FIELDIANA Geology Published by Field Museum of Natural History . • 1 1 THE MAMMALIAN FAUNA OF MADURA CAVE, WESTERN AUSTRALIA PART V: DIPROTODONTA (PART) i RNESTL. LUNDEL1US, JR. WILLIAM D. TURNBULL June 29, 1982 Publication 1332 l\^l n 10l a« f\EU> wustw *■* THE MAMMALIAN FAUNA OF MADURA CAVE, WESTERN AUSTRALIA PART V: DIPROTODONTA (PART) FIELDIANA Geology Published by Field Museum of Natural History New Series, No. 11 THE MAMMALIAN FAUNA OF MADURA CAVE, WESTERN AUSTRALIA PART V: DIPROTODONTA (PART) ERNEST L. LUNDELIUS, JR. Professor of Geological Sciences University of Texas at Austin Research Associate Field Museum of Natural History WILLIAM D. TURNBULL Curator. Fossil Mammals Field Museum of Natural History Committee on Evolutionary Biology University of Chicago Research Associate Texas Memorial Museum Accepted for publication November 21, 1979 June 29, 1982 Publication 1332 Library of Congress Catalog Card No.: 82-071140 ISSN: 0096-2651 PRINTED IN THE UNITED STATES OF AMERICA CONTENTS List of Illustrations vi List of Plates (SEM Photos) vi List of Tables vii Abstract 1 Introduction 1 Systematics 2 Lasiorhinus (Gray, 1863) 2 Lasiorhinus c.f. latifrons (Owen, 1845) 2 Material 2 Description 2 Discussion 12 Vombatidae 12 Genus and species indeterminate 12 Material 12 Phascolarctos Blainville, 1816 13 Phascolarctos cinereus (Goldfuss, 1817) 13 Material 13 Description 13 Discussion 15 Trichosurus Lesson, 1828 18 Trtchosurus vulpecula (Kerr, 1792) 18 Material 18 Description 18 Pseudocheirus Ogilby, 1837 20 Pseudocheirus peregrinus (Boddaert, 1785) 20 Material 20 Description 20 Discussion 20 Cercartetus Gloger, 1841 22 Cercartelus concinnus (Gould, 1845) 22 Material 22 Description 23 Discussion 29 Summary 30 Acknowledgments 30 Literature Cited 31 LIST OF ILLUSTRATIONS Figures Frontispiece: Maps of the Nullarbor Plain area ii 1. Lasiorhinus c.f. latifrons, right Ii 3 2. Lasiorhinus c.f. latifrons, maxillary fragment 5 3. Lasiorhinus c.f. latifrons, anterior moiety of an upper molar 6 4. Lasiorhinus c.f. latifrons, upper molar 7 5. Lasiorhinus c.f. latifrons, molar tooth 8 6. Vombatidae gen. and sp. indet., half of one moiety of a molar tooth 9 7. Phascolarctos cinereus, right M2 and left M4 14 8. Trichosurus vulpecula, left Mi 19 9. Pseudocheirus peregrinus, right M1 21 10. Cercartetus concinnus. Schematic outlines of 14 ramus fragments 24,25 Plates I. II. III. IV. Lasiorhinus c.f. latifrons 10 Lasiorhinus gen. and sp. indet 11 Montages of Phascolarctos cinereus and oiTrichosurus vulpecula and a stereopair of Pseudocheirus peregrinus 16,17 Cercartetus concinnus 26,27 LIST OF TABLES 1. Measurements and ratios of upper dentitions of Phascolarctos cinereus 13 2. Measurements of lower dentitions of Phascolarctos cinereus 15 3. Measurements of Trichosurus, Mi 20 4. Measurements on fossil and Recent specimens of Pseudocheirus peregrinus 21 vn ABSTRACT Five species of the Order Diprotodonta are reported from the deposits of Madura Cave. The vombatid Lasiorhinus c.f. latifrons is represented by teeth from mostly juvenile individuals from Levels 2 through 4-5 in Trenches 1, 3, and 4. Some indeterminate juvenile specimens may represent another vombatid taxon. They are present throughout the excavated sequence (7,500-38,000 years b.p.). Sev- eral specimens are noteworthy for the preservation of some of the original tooth crown features. Three teeth of Phascolarctos cinereus from a level in Trench 4 dated 20,000 y b.p. imply more mesic conditions than exist at present. One tooth each of Trichosurus vulpecula and Pseudocheirus peregrinus was re- covered from units dated 20,000 y b.p. and 15,600 y b.p., respectively. Today these taxa live in more humid environments than are found on the Nullarbor. Sixteen mandibles and eight isolated teeth of Cercartctus concinnus came from the Pleistocene units (15,000-38,000 y b. p.), but none were found in Unit 1, the Holocene unit. At present none of these taxa are found in the area of Madura Cave, although Lasiorhinus latifrons occurs in similar habitats in nearby South Australia and Cercartctus concinnus is found at the eastern and western edges of the Nullarbor Plain. Phascolarctos cinereus, Pseudocheirus peregrinus, and Trichosurus vulpecula are restricted to more mesic environments, and they occur only in the 15,000- to 20,000-y b.p. interval. INTRODUCTION This portion of the Madura Cave work continues the systematic section begun in Part I and continued in Parts II, III, and IV (Lundelius & Turnbull, 1973, 1975, 1978, 1981) and covers part of the Order Diprotodonta, specifically the Vombatidae {Lasiorhinus c.f. latifrons), Phascolarctidae (Phascolarctos cinereus), Phalangeridae (Trichosurus vulpecula and Pseudocheirus peregrinus), and Burramyidae (Cercartctus concinnus). Measurements and abbreviations used are either those in standard use or they have been outlined in one of the previous sections. The Hershkovitz (1971) summary is used for alternative dental terminologies. 2 FIELDIANA: GEOLOGY SYSTEMATICS (section continued) CLASS MAMMALIA (continued) Subclass Theria (continued) Infraclass Eutheria (continued) Cohort Marsupiata (sensu Turnbull, 1971; = Metatheria) (continued) Order Diprotodonta (Owen, 1866; revised Ride, 1964; Kirsch, 1968) Family Vombatidae Lasiorhinus (Gray, 1863) Lasiorhinus c.f. latifrons (Owen, 1845) Material Trench 1, 2V2 ft level (about top of Unit 2) PM 4782, left M4 or right M1 (fig. 5) Trench 3, Unit 2, Level ?, probably Level 1 TMM 41106-71, right Ii TMM 41106-72, right Ii (fig. 1) Trench 3, Unit 3, Level ?, probably Level 1 TMM 41106-52, fragment of the palatal portion of a left maxillary with sym- physis and alveoli for posterior lobe of M1 and for both lobes of M2; as- sociated with it are two teeth, one the posterior lobe of an M1 that appears to fit the alveolus and a right M1 at a very young age/ wear stage (fig. 2; pi. I-A) Trench 4, Unit 2, Level 1 PM 36833, half molar PM 36834, half molar (fig. 3) PM 36835, complete upper molar (fig. 4) TMM 41106-5021, one lobe of a molar Trench 4, Unit 2, Level 2 PM 36836, half a lobe of a molar, probably M2, M3, or M4 PM 36838, half molar tooth Trench 4, Units 4-5 TMM 41106-5022, an upper molar with one lobe heavily leached Trench 5, Unit 6 WAM 75.1.142, nearly complete molar at an early wear stage (pi. I-B) TMM 41106-5023, three quarters of a molar Description Most of the material from Madura Cave referred to Lasiorhinus c.f. latifrons represents juvenile individuals. Two relatively complete right lis show the characters stated by Merrilees (1967) to be diagnostic of that taxon (fig. 1). In cross section they are higher than wide, in contrast to Vombatus hirsutus in which the lis are wider than high. There is a stout enamel cover for the full length of the ventral (front) surface. This enamel band extends a short distance onto the medial and lateral sides of the tooth where it ends abruptly. The remainder of the tooth is covered by a thin layer of cementum that also extends over the enamel in places. B o 1 cm Fig. 1. Lasiorhinus c.f. latifrons, TMM 41106-72, a right Ii shown in dorsal (occlusal) and crown and root (A), mesial (B), and distal (C) views. Two cross- sectional outlines are also given. 4 FIELDIANA: GEOLOGY The wear surface is slightly concave anteroposteriorly and transversely, espe- cially near the tip. It makes a more acute angle with the ventral surface of the tooth than is seen in Vombatus. This acute angle is also seen in a Recent speci- men of Lasiorhinus (TMM M-943 A). The palatal portion of the maxillary (TMM 41106-52), a fragment that extends from the midline to the alveolar border, shows the abrupt widening behind M1 and parts of the alveoli for the posterior lobe of M1 and both lobes of M2 (fig. 2). Merrilees (1967) states that the least width of the palate between M1 alveoli of opposite sides is wider in Lasiorhinus than in Vombatus. The Madura Cave specimen measures 3.5 mm from the posterior lobe of M1 to the midline suture, which when doubled gives a total width of 7.0 mm. This is slightly narrower than any of Merrilees' specimens. Associated with the maxillary fragment in the matrix was one complete lobe of a molar whose shape and curvature fit the alveolus of the posterior lobe of M1. It may or may not be part of the same individual as the maxillary fragment. Isolated teeth of wombats are difficult to identify both as to taxon and as to position in the jaw, and in the case of molars, the difficulty even extends to whether they are upper or lower teeth. Merrilees (1967) has noted that the upper molars are usually more tightly curved, especially in Lasiorhinus, and that the anterior ones are more so than the posterior ones. He has also recognized the following criteria for distinguishing Lasiorhinus teeth from those of Vombatus in most cases: an inward (inflected) curvature of the enamel on the anterobuccal surface of the fore lobe of M3 and M4 of Lasiorhinus (it is also present on the M2 of one of our comparative specimens, TMM M-943 B); the more rounded apices of the lobes of the molars in Lasiorhinus; and the more rounded aspect of the anterolabial and posterolabial corners of the upper molars and the anterolingual and posterolingual corners of the lower molars. All of the Madura Cave speci- mens assigned to Lasiorhinus showed strong development of one or more of these features. All post-incisor teeth in the Madura Cave collection that have been so assigned are molars, and no premolars have been recognized. All the molars with sufficient wear and growth to have achieved the adult pattern show features typical of Lasiorhinus. They are open-rooted, bilobed, subtriangular prisms, with the apices of the lobes broadly rounded. The an- terolingual and posterolingual (for lowers) and anterolabial and posterolabial (for uppers) corners of the teeth are smoothly rounded curves (figs. 3, 4), not more sharply angled as in Vombatus. Specimen PM 36834 (fig. 3) shows the inward curve (or concavity) of the enamel on one side of a lobe, and PM 36836 shows a concavity on both sides of the same lobe. Enamel is absent from the lingual surfaces of the lower molars and the labial surfaces of the upper molars. A layer of cementum encases the tooth but is thickest in the "V" between the lobes and on those parts of the tooth that lack enamel. Where the enamel is absent the cement is in direct contact with the dentine and forms a distinct outer layer of the tooth of approximately the same thickness as the enamel. Often it mimics it closely. This is probably the reason for the statement in Stirton et al. (1967, p. 456) that "the enamel encircles the tooth." They referred to a premolar, but the condition is found to be common in all cheek teeth of all of our available specimens of modern Vombatus hirsutus and L. latifrons. Figure 5 shows one of the Madura Cave specimens that illustrates the condition very well, and Figure 6 shows the extent of the cementum. 0 5 mm Fig. 2. Lasiorhinus c.f. latifrons, TMM 41106-52, a maxillary fragment shown in palatal view. The broken posterior tooth half was associated and appears to fit the posterior alveolus for the Mi. -i-«o -■-O >.~a o 2 § c (Urn4' 111 iii so m .2 I "2 a o "9 ^ 3.0« Oi to X 01 ■£ "3 O C !E C 10 01 -g uj tu ° 2 3 ■St ti * 01 > u *J > « 5 en •g 8 *H — £ .2 ^ J3 C fa " C S c a .s s *3 * K ffl ® 2 oo .. u c oi « * p 773 .o o !§.CEu ■e a> o J* (J x • •£ tO *^ ammo .5 t; g a. -C O o °o .2 oi — , oi »1 ^ n . « 3 n m to .5 ai ■ Joe Hi eg *i ti 9 5:5 1cm Fig. 4. Lasiorhinus c.f. latifrons, PM 36835, an upper molar shown in crown (A); lingual (B); and labial, mesial, and distal (C) views. FIELDIANA: GEOLOGY Fig. 5. Lasiorhinus c.f. latifrons, PM 4782, a molar tooth (either LM4 or RM1, or RMi or LM4) shown in five views: crown (A), root (B), labial or lingual (C), and proximal or distal (D and E). The worn occlusal surface of the tooth usually has low ridges, one on each lobe, that extend nearly transversely across the teeth almost to the apex of each lobe. On specimen WAM 75.1.142 (pi. I-B), which is from a very young individ- ual, each of these low ridges originates at a shallow pit on the occlusal surface on the side of the tooth opposite the apices of the lobes. These pits appear to open into the pulp cavities and to be the sites of deposition of secondary dentine. Three teeth, WAM 75.1.142 (pi. I-B), TMM 41106-52 (pi. I- A), and TMM 41106-5023, are from very young individuals. They show a distinct taper toward the crown. In TMM 41106-5023, which is one of the younger, the enamel partly covers the crown, and wear has produced one small oblique facet that is in- clined toward the center of the tooth where the lobes coalesce. In one other tooth, TMM 41106-52, which is about at the same age/wear stage as TMM 41106-5023, the enamel has just gone from the crown, and some features of the Fie. 6. Vombatidae gen. and sp. indet., PM 36837, one half of one moiety of a molar tooth preserving the external surface on one side and the now polished longitudinal section from near the center of the moiety on the reverse side. A, The section exposing the filled pulp canal (P), the dentine (D), the enamel which is seen along the convex curve (E), and the superficial cementum covering (C) that can be seen as a thin dark band on the outer surfaces near the functional wear surface (FW). B, The outer surface of the tooth fragment (either its anterior or posterior surface) showing that the cementum cover (C) surrounded the moiety for much if not all of its length. It is broken away so that a wide expanse of the superficial surface of the enamel (E) is exposed. The enamel did not completely surround the moiety, but instead thinned to a sharp edge, forming an enamel dentine junction (E/D). Dentine (D) is also visible where the enamel is broken away along the side with the convex curvature. 10 FIELDIANA: GEOLOGY Plate I. Lasiorhinus c.f. latifrons. Scanning electron microscope (SEM) photographs of two specimens: A, TMM 41106-52, a montage of two exposures showing the occlusal surface of a right M1 at a young age/wear stage. B, WAM 75.1.142, the same for another molar tooth whose position is uncertain. underlying dentine surface can be seen. There is a ridge that runs transversely across the larger of the two lobes to connect two faint cusps. The smaller lobe has a truncated occlusal surface — a flat facet. In addition to the wombat materials that have been assigned to L. latifrons, an equal number of specimens are indeterminate. These are teeth or scraps that are probably also L. latifrons, but there is no good evidence upon which to base an assignment. They are listed below and will be discussed with the assigned specimens. Three of them are from juvenile individuals and preserve some crown features (pi. II). Plate II. Lasiorhinus gen. and sp. indet. SEM photographs of the occlusal surface of isolated teeth, each at an early age' wear stage. A, PM 36831, tooth row position uncertain. B, PM 36839, probably an M', montage of two SEM photos. C, WAM 75.1.143, tooth row position uncertain, montage of two SEM photos. 11 12 FIELDIANA: GEOLOGY Discussion Lasiorhinus was a member of the fauna of the region continuously between about 38,000 y b.p. and 7,500 y b.p., although it is absent today. None of the levels indicate any great abundance, but Unit 2 contains appreciably more specimens than any other unit; Units 1, 4-5, and 7 have each produced only a single specimen. The presence of Lasiorhinus is not surprising; it lives today on the eastern side of the Nullarbor Plain (east of Eucla). Vombatidae Genus and species indeterminate Material Trench 2, top 1 ft (= Unit 1, probably Levels 1 or 2) PM 4781, left M2 or Ms Trench 3, Unit 2, Level 1 TMM 41106-114, left M1 from a subadult individual TMM 41106-115, molar Trench 3, Unit 2, Level 2 PM 36831, molar at a very early age/wear stage (pi. II- A) Trench 4, Unit 2, Level 1 TMM 41106-5024, fragment of a right I1 Trench 4, Unit 2, Level 2 PM 36837, half molar (fig. 6) PM 36839, very early age stage molar, probably a LM1 (pi. II-B) WAM 75.1.144, half of a lobe of a molar TMM 41106-5025, one lobe of a molar Trench 4, Unit 7, Level 2 TMM 41106-5026, one lobe of a molar Trench 5, Unit 6 WAM 75.1.143, nearly complete molar at very early age stage (pi. II-C) PM 36840, molar fragment Several of these indeterminate specimens deserve brief comments: The extremely early tooth stages represented by PM 36831, PM 36839, and WAM 75.1.143 are of particular interest because they show traces of the original crown structure, which are illustrated (pi. II) by scanning electron micrographs (SEM). PM 4781 and TMM 41106-114 and TMM 41106-115 are at subadult age stages, judging by their sizes, proportions, and wear. They represent some of the older individuals in the collection. One specimen, PM 36832, is eroded in a peculiar manner. Much of its surface may have been removed by chemical leaching. PM 36837 is a longitudinally broken tooth, the broken surface of which has been ground and polished to show the nature of the dentine-pulp cavity and the enamel-cementum and dentine-cementum relationships (fig. 6). LUNDELIUS & TURNBULL: MADURA CAVE Family PHASCOLARCTIDAE Phascolarctos Blainville, 1816 Phascolarctos cine re us (Goldfuss, 1817) 13 Material Trench 4, Unit 2, Level 2 TMM 41106-628, right M1 (pi. I1I-A) TMM 41106-17, right M2 (fig. 7A, B) PM 36815, very worn left Ma (fig. 7C, D) This is the second record of this taxon from the Nullarbor Plain; the other record is also from Madura Cave (Milham & Thompson, 1976). That specimen came from the south passage and is apparently reworked from an earlier deposit than the unit in which it was found. Description The M1 is similar to the M1 of Recent specimens from New South Wales (FMNH 81524) and Victoria (TMM M2946and FMNH 112541) but differs in size. The Madura Cave specimen is smaller than each of the Recent specimens (table 1). It is at a nearly unworn stage, hence all crests, ridges, and crenulations are strong, but they do not appear to be any more developed than those on the Recent specimens. The right M2 shows the same differences from the Recent materials as does the M1. It too is smaller than the Recent specimens. Length measures of the M2s of the three Recent specimens available for comparison are shorter than the anterior width measures, whereas the M* lengths are greater than the anterior widths (table 1). In the Madura Cave specimen this ratio is 0.92, which is even lower than in any of the Recent M2s and is far from the proportions seen for the M*s. On this evidence, it is considered to be an M2 rather than an M \ One other Table 1. Measurements (in mm) and ratios of upper dentitions of fossil and Recent specimens of Phascolarctos cinereus. Recent Madura Cave TMM FMNH FMNH TMM TMM Variate M-2946 81524 112541 41106-625 41106-17 M'L 7.9 7.9 7.9 7.0 AW 8.2 7.6 7.9 7.2 PW 7.7 7.3 7.0 6.6 M2L 8.1 7.4 7.9 6.8 AW 8.3 7.5 8.1 7.4 PW 7.4 6.8 6.7 6.4 L/AW 0.976 0.987 0.976 0.92 M3L 7.9 7.4 7.5 AW 7.8 7.4 7.4 . . . . PW 6.7 6.2 6.4 . . . . L/AW 1.012 1.0 1.14 . . . . M4L 7.2 >7.0 7.1 . . . AW 6.9 6.9 5.8 . . . . PW 5.4 5.6 >4.7 . . . . 5 mm Fig. 7. Phascolarctos cinereus. TMM 41106-17, a right M2 shown in crown (B), labial (A), and lingual (C) views. PM 36815, a left M4 shown in crown (D) and labial (E) views. 14 LUNDELIUS & TURNBULL: MADURA CAVE 15 Table 2. Measurements (in mm) of lower dentitions of fossil and Recent specimens of Phascolarctos cinereus. Fossil Recent Labyrinth Cave Madura Cave TMM FMNH FMNH cast WAM PM Variate M-2946 81524 112542 69.4.1 36815 MiL 7.8 7.6 8.2 8.5 AW 5.0 4.6 4.8 5.6 . . PW 5.3 5.2 5.3 6.0 Ml 8.0 7.5 7.7 8.5 AW 5.4 4.8 5.2 5.5 PW 5.3 4.8 5.2 5.8 MjL 8.2 7.5 7.9 8.2 AW 5.3 + 4.9 5.2 5.3 PW 5.2 4.8 5.2 5.3 M. I 8.1 7.5 8.0 8.2 6.4 AW 5.1 4.8 5.4 5.0 >3.8 PW 4.5 4.5 4.8 5.0 >3.4 minor difference can be seen; the tooth has a nearly straight anterior edge compared with the Recent specimens in which it bulges forward slightly. The M» is badly etched, with most of its enamel surface destroyed. Wear had breached the tips of the major cusps and crests to expose dentine. The pro- toconid and hypoconid are deeply worn on their tops, and the enamel on the labial side of the whole tooth is gone. The exposed dentine of the labial side is polished, which could have resulted from either masticatory wear or post- depositional leaching. The tooth appears to be very elongate and narrow (table 2), but the appearance is exaggerated by the loss of the labial enamel surface and a similar but less extreme loss of enamel on the lingual side. The crenulations of the original enamel surface can be vaguely seen in the central valley. The pos- terior sides of hypoconid and entoconid and the entire postcingulum are bev- eled by wear as is typical of worn Mas in the living form. The metaconid and entoconid are higher and thinner and have better developed longitudinal crests than the labial cusps, as in the Recent comparative specimens and a fossil from Labyrinth Cave (Merrilees, 1969, WAM 69.4.1). The labial cusps, with their associated crests, form open selenes of about equal proportions as in the living form. The roots, especially the posterior, are inclined posteriorly, and both are eroded and polished. Discussion The Madura Cave material is slightly smaller than the three modem speci- mens of Phascolarctos cinereus we have for comparison (tables 1, 2). It is also slightly smaller than the modern sample reported by Bartholomai (1968) and the Pleistocene examples from Mammoth Cave (Glauert, 1910). Phascolarctos stirtoni from the Cement Mills limestone of southeastern Queensland differs from P. cinereus in being 25% to 30% larger and in some minor and variable dental characters (Bartholomai, 1968). Marshall (1973a) re- ports the presence of Phascolarctos remains referable to P. stirtoni from Pleis- tocene cave deposits at Narracoorte, South Australia. Plate III. SEM photo montages of: A, Phascolarctos cinereus, TMM 41106-628, a right M1 in posterolingually oblique view of occlusal surface. Top: Montage of all ten exposures. Bottom: Nine separate individual exposures are shown. 16 Plate HI. SEM photos of: B, Trichosurus vulpecula. PM 36830, a left Mi in posterodorsal oblique view; C, Pscudochcirus peregrinus, TMM 41106-5027, stereopair of a right M1 in crown view (rotate 90° for stereo viewing). 17 18 FIELDIANA: GEOLOGY The Madura Cave Phascolarctos is referable to P. cinereus on the basis of all characters of the material currently available. A larger sample would be neces- sary to determine whether the apparent small size is significant. Phascolarctos cinereus is found today in sclerophyll forest from Queensland to Victoria east of the Great Dividing Range (Marlow, 1958). According to Tyndale-Biscoe & Calaby (1975), its optimum habitat is tall open forest, but some of its range is in dry woodland, which implies that dense forest is not a prerequisite for its existence (Calaby, 1971, p. 87). Its presence in Pleistocene deposits of the Nullarbor Plain indicates somewhat moister conditions during the Pleistocene but not necessarily those of its optimum habitat. Phascolarctos cinereus is known from late Pleistocene deposits at Lake Menin- dee and Lake Victoria, southwestern New South Wales (Tedford, 1967; Mar- shall, 1973b), and from Holocene deposits in McEachern's Cave, southwestern Victoria (Wakefield, 1967). Madura Cave is almost exactly halfway between the Pleistocene localities in eastern Australia and those of southwestern Australia (Glauert, 1910; Merrilees, 1969; Archer, 1972). The age of the Madura Cave material (20,000 ± 430 y b.p.) supports the hypothesis put forth by many people (Serventy, 1951, 1953; Main et al., 1958; Gentilli, 1961) that faunal exchange between east and west took place across this area during the last glacial maximum. Family PHALANGERIDAE This family is represented by two teeth, one each of Trichosurus vulpecula and Pseudocheirus peregrinus. Trichosurus Lesson, 1828 Trichosurus vulpecula (Kerr), 1792 Material Trench 4, Unit 2, Level 2 PM 36830, left M, (fig. 8, pi. III-B) Description This tooth compares well with the available modern examples of T. vulpecula cf. hypoleucas from southwestern Australia, but it is slightly smaller. It is from a juvenile because it is virtually unworn; a comparably worn modern example has functional dP4S. Its trigonid is narrower than the talonid, and the protoconid stands higher than any other cusp. In crown view the protoconid is centrally located within the trigonid and is connected to a distinct metaconid that lies at the lingual edge of the crown. The metaconid stands nearly as high as the protoconid and entoconid, which are the highest cusps. There is no distinct paraconid: the anterior crest from the protoconid extends nearly straight for- ward to the rounded anterior point of the tooth. There are weak but distinct anterolabial and anterolingual cingula. The posterior cingulum is small. It swings down from high on the entoconid and is tied to a lower, stout hypoconid immediately labiad of the bottom of the "V" of the crest between hypoconid and entoconid. Although all of the available Recent Western Austra- lian comparative specimens show considerable variation among themselves in B h 0 5 mm Fig. 8. Trichosurus vulpecula, PM 36830, a left Mi shown in crown (A) and labial (B) views. 19 20 FIELDIANA: GEOLOGY Table 3. Measurements (in mm) of Trichosurus, Mi. L AW PW Specimen No. and taxon N Range Mean Range Mean Range Mean Madura Cave PM 36830, T. vulpecula 1 ... 4.40 . . . 3.00 . . . 3.25 Holocene and Recent, S.W. Australia TMM M-849, M-1743, M- 2031a-c, M-2032; TMM 40238-1, 41110-6 through 10, T. v. hypoleuca 12 4.70-5.30 5.12 2.75-3.30 3.02 3.25-3.80 3.53 FMNH 98783, T. v. vulpecula 1 5.30 3.15 3.35 FMNH 60936, T. v. johnstonii 1 5.30 2.90 3.45 USNM 221121, T. caninus 1 5.45 3.10 3.70 the stoutness of the cingulum and its connecting ridges, they have a more poorly developed posterior cingulum complex than is seen in the eastern forms (T. v. vulpecula, T. v. johnstoni, orT. caninus). In these the posterior cingulum is strongly developed. The Madura Cave specimen is somewhat smaller than Re- cent specimens from southwestern and southeastern Australia (table 3), although we have not had adequate samples from the latter area. It is most like the Recent specimens from southwestern Australia referred to T. v. hypoleucas. Comparative material of T. v . ruficollis was not available. Pseudocheirus Ogilby, 1837 Pseudocheirus peregrinus (Boddaert), 1785 Material Trench 3, Unit 2, Level ? (probably Level 1) TMM 41106-5027, right M1 (fig. 9, pi. III-C) Description The protocone, paracone, metacone, and hypocone of the Madura Cave specimen each has a simple crescentic form with no secondary ridges. The paraconule and metaconule are asymmetric. The long arm of the paraconule extends posteriorly into the central basin; the long arm of the metaconule ex- tends into the "V" formed by the junction of the posterior crest of the paracone and the anterior crest of the metacone. The enamel of the tooth is smooth. There are small secondary ridges from the hypocone and posterior crest of the pro- tocone. All of these characteristics are found in P. peregrinus, and the Madura Cave specimen is thus referred to that species. Dental measurements of the M1 from Madura Cave and of some fossil and Recent comparative materials are given in Table 4. Discussion Pseudocheirus peregrinus and Trichosurus vulpecula both have disjunct popula- tions in southwestern Australia and in eastern Australia today. Both have been Fig. 9. Pseudocheirus pcrcgnnus, TMM 41106- 5027, a right M1 shown in crown (A), labial (B), lingual (C), and anterior or mesial (D) views. Table 4. Measurements (in mm) on fossil and Recent specimens of Pseudocheirus pere- grinus and P. convolutor. Specimen No. Tooth L AW PW Madura Cave TMM 41106-5027 M1 4.00 3.10 3.30 Pleistocene, S.W. Australia TMM 41237-2 M' 4.70 3.60 4.00 TMM 41110-2 M' 4.70 3.70 4.00 Recent FMNH 98771 M1 4.30 3.50 3.90 W 3.80 3.10 MJ 3.90 2.90 21 22 FIELDIANA: GEOLOGY found in surficial deposits in other Nullarbor caves: from Webb's Cave (Lun- delius 1957, 1964) and Murraelellevan Cave (Lundelius & Turnbull, un- published data) and T. vulpecula from Cave N41 of Anderson (1964) and Weebubbie Cave (Lundelius & Turnbull, unpublished data). Trichosurus vul- pecula has been reported from Horseshoe Cave (Archer, 1974), but without stratigraphic context. As in the case of Cercartetus concinnus, these occurrences do not necessarily indicate present or recent historic occupancy of this region. They probably indicate their presence on some parts of the Nullarbor Plain after the time of their latest known record in the Madura Cave sequence (15,600 y b.p.) to judge by the C14 date of 3,300 y b.p. on a mummy of a thylacine from Murraelellevan Cave (see discussion of Cercartetus concinnus). The occurrence of these taxa in Unit 2 with other taxa characteristic of more mesic conditions than the present is consistent with their known distributions and habitat re- quirements. Family BURRAMYIDAE Cercartetus Gloger, 1841 Cercaertus Burmeister, 1837. Cercartus Gloger, 1841. Dromicia Grey, 1841. Grey's Australian Appendix, vol. ii, p. 407. Eudromicia Mjoberg, 1916. Cercartetus Iredale & Troughton, 1934. A checklist of the mammals recorded from Aus- tralia. Mem. Austral. Mus., 6:122. Cercartetus Wakefield, 1963. The Australian pigmy-possums. Victoria National, 80(4):99-116. Cercartetus concinnus (Gould), 1845 Dromicia concinna Gould, 1845. Proc. Zool. Soc, 1845, p. 2. Phalangista (D) concinna Waterhouse, 1846. Nat. Hist. Mus. U.I., p. 314. Phalangista (D) neilli Waterhouse, 1846. Nat. Hist. Mus. U.I., p. 315. Dromicia concinna Thomas, 1888. Cat. Marsup. Brit. Mus., p. 146. Material Trench 3, Unit 2, Level ? WAM 75.1.137, right ramus with alveoli for all teeth (fig. 10B) Trench 4, Unit 2, Level 2 PM 36817, edentulous left ramus with alveoli of Ii, molars, and most of inter- vening teeth (fig. 10A) PM 36818, edentulous left ramus with alveoli for all teeth (fig. 10B) TMM 41106-5006, edentulous right ramus with alveoli for incisors and M3. PM 36819, edentulous left ramus fragment with alveoli for Ii through M2 (fig. 10A) TMM 41106-5007, edentulous right ramus with alveoli for P4 through M3 (fig. 10B) Trench 4, Units 4-5 TMM 41106-3, edentulous right ramus with alveoli for all teeth (fig. 10B) TMM 41106-5001, right ramus with M, and alveoli for I,, and I3, P3.4, M2 (fig. 10B; pi. IV-D) TMM 41106-5002, edentulous left ramus with alveoli for I1-M2 (fig. 10A) LUNDELIUS & TURNBULL: MADURA CAVE 23 TMM 41106-5003, left I. (pi. IV-H) TMM 41106-5004, right M' (pi. IV- A) TMM 41106-5005, right M2 (pi. IV-B) TMM 41106-5008, edentulous right ramus with alveoli for all teeth (fig. 10B) PM 36823, left Mi (pi. IV-E) PM 36824, right Mi (pi. 1V-F) WAM 75.1.139, left M. (pi. IV-G) PM 36826, left Ii (pi. IV-I) PM 36820, edentulous left ramus fragment with alveoli for all teeth (fig. 10A) PM 36821, edentulous left ramus with alveoli for all teeth (fig. 10A) Trench 4, Unit 7, Level 2 PM 36827, right M2 (pi. IV-C) Trench 5, Unit 4? WAM 75.1.138, edentulous left ramus with alveoli for all teeth (fig. 10A) TMM 41106-5009, edentulous right ramus with alveoli for Ii and Mi-3 TMM 41106-5010, edentulous right ramus with alveoli for all teeth (fig. 10B) PM 36822, edentulous left ramus with alveoli for all teeth (fig. 10A) Comparative materials: FMNH 34721 Cercartetus concinnus from Tambellup, Western Australia TMM M 842 Cercartetus concinnus from Kybybolite, Western Australia TMM M 831 Cercartetus concinnus from near Bridgetown, Western Australia TMM M 832 Cercartetus concinnus from near Bridgetown, Western Australia Monash 977 Cercartetus concinnus locality unknown and casts of: AMNH 119720 Cercartetus concinnus from Margaret River area, Western Australia AMNH 119723 Cercartetus concinnus from Picton Junction, Western Australia AMNH 191027 Cercartetus caudata from Mt. Otto, New Guinea AMNH 191006 Cercartetus caudata from Mt. Wilhelm, New Guinea Comparisons of fossil and Recent C. concinnus with C. nanus, C. lepidus, C. caudatus were made by reference to Tumbull & Schram (1973). The four species of Cercartetus (sensu Wakefield, 1963) show that C. nanus and C. concinna both have only three molars, whereas C. lepidus and C. caudatus have the usual marsupial complement of four. Between C. nanus and C. concinnus there is a marked size difference in the P4: in the former, P4 is double rooted and is several times larger than that of C. concinnus. The Madura Cave sample con- forms to C. concinnus; the differences noted below between the fossils and the Recent specimens are considered to be intraspecific. Description The 16 ramus fragments have the same basic form but show minor variations (fig. 10). They indicate the normal lower tooth formula for Cercartetus concinnus of Ii 3. P3-4. M1-3. No P4S were recovered, but some variation is indicated by 24 25 Plate IV. Cercartetus concinnus. SEM photos of six of the nine teeth representing this taxon from Madura Cave: A, TMM 41106-5004, right M1; B, TMM 41106-5005, right M2; C, PM 36827, right M2; D, TMM 41106-5001, right M. in ramus fragment; E, PM 36823, left Mi; F, PM 36824, right Mi. 26 Plate IV. Cercartetus concinnus. SEM photos of three of the nine teeth representing this taxon from Madura Cave: G, WAM 75.1.139, left Mi; H, TMM 41106-5003, left li shown in several views; I, PM 36826, left li shown in several views. 27 28 FIELDIANA: GEOLOGY differences in size and form of alveoli for it, which may be very small and single (as in TMM 41106-3 and TMM 41106-5010), large and single but with a weak constriction (as in TMM 41106-5001, PM 36817, PM 36821, WAM 75.1.138, and WAM 75.1.137), or paired for a small double-rooted tooth (as in TMM 41106- 5008 and PM 36818). The major differences between the Madura Cave speci- mens and the modern comparative materials are the greater depth of the man- dible of the fossils, particularly in the region of the premolars and Mi, and the larger size of P3 and P4 as indicated by their alveoli. In the modern specimens the teeth are minute (about the size of the point of a pin). The M1 (pi. IV-A) is similar to that of the Recent specimens. It is a four- cusped, three-rooted tooth that is elongate anteroposteriorly. The size order of the cusps is paracone > metacone 3= protocone > hypocone. The paracone and metacone are high, laterally compressed cusps with anteroposterior crests from their apices. These cusps along with their crests and a tiny, laterally compressed parastyle and an even smaller metastyle form an almost straight ectoloph. The protocone and hypocone are lower than the labial cusps and are laterally com- pressed to a lesser degree than the labial cusps and are located on the lingual edge of the tooth. Both have anterior and posterior crests. A nearly straight anterior cingulum extends from the parastyle to a point immediately anterior to the end of the elongate central valley where a cingular cuspule is located. The cingulum then bends posterolingually and joins the anterior crest of the pro- tocone. At the posterolingual corner there is another cingular cuspule that is more pronounced in the fossil tooth than on any of the Recent specimens. An inconspicuous labial cingulum constitutes the stylar shelf. The anterolabial root supports the paracone, the anterolingual root the protocone, and the posterola- bial root the metacone. No root is associated with the hypocone, but the one for the metacone has a lingual expansion and doubtless provides support for both cusps. The M2s are four-cusped, three-rooted teeth (pi. IV-B, C). They are elongated, tapered posteriorly, and basined. The cusps in order of decreasing size are paracone > protocone 3= metacone > hypocone. The four major cusps are lo- cated on the perimeter of the tooth. The paracone and metacone are laterally compressed, crested, and stand higher than the lingual cusps. There is a para- style but no metastyle. The protocone is somewhat crescentic, with its low crests forming the arms of the crescent. The anterior crest extends labially to the parastyle with only a trace of the homologue of the cingular cuspule seen on the M1. The posterior crest of the metacone curves labially to a posterior lingual corner that is situated almost on the longitudinal axis to the tooth. This point may be the homologue of the posterior cingular cusp of the M1 because it is in the equivalent position and is joined by the oblique posterior crest of the hypocone. There is a small stylar shelf labial to the paracone and the anterior half of the metacone. This feature is somewhat variable in our comparative specimens. The lis are long, enamel-covered, slightly tapered cylinders that are weakly spatulate for the anterior third of their length (pi. IV-H, I). Ridges extend pos- teriorly from the tip on the ventromedial and dorsolateral edges. The lis of Cercartetus are less curved and less spatulate than those of Acrobates or Dis- tochoerus. The Mis (pi. IV-D-G) are similar to those of the modern specimens. Each has LUNDELIUS & TURNBULL: MADURA CAVE 29 three cusps that make up an anteriorly tapered, elongate crown. The anterior cusp is the protoconid; it is the only trigonid cusp and is the largest and highest cusp of the tooth. It occupies nearly the anterior half of the tooth. It has a somewhat triangular cross section and is crested along its posterolingual side from its tip to the midpoint of the lingual side of the tooth where it joins a short crest from the entoconid in the saddle between the two cusps. The entoconid is smaller than the hypoconid but it stands nearly as high. It has a weak posterior crest that extends toward the minute hypoconulid at the posterolingual corner of the tooth. The hypoconid is V-shaped in crown view. The anterior limb of the "V" is a crest, the cristid obliqua (or premetacristid, I'"b of Hershkovitz, 1971), that extends diagonally anterolingually to the base of the protoconid. The pos- terior limb swings posterolingually, then lingually to the hypoconulid. There is a little variation in the degree of development of the cusps on our comparative material and more variation in the fossils. On PM 36824 there is a trace of the metaconid near the top of the protoconid on its posterolingual crest. This specimen also has a faint anterior cingulum tied to the anterior crest of the protoconid. In PM 36823 all cusps are less prominent than in two of the other fossil examples, and WAM 75.1.139, which is the smallest tooth of all, has the weakest cusp development. Some of this variation may be due to differences in wear. Discussion Cercartetus concinnus is found living today as two disjunct populations, one (C. c. concinnus) in southwestern Australia with its eastern limit at Balladonia, and one (C. c. minor) in Western Victoria and South Australia with its western limit at Ceduna (Wakefield, 1963). Marlow's map (1962) shows a continuous range across the coastal strip of the Great Australian Bight, but lacks the Victo- rian and southeastern portion of the South Australian part of the range because Marlow's work preceded the Wakefield report. We assume that the strip along the Bight was an inference on Marlow's part because we can find no firm reports for the area between Ceduna and Balladonia. Further, Wakefield (1963, p. 100) is quite explicit on this point. He states, "The South Australian and Victorian population of the species is geographically isolated from that of Western Aus- tralia, and the two groups differ appreciably in morphology." Although apparently absent from the Nullarbor Plain today, it has been re- corded there in deposits in three other caves besides Madura Cave: Murraelel- levan Cave (Lundelius, 1957, 1963; Partridge, 1967), Thylacine Hole (Lowry & Merrilees, 1969), and Horseshoe Cave (N59; Archer, 1974). Unfortunately, there are no dates available for the occurrences reported by Lundelius. Although the material from Murraelellevan Cave was collected from the surface, it could be several thousand years old. That this is possible is demonstrated by a C14 date of 3,300 y b.p. on a mummified carcass of Thylacinus from Murraelellevan Cave (Partridge, 1967) and a CM date of 4,600 y b.p. on another mummified carcass from Thylacine Hole (Lowry & Merrilees, 1969). Cercartetus concinnus is represented in all of the fossiliferous Pleistocene units of Madura Cave. It is missing from Unit 1, the Holocene unit, and the absence is believed to be real and not just an accident of sampling, because large volumes of that unit were processed. It should be recalled that Unit 1 lies unconformably 30 FIELDIANA: GEOLOGY upon Unit 2 deposits dated at 15,600 y b.p. in Trench 3 and 18,900 y b.p. in Trench 4. Our only date for Unit 1 was for the top 1 foot at 7,500 y b.p. At face value, this suggests that Cercartetus was reasonably abundant during the last 15,000 to 20,000 years of the Pleistocene, was rare or extinct in the area during the early part of the Holocene, and reappeared in the later Holocene, only to die out again sometime between 3,300 y b.p. and historic times. According to Wakefield (1963), who has marshalled the most evidence about the habitat requirements of C. c. concinnus, "The Mundarda is abundant in Jarrah forests (Eucalyptus marginata), where there is an undergrowth of sclerophyllous shrubbery containing many of the Myrteaceae and Proteaceae." He describes these regions and states that C. c. concinnus also occurs in the Tuart forest (£. gomphocephala) in scrubby areas and that it is absent from the Karri forests (E. diversicolor) where rainfall exceeds 40 inches per annum. He further points out that in the drier parts of its range (conditions wherein some eucalypts are present to provide homes for Cercartetus and heath or mallee, with their abundant Myrteaceae and Proteaceae and other nectar producers, provide food) the environment is still adequate. Possibly, the increase in mallee scrub cover 4,000 to 6,000 y b.p. that was reported for Norina and Madura by Martin (1973) on the basis of pollen studies could have provided the minimum habitat base for the repopulation by Cercartetus for a time in the late Holocene. Sub- sequent to this peak in the ratios of Myrteaceae/chenopod-type pollens, the ratios are shown to drop, and Martin reports that from the period 4,000 to 5,000 y b.p. to the present the scrub cover has declined markedly. This history of change of the vegetation is consistent with the history of Cercartetus in this region in the light of its habitat requirements. SUMMARY The families Phascolarctidae, Phalangeridae, and Burramyidae are repre- sented in the Madura Cave deposits by relatively little material. All are from the Pleistocene units. The family Vombatidae is represented in both Pleistocene and Holocene units. All except Phascolarctos are known from Holocene deposits in other caves of the Nullarbor Plains, but only Lasiorhinus latifrons is extant in this area. The presence of Phascolarctos in the Pleistocene deposits indicates that the time of the last major interchange between the more humid areas of south- western and southeastern Australia was during the last glacial maximum. The presence of Trichosurus, Pseudocheirus, and Cercartetus in Holocene deposits in other cave deposits in this area indicates continued post-Pleistocene increase in aridity. The evidence for climatic change presented by these taxa appears to be consistent with that suggested by the Dasyuridae and Peramelidae in prior reports (Lundelius & Turnbull, 1973, 1975, 1978, 1981). ACKNOWLEDGMENTS In addition to those individuals mentioned in earlier parts of this faunal report we wish to thank Dr. Duncan Merrilees of the Western Australian Museum; the late Dr. Hobart Van Dusen of the American Museum of Natural History; Dr. David Johnston of the United States National Museum; Dr. A. K. Lee of Monash University, Melbourne, for the loan of material; Mr. Zbigniew Jastrzebski for illustrations; and Ms. Caroline Good and Ms. Elizabeth Moore for typing. LUNDELIUS & TURNBULL: MADURA CAVE 31 Financial support was provided by Field Museum of Natural History and the Geology Foundation of the University of Texas at Austin, and by Grants GB 975, GB 3729, GB 7662 from the National Science Foundation. LITERATURE CITED Anderson, E. G. 1964. Nullarbor expedition 1963-4. 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D., and Schram, F. R. 1973. Broom Cave Cercartetus, with observations on pygmy possum dental morphology, variation and taxonomy. Rec. Austral. Mus., 28(14): 437-464. Tyndale-Biscoe, C. H., and Calaby, J. H. 1975. Eucalypt forests as refuge for wildlife. Austral. Forestry, 38(2): 117-133. Wakefield, N. 1963. The Australian pigmy-possums. Vict. Nat., 80: 99-116. . 1967. Preliminary report on McEachern's Cave, Southwestern Victoria. Vict. Nat., 84(12): 363-383. Waterhouse, G. R. 1846. Natural History of the Mammalia. I. Marsupiata. London, Balliere, pp. 1-553. Field Museum of Natural History Roosevelt Road at Lake Shore Drive Chicago, Illinois 60605-2496 Telephone: (312) 922-9410