H E o "^ a 2 CO £ 2 £ H aj s '> s k, fa. C £ H c co tf u U -a c 03 03 C "o3 3 a o be q u CJ c T3 — ' t-- C c »H 03 '-" ,__, c U 03 u fa* .C _ 60 > « cfl "° •s c -^ eo CO ^ £ T3 's « CO c u r .O C8 (l) i—i > , ' > .-« £ 6 « — _ — be cri 05 SI . a 61 62 FIELDIANA: GEOLOGY, VOLUME 38 doubted dP4 of TMM 41106-368 suggests that the dP4 is variably retained and in some individuals is not replaced by a P4. The peg-like dP4 of Dasyuroides byrnei is similar in size and morphology to the tooth that usually occupies the P4 position in Dasycercus cristicauda (fig. 14A, B; table 17). One specimen in the Murraelellevan sample (PM 4880) has a tooth on its left side that resembles the P4 of Dasyuroides byrnei both in size (fig. 15E) and morphology. On its right side it has two teeth, one re- sembling the P4 of Dasyuroides byrnei in morphology but not in size and one resembling the usual single-rooted peg seen in the remainder of the Dasycercus cristicauda sample. The abnormal situation seen in this specimen, plus the undoubted replacement of a simple dP4 by a more complex P4 in Dasyuroides byrnei suggest that the usual single-rooted tooth in the P4 position in Dasycercus cristicauda is also a dP4. The M1 has the asymmetrical shape typical of the Mrs of other dasyurids. The metacone is the highest cusp, followed in order by the stylar cusp C, paracone, and protocone. The metacone and sty- lar cusp C are close together but separated by a deep groove. The stylocone is fused with the paracone, and a distinct parastyle with a short cingulum is present. The protocone is long anteropos- terior^; crests from it extend anteriorly around the metacone. A small protoconule is present. A very indistinct stylar cusp D can be seen close to the metastyle on unworn teeth. The M2 is triangular and slightly asymmetrical. The metacone is the tallest cusp, with stylar cusp C only slightly shorter, fol- lowed by the stylocone, paracone, and protocone. The labial edges of the major stylar cusps are almost in line, resulting in a nearly straight edge to the stylar shelf. The M3 is triangular with stylar cusp C located far lingual to a line that connects the stylocone and the metastyle. The metacone is the tallest cusp, followed by cusps C, the paracone, stylocone, and protocone. The parastyle is joined to an anterior cingulum that reaches lingually to the base of the midpoint of the paracone. There is no posterior cingulum. The protocone is less massive than in M1 or M2. Protoconule and metaconule are present just anterolingual and posterolingual to the bases of paracone and metacone respectively. The stylar shelf bears a small cusp C, and a cusp D is located near the metastyle. LUNDELIUS & TURNBULL: MADURA CAVE 63 The M4 is reduced posteriorly. The stylocone and paracone are the largest cusps. They are widely separated and joined by a slightly sinuous paracrista. The metacone is joined to the para- cone by a short, straight centrocrista. The protocone is a small, lingually rounded cusp located lingual to and lower than the paracone; its margins extend around the base of the paracone. An anterior cingulum is present from the stylocone to the midpoint of the paracrista. The stylar shelf is a posteriorly sloping surface with no cusps. Discussion. — The two dasyurids that are most similar to Dasyuroides byrnei are Phascogale tapoatafa and Dasycercus cris- ticauda: The distinctions between the upper dentitions of Dasyuroides byrnei and Dasycercus cristicauda have been dis- cussed in detail in the section dealing with the latter taxon. The upper dentition of Phascogale tapoatafa differs from that of Dasyuroides byrnei primarily in the larger size of the P4 and the M4. The other molars are much the same size. The P4 is al- ways present in P. tapoatafa and is sometimes absent in Dasyuroides byrnei. The P4 of Dasyuroides byrnei, when present, is shorter, lower crowned, and relatively less compressed laterally. The dP4 of P. tapoatafa is longer than that of Dasyuroides byrnei (fig. 15E). The M4 of P. tapoatafa is longer anteroposteriorly as a result of the greater development of the stylar area. The lower dentition of Dasyuroides byrnei can be distin- guished from that of Phascogale tapoatafa by the more reduced and laterally compressed trigonid of its Mi. This has been de- scribed in detail in the P. tapoatafa section. Dasyuroides byrnei can be distinguished from Dasycercus cristicauda by the rela- tively longer protoconid and larger paraconid of the Mi and a trend toward larger and more conical entoconids on Mi through M3. The lower canine of Dasyuroides byrnei differs from that of Dasycercus cristicauda in being relatively shorter, more procum- bent and recurved, with a greater expansion at its base. Dasyuroides byrnei is known as a living animal only from the area where Queensland, South Australia, and the Northern Territory come together. It is an occupant of desert grassland or desert steppe (Spencer, 1896; Marlow, 1962; Ride, 1970). Finlay- son (1961) has reported it from Charlotte Waters, Killalpaninna, and Birdsville. Its presence at Madura Cave demonstrates that it 64 FIELDIANA: GEOLOGY, VOLUME 38 had a much wider distribution in the past. Remains of this species are rare in the upper stratigraphic unit (Unit 1). It apparently was disappearing from this region at the time of the climatic change that is implied by the change in the character of the sedi- ments between units 1 and 2 (Lundelius and Turnbull, 1973). The waning number of Dasyuroides byrnei coincides with a small increase in size of Dasycercus cristicauda. These two species are quite similar in size and structure and, although no informa- tion is known to us about their competitive relationships where they occur together today, it is possible that the reduction in numbers of Dasyuroides byrnei somewhat lessened the restrictions on Dasycercus cristicauda and permitted an increase in size. INCERTAE SEDIS Dasycercus cristicauda or Dasyuroides byrnei Material. — Trench 3 Unit 2 PM 29352, left maxillary fragment with M1; alveoli for C, P2 3, dP4 , or P4 Description and Discussion. — This specimen cannot be un- equivocally assigned to either taxon. It is intermediate in size, and the one tooth has no morphological features that allow an assignment. Additional uncatalogued, edentulous ramus and maxillary fragments or isolated teeth which probably belong to one or the other of these taxa are: Trench 4, Unit 2, level 2 One maxillary, five ramus fragments, and approximately 50 isolated teeth. Trench 4, Unit 2, level 3 Four ramus fragments. Trench 4, Units 4-5 Two ramus fragments. Trench 4, Unit 7, level 1 One maxillary, nine ramus fragments, and one isolated lower molar. Trench 4, Unit 7, level 2 Five maxillary and 17 ramus fragments. LUNDELIUS & TURNBULL: MADURA CAVE 65 Trench 5, Unit 5 One ramus fragment and approximately 25 isolated teeth. Dasyurus Geoffroy, 1796 Dasyurus (Dasyurinus) geoffroyi Gould, 1841. Tables 25-27; Figures 18-19. Material. — Lower Dentitions Trench 3 Unit 2 WAM 74.9.42, right ramus fragment, edentulous, with al- veoli for Ma-4, (fig. 18 A2, B2) Unit 3 TMM 41106-44, right ramus fragment, edentulous, with al- veoli for M3-4 Trench 4 Unit 1 TMM 41106-477, left ramus fragment with alveoli for C-M2; M2 that may belong to the jaw. (fig. 18A3, A4, B3, B4) TMM 41106-519, right Ma TMM 41106-554, right lower C (fig. 18 B7) Unit 2, level 1 PM 30525, left M2 (probably goes with PM 30526) (fig. 18 A5, B5) PM 30526, left Ma (probably goes with PM 30525) (fig. 18 A6, B6) Unit 2, level 2 PM 30528, right Mi (fig. 18B8) PM 30529, broken left M? PM 30530, left M2 PM 30531, left M2 WAM 74.9.43, left M2 Units 4-5 WAM 74.9.44, right M2 PM 30541, left M2or3 PM 30542, left M2 PM 30545, right Mi PM 30546, right M2 66 FIELDIANA: GEOLOGY, VOLUME 38 Unit 7, level 1 PM 30555, left M2 Unit 7, level 2 PM 30556, left Ms WAM 74.9.45, right Mi Trench 5 Unit 5 or 6 PM 30562, left Ms Upper Dentitions Trench 3 Unit 2 TMM 41106-68, left M1 TMM 41106-69, right maxillary with M1; alveoli for C-P3, M23, (fig. 19A2, B2) PM 30520, left M1 Unit 3 PM 30521, right M1 Trench 4 Unit 1 PM 30522, broken right M1 PM 30523, left M3 Unit 2, level 1 PM 30524, left maxillary with M1; alveoli for P3, M23 PM 30527, right M3 (fig. 19 A5, B5) Units 4-5 PM 30533, broken right M1 PM 30534, left M1 PM 30535, left M1 WAM 74.9.46, right M1 WAM 74.9.47, broken right M2 PM 30538, right M2 PM 30540, right P3 (fig. 19A3, B3) PM 30543, right M1 PM 30544, right M2 (fig. 19A4, B4) PM 30547, left M1 PM 30548, left M1 Unit 7, level 1 PM 30553, right M1 PM 30554, right M2 LUNDELIUS & TURNBULL: MADURA CAVE 67 Unit 7, level 2 PM 30557, right M2 Trench 5 Unit 5 PM 30559, right M3 PM 30560, right M3 WAM 74.9.48, right M3 Description. — Mandibles: The fossil mandibular material closely resembles a Recent specimen of Dasyurus geoffroyi from southwestern Australia (FMNH 35329, fig. 18A1, Bl; 19A1, Bl) in all morphological characters, but is slightly smaller. Specimen TMM 41106-477 (fig. 18A3, B3), having alveoli for the canine P2, P3, and Mi, particularly demonstrates the similarity in size and arrangement of these teeth. A large mental foramen opens under Mi . The characters of the posterior end of the mandible are shown by specimen WAM 74.9.42 (fig. 18A2, B2). The anterior edge of the ascending ramus rises about 60° from the line of the tooth row, the angle of the masseteric fossa is approximately 45°, and the ventral border of the masseteric fossa curves dorsally in a posterior direction. The articular face of the condyle is oval and convex anteroposteriorly. The angular process has a ridge on the dorsal surface. The specimen is that of a young adult as shown by the position of the posterior alveolus of the M4 lingual to the an- terior border of the ascending ramus. This probably accounts for some of the size difference between the Madura Cave specimen and the Recent specimen. Lower Dentitions: No lower incisors of this taxon have been recognized. The lower canine is a simple curved tooth (TMM 41106-554, fig. 18B7). It is oval in cross-section for most of its length and is twisted medially toward the apex. The root con- stitutes more than half of the length of the tooth and is expanded just below the base of the crown. The medial surface at this point has a broad, shallow, concave groove. The posterior edge of the crown has a thin ridge from apex to base. A faint basal cingulum is present lingually; it extends anteriorly and downward to join a poorly defined ridge on the anterior edge of the tooth at a cingu- lar cuspule. The flattened lingual surface of the crown is continu- ous with that of the root. The Mi is a laterally compressed tooth with a reduced trigonid (PM 30528, fig. 18B8). The protoconid, by far the largest cusp of the trigonid, is laterally compressed with sharp ridges on both the 68 FIELDIANA: GEOLOGY, VOLUME 38 anterior and posterior edges. The anterior ridge connects the apex with a minute cingular cuspule and usually there is no paraconid. The anterior cuspule is variable in its size and relationship to other structures. In PM 30528 there is a small but distinct para- conid and a definite carnassial notch on the anterior ridge of the protoconid. In PM 30558 there is another small cingular cuspule in the position of the missing paraconid, but it is situated just lingual to the ridge. The posterior ridge of the protoconid connects the apex with the metaconid. The anterior cingulum is large but does not extend onto the base of the protoconid. The metaconid is variable. In WAM 74.9.45 and PM 30528, the metaconid is a small but discrete cusp, round in cross-section, that is located on the posterolingual flank of the protoconid. In PM 30545 the meta- conid is reduced to a small rounded cuspule in the same position. The talonid basin is oval with its long axis extending from the posterolabial corner of the protoconid to the groove between the entoconid and hypoconulid. The hypoconid is the largest cusp of the talonid, followed by the entoconid and hypoconulid. The hypo- conid is connected with the hypoconulid by a postmetacristid that turns abruptly posterior just anterior to the hypoconulid. A well- defined carnassial notch separates the premetacristid from the postparacristid on the labial side of the tooth. The postparacristid is a prominent ridge on the posterior side of the protoconid. The entoconid is large, oval to round in cross-section, and separated from all other cusps by rounded grooves. The posterior cingulum does not have the usual ridge structure but is reduced to a smooth bulge and cuspule on the posterolabial corner of the tooth. The M2 and M3 are similar in size and morphology and will be described together (fig. 18A4, A5, A6; B4, B5, B6). The major difference between these two teeth is the greater lateral compres- sion of the trigonid of the M2 and its smaller paraconid and meta- conid relative to the protoconid. In both the M2 and M3 the trigonid is well developed. The protoconid is the largest trigonid cusp, followed by the paraconid and metaconid. The lingual sur- face of the protoconid has a broad, rounded ridge from apex to base. The paracristid and epicristid have strong carnassial notches. The parastyle is weak on most teeth and is separated from the anterior cingulum by a notch into which fits the hypo- conulid of the tooth in front. The talonid basin is round. The largest talonid cusp is the hypoconid, followed by the entoconid and hypoconulid. A strong carnassial notch separates the post- paracristid from the premetacristid. As in the Mi, the postpara- 69 70 FIELDIANA: GEOLOGY, VOLUME 38 cristid is a prominent ridge on the posterior face of the protoconid. The entoconid is large, oval in cross-section at its base, round at its apex. It is not joined to any other cusps by ridges. The hypo- conulid is the posterior extremity of the tooth. It is not joined to the posterior cingulum. The anterior cingulum is broad but is not joined to the parastylid and does not extend onto the labial sur- face of the tooth. The posterior cingulum is variable in its form; it may be ridge-like or it may be a rounded bulge, with or without a cuspule. It does not reach the hypoconulid or extend onto the la- bial surface of the hypoconid. There are no Mi's in the Madura Cave material. Maxillaries and upper dentitions: The only identifiable skull material of Dasyurus geoffroyi from Madura Cave are parts of two left maxillaries. One specimen (TMM 41106-69, fig. 19A2, B2) has an M1 and alveoli for C-P3 and M23. The other (PM 30524) has an M1 and alveoli for P3 and M2 3. These two speci- mens show the infraorbital foramen to be large and situated over the M1. There is no well-defined fossa on the lateral surface of the maxillary into which the foramen opens as in Dasycercus . No upper incisors, canines, or P2's have been identified for this taxon. One right P3 (PM 30540, fig. 19A3, B3) is present. It is a double-rooted tooth with a large cusp on its anterior half. Its greatest width is posterior to the apex of the main cusp where the cingular area is expanded on the lingual side. A cingulum is present on each side of the tooth, from the base of the main cusp to the posterior end of the tooth. A small cuspule marks the junc- tion of this posterior cingulum with a ridge on the posterior edge of the main cusp. An anterior cingulum is present that does not join the posterior one. This is similar to the P3 in the Recent specimen of Dasyurus geoffroyi. The M1 is a triangular tooth with the protocone extending as far anteriorly as the paracone (fig. 19A2, B2). The metacone is the tallest cusp, followed by cusp C, the paracone, and the proto- cone. The paracone and stylocone are fused. The parastyle is small and may be either rounded or bladelike. It is connected to the anterior cingulum. The metacone and cusp C are approxi- mately the same size, and, although close together, are separated by a deep, rounded groove. The premetacrista and a parallel crest on the anterior face of cusp C are variably developed. They ap- proach, but do not join, a postparacrista and a stylocrista from the paracone. Stylar cusps B and C may or may not be present. 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Cm 3 X u e cu CU a 'X c CD CO £2 C eg U E n M a CO *j ■M .cc£ H oo U5 ._f cu 9 CC C X CO C o E od m -r X x M £ 2 o CO c -a * C35 Cu > •^ cu *- CU oi -r X 9 b= '"t O" _o t~ P3 CN ►JJ 2 :« s Cu Cm c ,80 < Cm o 0) !ss a CO "o [- c CC TC cu £ c CO M o -TC TC » CO is 3 -a a * > "5 cj != i- c ■~ c CU eg w 0 d CO CO CO c '2 CJ eg c '2 CO c E S c > 0 CU E TC i. a eg o M a> £ o 9 is (S o a. '-. c 3j _c CO cu X 77 78 FIELDIANA: GEOLOGY. VOLUME 38 shows the fossil to have a somewhat more discrete facet pattern. (This may be the result of diet — the Recent specimen was cap- tive in a zoo for part of its life.) The single piece of an Mi (TMM 41106-548, fig. 20A4, B4) is the anterior half of a left tooth. It is from a young individual and is similar to the Mi of the Recent FMNH 57801. The break goes through the apex of the protoconid which is high and is situated over the center of the space between the two roots. An anterior crest connects the apex to the anterior cingulum. The cingulum is sharp around the front half of the tooth. As in the Recent speci- men, the protoconid is slightly recurved medially, and there is no paraconid. All other lower molars represent young individuals as the teeth are virtually unworn. The description is based mainly on the complete tooth (PM 30568). The tallest trigonid cusp is the protoconid, followed by the paraconid and metaconid. The apex of the protoconid is high and lies well within the posterior third of the tooth. The anterolabial face of the tooth is high, almost flat, and consists of a sharp, thin crest — the paracristid (F) which runs forward and down toward the base of the trigonid where it meets a posterolabially directed crest from the lower and promi- nent paraconid. There is a deep carnassial notch where the two parts of the paracristid meet. The epicristid takes the form of a steep posterior crest from the apex of the protoconid. It runs directly toward the small metaconid at the base of the trigonid but is separated from it by a short cleft. The posterior side of the metaconid joins the cingulum which swings postero-labially, then labially, to meet a posterior cingular cuspule just linguad to the small hypoconid, which it does not join. Instead it sends a minute, but distinct crest anterolabially forward to the base of the proto- conid. A tight shallow valley separates this ridge from the hypo- conid except at one point midway along its course to the proto- conid. A rounded, shallow basin with a small isolated cuspule within it is thus delimited between the metaconid and the hypo- conid. The hypoconid is elongated and is also tied to the base of the protoconid by a crest, just labiad the other crest. The hypo- conid is notably smaller than its counterpart in the Recent FMNH 57801 (where it is smaller than that of either Mi or M2). The cingulum rises slightly at the posterior edge of the tooth near the hypoconid in a cuspule that could be considered a hypo- conulid. There is a strong, anteriorly rising cingulum antero- labial to the paraconid. A small notch separates the highest point LUNDELIUS & TURNBULL: MADURA CAVE 79 of this cingulum from a minute parastylid, but this notch prob- ably served to contact the cingulum of the preceding tooth, for the Recent specimen has Nk's with no trace of a hypoconulid. The posterior cingulum is weakly developed along the lingual side of the base of the protoconid. It extends back and around the poste- rior edge of the tooth to disappear along the side of the posterior root beneath the junction of the hypoconid and protoconid on the labial side. Behind the carnassial notch of the paracristid and parallel to it there is an elongate shallow groove on the labial surface of the protoconid — a feature that appears to distinguish M2 and M3 of Sarcophilus from those of Thylacinus. Upper Dentitions: The I4 (fig. 21 A3, B3) has a worn crown, reduced apparently to about half its original height, and a bulb- ous root expansion typifying advanced age. The crown is rather triangular in occlusal view. A central main cusp has anterior and posterior crests that connect it to the weak cingula. The labial surface is smoothly convex and truncated by wear; the lingual surface is heavily worn in a deep bevel that cuts through the posterolingual cingulum. There is a small but distinct cingular cuspule at the posterior end of the main crest. Aside from differ- ences in wear the fossil material is similar to the Recent speci- men used for comparison (FMNH 57801, fig. 21A1, Bl). The upper canines are both heavily worn, and like the I4 have a bulbous root condition typical of old age. At least half of the crown of both teeth is worn away so that the cusp is truncated below midheight. On PM 30565 the anterior and posterior faces have the enamel worn away as well; on PM 30573 (fig. 21A2, B2) wear on the posterior face did not breach the enamel. The only enamel in PM 30565 is on the labial and lingual sides. On the labial side the enamel retains a weak convexity; on the lingual side it is almost flat. On both canines there is a constricted ap- pearance at the junction of crown and root, apparently the result of the bulbous root condition with some enhancement from wear along the posterior edge just beneath the crown. The P3 (fig. 21A4, B4) is an unworn tooth consisting of a blunt, nearly conical main cusp with a surrounding basal cingulum that is heaviest behind. There is a weak crest from the apex of the cusp up its posterolateral side that just reaches the cingulum. The two left M2's are similar. PM 30578 consists of only the anterior face of the tooth, including part of the protocone, about half of the paracone, the parastyle, and the anterior cingulum. CO CO c« <; u 3 «- CO "^ S£ 0) bo ■$£ §s .2 fa *-l ^ 5 w O Q. «2 * CO ,-h bo o "C 00 o co .2; to fa o I* as 0) 8. 1 o. > 3 _ CO ^ cp a> a co ^ o •* ■ 2 co — •2 5 £ fc fa * 03 w £ .2 la co CO 6 a 0) o D--C co co 80 LUNDELIUS & TURNBULL: MADURA CAVE 81 PM 30570 (fig. 21A5, B5) is complete and unworn. All three roots are well formed although not closed off. This tooth is triangular in crown view. The protocone is low, crested, and juts in toward the palate somewhat more than does its counterpart in the Recent specimen (FMNH 57801) so that there is a deeper notch in the anterior margin. The cresting of the protocone is V-shaped, and the anterior crest or protoloph (III') runs from the tip of the cusp parallel to the front edge of the tooth to the anteromedial edge of the base of the paracone. The posterior crest, plagiocrista, or metaloph (IV) extends diagonally back and labially to the base of the metacone and then rises abruptly to meet the cingulum. The paracone is high with posterolingual and anterolabial crests. The former crest goes to the base of the metacone; the latter connects the paracone to the stylocone. The metacone is the highest cusp and is also crested. A short crest swings labially from its apex to join immediately the compressed stylar cusp C. The posterior crest from the metacone extends straight back and then postero- laterally to a lightly-defined metastylar corner of the tooth. In the metastylar area a flattened and weakly crested stylar cusp D forms the main feature of the rear part of the stylar shelf between stylar cusp C and the vestigial metastyle. The paracone and metacone, each with its associated stylar cusps and crests, there- by form two shallow basins posterolateral^ inclined to their re- spective cusps. There is an anterior cingulum that descends to a weak parastyle. All of these features agree closely with the M2 of the Recent specimen. Discussion. — The material of Sarcophilus from Madura Cave is referred to S. harrisii on the basis of size and morphology. It is much smaller than that of S. laniaris and is either smaller than or equal to the smaller specimens of a Recent sample from Tas- mania and late Pleistocene specimens from the west coast area of Australia (table 28). Sarcophilus is known from three other localities on the Nul- larbor Plain, Balladonia (Glauert, 1912; Merrilees, 1968), Webbs Cave (Lundelius, 1963), and Cocklebiddy Cave (Merrilees, 1968). These occurrences are not well dated but are late Pleistocene or post Pleistocene in age. The Madura Cave material is associated with C-14 dates that demonstrate its presence on the Nullarbor Plain from about 20,000 to about 7,500 years before present. Some of the other occurrences are from superficial deposits and may be younger. A discussion of the distribution of this animal on the mainland is given by Calaby and White (1967). The burned 82 FIELDIANA: GEOLOGY, VOLUME 38 specimen is one of the few evidences uncovered to date that might indicate the presence of man at Madura Cave. Remains of Sar- cophilus have been found in other localities in circumstances that indicate that it was an aboriginal food item (Gill, 1953; Mulvaney et al., 1964; Calaby and White, 1967; Dortch and Merrilees, 1971). Thylacinidae Boneparte, 1838 Thylacinus (Temminck, 1824, 1827) Thylacinus cynocephalus (Harris, 1808). Table 29; Figures 22-23. Material. — Trench 3 Unit 2 TMM 41106-59, left P3 (fig. 23A2, B2) PM 30576, posterior half of right P4 (fig. 23A3, B3) WAM 74.9.51, right Mi (or M2), broken (fig. 22A3, B3) Trench 4 Unit 2, level 1 PM 30579, broken right M2 or 3 (fig. 22A2, B2) Description. — The material of Thylacinus is adequate only to demonstrate its presence in the fauna. Lower Dentitions: WAM 74.9.51 is the posterolabial face of a right lower molar. It may be considered an Mi because of the form of the posterior cingulum, the close spacing of the roots, the plunging crown rim over the posterior root, and the flatness of the protoconid. These features resemble the Mi's of the two Recent specimens available (FMNH 81522 and Princeton University, PU Ost 568 — figs. 22A1, Bl; 23A1, Bl). If it is an Mi, size is the most distinct difference, the fossil being some 15 per cent larger than the Recent specimens. On the other hand, the fossil tooth fragment compares favorably in size with the equivalent parts of M2 and M3, the fossil being quite close in size to M2. The proto- conid is gently convex labially, the flat hypoconid about two- thirds the height of the protoconid, and there is a weak postero- labial cingulum. PM 30579 is the anterior two-thirds of a right M2 or 3. The protoconid is much more massive and stands almost twice as high as the paraconid. There is a heavy, oblique wear facet along the entire paracristid that extends down to the cingulum. The cin- gulum is truncated by another facet. The anterior root is broad 83 84 FIELDIANA: GEOLOGY, VOLUME 38 and grooved behind and narrows somewhat anteriorly. The para- conid appears to have been chipped in life and the broken edges subsequently worn. The carnassial notch is nearly worn away. The fragment represents a tooth that appears to have been slightly more massive than the modern counterpart. Upper Dentitions: TMM 41106-59 is a complete left P3 (fig. 23A2, B2). It is elongate and nearly elliptical in crown view, with a single, centrally located, prominent cusp. A weak crest extends forward from the apex of the cusp to an anterior cingular cuspule. A slightly stronger posterior crest runs almost straight from the apex to the base, where it flattens out and extends slightly la- bially in a backward sweep across the heel to a pronounced pos- terior cingular cuspule. It is similar to our modern comparative materials as far as wear differences permit comparison. PM 30576 is the posterior half of a right P4. It is quite worn but has the rear half of the central cusp with a posterior crest and a rather bulbous posterior cingular cuspule, both heavily worn. Discussion. — Although only four fragmentary specimens of the thylacine were recovered at Madura Cave, they demonstrate the presence of this taxon in Unit 2. Remains ofThylacinus have been reported from several caves on the Nullarbor Plain (Cook, 1963; Lowry and Lowry, 1967; Partridge, 1967; Lowry and Mer- rilees, 1969; and Archer, 1974). Ride (1964) compared fossil Thylacinus from eastern and western Australia. His data show that the western Australian sample has smaller mean values for most measurements than those for the fossil sample from eastern Australia or the modern sample from Tasmania, although there is extensive overlap be- tween all of the samples. Ride does not give quantitive data for the two teeth (P3 and Ma) that are measurable in the Madura Cave sample. A compari- son of the size of these two specimens with three Recent specimens and one fossil from Wellington Caves (table 29) shows the same patterns outlined by Ride for other measures on the basis of much larger samples. Family Myrmecobiidae Jones, 1923 Myrmecobius (Waterhouse, 1836) Myrmecobius fasciatus (Waterhouse, 1836). Figures 24-25. 85 86 ^ I Q > V 1 * .£ T3 — C CO « C > — U CO •■"" c c o o ^ — <■> to .S j= c <^ 03 o c « .2 *- T3 o - a * ;3 3 > s CO 13 J- J3 *J — % is ►J -l-> 6 c 4) £ CO DC re CO s_ s •— fc CO 3 PQ £ od < >-> — 03 CD 5: •-J a CD o > •■/-. CO •a O co CO 2 hi c CJ 3 — CO •P. £ c t^ IO 00 -M CO d CO £ 1— 1 87 88 FIELDIANA: GEOLOGY, VOLUME 38 Material. — Trench 4 Top of Unit 1 TMM 41106-687, left ramus, edentulous, with alveoli for posterior four teeth (fig. 25C, D). Description . — This is the only specimen of this taxon in the Madura Cave deposits. The specimen consists of the posterior part of the mandible with the alveoli of the posterior four teeth, appar- ently M2-5 judging by the three Recent specimens used for com- parison. In the Recent specimens (fig. 24A, B, C, D; 25A, B) the variation in the lower tooth formula is as follows: M. fasciatus fasciatus, FM 35259 3:I:3:5~ M. fasciatus fasciatus, FM 36049 3:1:3:5 or 6 M. fasciatus (?rufus), FM 19982 3:1:3:4 or 5 The angular process and most of the ascending ramus are bro- ken away in the fossil. The dorsal and ventral margins of the horizontal ramus are parallel. The small alveoli indicate that each tooth had two roots that were aligned parallel to the long axis of the mandible. There is a broad, rounded groove on the lingual side of the mandible that extends forward from just below the inferior dental foramen. The mandibular foramen is round, about 1 mm. in diameter, and opens posteriorly about 4.5 mm. posterior to and below the last tooth. The fossil compares well with the Field Museum Recent specimens, but shows the following minor differences: 1. the fossil ramus is slightly thicker and more massive, 2. the fossil specimen has a much larger nutrient foramen lo- cated behind the last molar on the lingual crest of the coronoid process, i.e., about halfway between the last molar and the man- dibular foramen, and 3. in the fossil this crest and those that bound the masseteric fossa are heavier and more distinct. Discussion. — Modern Myrmecobius has a spotty distribu- tional record: the south part of Western Australia, widely sepa- rated areas of South Australia, and southwestern-most New South Wales. This pattern indicates that the present populations are remnants of a formerly more continuous distribution across the southern part of Australia. This is the first record of it, fossil LUNDELIUS & TURNBULL: MADURA CAVE 89 or Recent, on the Nullarbor Plain, and is, as far as we know, the only fossil record to date. Most accounts of its distribution (Mar- low, 1962; Troughton, 1962; Ride, 1970) indicate that its occur- rence in the more arid parts of Australia is north of the Nullarbor Plain. Order INCERTAE SEDIS Thylacoleonidae Gill, 1872 Thylacoleo Owen, 1859 Thylacoleo sp. Figures 26-27. Material. — Trench 4, Unit 1 TMM 41106-516, right P4 (fig. 26A, B, C). Description . — The right Pi is the only indication of this species from Madura Cave. The crown of the tooth is complete. The roots, which appear to have been stout, are broken off just below the enamel line. This specimen agrees in every respect except size with the description of Woods (1956). The crown has vertically ridged enamel, especially on the lingual surface. Anteriorly one of these ridges is enlarged to form a prominent, rounded ridge. The anterior edge of the tooth is sharp. There is a narrow wear facet along the labial side of the cutting edge. Another broader wear facet is pre- sent on the labial side near the ventral margin of the crown. There is a small interdental facet on the posterior end where it comes in contact with the Ml. The length is 33.8 mm.; anterior width, 10.9 mm.; posterior width, 12.0 mm. Discussion . — Remains of Thylacoleo have been found over a wide area of southern and eastern Australia and from Tasmania (Gill, 1954). No comprehensive study of this animal on a continent-wide basis has been attempted, but size varies geo- graphically (Glauert, 1912) and temporally (Woods, 1956; Barth- olomai, 1962). A comparison of the Madura Cave specimen with samples from Wellington, New South Wales, and Kurramulka, South Australia shows it to be considerably smaller than all specimens in either of the other samples (fig. 27). It is also shorter than any of four specimens of T. crassidentata from the Chinchilla sand of Queensland, reported by Bartholomai (1962), although it resembles this species in the strong curvature of the cutting edge. The location of the maximum width of the Madura specimen above Fig. 26. Thylacoleo species indet., from Madura Cave. TMM 41106-516. Right P4 shown in A, lingual, B, crown, C, labial views. 90 LUNDELIUS & TURNBULL: MADURA CAVE 91 the anterior part of the posterior root appears to be another point of resemblance to T. crassidentata . However, an Australian Museum specimen from Wellington Cave (F 16617) also shows a distinct broadening over the anterior part of the posterior root. In size it resembles the specimen from Balladonia reported by Glauert (1912) to be 72 to 80 per cent of the size of the type. This is close to the difference in size between the Madura Cave specimen and the material from Wellington and Kurramulka caves (fig. 27). Small examples of Thylacoleo are not restricted to Western Australia. A juvenile specimen in the Australian Museum (F 41447) from a cave deposit at Geurie, New South Wales is very close to the size of the Madura Cave specimen (fig. 27). The P4 of the Geurie specimen is not completely erupted thus preventing a determination of the position of its maximum width. As a compre- hensive study based on more complete material is needed to estab- lish the significance of these size differences, the Madura Cave specimen is not assigned to a species. The presence of this taxon in the Madura Cave deposits is ex- pected from a distributional standpoint, but its association with a relatively recent C-14 date (7,470 + 120BP, Lundelius and Turnbull, 1973) deserves comment. This is somewhat younger than most terminal dates on extinct genera in Australia (11,000 to 14,000BP, Martin, 1967; 20,000 to 25,000 BP, Marshall, 1974), al- though a few younger dates have been reported (Martin, 1967). It is possible that the Madura Cave specimen has been eroded from older deposits at the mouth of the cave and redeposited, and one aspect of its preservation (color) suggests this. A C-14 date applic- able to Thylacoleo of the same general age (6570+100 BP) reported from Lake Menindee, New South Wales by Tindale (1957) is now thought to be too young (Tedford, 1967, pp. 17-18). The Madura Cave date then remains the youngest date on Thylacoleo , if its as- sociation within Unit 1 is to be trusted. We are reluctant to accept this even though the specimen undoubtedly came from Unit 1, for its red color suggests redeposition from a deeper unit to which the Unit 1 date would not apply. o 7\» \ zzoei ainn / • 1 • •\ / • ' i » \ ?6fZ=l / < ( 7 xc / • E061AWN o v v "V ■ \ x °\ • • • \ U6U 8l6li 0 \ \ • o\ \ — o \ seum) ali an Mus.) 56(Q'l'd,Mus F 41447 ) - 62(QTd,Mus • \ • \ • o] / « J ngton( Australian Mu ;oorte Caves (So. Austr arling Downs-Woods ■ ive TMM -41106-5KS W. ( Austral ian Mus. • ict or ia [Nat. Hist.) ;hinchilla-Barthalomai ' 0 , / • ^/ C9?Cd f96ZJ 1961=* 0) X 01 c ■ 0 r- • d ra Z X a) c 0) o hj O Q UJ 0) X ai c to u h D Madura C: Seurie, N S Museum Museum identatus C 0J6S1AWN a 09 hll- 6 NMV=Nat x = British O = T. crass 0 hi i m yoiaaiNV 92 ENVIRONMENTAL INTERPRETATION The environmental implications of the larger carnivorous mar- supials are poorly understood. Thylacoleo carnifex is not known as a living animal, and its habitat and other requirements are un- known. Thylacinus cynocephalus and Sarcophilus harrisii are ex- tant only in Tasmania, but they had a wide distribution in Aus- tralia during the Pleistocene and early Recent (Ride, 1964; Calaby and White, 1967). Thus their presence in an assemblage of fossils provides little information on environmental conditions. The smaller species, all of which are extant, are better en- vironmental indicators. Three species, Antechinus flavipes, Phas- cogale calura , and Parantechinus apicalis , are not known from the Recent fauna of the Nullarbor Plain but are found today in more humid regions to the east and west. This indicates the presence in this region of more humid conditions in the past, which is in agree- ment with the interpretations of Main et al. (1958) and Serventy (1951) based on the distribution of living organisms. One species lives today in the area of the junction of South Australia, Queensland, and the Northern Territory where it inhabits desert grassland (Marlow, 1962). The change that lead to its disappearance from the Nullarbor Plain is uncertain. Opposite: Fig. 27. Bivariate graph comparing proportions of P4's of various specimens of Thylacoleo with one another and with the Madura Cave specimen. The Wel- lington Cave sample of T. carnifex included the following Australian Museum specimens: MF 21, MF 82-3, MF 116, MF 388, 575, MF 734, F10835, F 16454-5, F 16458-60, F 16606, F 16614, F 16617-9, F 16623-7, F 18659, F 18884, F 19370, F 31035, and F 37691-2. The Naracoorte Caves and other specimens of T. carnifex in the South Australian Museum included 11 specimens measured in 1964. Here maximum widths were measured, not necessarily AW, for the following: P50, P89-91, P94, P96-7, P105, Pill, and P13719-20. The British Museum specimens are mostly considered to be T. carnifex, but it is possible that one or two T. cras- sidentatus are among them. They are: BMNH M39, M2572, M3570, M3665, BMNH 39995, BMNH 42516, BMNH 46835, and one uncatalogued specimen. 93 94 FIELDIANA: GEOLOGY, VOLUME 38 The decrease in taxonomic diversity of the insectivorous- carnivorous marsupials indicates the disappearance of some habitats. There are 14 taxa of insectivores and carnivores (13 Mar- supicarnivora and one incertae sedis — Thylacoleo) represented in the deposits of Madura Cave. If the very specialized, atypical Myr- mecobius is omitted, the total is 13. This is greater than the number known from the Recent fauna of the Nullarbor Plain (se- ven) or from the Recent fauna of any area of Australia. Another difference is the coexistence in the Pleistocene fauna of species that are now allopatric. Antechinus flavipes, Phascogale calura, and Parantechinus apicalis are not found today in the same area with Dasycercus cristicauda or Dasyuroides byrnei. Similar phenomena have been observed in North America (Hibbard, 1960; Guilday et al., 1964; Dalquest, 1965; Lundelius, 1967; 1974;) and in Europe (Kowalski, 1967). In those areas the disappearance of these associations has been interpreted as indicating a decrease in the equability of the climate. The changes in the carnivorous and insectivorous marsupial fauna from the Nullarbor Plain suggest a change to a lower and less reliable rainfall from the late Pleis- tocene and Holocene to Recent. ACKNOWLEDGEMENTS In addition to those persons and institutions listed in Part I (pp. 32-33) and Part II (pp. 93-94) of this faunal report, we wish to offer our thanks to Dr. Perenyi for the artwork, and to Field Museum and University of Texas Geology Foundation for continuing sup- port. For the loan of specimens, we thank the following institu- tions: American Museum of Natural History, Australian Museum, British Museum (Natural History), Princeton University Museum, South Australian Museum, United States National Museum, and Western Australian Museum. REFERENCES Archer, M. 1974. New information about the Quaternary distribution of the thylacine (Marsupialia; Thylacinidae) in Australia. Jour. Roy. Soc. Western Austral., 57, no. 2, pp. 43-50, figs. 1-2. Bartholomai, A. 1962. A new species of Thylacoleo and notes on some caudal vertebrae of Palor- chestes azael. Mem. Queensland Mus., 14, no. 1, pp. 33-40, figs. 1-4. Bensley, B. A. 1903. On the evolution of the Australian Marsupialia; with remarks on the relationships of marsupials in general. Trans. Linn. Soc, London, Ser. 2, Zool. 9, pp. 83-216, figs. 1-6, pis. 5-7. BOITARD, M. 1841, "1842." Pantheon Populaire. Le Jardin des Plantes. G. Barba, Paris; Roe Lockwood and Son, N.Y. 204 pp. Calaby, J. H. 1966. Mammals of the upper Richmond and Clarence Rivers, New South Wales. Commw. Sci. Industr. Res. Org., Austral. Div. Wildl. Res., Tech. Pap. 10, 55 pp. Calaby, J. H. and C. White 1967. The Tasmanian devil {Sarcohilus harrisii) in Northern Australia in Re- cent times. Austral. Jour. Sci., 29, no. 12, pp. 473-475, figs. 1-2. Cook, D. L. 1963. Thylacinus and Sarcophilus from the Nullarbor Plain. W. Austral. Nat., 9, pp. 47-48. Dalquest, W. W. 1965. New Pleistocene formation and local fauna from Hardeman County, Texas. Jour. Paleo., 39, no. 1, pp. 63-79, figs. 1-2. Dortch, C. E. and D. Merrilees 1971. Human occupation of Devil's Lair, Western Australia during the Pleis- tocene. Archeol. Phys. Anthropol. Oceania, 8, no. 2, pp. 89-114. FlNLAYSON, H. H. 1961. On central Australian mammals. Part IV. The distribution and status of central Australian species. Rec. S. Austral. Mus., 14, no. 1, pp. 141-191, fig. 1. 95 96 FIELDIANA: GEOLOGY, VOLUME 38 Geoffroy Saint-Hilaire, E. 1796. Mag. Encyclop., ser. 2, 3, p. 469. Geoffroy Saint-Hilaire, E. and G. Cuvier 1837. Histoire naturelle des mammiferes, 4, pt. 60, p. 6. Gill, E. D. 1953. Distribution of the Tasmanian devil, the Tasmanian wolf, and the dingo in S. E. Australia in Quaternary time. Vict. Nat., 70, pp. 86-90. 1954. Ecology and distribution of the extinct giant marsupial, Thylacoleo. Vict. Nat., 71, pp. 18-35. Glauert, L. 1912. Fossil marsupial remains from Balladonia in the Eucla division. Rec. W. Austral. Mus., 6, pp. 47-65, pis. 6-8. Gould, J. 1841. Observations on Dasyurus maugei and D. viverrinus of Geoffroy, and de- scription of a new species. Proc. Zool. Soc, London, 1840, p. 151. 1844. Exhibition and character of a number of animals transmitted from Au- stralia by Mr. Gilbert. Proc. Zool. Soc, London, 1844, pp. 103-107. Gray, J. E. 1842. A new species of tapering-tailed phascogale in the collections of the British Museum. Ann. Mag. Nat. Hist., ser. 1, 9, p. 518. Guilday, John E., Paul S. Martin, and Allen D. McCrady 1964. A Late Pleistocene cave deposit in Bedford County, Pennsylvania. Bull. Nat. Speleo Soc, 26, no. 4, pp. 121-194. Guiler, E. R. and R. W. L. Heddle 1974. The eruption and growth of teeth in the Tasmanian Devil, Sarcophilus harrisii (Marsupialia: Dasyuridae). Pap. Proc. Roy. Soc. Tasmania, 108, pp. 137-140, 1 fig., 1 pi. Harris, G. P 1808. Description of two new species of Didelphis from Van Diemen's Land. Trans. Linnean Soc, London, 9, no. 11, pp. 174-178, pi. 19. Hibbard, Claude W. 1960. An interpretation of Pliocene and Pleistocene climates in North America. Mich. Acad. Arts Sci. Letters, 62nd Ann. Rept. (1959-60), pp. 5-30. Jones, F. Wood 1923. The mammals of South Australia. Part 1. The monotremes and the car- nivorous marsupials. Handbooks of the Flora and Fauna of South Australia, Adelaide., pp. 1-131, figs. 1-88. KOWALSKI, K. 1967. The Pleistocene extinction of mammals in Europe, pp. 349-364, 6 text figs. In Martin, P. S. and H. E. Wright, Jr., eds., Pleistocene extinctions, The search for a cause, vol. 6, Proc. VII, Congr. Int. Assn. Quaternary Res., Nat. Acad. Sci., Yale Univ. Press, New Haven and London. LUNDELIUS & TURNBULL: MADURA CAVE 97 Krefft, G. 1867. On the classification of the small Dasyuridae of Australia, with descrip- tions of two new genera and one new species. Proc. Zool. Soc, London, 1866, pp. 431-435, pi. 36. Kurten, B. 1965. The Carnivora of the Palestine caves. Acta Zool. Fenn., no. 107, pp. 1-74, 11 figs., 4 pis. Lowry, D. C. and J. W. J. Lowry 1967. Discovery of a Thylacine (Tasmanian Tiger) carcase in a cave near Eucla, Western Australia. Helictite, -5, pp. 25-29. Lowry, J. W. J. and D. Merrilees 1969. Age of the described carcase of a Thylacine (Marsupialia, Dasyuroidea) from Thylacine Hole, Nullarbor Region, Western Australia. Helictite, 7, pp. 15-16. Lundelius, E. L., Jr. 1963. Vertebrate remains from the Nullarbor Caves, Western Australia. Jour. Roy. Soc. W. Austral., 46, part 3, pp. 75-80. 1967. Late-Pleistocene and Holocene faunal history of Central Texas, pp. 287- 319, figs. 1-11. In Martin, P. S. and H. E. Wright, Jr., eds., Pleistocene ex- tinctions, The search for a cause, vol. 6, Proc. VII, Congr. Int. Assn. Qua- ternary Res., Nat. Acad. Sci., Yale Univ. Press, New Haven and London. 1974. The last fifteen thousand years of faunal change in North America, pp. 141-160. In Black, C. C, ed., History and prehistory of the Lubbock Lake site, Mus. Jour. (Texas Tech. Univ. Lubbock), 15. Lundelius, E. L., Jr. and W. D. Turnbull 1973. The mammalian fauna of Madura Cave, Western Australia. Part I. Field- iana: Geol. 31, no. 1, pp. 1-35, figs. 1-13, tables 1-2. 1975. The mammalian fauna of Madura Cave, Western Australia. Part II. Fieldiana: Geol. 31, no. 2, pp. 37-117, figs. 1-21, tables 1-15. MacLeay, W. S. 1841. (1842). Notice of a new genus of Mammalia discovered by J. Stuart, esq., in New South Wales. Ann. Mag. Nat. Hist., 8, ser. 1, no. 51, art. XXXIX, pp. 240-243, pi. VII. Main, A. R., A. K. Lee, and M. J. Littlejohn 1958. Evolution in three genera of Australian frogs. Evolution, 12, pp. 299-304. Marlow, B. J. 1958. A survey of the marsupials of New South Wales. Commw. Sci. Industr. Res. Org., Austral., Wildl. Res. 3, no. 2, pp. 71-114, figs. 1-34. 1962. Marsupials of Australia. Jacaranda Pocket Guides, Jacaranda Press, Brisbane, pp. 1-141. Marshall, L. G. 1974. Late Pleistocene mammals from the "Keilor cranium site," Southern Vic- toria, Australia. Mem. Nat. Mus. Victoria, 35, pp. 63-86, 11 figs. 98 FIELDIANA: GEOLOGY, VOLUME 38 Martin, P. S. 1967. Pleistocene overkill, pp. 75-120, figs. 1-4. In Martin, P. S. and H. E. Wright, Jr., eds., Pleistocene extinctions, The search for a cause, vol. 6, Proc. VII Congr. Int. Assn. Quaternary Res., Nat. Acad. Sci., Yale Univ. Press, New Haven and London. Merrilees, D. 1968. Man the destroyer: late Quaternary changes in the Australian marsupial fauna. Jour. Roy. Soc. W. Austral., 51, no. 1, pp. 1-24, figs. 1-3. Meyer [initials not known] 1793. Systematischsummarisch Uebersicht der neusten zoologischen Entdecku- gen in Neuholland, pp. 28, 281 (fide Tate, 1947, p. 135). Mulvaney, D. J., G. H. Lawton, C. R. Twidale; with special reports by N. W. G. Macintosh, J. A. Mahoney, and N. A. Wakefield 1964. Archaeological excavation of Rock Shelter No. 6, Fromm's Landing, South Australia, Proc. Roy. Soc. Vict., 77, pp. 479-516. Owen, R. 1859. XVI. On the fossil mammals of Australia. Part I. Description of a muti- lated skull of a large marsupial carnivore (Thylacoleo carnifex, Owen), from a calcareous stratum, eighty miles S. W. of Melbourne, Victoria. Phil. Trans. Roy. Soc, 149, no. 16, part 1, pp. 309-322, pis. 11-15. Partridge, J. 1967. A 3,300 year old Thylacine (Marsupialia: Thylacinidae) from the Nullar- bor Plain, Western Australia. Jour. Roy. Soc. W. Austral., 50, pp. 57-59. Peters [initials not known] 1875. Sitzber. Gesellsch. Naturf. Fr., Berlin, p. 73, (New Name for C haetocercus Krefft.) (fide Tate 1947, p. 141). Ride, W. D. L. 1964. A review of the Australian fossil marsupials. Jour. Roy. Soc. W. Austral., 47, pt. 4, no. 16, pp. 97-131, figs. 1-13. 1970. A guide to the native mammals of Australia. Oxford Univ. Press, Mel- bourne, London, Wellington, N.Y. i-xiv, pp. 1-149, figs. 1-62 by Ella Fry. Serventy, D. L. 1951. The evolution of the chestnut-shouldered wrens Malurus). Emu, 51, pp. 113-120. Siegal, S. 1956. Nonparametric statistics for the behavioral sciences. McGraw Hill, New York, pp. i-xvii, 1-312. Smith, M. J. 1972. Small fossil vertebrates from Victoria Cave, Naracoorte, South Australia. II. Peramelidae, Thylacinidae and Dasyuridae (Marsupialia). Trans. Roy. Soc. S. Austral., 96, Part 3, 125-137, figs. 1-8. LUNDELIUS & TURNBULL: MADURA CAVE 99 Spencer, B. 1896. [Preliminary notice of Dasyuroides byrnei]. Proc. Roy. Soc, Vic, N.S., 8, p. 6. 1896. Report on the work of The Horn scientific expedition to central Australia, 2, Zoology, pp. 36-40, pis. 3 and 4 (figs. 1-4). Tate, G. H. H. 1947. On the anatomy and classification of the Dasyuridae (Marsupialia). Re- sults of the Archbold Expeditions. No. 56. Bull. Amer. Mus. Nat. Hist., 88, art. 3, pp. 97-156, text-fig. 1. Tedford, R. H. 1967. The fossil Macropodidae from Lake Menindee, New South Wales. Univ. Calif. Pubs., Geol. Sci., 64, i-v, pp. 1-165, figs. 1-32, plates 1-4. Temminck, C. J. 1824, 1827. Monographies de Mammalogie, 1, pp. xxiii, 23, 60, 267 (Thylacinus) also xxiii, 23, 56 (Phascogale). (fide Tate, 1947 pp. 118, 135). Thomas, O. 1888. Catalogue of the Marsupialia and Monotremata in the collection of the British Museum (Nat. Hist.), London. Taylor & Francis, London, I-XIII, 401 pp. TlNDALE, N. 1957. Culture succession in southeastern Australia from late Pleistocene to the present. Rec. S. Austral. Mus., 13, pp. 1-49. Troughton, E. 1962. Furred animals of Australia. 7th ed. Angus and Robertson, Sydney, Lon- don, Melbourne, Wellington, xxxii pp., 376 pp., pis. I-XXV. Turnbull, W. D. 1970. The Trinity therians: their bearing on evolution in marsupials and other therians, chap. 9, pp. 151-179, figs. 1-6. In Dahlberg, A. A., ed., Dental mor- phology and evolution, Univ. of Chicago Press, Chicago, London. Wakefield, N. A. and R. M. Warneke 1967. Some revision in Antechinus (Marsupialia) -2. Vict. Nat., 84, pp. 69-99, figs. 1-9, pi. 3. Waterhouse, G. R. 1836. Description of a new genus (Myrmecobius) of mammiferous animals from New Holland, probably belonging to the marsupial type. Proc. Zool. Soc, London, 1836, pp. 69, 70, 131 (preliminary description). Trans. Zool. Soc. London, 1841, pp. 149-154, pis. 27, 28 (extended description). 1838. Characters of some new species of the genera Mus and Phascogale . Proc Zool. Soc, London, 1837, pp. 75-77. Woods, J. T. 1956. The skull of Thylacoleo carnifex. Mem. Queensland Mus., 13, pt. 2, pp. 125-140, figs. 1-6. o o o o o o o ^ ^ o a c o > ee 1-4 0 0 »hJSJ3jsjs3J33-j5? ,3? 100 TABLE 2. Numerical data on upper dentitions of Recent specimens of Antechinus flavipes compared with the small sample of A. stuartii adusta, as in Table 1. Antechinus stuartii adusta A.f. flavipes A.f. rufogaster A.f. leucogaster FM 60955 - FM 60960 FM 35337 FM 104787 FM 18889 Sample Observed Variate Size Range Mean M1"* 6 6.39-6.72 6.59 6.48 7.18 6.30 M1"3 5 5.73-5.99 5.83 5.70 6.34 5.63 L.up.Pm 6 2.84-3.17 3.30 3.10 3.08 3.31 P2L 6 .81- .91 .87 1.02 .85 .87 W 6 .61- .66 .64 .61 .66 .61 P3L 6 1.01-1.12 1.07 1.17 1.10 1.05 W 6 .66- .76 .71 .73 .73 .68 PAL 6 1.12-1.22 1.16 1.24 1.22 1.18 W 6 .71- .86 .77 .61 .73 .76 ^L 6 2.05-2.23 2.17 2.20 2.39 2.09 W 6 1.52-1.67 1.59 1.51 1.61 1.48 M*L 5 2.00-2.10 2.05 2.05 2.32 2.05 W 5 1.93-2.13 2.02 1.81 2.20 1.75 M3L 5 1.77-1.88 1.80 1.78 1.95 1.85 W 5 2.13-2.33 2.24 2.07 2.32 2.03 M«L 6 .63 .74 .67 .85 .73 .76 W 6 2.03-2.13 2.10 1.98 1.95 1.82 TABLE 3. Numerical data on upper and lower dentitions of Antechinus flavipes Sample Variate Size M1"* 15 M1"3 15 L.up.Pm 15 P3L 14 ^1-4 15 L . 1 ow . Pm 14 P, L 14 leucogaster from Western Australia. Observed Range Mean + Standard Error Standard Deviation Coefficient of Varia- tion (Z) 6.17-6.71 6.37+.030 .149 2.34 5.32-5.88 5. 58+. 027 .133 2.38 2.80-3.61 3. 16+. 044 .219 6.94 1.01-1.33 1.11+.021 .079 7.14 6.64-7.20 6. 89+. 034 .172 2.49 2.80-3.57 3.22+.057 .213 6.62 .75-1.03 .92+. 026 .097 10.59 Specimens included are: WAM - M194, M279, M894 , M1159, M1334, M1401, M1411, M1695, M1759, M1822, M2037, 7050, 9035, BMNH 41.12.44; and FMNH 18889. 101 TABLE 4. Numerical data on upper and lower dentitions of Antechinus f lavipes from Madura Cave. Mear + Coefficient Sample Observed Stanc ard Standard of Varia- Variation Size Range Error Deviation tion (%) P4 L 2 .96-1.16 1.06 -- - W 3 .56- .71 .63 - - M L 4 1.52-1.62 1.57 - - AW-E 4 .76- .81 .80 - - PW 4 .86- .91 .88 - - M2 L 8 1.67-1.88 1.78 + .03 .088 4.9 AW-E 8 1.01-1.17 1.096+.02 .052 4.7 PW 8 .99-1.06 1.026+.009 .026 2.5 M3 L 8 1.83-1.93 1.89 + .011 .032 1.7 AW-E 8 1.14-1.22 1.18 + .008 .023 1.9 PW 8 .91-1.01 .98 + .012 .034 3.4 M4L 9 1.62-1.90 1.74 + .032 .095 5.5 AW-E 9 1.01-1.17 1.04 + .018 .053 5.1 PW 8 .30- .71 .47 + .056 .158 33.3 M3L 2 1.74-1.88 1.81 - -- W 2 2.07-2.08 2.08 — — 102 TABLE 5. Numerical data on lower dentitions of Phascogale calura from Unit 1, Madura Cave. Variate Sample Size ^1-4 3 L.low.Pm 3 P2L 3 W 3 P3L 2 w 2 PAL 3 W 3 M1L 3 AW-E 3 PW 3 M2 L 2 AW-E 2 PW 2 M3L 5 AW-E 5 PW 5 M4L 5 AW-E 5 PW 5 Observed Range 7.52-7.66 3.36-3.57 1.24-1.32 .85- .93 1.27-1.29 .88-1.07 .63- .66 1.78-1.88 .90-1.07 .95-1.00 2.15-2.20 1.15-1.20 2.07-2.32 1.27-1.37 1.05-1.20 1.88-1.93 1.07-1.17 .49- .57 7.58 3.47 1.28 .89 1.28 .88 .98 .64 1.81 .98 .98 2.18 1.24 1.18 2.18 1.33 1.15 1.90 1.11 .52 103 TABLE 6. Numerical data on lower dentitions of Phascosale calura from Units 2 and 3, Madura Cave. Sample Size Observed Range Mean + Standard Error Standard Deviation Coefficient of Varia- tion (%) *l-4 1 — 7.70 P2L 1 ~ 1.22 W 1 - .91 P3L 1 - 1.37 W 1 ~ .95 P4L 2 .93-1.05 .99 W 2 .66- .80 .73 M1L 4 1.73-1.92 1.83 AW-E 4 .91-1.14 1.01 PW 4 .96-1.06 1.00 M2L 7 2.00-2.29 2. 17+. 039 AW-E 8 1.22-1.37 1. 31+. 016 PW 8 1.15-1.37 1. 24+. 027 M3L 9 2.08-2.36 2.20+.033 AW-E 9 1.27-1.44 1. 34+. 022 PW 9 1.06-1.24 1. 15+. 025 M4L 9 1.77-2.13 1. 96+. 043 AW 9 1.06-1.22 1. 14+. 020 PW 9 .37- .66 .5O+.029 .102 4.72 .044 3.38 .075 6.06 .099 4.48 .065 4.83 .074 6.44 .129 6.59 .059 5.17 .086 16.96 104 TABLE 7. Numerical data on lower dentitions of Phascogale calura from Units 4 and 5, Madura Cave. Varlate Sample Size "*l-4 1 P3L 1 W 1 P4 L 3 W 3 ^L 4 AW-E 4 PW 4 «2L 3 AW-E 3 PW 3 M3L 7 AW-E 7 PW 7 M4 L 6 AW-E 6 PW 6 Observed Range .93-1.18 .61- .68 1.59-1.94 .91-1.07 .91-1.14 2.16-2.28 1.22-1.33 1.14-1.37 1.98-2.34 1.26-1.37 1.06-1.28 1.86-2.15 1.14-1.22 .44- .61 8.08 1.33 .84 1.05 .64 1.81 1.01 1.07 2.24 1.28 1.27 2.22 1.33 1.13 1.99 1.16 .51 Numerical data on dentitions of Recent samples of Phascogale calura from Western Australia. ' Variate L. upper molars Length M1"3 P2"4L P4L M1-4L P2-4L P, L Sample Size 16 17 16 16 17 16 15 Observed Range Mean + Standard Error Standard Deviation 6.74-7.30 7. 06+. 036 .142 6.01-6.79 6. 27+. 044 .183 2.94-3.70 3.28+.056 .224 .95-1.40 1.12+0.28 .111 7.09-8.50 7. 69+. 078 .321 3.03-3.87 3. 46+. 058 .230 .87-1.08 .98+. 019 .074 Coefficient of Variation (Z) 2.01 2.91 6.83 9.91 4.17 6.65 7.60 'Based on the following specimens: W.A.M. M332, M1246, M2864, M2292, M1277, M866, M2581, M1226, M1262, M1106, M1089, M2532; FMNH 36052; BMNH 83.10.19.11, 35 (or 36). 12. 5.1, 7.7.18.14. 105 TABLE 9. Numerical data on upper dentitions of Phascogale calura from Madura Cave. W M2 L W M3 L W M4L W WAM 74.9.16 TMM 41106-739 TMM 41106-738 PM 25580 PM 36052 — - 2.32 - 2.27 ~ - 1.66 - 1.54 — - - 2.43 2.20 - - " 2.17 1.98 2.17 2.07 - ~ 1.98 2.15 2.27 " " 2.20 — .81 " - .90 2.00 2.15 TABLE 10. Numerical data on lower dentitions of Phascogale tapoatafa from Madura Cave. Observed Range Mean 11.46 4.78 1.93 1.32 2.18 1.34 1.22-1.37 1.30 .91-1.06 .99 2.69-2.94 2.83 1.67-1.77 1.70 1.47-1.57 1.53 3.24-3.29 3.27 1.77-1.93 1.86 1.72-1.80 1.76 3.14-3.19 3.17 1.93-2.03 1.98 1.77-1.93 1.85 2.69-2.84 2.77 1.67-1.77 1.72 Sample Variate Size ™l-4 1 LP2-4 1 P2L 1 W 1 P3L 1 W 1 P4L 2 W 2 Ml L 4 AW-E 3 PW 3 M2 L 3 AW-E 3 PW 3 M3L 2 AW-E 2 PW 2 M. L 4 2 AW-E 2 PW 2 .76- .91 .84 106 TABLE 11. Numerical data on lower dentitions of Phascogale tapoatafa from Wedge's Cave. Sample Size 2-4 P., L P4L W M1L AW- E PW ^L AW- E PW M3 L AW- E PW M. L 4 AW- E PW Observed Range 11.17-11.87 1.27-1.37 1.01-1.06 2.69-3.04 1.57-1.67 1.52-1.62 3.19-3.42 1.85-1.98 1.72-1.88 3.19-3.29 1.90-2.03 1.57-1.67 2.58-2.74 1.67-1.75 .76- .81 Mean 11.48 5.52 2.28 1.72 2.28 1.62 1.34 1.03 2.86 1.64 1.57 3.28 1.90 1.83 3.22 1.95 1.64 2.65 1.71 .78 TABLE 12. Numerical data on upper and lower dentitions of a Recent sample of Phascogale tapoatafa. Sample Observed Mean + Standard Coefficient of Variate Size Range Standard Error Deviation Variation (I) LM1-4 15 9.3-10.9 10.24+. 12 .458 4.47 LM1"3 15 8.0-9.6 9. 02+. 11 .438 4.85 LP4 15 1.2-2.0 1.77+.05 .202 11.45 LMl-4 15 10.4-12.0 11. 27+. 12 .474 4.21 L P, 15 2.0-2.4 2. 25+. 04 .136 6.02 'Based on following specimens: BMNH 7.7.18.13; 44.2.15.19; 46.4.4.89; 42.6.29.5; 41.1240; 53.10.22.18; 42.6.29.4; 54.10.24.20; 41.1239; 44.7.9.8; 46.4.4.63; 43.8.12.27; 53.10.22.17; 4.1.3.101; TMM M-838. 107 TABLE 13. Numerical data on lower dentitions of Parantechinus apicalis from Madura Cave and Hastings Cave. Length M M1L AW AW-E PW M. L 4 AW AW-E PW Madura Cave PM 25330 2.13 1.09 1.42 1.22 2.54 1.37 1.47 .56 Hastings Cave1 Sample Size Observed Range Mean 9 8.48-9.45 8.81 9 2.03-2.38 2.16 9 .92-1.04 .97 9 1.12-1.27 1.18 9 1.01-1.12 1.06 12 2.13-2.33 2.22 12 1.20-1.35 1.29 12 1.19-1.37 1.29 12 .48- .61 .53 'Based on the following specimens: FM PM 7268-7270; 13329-13337. TABLE 14. Numerical data on dentitions of a Recent sample of Parantechinus apicalis from Western Australia. Variate Sample Size L.up.Pa 4 LP4 4 L. up. molar 4 LM1"3 4 L.lw.Pm 4 LP4 4 L. lw. molar "l-4 4 Observed Range 3.40-3, .65 .50- - ,68 7.57-8, .43 6.66-7 .47 3.47-4 ,13 .32- ,55 8.60-9 ,53 Mean 3.56 .61 7.99 7.04 3.65 .46 8.98 Baaed on B.M. 44.9.30.6; 48.1.27.27; 46.4.4.90; 44.6.15.8. 108 TABLE IS. Numerical data on lower dentitions of Dagycercua crlsticauda from Unit Sample L of Madura Cave. Observed Variate Size Kaan Mean Canine- Ant- -Post Diameter 5 1.12-1.57 1.30 Length premolars 4 3.17-3.48 3.30 P2L 5 1.52-1.67 1.62 P2W 5 0.91-1.06 1.00 P3L 5 1.67-2.08 1.84 p3w 5 1.01-1.17 1.09 Length molars 2 9.6-10.03 9.82 HjL 5 2.23-2.33 2.28 Mx AW-E 5 1.27-1. 47 1.38 Mx PW 5 1.09-1.32 1.20 M2 L 6 2.35-2.77 2.64 Mj AW-E 6 1.32-1.72 1.55 Mj PW 6 1.27-1.57 1.44 fL L 3 2.69-2.89 2.81 M3 AW-E 3 1.80-1.88 1.83 Rj PW 3 1.47-1.57 1.53 M,L 3 2.59-2.69 2.61 M, AW-E 3 1.52-1.72 1.64 M, PW 3 0.56-0.66 0.61 109 TABLE 16. Numerical data on lower dentitions of Dasycercus cristicauda from Units 2-7 of Madura Cave. Sample Observed Mean + Standard Coefficient of Variate Size Range Standard Error Deviation Variation (%) L.Pm 1 - 3.53 - — P2L 1 ~ 1.77 - - P2W 1 - 0.91 ~ " P3L 2 1.42-1.77 1.60 - ~ P3W 2 .86-1.06 .96 - - M1L 4 1.98-2.18 2.07 " -- >L AW-E 4 1.32-1.47 1.37 - - M PW 4 1.06-1.37 1.19 " - M2 L 11 2.48-2.59 2. 52+. 01 .048 1.34 M AW-E 11 1.42-1.72 1.53+.02 .077 5.02 M2 PU 11 1.32-1.57 1.4O+.02 .076 5.44 M3L 11 2.53-2.74 2. 63+. 02 .076 2.95 M AW-E 11 1.62-1.82 1.69+.02 .065 3.84 M3 PW 11 1.32-1.52 1.39+.02 .062 4.44 M, L 4 8 2.18-2.64 2. 38+. 05 .145 6.10 M, AW-E 4 8 1.37-1.62 1.48+.03 .073 4.93 M. PW 8 0.36-0.56 0.45+.02 .063 13.95 110 I t» ,-( 01 C -I ri C J <^ eg c i H N 3 -J r 0 00 u ^ 2 n c 1 00 00 V H ■^ aj tr. in I I I r- ,-1 l r* oo C4 ^ o* CM CM CM O ti rv o -T m Cft 00 00 e i) ~5 Is 1c 1b ^ 111 TABLE 18. Results of Mann-Whitney text comparisons of measuremnts of lower and upper dentitions of Dasycercus crlstlcauda samples from Unit 1 and Units 2-7. (Two tailed tests) Sample Mean Size U P L. lower premolars Unit 1 3.30 4 0.0 .4 Units 2-7 3.53 1 P L Unit 1 Units 2-7 1.62 1.77 5 1 0.0 .334 P. W Unit 1 Units 2-7 1 0.91 5 1 0.5 .334-. 666 P L Unit 1 Units 2-7 1.84 5 7.5 >.5 1.60 2 P, W Unit 1 Units 2-7 1.09 5 8.0 >.5 0.96 2 VL L Unit 1 Units 2-7 2.28 2.07 5 4 0.0 .016 M AW-E Unit 1 Units 2-7 1.38 5 1.37 4 11.5 >.5 VL PW Unit 1 Units 2-7 1.20 1.19 5 4 12.0 >.5 M. L Unit 1 Units 2-7 2.64 2.52 6 11 11.0 <.05 M AW-E Unit 1 Units 2-7 1.55 1.53 6 11 28.0 >.l M PW Unit 1 Units 2-7 1.44 1.40 6 11 26.0 >.l M L Unit 1 Units 2-7 2.81 3 3.0 .05 2.63 11 M AW-E Unit 1 Units 2-7 1.83 1.69 3 11 1.5 .02-. 05 M PW Unit 1 Units 2-7 1.53 1.39 3 11 1.5 .02-. 05 M, L Unit 1 Units 2-7 2.61 2.38 3 8 2.0 .048 M, AW-E Unit 1 Units 2-7 1.64 1.48 3 8 1.5 .042-. 048 112 TABLE 19. Numerical data on lower dentitions of Dasyuroldes byrnei from Unit 1 of Madura Cave. Variate PM 30504 PM 30507 M2L 2.99 - M, AW-E 1.88 - M2 PW 1.77 -- M3L 2.94 " M AW-E 1.93 1.98 M, PW 1.62 — TABLE 20. Numerical data on lower dentitions of Dasyuroides byrnei from Units 2-3 of Madura Cave. 1-4 P., L 11.01 8.32 3.82 P3 " Ml L Mx A\ M, PW 1.37-1.62 1. 49+. 015 .069 4.62 Sample Observed Size Range 15 9.99-11.51 5 1.52-1.98 5 0.91-1.12 13 1.88-2.13 14 1.04-1.27 22 2.48-2.91 21 1.67-1.98 22 1.37-1.62 39 2.64-3.19 39 1.67-1.99 39 1.49-1.93 46 2.58-3.19 46 1.67-2.03 46 1.47-1.82 39 2.38-2.89 38 1.57-1.77 39 .41- .81 1 M2 L M2 AVi M2 PV M3 L M AV M3 PV M. L 39 2.38-2.89 2. 67+. 020 .122 4.56 4 — M. AW-E 38 1.57-1.77 1. 65+. 011 .065 3.96 4 — M, PW 39 .41- .81 .59+. 014 .089 15.02 Mean + Standard Error Standard Deviation 10.89+. 118 .455 1. 78+. 087 .196 1. 00+. 037 .083 1. 98+. 021 .076 1. 15+. 020 .073 2. 77+. 025 .119 1. 88+. 027 .123 1. 49+. 015 .069 2. 99+. 019 .116 1. 82+. 012 .073 1. 72+. 013 .082 2. 97+. 017 .114 1. 88+. 011 .072 1. 64+. 018 .125 2. 67+. 020 .122 1. 65+. 011 .065 .59+. 014 .089 Coefficient of Variation (%) 4.17 113 TABLE 21. Numerical data on lower dentitions of Dasyuroides byrnei from Units 4-5 of Madura Cave. Sample Va riate M3 L M3 AW- E M3 PW M4 L M4 AW- E M, PW 2.91 1.81 1.37-1.62 1.55 4 2.43-2.81 2.67 3 1.57-1.77 1.65 5 0.48-0.91 0.64 TABLE 22. Numerical data on lower dentitions of Dasyuroides byrnei Observed Range 2 71- -3 04 1 67 -1 88 1 37- -1 62 2 43- -2 81 1 57- -1 77 0 48- -0 91 Mean from Units 6-7 of Madura Cave. : Mean + Sample Observed Standard Standard Coefficient of Variate Size Range Error Deviation Variation (%) P3 L (R) 3 1.88-2.03 1.98 " " P3 W (R) 3 1.12-1.16 1.13 - " Mx L (RL) 4 2.74-3.04 2.89 " " M AW-E (RL) 4 1.77-1.98 1.86 " " M PW (RL) 4 1.47-1.79 1.58 - " M2 L (R) 7 2.94-3.09 3. 03+. 025 .067 2.21 M AW-E (R) 8 1.77-1.93 1. 82+. 021 .058 3.19 M2 PW (R) 8 1.67-1.82 1. 75+. 016 .046 2.61 M3 L (R) 10 2.84-3.19 2. 99+. 030 .094 3.14 M AW-E (R) 11 1.79-1.98 1. 90+. 018 .058 3.06 M PW (R) 10 1.52-1.77 1. 65+. 027 .084 5.10 R based on rights only; RL based on rights and lefts combined. 114 TABLE -3. Numerical data on upper dentitions of Dasyuroides byrnei from Units 2 -3 of Madura Cave. Mean + Sample Observed Standard Standard Coefficient of Variate Size Range Error Deviation Variation (Z) P2L 2 " 1.47 -- " p2w 2 0.91-0.96 0.94 - -- P3L 5 1.52-1.69 1.60 - -- P3W 5 0.96-1.08 1.02 -- -- P4L 15 1.01-1.47 1. 23+. 038 .!4h 11.87 P4W 13 0.76 0. 87+. 015 .054 6.21 M1-4 7 9.62-9.92 9.74 — — M1"3 11 8.24-8.92 8. 63+. 060 .120 2.31 MXL 32 2.89-3.40 3. 11+. 019 .108 3.47 M1 W 31 2.03-2.28 2. 14+. 012 .068 3.15 M2L 34 2.89-3.24 3. 06+. 01 5 .085 2.78 M W 32 2.64-2.94 2. 76+. 015 .088 3.18 M3L 26 2.53-2.89 2. 74+. 019 .094 3.44 3 M W 28 2.64-3.09 2.87+.021 .110 3.84 M4L 11 0.96-1.16 1. 04+. 019 .062 5.96 M4W 11 2.25-2.56 2. 44+. 034 .113 4.63 TABLE 24. Numerical Dasyuroid data on ;s bryne dentitions of a Recent sam Variate Sample Size Observed Range Mean M1-4L 6 10.96-11.39 11.13 P2-3L 6 3.76- 4.44 4.14 M^L 6 9.76-10.13 9.96 MX-3L 6 8.60- 9.07 8.79 P2"4L 6 3.86- 4.57 4.18 P4L 6 0.46- 1.34 0.98 115 TABLE 25. Numerical data on dentitions of Dasyurus geof froyi from Madura Cave. Sample Observed Variate Size Range M2 L 9 4.79-5.40 M2 AW 10 2.43-2.96 M2 PW 10 2.43-2.96 M3 L 3 5.24-5.70 M3 AW 4 3.12-3.34 M PW 3 4 2.58-2.93 M1 L 12 4.94-5.85 M1* 13 3.04-3.66 M2 L 5 5.32-5.93 M2W 4 3.99-4.33 M3 L 5 5.09-5.78 M3 W 4 4.64-5.12 Mean + Standard Error Standard Deviation Coefficient of Variation (%) 5.12 - - 2. 80+. 059 .188 6.71 2. 68+. 05 3 .167 6.24 5.42 - ~ 3.19 -- - 2.77 - - 5. 54+. 074 .258 4.65 3.41+.046 .167 4.89 5.53 - - 4.15 - " 5.32 - - 4.89 — — 116 05 u ~ Fi 9 5 - --» OJ <£> O £ t-| r* N0 t*> 1-* r>| p«* \0 O so ih r* ■ 1 a. n E - T 00 00 r- r^ o I CM r*> ^H C nj o o H vO C -J -H M .H NO 1 I 1 « 1 1 1 ri to u X — ' a. c a o «J c 00 on i e •H •3 o •& -* 00 CO 1 1 1 1 w| 01 3 X E nO 2 CC 01 > a u 2 a cn in m o l^ ^ on o ON cc 1 01 1 00 X CO •o E 01 3 R m m 00 m NO ON £ IX in -» ,_; NO J, ON 01 1 r~ > on m a •& (M o a a u in T •* o a. to CJ NO cm" d 3 OS e I i 1 1 e o I i o o -o> ~T z 3 NO 3 ON z ^ H m CM *» ^r l"> O 1*1 p^ o O o m n"l •-I ON GO CN| NO 00 - O 1 X 10 *» o ^C ,_! ON 01 O m > a m on ■ u O • o •H a 0£ ki |4 no *n T 3 NO < O 1 1 1 1 £ "O o 1 1 1 1 « m 3 ao «i| x O X « -* E z _ ^ CO o IX ON *» On « »-• > r*N 0. a. r"l <"1 X "V: °x 119 TABLE 29. Numerical data on dentitions of Thylacinus cynocephalus Madura Cave Length P Width P Width M PM 30579 " " 6.68 TMM 41106-59 8.10 3.70 - Recent Pu ost 568 7.60 3.77 6.40 USNM 38801 8.26 3.95 6.70 FM 81522 7.60 3.54 6.50 Wellington Caves PM 1635 — — 7.80 120 Publication 1290