to CO o> CO LU CO UNiVtrfSlTY OF JLLINOIS LIBRARY AX URBANA-CHAMPAIGIM GEOLOGY FIELDIANA Geology Published by Field Museum of Natural History New Series, No. 6 THE MAMMALIAN FAUNA OF MADURA CAVE, WESTERN AUSTRALIA PART IV ERNEST L. LUNDELIUS, JR. WILLIAM D. TURNBULL 4MW1981 February 27, 1981 Publication 1315 THE MAMMALIAN FAUNA OF MADURA CAVE, WESTERN AUSTRALIA PART IV FIELDIANA Geology Published by Field Museum of Natural History New Series, No. 6 THE MAMMALIAN FAUNA OF MADURA CAVE, WESTERN AUSTRALIA PART IV 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 January 3, 1979 February 27, 1981 Publication 1315 Library of Congress Catalog Card Number: 80-68335 US ISSN: 0096-2651 PRINTED IN THE UNITED STATES OF AMERICA CONTENTS List of Illustrations vi List of Tables vii Abstract 1 Introduction 1 Systematics 2 Perameles Geoffrey, 1803, 1804 2 Perameles bougaittvillei Quoy and Gaimard, 1824 3 Material 3 Descriptions 14 Discussion 28 Isoodon Desmarest, 1817 31 Isoodon obesulus (Shaw and Nodder, 1797) 31 Material 31 Descriptions 32 Discussion 43 Chaeropus Ogilby, 1838 44 Chaeropus ecaudatus Ogilby, 1838 44 Material 44 Descriptions 44 Discussion 52 Macrotis Reid, 1837 52 Macrotis lagotis (Reid, 1837) 52 Material 52 Descriptions 55 Discussion 65 Summary 69 Acknowledgments 70 References 70 LIST OF ILLUSTRATIONS Frontispiece. Maps of the Nullarbor Plain area ii 1. Perameles bougainvillei , a subfossil specimen from the surface of Murraelellevan Cave 15 2. Perameles myosura notina Thomas = P. bougainvillei 16, 17 3. Perameles myosura notina Thomas = P. bougainvillei 18 4. Perameles bougainvillei from Madura Cave 19 5. Perameles bougainvillei from Madura Cave 21 6. Perameles bougainvillei from Madura Cave 22 7. Perameles bougainvillei from Madura Cave 23 8. Bivariate graphs of the cheek teeth of Perameles bougainvillei 26, 27 9. Bivariate graphs of isolated M 2s and M3s of Perameles bougainvillei 28 10. Isoodon obesulus affinis (Waterhouse) from Wynyard, Tasmania 34, 35 11. Isoodon obesulus from Madura Cave 36 12. Bivariate graphs of length vs. anterior width measurements of upper and lower molars of Recent and fossil specimens of Isoodon obesulus 38 13. Isoodon obesulus from Madura Cave 40, 41 14. Chaeropus ecaudatus, sub-Recent fossil specimens from several other Nullarbor Caves 45 15. Chaeropus ecaudatus, sub-Recent fossil specimens from several other Nullarbor Caves 46, 47 16. Chaeropus ecaudatus from Madura Cave 49 17. Bivariate graphs showing plots of length vs. anterior width measures for each of the molar teeth of sub-Recent specimens of Chaeropus ecaudatus from several Nullarbor Caves 51 18. Macrotis lagotis (Reid), a Recent juvenile specimen from Woyaline Wells ... 56, 57 19. Macrotis lagotis specimens from Madura Cave 59 20. Macrotis lagotis from Madura Cave 62, 63 21. Bivariate graphs of length vs. anterior width for the upper and lower molars of Macrotis lagotis from Madura Cave 66 LIST OF TABLES 1. Numerical data on some dental measurements of Perameles bougainvillei from units 1 and 2 of Madura Cave 29 2. Numerical data on some dental measurements of Perameles bougainvillei from units 4-5 of Madura Cave 30 3. Results of "t" tests of some dental measurements of Perameles bougainvillei from units 1 and 4-5 of Madura Cave 30 4. Results of "t" tests on means of some mandibular and dental measurements of Perameles bougainvillei from Victoria Cave and Fromm's Landing 30 5. Numerical data on dentitions of various fossil and Recent samples of Isoodon obesulus 42 6. Numerical data on dentitions of two samples of Chaeropus ecaudatus 50 7. Numerical data on upper dentitions of fossil and Recent samples of Macrotis lagotis 67 8. Numerical data on lower dentitions of fossil and Recent samples of Macrotis lagotis 68 ABSTRACT The peramelids represented in the Madura Cave deposits are Perameles bougainvillei , Isoodon obesulus, Chaeropus ecaudatus, and Macrotis lagotis. Perameles bougainvillei is found in abundance in all stratigraphic units. Specimens from unit 1 (7420 b.p. and younger) show no qualitative differences from those in units 2 through 4-5 (15,000-22,000 b.p.). Post-Pleistocene reduc- tion in size is indicated by significant differences in three of nine dental mea- sures tested (P < .05, one sided "t" test). Chaeropus ecaudatus and Macrotis lagotis are represented in all units and are known from the Nullarbor Plain in historic times. No morphological changes in these species are seen in the period of time represented by the Madura Cave deposits. Isoodon obesulus is present in all units except unit 7. Its absence from unit 7 is possibly a sampling accident. The recovered specimens are intermediate in size between specimens of/, obesulus from southwestern Australia and Victoria and specimens of /. auratus, but they are closer to the former. No morphological differences are seen between the specimens from units 4-5 and unit 1. This taxon is not known to have been present on the Nullarbor Plain in historic time. Its presence in early Holocene deposits suggests a change to drier climatic conditions in this region during this time. INTRODUCTION This section of the Madura Cave work continues the systematic section begun in Part I (1973) and continued in Parts II (1975) and III (1978), in which 14 taxa of the Dasyuroidea and one of Thylacoleonidae were covered. Here we consider the Order Peramelina, which is represented in the Madura Cave deposits by at least four taxa: the very abundant Peramelus bougainvillei and smaller numbers of Isoodon obesulus, Chaeropus ecaudatus, and Macrotis lagotis. Madura Cave is located on the Roe Plain 6 miles (9.6 km) south of the settle- ment of Madura, 110 miles (177 km) west of Eucla (see frontispiece). The cave system consists of a shallow oval doline whose long axis is oriented NW-SE, with two tunnels extending outward from its margins. One tunnel extends southwestward from the doline's southern end, the other northwestward from its northern end. No excavations were carried out in the southern tunnel. The northern tunnel, which is much larger than the southern tunnel, has a floor 8-10 ft lower than the surface of the doline and a gentle gradient toward the back of the cave. Five trenches were dug in the northern tunnel, two by Lundelius in 1955 and 2 FIELDIANA: GEOLOGY three by us in 1964. The stratigraphic sequence in trenches 3 and 4, which produced the most faunal material, consists of the following: a top unit (unit 1 of both trenches) of loose, gray brown silt with many limestone fragments, abundant small bones, and organic material. This unit is separated from the underlying material by an irregular surface. A C-14 date (tx 1146) of 7,470±120 b.p. was obtained from the top foot of this unit in trench 4. A sequence of red, clayey silts, sands, and limestone powder underlies unit 1. The following C-14 dates were obtained: Trench 3 Unit 2, upper 1 ft, 15,600±250 years b.p. (tx 1145) Unit 2, lower 1 ft, 22,400±580 years b.p. (tx 1142) Trench 4 Unit 2, upper 6 inches, 18,990±220 years b.p. (tx 1140) Unit 2, 6-12 inches, 20,000±430 years b.p. (tx 1141) Units 4-5, 22,220±570 years b.p. (tx 1144) Unit 7, lower 1 ft, exposed: 37,880±3,520 years b.p. (tx 1143) Measurements and abbreviations used are either those in standard use or they have been outlined in one of the prior sections. The dental terminology has recently been summarized by Hershkovitz (1971), and the reader is referred to that work as containing the most complete compilation of terms and abbrevia- tions available. Those we use are all listed there. The diagrams below indicate the manner used in taking the dental measures: L (= length), AW (= anterior width), PW (= posterior width). They also show some of the Hershkovitz sym- bols for certain of the stylar cusps and segments of the eocrista(id) that are convenient designations. Major cusp abbreviations follow the Cope-Osborne standard. PA'd ME'd SYSTEMATICS (section continued) Class Mammalia (continued) Subclass Theria (continued) Infraclass Eutheria (continued) Cohort Marsupiata (sensu Turnbull, 1971: = Metatheria) (continued) Order Peramelina (Gray, 1825; revised Ride, 1964) Family Peramelidae (Waterhouse, 1838, 1841) Perameles Geoffroy, 1803, 1804 LUNDELIUS & TURNBULL: MADURA CAVE 3 Perameles bougainvillei Quoy and Gaimard, 1824 Material Trench 1, top 1 ft Lundelius (1963) listed P. bougainvillei as occurring in this level, but he gave no catalogue numbers, and we've not been able to locate specimens so assigned. Apparently they have been misplaced. Trench 1, top 30 inches PM 26183, right ramus with M3 and alveoli of all other cheek teeth PM 26184, left ramus with M3 and alveoli of P4-M2 and M4 PM 26185, edentulous left maxillary fragment with alveoli of premolars and M1"2 Trench 2, 2V2 ft below surface PM 25223, edentulous left ramus fragment with alveoli of P4-M4 PM 25225, left ramus fragment with M2-3 and alveoli of Mi and M4 PM 25226, right ramus fragment with M4 and alveoli of M2-3 Trench 3, Unit 2, Level 1 TMM 41106-68, left mandible with P4 and alveoli of rest of cheek teeth TMM 41106-69, left ramus fragment with P4 and alveoli of Pi -3 and Mi -3 TMM 41106-70, right mandible with M3-4 and alveoli of h and rest of teeth to Ma TMM 41106-71, right ramus fragment with M4 and alveoli of M1-3 TMM 41106-72, right ramus fragment with C, P3-4 and alveoli of b. Pi, M2-3 TMM 41106-73, right ramus fragment with broken M4 and alveoli of M2-3 TMM 41106-74, right ramus with M3-4 and alveoli of premolars and Mi -2 TMM 41106-75, right ramus with M4 and alveoli of C-M3 TMM 41106-76, left ramus fragment with Mi -3 and alveoli of b through P4 TMM 41106-77, left mandible with C, Pi or 2, P4, Mi, M3-4 and alveoli of incisors and intervening teeth (fig. 6A, B) TMM 41106-78, left mandible with P4 and M4 and alveoli of C and rest of cheek teeth TMM 41106-79, right mandible with M3-4 and alveoli of C-M2 TMM 41106-80, right ramus fragment with M3 and alveoli of I2-M2 TMM 41106-81, left ramus fragment with P4 and alveoli of C-P3, M1-2 TMM 41106-82, left ramus fragment with M4 TMM 41106-363, right maxillary with P3"4, M1, alveoli of rest of cheek teeth (fig. 4A, B) TMM 41106-2753, left mandible with C-M4, (P3 broken), alveoli of I1-3 (fig. 7A, B, C) PM 13339, edentulous right maxillary with alveoli for P2, M4 PM 13340, right maxillary with M23, alveoli for P^M1 and M4 (fig. 4C, D) PM 13341, left maxillary with P3, alveoli for P2, P4, M1 PM 13342, right maxillary with M2, alveoli for M3'4 PM 25584, right ramus fragment with C PM 25585, left ramus fragment with parts of C and Pi or 2 PM 26990, left ramus fragment with M4 and alveoli of M3 PM 26991, right ramus fragment with P4 and alveoli of C-P3 PM 26992, right ramus fragment with broken P4 and alveoli of P1-3, M1-3 4 FIELDIANA: GEOLOGY Trench 3, Unit 2, Level 1 (continued) PM 26993, left ramus fragment with Ms and alveoli of M2 and M4 PM 26994, left ramus with M4 in crypt, alveoli of P3-M3 PM 26995, right ramus fragment with P4, alveoli of C-P3 and M1-3 PM 26996, left ramus fragment with P3-4, alveoli of Mi PM 26997-8, two right Mis PM 26999-7002, four left Mis PM 27003-23, 21 right M2S or Mss PM 27024-40, 17 left M2S or Mss PM 27041-3, three left M4s PM 27044-5, two right M4S PM 27046-9, four right MJs PM 27050-4, five left M*s PM 27055-75, 21 left M2s or M3s PM 27076-90, 15 right M2s or M3s PM 27091-2, two left M4s uncatalogued, ca. 100 edentulous ramus and maxillary fragments uncatalogued, two specimens with label lost, probably this level uncatalogued, one vial with hundreds of uncatalogued premolars Trench 3, Unit 2, Level 2 TMM 41106-43, IPerameles cranium TMM 41106-322, left M1 TMM 41106-2755, right M4 TMM 41106-2756-9, four right M^ WAM 74.9.52-.53, two left M2s or M3s WAM 74.9.54, right M4 WAM 74.9.55, left Mi PM 27093-112, 20 left M2s or M3s PM 27113-5, three right M4S PM 27116-7, two left Mis PM 27118-20, three right Mis PM 27121-9, nine right Mas or Mss PM 27130-2, three premolars PM 27133-40, eight right M2s or M3s PM 27141-52, 12 left M2S or Mss PM 27219, left M4 Trench 3, Unit 2, Level 4 TMM 41106-399, right ramus fragment with M4, coronoid, and condyle TMM 41106-400, right ramus fragment with M3-4, coronoid, and condyle TMM 41106-402, right ramus fragment with M3-4, alveoli of M1-2, coronoid, and condyle TMM 41106-405, right ramus fragment with P4, alveoli of C-P3 TMM 41106-408, right ramus fragment with P2 and P3, alveoli of P4-M2 TMM 41106-414, right maxillary fragment with M1, alveoli of P3"4 PM 27162, left maxillary fragment with M2 or 3 PM 27163, left maxillary fragment with M1, alveoli of M2 PM 27164-5, two edentulous left maxillary fragments PM 27166-71, five edentulous right maxillary fragments LUNDELIUS & TURNBULL: MADURA CAVE 5 Trench 3, Unit 3 TMM 41106-38, left ramus with alveoli of C-M3 TMM 41106-39, left ramus with alveoli of P2-M4 TMM 41106-40, left ramus with alveoli of M1-M4 TMM 41106-41, right ramus with P2, alveoli of P3-M3 TMM 41106-42, right ramus with alveoli of M3, M4 TMM 41106-2764, left ramus with M4, alveoli of M2-3 TMM 41106-2765, right ramus with M4, alveoli of M3 TMM 41106-2766, left ramus with M2 or Ms, alveoli of P4-M1 or M1-2 TMM 41106-2754, left ramus alveoli of M1-M4 TMM 41106-2760, edentulous left maxillary TMM 41106-2761, left maxillary fragment with M3 and alveoli of M2 and M4 TMM 41106-2762, right M1 uncatalogued, two edentulous ramus fragments Trench 4, Unit 1, Level 1 TMM 41106-475, right M1 TMM 41106-507, 10 edentulous mandibular fragments TMM 41106-508, left maxillary fragment with P3, alveoli of P2, P4, and M1 TMM 41106-509, left Ps TMM 41106-512, right maxillary fragment with M1"4 (fig. 4E, F) TMM 41106-513, right maxillary fragment with M34 TMM 41106-514, left ramus fragment with P4, alveoli of Ps, and Mi TMM 41106-515, premolar TMM 41106-518, edentulous left maxillary fragment with palatal bar and foramina, alveoli of P1, P3, and P4, and M1. P4 is within crypt. TMM 41106-549, left M3 TMM 41106-550, left M1 TMM 41106-678, left M2 TMM 41106-5089, edentulous maxillary fragment TMM 41106-5090, four edentulous maxillary fragments TMM 41106-5091, two right M2s TMM 41106-5092, right M3 TMM 41106-5093, three edentulous right ramus fragments TMM 41106-5094, two edentulous left ramus fragments TMM 41106-5095, three left M2S TMM 41106-5096, left M1 TMM 41106-5097, right M1 PM 27153-4, two left M2s PM 27155, left M3 PM 27156, left M2 PM 27157-8, right and left M*s PM 27159-61, three right M2s PM 27172-4, three left M2S PM 27175, left Ms PM 27176, right M4 talonid PM 27177-8, two right Mis PM 27179-80, two left M* fragments PM 27181-3, two left Mis and one right Mi 6 FIELDIANA: GEOLOGY Trench 4, Unit 1, Level 1 (continued) PM 27184, broken right M2 or 3 PM 27185, right Mb PM 27186, broken right Mb or 3 PM 27187-8, two right Mi 8 PM 27189, right M1 PM 27190, right M1-2- or 3, probably M2 PM 27191-2, left and right MJs PM 27193, premolar PM 27194-5, edentulous left and right maxillary fragments PM 27196, edentulous ramus fragment PM 27197, left Ms PM 27198-9, right and left M2S PM 27200, premolar PM 27201, left ramus fragment with M4 in crypt, alveoli of M3 PM 27202, edentulous right mandible PM 27203-6, four edentulous left ramus fragments PM 27207, one left, one right edentulous maxillary fragment PM 27208, right P PM 27209, left M3 PM 27210, left Mi PM 27211, left Ms PM 27212-14, three left M2S PM 27215, left Ms PM 27216, broken left M2 or 3 PM 27217, left M3 PM 27220-21, two right Mis PM 27222, right M2 PM 27223, right M3 PM 27224, left M2 PM 27225, left Ms PM 27226-7, two left M2S PM 27228, left Ms PM 27229-30, two right Mss PM 27231-6, six right M2S Trench 4, Unit 2, Level 1 PM 27237, right mandible with Mt, alveoli of other teeth PM 27238, right mandible with P4, M3-4, alveoli of rest of teeth (fig. 5C, D) PM 27239, left ramus fragment with M3, alveoli of premolars, and other molars PM 27240, left mandible edentulous except for parts of C and P4 PM 27241, left ascending ramus fragment with coronoid, condyle, and angular process PM 27242, right mandible with P2, Pa, M3-4, alveoli of other teeth (fig. 6C, D) PM 27243, left ramus fragment with M4 in crypt, alveoli of P4-M3 PM 27244, left maxillary fragment with M2 or M3 PM 27245, edentulous left ramus with alveoli of P3-M4 PM 27246, edentulous right ramus fragment with alveoli of M3-4 LUNDELIUS & TURNBULL: MADURA CAVE 7 Trench 4, Unit 2, Level 1 (continued) PM 27247, edentulous right ramus fragment with alveoli of P3-M3 PM 27248, edentulous right ramus fragment with alveoli of M4 PM 27249, edentulous left ramus fragment with alveoli of M2-3? PM 27250, edentulous right ramus fragment with alveoli of C-P4 PM 27251, edentulous left ramus fragment with alveoli of C-P2 PM 27252, edentulous right ramus fragment with alveoli of P3-M1 PM 27253, edentulous left ramus fragment with alveoli of Pi -4 PM 27254, edentulous ramus fragment with alveoli of premolars PM 27255, left ramus fragment with P2, alveoli of P3-4 PM 27256, edentulous left maxillary fragment with alveoli of P3-M3 PM 27257, left ramus fragment with Pi, alveoli of b, C, P3 PM 27258, edentulous left mandible with coronoid, condyle, and part of angular process PM 27259, edentulous right maxillary fragment with alveoli of P3, dP4, M1 PM 27260, edentulous left maxillary fragment with alveoli of cheek teeth PM 27261-8, eight right M»s PM 27269-75, seven left M2s or M3s PM 27276, edentulous left ramus fragment with alveoli of I3-M2 PM 27277, left ramus fragment with P2 and P3, alveoli of P4-M2 PM 27278, edentulous left ramus fragment with angle, coronoid, and condyle PM 27279, edentulous right ramus fragment with alveoli of Mm PM 27280, left ramus fragment with P4, alveoli of I3-P3 PM 27281, edentulous left ramus fragment with alveoli of Ms-* PM 27282, edentulous right ramus fragment with alveoli of C-M2 PM 27283-5, three edentulous right ramus fragments with alveoli of M3-4 PM 27286, edentulous right ramus fragment with alveoli of P2-M3 PM 27287, edentulous left ramus fragment with alveoli of C-P3 PM 27288, edentulous right ramus fragment with alveoli of M3-4, portion of angle PM 27289, edentulous left ramus fragment with alveoli of P1-3 PM 27290, edentulous right ramus fragment with alveoli of premolars PM 27291, edentulous left ramus fragment with alveoli of M4 PM 27292, edentulous left ramus fragment with alveoli of P4-M2 PM 27293, edentulous left ramus fragment with alveoli of premolars PM 27294, right maxillary fragment with M4, alveoli of P4-M3 PM 27295, left Mi PM 27296-7, two premolars PM 27298, edentulous right maxillary fragment with alveoli of P4, M1 PM 27299-300, two right M2s or M3s PM 27301-7, seven left MJs PM 27308-12, five left Mis PM 27313-9, seven right Mis PM 27320-37, 18 left M2S or Mss PM 27338-63, 26 M2s or M3s PM 27364, left M4 PM 27365-8, four left M4s PM 27369-99, 31 right M2S or Mss 8 FIELDIANA: GEOLOGY Trench 4, Unit 2, Level 2 TMM 41106-713, right M2 or 3 TMM 41106-2767, premolar TMM 41106-2768, left M2 or 3 TMM 41106-2769, partial left upper molar TMM 41106-2770, left P4 TMM 41106-2771, left mandible with M3.4, alveoli of rest of teeth (fig. 5 A, B) TMM 41106-2772, edentulous left ramus with coronoid, condyle, angle, and alveoli of P4-M4 TMM 41106-2773, left ramus fragment with M4 in crypt, alveoli of M3 TMM 41106-2774, edentulous left ramus fragment with alveoli of Mm TMM 41106-2775, edentulous left mandible with alveoli of all teeth TMM 41106-2776, left maxillary with P3, alveoli of rest of cheek teeth TMM 41106-2777, left maxillary fragment with P3 TMM 41106-2778, two right maxillary fragments, one with P1, the other with P4, probably same individual TMM 41106-2779, left ramus fragment with M3, alveoli of M1-2 TMM 41106-2780, left ramus fragment with P3, alveoli of I3-P1 TMM 41106-2781, right ramus fragment with M4, alveoli of P3-M3 TMM 41106-2782, edentulous right ramus fragment with alveoli of all cheek teeth TMM 41106-2783, edentulous right ramus fragment with coronoid, condyle, angle, and alveoli of M4 TMM 41106-2784, edentulous right ramus fragment with broken Pi, P4, Mi, alveoli of C-P3 TMM 41106-2785, right ramus fragment with M4, alveoli of P3-M3 TMM 41106-2786, right ramus fragment with P3, alveoli of P4-M2 TMM 41106-2787, edentulous right ramus fragment with alveoli of M3-4, coronoid, and condyle TMM 41106-2788-95, eight left Mis TMM 41106-2796, right Mi fragment TMM 41106-2797-804, eight right M»s TMM 41106-2805, right M1 fragment TMM 41106-2806-10, five left M4S TMM 41106-2811-3, three right M4s TMM 41106-2814-6, three left M4s TMM 41106-2817-9, three left M*s TMM 41106-2820-7, eight left M2s or M3s TMM 41106-5098, 20 edentulous ramus fragments WAM 75.1. 1-.5, five left M*s WAM 75.1. 6-. 11, six right Mis WAM 75.1. 12-. 16, five left M2s or M3s PM 27400-425, 26 right Mas or Mss PM 27426-7, two right M2S or M3S PM 27428-452, 25 right M2s or M3s PM 27453-460, eight right M4S PM 27461, right M2 or M3 PM 27462-9, eight left M2s or M3s LUNDELIUS & TURNBULL: MADURA CAVE Trench 4, Unit 2, Level 2 (continued) PM 27470-3, four left M»s Pm 27474-8, five right Mis PM 27479-517, 39 left M2S or M»s PM 27518-96, 79 premolars uncatalogued, 46 incisors or canines Trench 4, Unit 2, Level 3 PM 27597-9, three left M»s PM 27600-1, two left M2s or M3s PM 27602-4, three right M2s or M3s PM 27605-8, four right Mis PM 27609-10, two left M2S or M3S PM 27611, left lower molar fragment PM 27612-3, two premolars PM 27614-21, eight right M2S or M3S uncatalogued, six edentulous maxillary fragments uncatalogued, 21 edentulous ramus fragments Trench 4, Units 4-5 TMM 41106-718, right M2 or 3 TMM 41106-728, trigonid of left Mx TMM 41106-3001-85, 85 right Mis TMM 41106-3086-171, 86 left Mis TMM 41106-3172-221, 50 left M4s TMM 41106-3222-72, 51 right Mas TMM 41106-3273-312, 40 left M2S or Mas TMM 41106-3313-41, 29 left M*s TMM 41106-3342-58, 17 left M4s TMM 41106-3359-90, 31 right M4s TMM 41106-3391-444, 54 right MJs TMM 41106-3445-65, 21 right M2s TMM 41106-3466-99, 34 right M3s WAM 75.1.17, left ramus with Mi WAM 75.1.18, right ramus with Ma-» WAM 75.1.19, left mandible, edentulous except for C WAM 75.1.20, edentulous left mandible WAM 75.1.21-.26, six left Mis WAM 75.1.27-.32, six left Mas WAM 75.1.33-.38, six right Mas WAM 75.1.39-.50, 12 left M2S or Mas (probably M2) WAM 75.1.51-.62, 12 left M2S or Mas (probably Ma) WAM 75.1.63-.68, six left M»s WAM 75.1.69-.72, four left M4s WAM 75.1.73-.77, five right M4s WAM 75.1.78-.82, five right M*s WAM 75.1. 83-. 92, 10 right M3s (measured) WAM 75.1.93-.99, seven right M2s WAM 75.1.100-.103, four right M2s or M3s WAM 75.1. 104-. 110, six right M2s WAM 75.1.111, right M2 or Ma 10 FIELDIANA: GEOLOGY Trench 4, Units 4-5 (continued) WAM 75.1. 112-. 114, three right M*s WAM 75.1.115-.117, two right P4s and a left P4 PM 13401-20A, 40 right M2S or Mas PM 26318-9, two left Mis PM 26330, left M2 PM 27622, right ramus with Ps PM 27623, left ramus with Px PM 27624, premolar PM 27625, trigonid, left Mx PM 27626, edentulous right ramus PM 27627, edentulous left ramus fragment, alveoli of M3-4 PM 27628-9, two edentulous right ramus fragments, alveoli of M3-4 PM 27630, edentulous right ramus fragment, alveoli of P2-M1 PM 27631, edentulous left ramus fragment, alveoli of P2-M1 PM 27632, edentulous right ramus fragment, alveoli of P2-M1 PM 27633, edentulous left ramus, alveoli of cheek teeth PM 27634, edentulous right maxillary, alveoli of M1"4 PM 27635, left M4 PM 27636, right M4 in ramus fragment PM 27637, left M4 in crypt, ramus angle, alveoli of M2-3 PM 27638, left P3 in ramus fragment PM 27639, edentulous right ramus PM 27640, left edentulous ramus, alveoli of M1-4 PM 27641 (A-G), seven edentulous right ramus fragments PM 27642 (A-H), -43 (A-F), and -44 (A-F), 20 edentulous right ramus frag- ments PM 27645 (A-E) and -46 (A-E), 10 edentulous left ramus fragments PM 27647 (A-E) and -48 (A-F), 11 edentulous left ramus fragments PM 27649-675, 27 right M2S or Mss PM 27676-89, 14 left M2S or M3s PM 27690, right M2 or 3 PM 27691-739 and PM 27741-52, 61 left M2S or M3s PM 27753-7, five right M2S or Mss PM 27758, left M2 or M3 PM 27759-85, 27 right M2S or Mss PM 27786-801, 16 right M4s PM 27802-22, 21 right Mis PM 27825-30, six left M4S PM 27831-46, 16 left M*s PM 27847-66, 20 left Mis PM 27867, right M4 PM 27868-92, 35 right M*s PM 27893-902, 10 premolars PM 27903-8, six right M2s or M3s PM 27909-15, seven left M4s PM 27916-28, 13 right M4S PM 27929-30, two left M4s PM 27946-68, 23 left M2S (measured) LUNDELIUS & TURNBULL: MADURA CAVE 11 Trench 4, Units 4-5 (continued) PM 27969-80, 12 left Mas or Mas (probably Mas) PM 33501-63, 63 left Mas or Mas (probably Mas) PM 33564-610, 47 left Mas or Mas (probably Mas) (33564-89 measured) PM 33611-24, 14 left P4s PM 33625-8, four left Mis PM 33629 (A-Z), PM 33630 (A-AA), and PM 33631 (A-M), 66 right M2s or M3s PM 33632-58, 27 left MJs PM 33659 (A-C), three broken left M»s PM 33663-92, 30 left M2s or M3s PM 33693-732, 40 left M2s PM 33733-6, four right P4s PM 33737-69, 33 left M2s PM 33770 (A-EE), 31 broken left M2s or M3s PM 33771-2, two left M*s PM 33773-872, 100 left M3s PM 33873 (A-Y), PM 33874 (A-XX), and PM 33875 (A-N), 89 indet. pre- molars (no P4s) PM 33876 (A-L), PM 33877 (A-N), and PM 33878 (A-W), 74 indet. pre- molars PM 33879 (A-FFFFF), 110 indet. premolars uncatalogued, —400 isolated premolars PM 33880 (A-AA), -81 (A-Z), and -82 (A-M), 56 broken right Mas or Mas PM 33883-7, five right Mis PM 33888-91, four right Mis or Mas, (probably Mis) PM 33892-4054, 163 right Mas or Mas PM 34055-6, two left M4S PM 34057-64, eight right P4s PM 34065-84, 20 right M2s or M3s PM 34085-107, 23 right M2s (measured) PM 34108 (A-Z), 26 broken left MxS PM 34109 (A-U) and PM 34110 (A-U), 42 broken Mxs PM 34111-26 (A-P), 16 right M3s (measured) PM 34127-37, 11 right M's PM 34138-59, 22 right M2s or M3s PM 34160 (A-ZZZ), 78 edentulous left ramus fragments PM 34161 (A-DDDD), 82 edentulous right ramus fragments PM 34162, five broken right MxS PM 34163, broken Mx PM 34165, right mandible with M4 in crypt PM 34166, right mandible with M4 PM 34167, right mandible with P3, Ma-3 PM 34168 (A-B), two broken left MxS PM 34169, left maxillary with M4, alveoli of P4-M3 PM 34170, premolar PM 34171 (A-B), fragments of two premolars PM 34173 (A-H), eight upper incisors 12 FIELDIANA: GEOLOGY Trench 4, Units 4-5 (continued) PM 34174 (A-C), three lis or hs PM 34175 (A-F), six left las PM 34176 (A-M), 13 right las Trench 4, Unit 7, Level 1 TMM 41106-707, right M2 or 3 WAM 75.1.119, right NU WAM 75.1.120-1, two left Ivhs or Mas WAM 75.1.122, right M2 or Ma PM 34177, right Ma PM 34179, left Mi PM 34180, edentulous right maxillary fragment with alveoli of P3 PM 34181, edentulous right ramus fragment with alveoli of M4 PM 34182 (A-B), two edentulous left ramus fragments PM 34183-94, 12 left M2S or Mas PM 34195, right Mi PM 34196-9, four left M*s PM 34200-3, four right M3s PM 34204-8, five right M2s PM 34209, right M2 or M3 PM 34210, right Ma PM 34211-2, two left M2s PM 34213-4, two right M*s PM 34215-8, four left M3s PM 34219-31, 13 right M2S or Mas Trench 4, Unit 7, Level ? PM 34232, right ramus fragment with M4 in crypt PM 34233, left P4 PM 34234, right ramus fragment with M3-4 PM 34235, edentulous right ramus PM 34236 (A-B), C-G, two left and five right edentulous ramus fragments Trench 4, Unit 7, Level 2 TMM 41106-730, broken left Ma WAM 75.1.123, left Mi PM 34237-9, three right Mis PM 34240-8, nine left Mis PM 34249, left ramus fragment with P4 in crypt PM 34250, (A-B, C-H), two right and six left edentulous ramus fragments PM 34253-4, two left M4S PM 34256-9, four left M*s PM 34260-4, five right M*s PM 34265-7, three right M3s PM 34268-9, two right M2s PM 34270-5, six left M2s PM 34276-8, three left M3s PM 34279-80, two left M2s or M3s PM 34281, left M4 PM 34282-5, four right M4S LUNDELIUS & TURNBULL: MADURA CAVE 13 Trench 4, Unit 7, Level 2 (continued) PM 34286-94, nine left Mas or Mas PM 34295, left Mi PM 34296 (A-C), three broken left Mas or Mas PM 34297-314, 18 right Mas or Mas PM 34315, broken right M2 or Ma PM 34316, right Mi Trench 5, Unit 4 PM 34317, right P4 PM 34318, right M1 PM 34319, right M2 PM 34320, right M3 PM 34321-2, two left MJs PM 34323-5, three left M2s PM 34326, left M3 PM 34327 (A-B), two edentulous right ramus fragments PM 34328 (A-E), five edentulous left ramus fragments PM 34329 (A-D), four edentulous left maxillary fragments PM 34330 (A-D), four edentulous right maxillary fragments PM 34331, left M4 Trench 5, Unit 5 TMM 41106-692-3, edentulous right and left rami, respectively PM 26332, broken right M1 or M2 PM 34332 (A-B), right and left maxillary fragments, each with P3 PM 34333, edentulous left maxillary fragment PM 34334 (A-C), three edentulous left ramus fragments PM 34335, edentulous right ramus fragment PM 34336, right cranial fragment PM 34337-9, three left Mis PM 34340-1, two right Mis PM 34342-8, seven right M2S or M3S PM 34349-54, six right Mas PM 34355, left M4 PM 34356, broken left M1- 2- or 3 PM 34357, broken right M2 or M3 PM 34358-64, seven left M2S or Mas PM 34365, left Ma or M2 PM 34366, left Pi? PM 34367-75, nine left P3s PM 34376-9, four left P4S or Pas PM 34380, left P4 PM 34381-3, three right P4S or Pas PM 34384-7, four left M2S or Mas PM 34388-9, two left MJs PM 34390-95, six left M3s or M2s PM 34396-7, three left M3s PM 34398, left M4 PM 34399-402, four right M*s PM 34403-6, four right M2s 14 FIELDIANA: GEOLOGY Trench 5, Unit 5 (continued) PM 34407-8, two right M2s or M3s PM 34409-11, three right M3s PM 34412, right M4 PM 34413-4, two right P4s Trench 5, Unit 6 PM 34415-6, two left M2S or M3S The following Recent material was used for comparison: Perameles myosura notina Thomas, 1922 (= P. bougainvillei notina), A. 377-823, A. 377-822, A. 377- 821 (9.2.181; fig. 2), A. 377-82 (9.2.182; fig. 3), from the Odontological Museum of the Royal College of Surgeons, London; TMM 40235-2 from Murraelellevan Cave (fig. 1). Descriptions Cranium. — One specimen (TMM 41106-143), a partial cranium consisting of sutured frontal bones with no unusual features, and a number of maxillaries have been recovered. The maxillaries show the infraorbital foramen opening into a broad depression above the anterior part of M1. Within the depression, there is a foramen for a canal that extends anteriorly into the maxillary. The lateral side of the maxillary immediately ahead of the junction of the jugal is excavated. The palatal part of the maxillary of TMM 41106-363 (fig. 4 A) shows part of both anterior and posterior palatal vacuities. The bridge between the vacuities is located opposite P3 and is a thin strut of bone. None of our speci- mens preserves the complete perimeter of either vacuity, and so we cannot comment on the anterior limit of the anterior or the posterior limit of the pos- terior, or the widths of either. It is clear that the posterior palatal vacuities are large as in specimens from the surface of Murraelellevan Cave. Large posterior palatal vacuities are a characteristic of the Perameles bougainvillei group, ac- cording to Freedman (1967, p. 163). Upper dentitions. — None of the recognized cranial fragments includes pre- maxillaries, and we have not attempted to characterize the isolated premaxillary fragments or upper incisors because of uncertainties of identification. The same applies to P1. Several specimens of maxillaries do preserve part of the premolar tooth series, and a number have alveoli for all cheek teeth, but noP's are in position. Therefore, we begin our description with P3. The P3 is a double-rooted tooth with a variable shape. One specimen (TMM 41106-363; fig. 4A) has a P3 that is compressed laterally. There is a prominent principal cusp located at midlength. Low but distinct cingular cuspules are present, one at each end of the tooth. The posterior end is slightly wider than the anterior and has an indistinct cingulum associated with the posterior root. It is separated from the P1 by a short diastema. Another specimen (TMM 41106- 508) has a wider P3, particularly posteriorly where a posterior cingulum encir- cles the posterior end of the tooth. A cingular tubercle is present at the pos- terolingual corner of the tooth, removed from the posterior cuspule. The di- astema separating P1 and P3 is shorter than in TMM 41106-363. The P4 is two rooted with a triangular outline in crown view. There is a large centrally located principal cusp that is oval in cross section and slightly recurved LUNDELIUS & TURNBULL: MADURA CAVE 15 Fig. 1. Perameles bougainvillei, TMM -40235-2, a subfossil specimen from the surface of Murraelellevan Cave, Nullarbor Plain, Western Australia. A & B, Left lateral and ventral views of the facial portion of the skull and its dentition. C & D, Lingual and crown views of the right ramus and its dentition. posteriorly. (lingular cusps are present on the anterior and posterior ends of the tooth. A prominent lingual cingular shelf with a cuspule is present. It varies considerably in size and position. In specimen TMM 41106-363, it is located lingual to the main cusp, whereas in TMM 41106-2778, it is smaller and located posterolingual to the main cusp. The anterior cingular cuspule is barely dis- cernible on the latter specimen. The upper molars have the basic tribosphenic form modified by the addition of a prominent hypocone (except for M4) and by crown hypsodonty on the lingual side related to the protocone and hypocone. Also, they have a prom- ^ o U >-> o OS 6 3 ■a c <3 .e S j S« a *1 .22 § c ~V 5 c 5 o t J Q, - . S 60 3 16 o T3 s ■a c D u 17 Fig. 3. Perameles myosura notina Thomas = P. bougainvillei. Specimen 9.2.182, a juvenile, Odontological Museum, Royal College of Surgeons, London. A & B, Left lateral and ventral views of the facial portion of the skull with its dentition. C & D, Lingual and crown views of the left ramus and its dentition. 18 LUNDELIUS & TURNBULL: MADURA CAVE 19 Fig. 4. Perameles bougainvillei from Madura Cave. A, TMM 41106-363, a right maxillary with P3-M1 shown in lateral and crown views. Specimen is from Trench 3, Unit 2, Level 1 . B, PM 13340, a right maxillary with M2"3 heavily worn, shown in lateral and crown views. Specimen is from Trench 3, Unit 2, Level 1. C, TMM 41106-512, a right maxillary fragment with all molars, shown in the same standard views. Specimen is from Trench 4, Unit 1, Level 1. inent stylar shelf with cusps that connect to the paracone (except M1) and metacone (except M4) by crests so that a characteristically pinched, double V- shaped ectoloph results. They are all three-rooted teeth. Many of these crown features may be eroded by wear, and it is a common occurrence to find speci- mens in which all crown features are gone (fig. 4B). The M1 is approximately square in occlusal outline, with the parastyle and metastyle projecting anteriorly and posteriorly, respectively, along the labial edge. The metacone is the most prominent cusp, but it is not nearly as massive as the less prominent protocone. It is joined to the metastyle and to stylar cusp 1 (ectostyle, see Hershkovitz, 1971) to form a prominent triangle. The paracone is usually joined only to cusp k and consequently it does not usually form the apex of a triangle. Rarely does M1 have a weak crest to the parastyle. The protocone 20 FIELDIANA: GEOLOGY and the hypocone are crescentic, and the protocone is the much larger of the two. The posterior crest of the hypocone extends around onto the posterior face of the metacone and rises towards the roots. A deep ectoflexus notch separates stylar cusps k and 1. The M2 and M3 are similar and will be described together. They both take the form of a truncated triangle in occlusal outline (fig. 4C), with the lingual side slightly shorter than the labial, although M3 tends to be somewhat more trian- gular, with its lingually bulging protocone and more reduced hypocone. The metacone is the most prominent cusp, and its relationship to the stylar cusps is the same as with M1. The paracone joins stylar cusp k and the parastyle by its crests. Protocone and hypocone are crescentic, the latter being much reduced in M3. Other differences between M2 and M3 are related to the positioning of the stylar cusps; the parastyle and metastyle being located farther labially with respect to cusps k and 1 in M3 than in M2 where all stylar cusps are in a straight line. The M4 is reduced, especially posteriorly (fig. 4C). It is a triangular tooth, the longest edge being the transversely oriented anterior one. The paracone, which is the largest cusp, is located near the lingual edge of the tooth and is connected to the parastyle by a long paracrista (I'). A short postparacrista — premetacrista (I'" a & b) or centrocrista (I'") — extends posteriorly from the paracone and turns abruptly linguad at the posterior end of the tooth. The labial edge of the stylar area is almost straight from the parastyle to the posterior end of the tooth where it too turns abruptly linguad and makes a hooklike projection. This posterior cusp at the junction of the back of the stylar shelf and the centrocrista (I'") could be a fused metacone-metastyle, but its homology is quite uncertain. Several small tubercles are present on the stylar shelf. The protocone is small and cres- centic; its crests encircle the lingual part of the base of the paraconid. Mandible. — The horizontal ramus of the mandible is slender (figs. 5-7). The ventral margin is convex beneath the molars and the P4 and concave from there to the anterior end.1 The dorsal edge of the horizontal ramus is twisted to the outside anteriorly. The symphysis is long, extending from the region of P1-P3 to the anterior end of the ramus. The symphysial area is concave, rugose, and has one prominent foramen. The lingual surface has a broad, shallow groove in the region underlying the molars. The major mental foramen is located under the P2. Other small foramina are variably located both anterior and posterior to the large one. The ascending ramus rises at a low angle to the tooth row (less than 45°). Its anterior margin has a complex outline. Only the relatively short coronoid pro- cess rises steeply (at about 80°), and its tip is hooked posteriorly. The lower half of the ascending ramus is convex, and its upper half is concave. The masseteric fossa is narrow and has essentially the same width throughout its length. The posteroventral margin of the masseteric fossa is marked by a distinct ridge that parallels the anterodorsal margin. The ventral margin is broadened laterally to form a short shelf with a tubercle above the base of the angular process. The angular process tapers rapidly and evenly to a point. Its inner surface is con- cave. The mandibular foramen lies in a broad depression above the anterior part of 1In one specimen, TMM 41106-177 (fig. 6A, B), there is a prominent, pointed projection beneath P3, which is an abnormality that probably resulted from an injury. LUNDELIUS & TURNBULL: MADURA CAVE 21 >-^aA0L. Fig. 5. Perameles bougainvillei from Madura Cave. A & B, TMM 41106-2771, a left mandi- ble with M3.4, alveoli of the rest of the dentition shown in lingual and crown views. Trench 4, Unit 2, Level 4. C & D, PM 27238, a right mandible with P„, M3.4, and alveoli of the other teeth, shown in lingual and crown views. Trench 4, Unit 2, Level 1. the base of the angular process. The medial surface of the ascending ramus is smooth and slightly convex. The condyle is high with respect to the tooth row and is close to the coronoid process. Its articular surface is oval in outline but is slightly concave trans- versely and convex anteroposteriorly. There are diastemata between b and C, between C and Pi, and a very short one between Pi and P3. That between h and C is usually ridged (sometimes double ridged) and sharp, and the ridge is turned labially. That between the C and Pi is usually double ridged, but occasionally is only single ridged. The double ridges tie to the labial and lingual borders of the alveolus for the canine. Lower dentitions. — No Madura Cave specimen has an incisor in place. There are a great many isolated incisor teeth, but we cannot be certain about their taxonomic assignments. A number of bs that in the relatively unworn state are strikingly mitten-shaped most probably belong to this taxon because they com- pare closely with a sub-Recent specimen from Murraelellevan Cave (see figs.lC, D; 2C, D; 3C, D). The alveolar pattern within the constricted anterior end of the 73 <1> Ua i^ 1 s < C H 5 -fi "3 "* &0 > ^ « J 2^2 ii CN -2 ON C ^^D £Q | I* J ii u ^ • ui c 0) (U <£ - G q3 ~ o a §•3 J •5 2 9 «fi Z* «o«^ s ^ c o a ■ !? 01 «T 4* -S* ij *^ v 5 °^ « 01 C-, »j •£ v* a,? 3 vb °~ • X &H H fll i fc §12 "3 S 22 ■B G | i I o> 3 « w CO* =8 3* si > .2 U to 3 c 6~ Jl - 2 fi < 3 _ IP* c -£ J* c a. o 2* 23 24 FIELDIANA: GEOLOGY horizontal ramus shows a pinched condition of the roots whereby h is con- sistently squeezed up into a higher position than Ii, which is always tight against the symphysis, and h, which is low and farthest laterally for most of the length of its root. The lower canine is relatively short anteroposteriorly, low, and very com- pressed laterally. The anterior edge is rounded, and there is no anterior cingular cuspule. The posterior edge is narrow but not sharp in the unworn state and terminates ventrally in a posterior cingular tubercle. Wear occurs at the top and dorso-posterolingual side of the tooth and results in shifting the posterior ridge labially. No sexual dimorphism is apparent, either in the few specimens with canine teeth in place or in the many jaws with empty canine alveoli. Mea- surements of maximum anterior-posterior length at crown-root junction (nor- mal to ramus) for the only specimens with canines in place and for the as- sociated alveoli are: Crown Alveolar length length Specimens (mm) (mm) WAM 75.1.19 2.43 3.65 TMM 41106-172 2.60 3.65 TMM 41106-177 2.43 3.50 The two anterior lower premolars, here designated Pi and P3, are much alike. Each is double rooted, compressed laterally, and has its principal cusp located in the anterior half of the tooth and turned slightly lingually at its apex. They have anterior and posterior cingular cuspules. The anterior edge of the principal cusp of each is rounded, but the posterior edge has a thin sharp ridge in the unworn state. The posterior ridge that joins the posterior cuspule is rapidly modified by wear, most heavily on the lingual side of the posterior edge of the principal cusp. Both teeth have a faint posterior cingulum. The P3 is the longest premolar. The P4 is approximately the same length as the P3 but is less compressed laterally. Its principal cusp, which is located anterior to the mid-length, is relatively higher and wider than that of Pi or P3. Its lingual surface is flat, but the labial surface is strongly convex. Like the Pi and P3, the anterior edge of the principal cusp of P4 is rounded, and the posterior edge is ridged. There are anterior and posterior cingular cuspules. The posterior one is joined by the ridge on the posterior edge of the principal cusp. A strong posterior cingulum extends over the posterior root on both sides. Wear rapidly removes the ridge on the posterior edge of the principal cusp, and in late wear stages the cusp is reduced to the level of those of Pi and P3. All of the lower molars have a slightly modified tribosphenic pattern, and with the exception of the Mi are rather rectangular in crown view. All have tooth base hypsodonty (White, 1959) but to a lesser extent than in the other peramelids in the Madura Cave assemblage (Isoodon, Chaeropus, and Macrotis). It is strongest under the labial cusps. The paracristid and epicristid in all molars lack a carnassial notch. The trigonid of the Mi is distinctly narrower than the talonid, with a pro- toconid that has a more rounded, less protruding lingual outline. The other trigonid cusps are smaller and more tightly spaced than in the more posterior molars. The protoconid and metaconid of Mi are the same size, but the pro- LUNDELIUS & TURNBULL: MADURA CAVE 25 toconid is slightly higher in an unworn tooth. The paraconid is smaller and lower than the others and shows no tendency to be laterally compressed as is the case in many dasyurids. The hypoconid and entoconid are the main cusps of the talonid. They are approximately the same height but are differently shaped. The entoconid is circular in cross section when unworn and is not connected to any other cusp. It has an upstanding, high conical shape initially, and wear on its labial surface alters the cross section to an oval shape. The hypoconid is crescentic and is connected to the posterolingual part of the base of the pro- toconid and to the hypoconulid. The hypoconulid is low and projects backward. There is no posterior cingulum and usually only a trace of an anterior one. Sometimes the anterior cingulum is distinct but small, sometimes absent. A small cingulum occupies the notch between protoconoid and hypoconid near the base of the former. The M2 and M3 are similar in size and morphology. Both have a trigonid that is wider than long, as a result of the far lingual placement of the protoconid and the closely spaced paraconid and metaconid. The protoconid and metaconid are equal in size and height; the paraconid is smaller. The talonid of both M2 and Mb is like that of Mi in size and arrangement of the hypoconid and entoconid. The hypoconulid of M2 is smaller than that of the Mi, and that of M3 is virtually gone. In M2, the posterior crest of the hypoconid joins the hypoconulid as in the Mi. In the M3, this crest either joins the vestige of the hypoconulid or it dis- appears at the posterior side of the entoconid. Both teeth lack a posterior cin- gulum, and each has the small labial one between protoconid and hypoconid. The M2 and M3 differ from one another and from Mi and M4 in the size of the anterior cingulum. It is weakest on Mi and becomes progressively larger toward M4. The trigonid of the M2 is narrower than that of the M3. This character, which is easily observed in specimens with the teeth in jaws, is of little use in distinguishing isolated M2S and M3S because the variations in absolute size and extent of wear are great enough to cause broad overlap of measurements. The M4 resembles the M2 and M3 in the arrangement of its cusps, but it is narrower, and the relative sizes are different. The protoconid is the largest cusp, and the paraconid and metaconid are equal in size. The talonid is reduced in comparison with the other molars. A distinct basin is bordered by a tall conical entoconid and a crescentic hypoconid, the crests of which go to the anterior and posterior sides of the base of the entoconid. No hypoconulid is present. As a result of the failure of the cristid obliqua to join the epicristid, the talonid basin is joined to the trigonid only by the hypoflexid at the lingual edge of the tooth. The area labial to the hypoflexid slopes downward, labially into another basin that is bordered labially by the cingulum that joins the protoconid and hypoconid. Fossil bandicoots lose their teeth more readily than most other mammals. Hence, a high proportion of isolated teeth is encountered. The more distinctive first and last molars cause no problems of identity, but M2s and M3s are so similar to one another (as are M2S and M3s) that their separation is far less certain when they are recovered as isolates. The M2s can be distinguished from M3s most of the time by the more nearly equal-sized protocone and hypocone; also the M3 has a smaller hypocone than the M2. The stylar cusps of the M2s tend to be arranged in a straight line, whereas in the M3s the parastyle and metastyle tend to be located labial to stylar cusps k and 1. Standard measurements fail to separate the M2s and M3s (fig. 8). LENGTH B /• _ / • ^-r — ' ° / % V\ % • o o • / \* / • * ° % • \ ° \ • % • o • • • 0 •/ O */f o \ ° o o — .• • ^^^ L E NGTH Fig. 8. Bivariate graphs of the cheek teeth of Perameles bougainvillei. A & B, Plots of length vs. anterior width and length vs. posterior width, showing extensive overlap of M2s and M3s. Initial determination based on visual criteria. All are from Trench 4, Units 4-5. Dots indicate M2s, circles M3s. Measurements are in millimeters. 26 D ) M3 P4 —^3 Fig. 8. C & D, Length vs. width plots of tooth measures for identified cheek teeth from Madura Cave. Means for each adjacent tooth are connected to form a pattern. In C, the upper cheek teeth are shown, in D, the lowers. For the premolars, the ordinate measure is the greatest width perpendicular to tooth length; for the molars, it is anterior width. 27 28 FIELDIANA: GEOLOGY In crown view, the M2S differ from the Nfos in having talonids that are rela- tively wider than the trigonids. Scatter diagrams of length versus anterior width and length versus posterior width show no separation (fig. 9B, C), but scatter diagrams of anterior width versus posterior width and length versus AW/PW confirm the visual observation, although the clouds of points almost touch (fig. 9 A, D). Discussion Perameles bougainvillei (following Tate, 1948, and Wakefield, 1966, including P. myosura, P. fasciata, P. notina, and P. eretniana) is widely distributed through the drier parts of southern and western Australia (Shortridge, 1909; Jones, 1924; Troughton, 1962; Marlow, 1962; Freedman, 1967; Ride, 1970; Brooker, 1977). Abundant remains of P. bougainvillei that are found in surface deposits of caves A — 7^^" — * 7" / " " .* Z° ^N- / • \ Vf o % % i I — — *7^ / .15 M3L AW PW 1.726 .164 .737 28 29 28 P < .05* .40 < P < .45 .20 < P < .25 Significant at .05 level. *Means identical. Table 4. Results of "t" tests on means of some mandibular and dental measurements of Perameles bougainvillei from Victoria Cave, South Australia, and Fromm's Landing, South Australia (based on data from Smith, 1972, table 5). Degrees of Probability Measurement "t" freedom (one-sided test) Length M1"4 2.497 6 P < .025* Length ascending ramus 1.883 32 .05 > P > .025* Mandible breadth at M2 1.263 41 .15 > P > .10 Mandible height at M2 0.741 38 .25 > P > .20 P4L 0.768 11 .25 > P > .20 Mi L 0.067 5 P > .9 PW 2.473 6 .05 > P > .025* M2L 3.875 11 .005 > P > .0005* PW 1.420 10 .1 > P > .05 LUNDELIUS & TURNBULL: MADURA CAVE 31 Table 4. Continued Measurement //j// Degrees of freedom Probability (one-sided test) MsL 0.559 14 .30 > P > .25 PW 0.737 14 .25 > P > .20 M4L 1.364 15 .1 > P > .05 PW 0.131 15 .45 > P > ,40 'Significant at .05 level. Both of these situations indicate a post-Pleistocene reduction in size in Perameles bougainvillei , although the magnitude of the change was less than that reported for larger animals (Marshall & Corruccini, 1978). Isoodon Desmarest, 1817 Isoodon obesulus (Shaw and Nodder, 1797) Material Trench 2, 2Vi ft PM 25224, left ramus with Ma and alveoli of P4-M2 and M4 (fig. 11G) Trench 4, Unit 1, top 1 ft TMM 41106-568, fragment of upper molar TMM 41106-669, right M3 TMM 41106-672, left M2 (fig. 11B) TMM 41106-673, left M3 (fig. 11D) TMM 41106-675, left M2 TMM 41106-677, right M2 or 3 WAM 75.1.124, left maxillary fragment with M3 PM 26331, left ramus fragment with M4, juvenile PM 26362, very worn lower molar, probably Mi Trench 4, Unit 2, Level 1 TMM 41106-708, right M3 or 2 Trench 4, Unit 2, Level 2 TMM 41106-705, left M1 (fig. 11A) TMM 41106-706, left M2 or 3 TMM 41106-710, left M2 TMM-41 106-711, right M3 TMM-41106-712, left M2 or 3 PM 26307, left Mi (fig. 11F) PM 26334, left ramus with M4 and condyle (fig. 11H) Trench 4, Units 4-5 TMM 41106-709, right M2 TMM 41106-714, right M2 or 3 TMM 41106-715-6, two right M's TMM 41106-717, 19-20, three right M3s TMM 41106-721, right M3 (fig. 11C) TMM 41106-722, left M3 TMM 41106-723, left M2 32 FIELDIANA: GEOLOGY Trench 4, Units 4-5 (continued) TMM 41106-724, left M2 or 3 TMM 41106-725, left M2 TMM 41106-726, left M2 or 3 TMM 41106-727, left M2 WAM 75.1.125, left M2 or 3 PM 26308, left M2 or 3 PM 26309, talonid of a left lower molar PM 26310, right M2 or 3 PM 26311, left M2 or 3 PM 26312, right M4 PM 26313, left M4 (fig. Ill) PM 26314, left M2 PM 26315, fragment of left M2 or 3 PM 26316, right M1 PM 26317, left M2 or 3 PM 26320-3, four right Mis PM 26324, left M2 or 3 PM 26325, left M3 PM 26326, right M2 PM 26327, left M4 PM 26328, left lower molar PM 26329, left M2 PM 26962, left M4 (fig. HE) PM 27740, left M2 or 3 PM 30552, edentulous right ramus PM 33660-2, three left MJs Trench 5, Unit 5 WAM 75.1.126-7, two broken right lower molars Descriptions Upper molars. — The upper molars have all the characters seen in extant speci- mens of Isoodon obesulus (fig. 10A-C). The first through the third molars are nearly square in occlusal view with moderate tooth base hypsodonty (White, 1959), especially in that part of the tooth that supports the protocone and hypocone. As with most peramelids, the labial part of the tooth is offset from the lingual part so that it extends farther ventrally (or where seen in palatal view appears to be raised above the lingual side). The M1 (fig. HA, TMM 41106-705) is slightly rectangular in occlusal view. The anterior end is somewhat narrower than the posterior end. The wear facets indicate that the protocone is slightly taller than the hypocone in an unworn tooth, although wear rapidly brings them to the same level. They are joined at a higher level than the central basin. The protocone merges with an anterior lingual cingulum that extends across the anterior face of the tooth and nearly joins a smaller cingulum from the parastyle. A similar but larger postcingular ridge extends from the hypocone across the posterior face of the tooth nearly to the posterolabial corner. LUNDELIUS & TURNBULL: MADURA CAVE 33 The paracone is joined to the weak stylocone by a curved ridge that extends on to cusp k. The ridge joining the paracone and parastyle is sometimes faintly visible on the anterior face of the paracone (TMM 41106-705, -716, and PM 26316). The parastyle is much smaller than in the posterior molars. The metacone is joined to the anterolingual base of the mesostyle and to the meta- style to form a triangle as in M2 and M3. A smaller stylar cusp is present anterior to the metastyle in TMM 41106-705, -715, but is absent in -716 and PM 26316 (probably as a result of wear in the latter). The metastyle is located farther posterolabially than in most specimens of M2 and M3. In all of these features, the Madura Cave specimens conform to the comparative specimens available to us [TMM M-851, FMNH 98899 (fig. 10A, B), both Recent specimens; and PM 25506, -7, and -9, all Pleistocene to post-Pleistocene specimens from caves in the Margaret River area of Western Australia]. The M2 and M3 of J. obesulus are so alike that separation of isolated teeth is uncertain. They either lack a stylocone or have only a weakly developed one. They are square in contrast to those of Perameles, in which the labial edge is longer than the lingual edge (fig. 11B, C; TMM 41106-672, an M2; and -721, an M3). The protocone and hypocone are crescentic when unworn and are joined above the level of the central basin. The protocone is slightly higher than the hypocone. In some specimens (TMM 41106-673, -709, -710, and -721), a low, indistinct ridge extends from the junction of the protocone and hypocone into the central basin where it disappears. This area of the tooth is flat in the other Madura fossils and in our comparative specimens. A ridge extends anteriorly from the protocone for a variable distance around the paracone. It disappears on the anterior face of the tooth near the base of the parastyle in TMM 41106-669 and -672. In TMM 41106-673 (fig. 11D) and -719, it joins the parastyle. A similar ridge usually extends posteriorly from the hypocone around the metacone almost to the posterolabial corner of the tooth. In TMM 41106-669, -712, and -720, this ridge disappears about halfway across the posterior face of the tooth. The labial two-thirds of the occlusal surface is occupied by two triangles whose apices are the paracone and metacone. The paracone is not as tall as the metacone and is joined to the apex of the parastyle and the base of cusp k by the eocrista (I'y and I'"a, respectively). Cusp k has a crest that extends toward crest I'y in TMM 41106-673. It meets and joins a very weak crest from I'y in the intervening valley. The metacone is joined to the base of the mesostyle (the cusp lettered 1) by the crista (I'"b) and to the apex of the metastyle by a crista (I"). A small stylar cusp (m) is present anterior to the metastyle. Cusp k and the mesostyle (1) are approximately the same size, are larger than the parastyle and metastyle, and are weakly connected at their base. Cusp k is oval in cross section, and the mesostyle (1) is circular. In an attempt to find a way of distinguishing M2s from M3s, and M2S from M3s, a number of bivariate graphs were drawn, based upon the usual dental measures, none of which gave plots without some degree of overlap of the clouds of points. Figure 12 shows two of these in which length vs. anterior width are plotted for four small samples of Recent and fossil specimens of /. obesulus. The most successful plot is shown in (A) wherein M2s and M3s can be separated about three-fourths of the time. The sort of plot shown in (B) for the same measures for the lower teeth is more typical, however, and illustrates the 01 1> i „ s 19 h h ■2 £"2 .SP fc 6 £ ■£ — ■ » T3 I 4 '55 o < £ aj I'll •^ c 1 £ » J PX " ■» >.J 00 £ 1 CTv § I PC f i fc:3c o% -s s I 5 « o 5 81 « a> -« sx c I5 Jl I a5 I i c ■ 2 S £ e J 5S r1 o 5 in I o -a 3 rH 6 qj 34 > (A 8 1 I 3 W Q 35 OB. B 36 LUNDELIUS & TURNBULL: MADURA CAVE 37 need to find some discrete features or other kinds of dental measures than these standard ones if positive separations are to be achieved. Similar results were obtained with the Madura Cave teeth (fig. 13). The M4 differs from the other upper molars in the extreme reduction of the posterior cusps (fig. HE; PM 26962). It is ovoid in occlusal view. There are three principal cusps — protocone, paracone, and cusp k. The paracone and protocone are crescentic cusps with prominent anterior and posterior ridges. Those of the paracone both reach the labial edge of the tooth. The anterior one joins a small, indistinct stylocone-parastyle at the labial edge of the tooth. The posterior ridge of the protocone reaches the labial edge of the tooth, but the anterior one does not. Cusp k is a conical cusp that is situated on the labial edge of the tooth between the ends of the ridges of the paracone. There is no trace of a posterior cusp, unlike the situation in most of the Recent specimens we have examined, which have a small posterolabial cusp. Mandible. — Three partial mandibles oilsoodon are present in the Madura Cave collection. All represent the posterior part of the mandible. The horizontal ramus is slender and relatively shallower than that of dasyurids of comparable size. It is deeper and wider than that of Perameles. The angular process is directed inward and posteriorly, is broad basally, and tapers rapidly to a slen- der termination. The mandibular foramen is located above the anterior part of the base of the angular process. The ascending ramus is broad. Its anterior border rises steeply, forming an angle of more than 45° with the line of the tooth row (fig. 11G, PM 25224). In this character, Isoodon resembles the dasyurines, Chaeropus and Macrotis, and dif- fers from Perameles in which this angle is less than 45°, and the masseteric fossa is narrow. The ventral margin of the masseteric fossa is marked by a prominent shelf that is expanded laterally over the base of the angular process. The condyle is located closer to the coronoid process than to the angular process. The articular surface of the condyle is broadly oval in outline (fig. 11H; PM 26334). It has no curvature transversely but is gently convex antero- posteriorly. Lower molars. — All lower molars have well-developed trigonids but lack car- nassial notches on the major crests. They have tooth base hypsodonty especially beneath the hypoconid. The trigonid is higher than the talonid on all molars. The Mi is an elongate tooth that tapers anteriorly (fig. 11F; PM 26307). The trigonid is distinct but relatively smaller than in M2-M2-4. The protoconid and metaconid are subequal, and the paraconid is distinctly smaller and lower. There is no anterior cingulum, but there is a slight concavity (on the anterolat- eral side of the tooth) between the protoconid and paraconid. Opposite: Fig. 11. Isoodon obesulus from Madura Cave. A, Crown and labial views of TMM 41106- 705, a left M1. B, Labial, crown, distal (posterior), and lingual views of TMM 41106-672, a left M2. C, Proximal (anterior), labial, crown, and lingual (inverted) views of TMM 41106- 721, a right M3. D, Crown and labial views of TMM 41106-673, a right M3. E, Distal, lingual, proximal, and crown views of PM 26962, a left M4. F, Crown and labial views of PM 26307, a left M, . G, Lingual and dorsal views of PM 25224, a left ramus with M3, and alveoli of P4-M2 and M4. H, Labial and dorsal views of PM 26334, a left ramus fragment with M». I, Crown, labial, and lingual views of PM 26313, a left M4. 2.50 AT^ 1 ° \ zJ / //b a±// m\ e y ^AA A- £± / 1.50 L E N e T I RECENT TASMANIA RECENT SW. AUSTRALIA FOSSIL SW. AUSTRALIA HASTINGS CAVE M?- M% M% M$ O /S. Fig. 12. Bivariate graphs of length vs. anterior width measurements of upper and lower molars of Recent and fossil specimens of Isoodon obesulus. In A, the bivariate plots for the upper molars show a clear separation for MJs and M4s, but only a partial separation between M2s and M3s. In B, similar plots for the lower molars show extensive overlap between adjacent teeth. Straight lines connect the midpoints of the plots for adjacent teeth. Mea- surements are in millimeters. 38 LUNDELIUS & TURNBULL: MADURA CAVE 39 The talonid is broader than the trigonid. The entoconid is the highest cusp. It is subcortical in shape and occupies almost all of the lingual edge of the talonid. The hypoconid is slightly lower than the entoconid but is more massive because of the underlying tooth base hypsodonty. The hypoconulid is the smallest of all the cusps and the most subject to erasure by wear. In an unworn or little-worn tooth, it projects posteriorly behind the entoconid in line with all the other lingual cusps. The hypoconid is tapered toward its apex so that in an unworn or little-worn tooth its crests make a tight V. With wear, this V becomes more open. The cristid obliqua (I'"b) connects the hypoconid to the midpoint of the base of the epicristid (II) of the trigonid. The postmetacristid (I") connects the hypoconid to the hypoconulid. In an unworn tooth, this ridge has a slight posterior curvature but becomes straight with wear. There is a small cingulum in the reentrant between the protoconid and hypoconid. It may bear a small cuspule. In an unworn specimen, this cingulum is located halfway between the top of the cristid obliqua and the enamel line between the roots. The M2 and M3 are so similar in both size and morphology (fig. 11G) that they cannot be reliably separated and are therefore treated together. Both are similar to the Mi in general morphology. They can be distinguished from the Mi by the presence of a prominent bulging anterolabial cingulum, a more massive trigonid, and larger size. The most consistent difference between M2 and M3 seems to be the degree of development of the hypoconulid: larger on M2 than M3. Since this feature is reduced by wear, it is best seen in unworn specimens (PM 26325, 26329). Recent specimens with all teeth in place show a gradual decrease in development of the hypoconulid from Mi through M3. The lingual cusps of the M2 and the M3 do not fall on a straight line as they do in Perameles, but form a curved line, with the paraconid and hypoconulid located labial to the metaconid and entoconid. The M4 differs from the M2-3 primarily on the structure of the talonid and its slightly greater length. The trigonid of the M4 has the same construction as that of the M2-3 but is slightly more massive (fig. 11H, I; PM 26334 and PM 26313). The talonid is elongate but is otherwise reduced. The hypoconid and entoconid are the two principal cusps. The entoconid is higher, but the hypoconid is more massive and crescentic. The connection to the trigonid is variable, ranging from no ridge through a broad, rounded ridge, to a weakly developed, but sharp, ridge. The hypoconulid may be present or absent. Where present, it is small and below the notch between the hypoconid and entoconid. The cingulum between the protoconid and hypoconid is larger than on the other molars. It has a variable number of cuspules and isolates a small basin. Comparisons. — A comparison of the Madura Cave specimens with Recent and fossil specimens from southwestern Australia shows them to be similar in mor- phology but different in size (table 5). The means of the measures of the Recent southwestern specimens are larger than those of the Madura Cave specimens, but there is overlap in many measures. A comparison of the Madura Cave material with specimens from Hasting's Cave, Western Australia (TMM 40236-3, -7, -8; PM 7272-81, PM 25510-15) shows that the remains from these two localities are alike both in size and morphology. The smaller size of Isoodon from the drier areas of Australia has been noted in other studies. Wakefield (1964, pp. 494-498) reported that Isoodon from B 3.00 LENGTH Fig. 13. lsoodon obesulus from Madura Cave. In A & B, bivariate plots of molar tooth measures of length vs. anterior width and length vs. posterior width are shown. The solid symbols designate teeth whose positions are known with certainty; the x symbols indicate isolated M2s and M3s, which we cannot distinguish from one another with certainty. 40 2.50 3.00 3.50 4.00 LENGTH D 2.50 2.50 X=ISOLATED M2ORM3 X /^/ x A. / M1 ^ x /. / \o * * \ y' X \ ♦ \ ♦ ♦ \ \ ■ ■) (■ ■ , 2.50 3.00 3.50 4.00 LENGTH Fig. 13. In C & D, similar plots for the lower molars show extensive overlap of M 2s and M3s. Isolated teeth indicated by x. Straight lines connect the midpoints of the plots for adjacent teeth. 41 i-S S^S in O r-i co cm cm co co co co co ro cm cm co i-i cm co cm cm I I I CO tN CM WIN h > J; u s ~ S at 3 01 ■** 9 vO If) O n vO 1-1 CJ M M | | 1 ANN ■>* CM 1 ro ON in cn in 1 nan ' in CM (N 1-1 r-i t-h Nno in on in 1— 1 rt rtN CO CM CM CTl r-i n t-* rv h- in in in 6^ 3 3 •a o < & 01 Of) *5 « Ol (0 § A ll 2 as O 01 0 u. > M C\ ■* 'l' co rM rM ro cm ro ro ro co CM CM ro cj cm ro ro CO ro ro ro CM no rsl in GO CM 00 o M3 O mo cm is. cm in cm no CO ri CM CO CM CM co cm cm co cm cm ro ro cm Z cMCMcMcocococororo I I I CO CM tN ^ fN i-i I I CM CN CM n n J- 3 3 O U5 < c E 01 u (/I 01 01 cs s Jg COCMCMCOCNicMCOCOCOtNCMCOricMCOCMCNI S» ONcotv-TfooNvorxcooovoorscooo^o r 01 NvOOO rtlNM OlD* \OJ> N0\0 N'fri C &p co cm cm co coco co co ro cm cm co i cm cm ro cm ro u a nOi-iOnOOncOnOOOnOcOt-i'3,nOcOi-iOn JS & ^N* rjinvfi (J;NM (NN ONrH inNin O CMCMrMCOCMCMCMCOcOrMrNCOi-HCMCOCMCM Z cOCOCOCOCOCOCMCMCMCMCMCNCMCMCMfMCM I CO CM CM ^ CM CM CO CM CM ■<* CN CM CO CM CM CO CM CM CM CM CM CM CM CO CM CM CO CM CM On -^ CM CM CM rl CM 13 inoNoooNNonvooNO §fo> "-J""*. NOCMONOOrlO U OC rO rM rM CO CM CO CO CO CO « E ricM-^r^ONOOCOCOON o 00 "*J NO CO CM CM CM csi CN vc i no in CM CM CM i-i COCMCM CO CM CM COCMCM COCM ro CM CM ro CM CM ro CM T— O CM CM a CN CM CM NO CM o CM GO 00 >* COCMCM COCMCM COrli-l Tt^n* oooooo ononon «h rH minin HHH ononon cmcmcm ininin •c > 2 < e- J < Pi hJ < Cu hJ < d hJ < C-H 1^ _1 < &H hJ < 6. 42 LUNDELIUS & TURNBULL: MADURA CAVE 43 Holocene deposits in the Fromm's Landing shelters are smaller than the modem ones from Western Victoria, but he gave no measurements. Small Isoodon from the arid zone of Australia were referred to /. auratus by Finlayson (1961). Mar- shall (1973) found no overlap in size in 13 dental measures of two samples of these small-sized Isoodon from central and Western Australia and a sample of /. obesulus from Victoria. His single specimen from a Holocene deposit at Lake Victoria falls within the range of the central Australian sample and accordingly was assigned to 1. auratus. The samples from Madura Cave and Hastings Cave are intermediate in size between those of /. auratus and I. obesulus, but are closer to the /. obesulus samples from southwestern Australia and Victoria (table 5). The significance of these size differences is not clear. Tate (1948) considered /. auratus to be a subspecies of/, obesulus. Marshall (1973) followed Finlayson (1961) and consid- ered /. auratus to be a separate species. A revision of the genus is needed and is rendered more difficult in the face of the new evidence of the Madura and Hastings Caves samples. The Madura Cave material is referred to /. obesulus on the basis of the closer resemblance to modern samples of that taxon. Discussion Isoodon obesulus is found today in southwestern and southeastern Australia, Nuyts Archipelago, Tasmania, and Cape York Peninsula. Isoodon macrourus is found along the east coast of Australia, Arnhem Land, and north Kimberley. Isoodon auratus is found in northwestern Australia, central Australia, and Bar- row Island (Ride, 1970). The record of /. obesulus in the deposits of Madura Cave is close to halfway between its westernmost record in South Australia (Kan- garoo Island) and its easternmost record in Western Australia. Its presence in units 1, 2, and 4-5 demonstrates that it was an element of the faunas of terminal Pleistocene and early Holocene times on the Nullarbor Plain and that its dis- appearance from the area is relatively recent. Its absence from unit 7 may be an accident of sampling because the total amount of material recovered from that unit is small. We have been unable to find a definite record of its occurrence on the Nullarbor Plain within historic times. It is found today in situations that are considerably more mesic than exist now on the Nullarbor Plain, but it is also found in ones with xeric conditions similar to those of the Nullarbor (and with similarly erratic rainfall). Merrilees (1967) has pointed out that the broad distribution of Isoodon with respect to rainfall in Western Australia throws doubt on its use as an indicator of climatic change. Heinsohn (1966) in a study of Perameles and Isoodon in Tasmania found that Isoodon kept to brushy areas even when feeding, whereas Perameles fed in more open situations. This suggests that the disappearance of Isoodon from the Nullarbor Plain in the last 7,000 years was caused by the disappearance of the appropriate vegetation, not by increasing aridity as such, although the change in plant cover is almost certainly a result of increasing aridity. Palynological studies at Madura Cave indicate a decrease in the mallee cover from about 5,000 to 6,000 years b.p. to present (Martin, 1973). This change is attributed to a combination of increased aboriginal fire pressure and a decrease in rainfall. The climatic interpretations based on the disappearance of Isoodon are consistent with the pollen record. 44 FIELDIANA: GEOLOGY Chaeropus Ogilby, 1838 Chaeropus ecaudatus Ogilby, 1838 Material Trench 3, Unit 2 TMM 41106-695, right M2 (fig. 16A) Trench 3, Unit 3 TMM 41106-2763, edentulous right ramus fragment with alveoli of pre- molars (fig. 16D) Trench 4, Unit 1, Level 1 TMM 41106-674, right M2 or 3 PM 27218, right M2 or 3 (fig. 16B) PM 36828, right M2 or 3 (fig. 16F) Trench 4, Unit 2, Level 1 TMM 41106-670, right M2 or 3 TMM 41106-671, right M2 or 3 Trench 4, Units 4-5 TMM 41106-683, right M2 or 3 (fig. 16E) WAM 75.1.140, left M2 (fig. 16C) Trench 4, Unit 7, Level 2 WAM 75.1.141, right M2 or 3 TMM 41106-731, left M2 TMM 41106-732, left M2 PM 26333, left M2 or 3 Descriptions This is the rarest bandicoot in the Madura Cave fauna, with only 13 speci- mens recognized as representing the taxon. The molar teeth are quite unmistak- able because of their disproportionate labial and lingual hypsodonty. This con- dition gives the molars a characteristic diagonally sloping crown surface and renders them easily recognizable. We have no Recent comparative material, hence comparisons are made with more complete sub-Recent fossils from other Nullarbor Caves, including: a right maxillary with M1 from Weebubbie Cave (fig. 14A, PM 36806); a right maxillary fragment with M1"3 (fig. 14C, TMM 41209-572) and a left maxillary fragment with M14 (fig. 14B, TMM 41209-885), both from Webb's Cave; a right mandible with Pi, P4, and M3-4 from a cave between Kestrel #1 and #2 (fig. 15A-C, PM 36807); and a right mandible with Pi and M2-4 from Webb's Cave (fig. 15D-F, TMM 41209-573). A characteristic of Chaeropus upper molar teeth is their shallow root condi- tion. This makes them especially prone to dropping out after death. The alveoli of these teeth are distinctive too: the labial pair are deepest, but are very com- pressed anteroposteriorly, whereas the lingual two are fused into one, which is much expanded to accommodate the nearly completely fused roots supporting the paracone and hypocone (fig. 14 A). A faint ridge marks the line of fusion and gives a slight constriction to the oval alveolar outline. In the mandible, the alveoli of the molars are deeper to accommodate the Fig. 14. Chaeropus ecaudatus, sub-Recent fossil specimens from several other Nullarbor Caves. A, PM 36806, a specimen from Weebubbie Cave (north of Eucla, Western Australia) that consists of a right maxillary with M1 and that preserves the alveoli of the other molars, the premolars, and the canine. It is shown in labial and palatal views. B, TMM 41209-885, a left maxillary fragment with M1'4 from Webb's Cave, shown in labial and crown views. C, TMM 41209-572, a right maxillary fragment with M1"3 from Webb's Cave, shown in labial and crown views. 45 46 g 8 47 48 FIELDIANA: GEOLOGY long, tapered roots of the molar teeth. The labial side of the alveolar margin is markedly lower than the lingual side (fig. 15C, F), and the ramus is deeper than in Perameles. The ascending ramus is wide from front to back, very different from its narrow condition in Perameles, and is much more like that in Isoodon. Upper molars. — The upper molars are square in crown view and have a peculiar sort of tooth base hypsodonty, especially that part of the tooth under- lying the protocone, hypocone, paracone, and metacone. The stylar area of the tooth is significantly less hypsodont. No IVPs or M4s are present in our Madura Cave sample. The M2 and M3 are very similar (table 6; fig. 17A) but usually can be distin- guished by features of the stylar shelf. Two specimens from Webb's Cave (TMM 41209-572, -885; fig. 14B, C), which have M1"3 and M1"4, respectively, show that M2 has the metastyle more labially located than does M3. This results in re- versed asymmetries in the shape of the ectoloph, especially in little- worn teeth. In M3, the metastyle protrudes farther labially than the other cusps; in M2, the parastyle is the more labial cusp. These specimens also show that the M2 is further characterized by a rounded ridgelike extension of stylar cusp 1 into the V formed by the ridges of the metacone. If these specimens are typical, it seems unlikely that any M3s are present in the Madura Cave collection. TMM 41106- 695 (fig. 16A) is used to illustrate the upper molars. The protocone is crescentic when unworn. It is higher and more extensive than the hypocone. Anteriorly, the protocone extends rootward onto the an- terior face of the tooth. Posteriorly, it extends around the base of the paracone and into the central basin between the paracone and the metacone. The hypocone is a flattened column that expands toward its base. It is fused to the posterior limb of the protocone nearly to its apex. When worn, the posterior part of the protocone that projects labially into the curved valley appears to be a spur from the joined protocone and hypocone. Posteriorly, the hypocone ex- tends rootward to the posterolingual corner of the tooth. There is a broad but prominent vertical groove on the lingual face of the tooth that separates the protocone and hypocone. This groove is better developed in Chaeropus than in Isoodon. The paracone and metacone are very high, prominent cusps that, as in Isoodon, are the apices of V's that open on the labial border of the tooth. The metacone stands higher than the paracone. The paracone is joined to the para- style (a) via the stylocone (j) by a straight eocrista (preparacrista or I'x). A straight postparacrista (I'"a) joins the paracone with the posterior side of the base of cusp k. The metacone is joined to the metastyle by a postmetacrista (I") and to the anterior side of the base of the mesostyle (cusp 1) by a premetacrista (eocrista or I'"b). A distinct, small stylar cusp m is present on TMM 41106-695 and PM 27218 (fig. 16A, B). It would be rapidly removed by wear and therefore is not distin- guishable on worn teeth. As in Isoodon, the cusp k and the mesostyle (cusp 1) are about the same size and are larger than the parastyle and metastyle. Both cusps, k and 1, are oval in cross section with their long axes oriented nearly transversely. Cusp k gives rise to a ridge that extends lingually from its apex for a short distance between the arms of the paracone crests, turns posteriorly, and joins the postparacrista. The mesostyle gives rise to a ridge that extends deeply into the V between the crests JL£m_ Fig. 16. Chaeropus ecaudatus from Madura Cave. A, TMM 41106-695, a right M2 shown in labial, crown, anterior (mesial), and lingual views. B, PM 27218, a right M2 or M3 shown in labial, crown, anterior (mesial), lingual, and posterior (distal) views. C, TMM 41106-684, a left M2 shown in labial, crown, anterior (mesial), and lingual views. D, TMM 41106-2763, an edentulous right ramus fragment with the alveoli of the three premolars shown in dorsal and lateral (labial) views. E, F, TMM 41106-683 and -694, two right M2s or M3s shown in lingual, crown, anterior (mesial), and labial views. 49 50 FIELDIANA: GEOLOGY Table 6. Numerical data on dentitions of two samples of Chaeropus ecaudatus. Other Nullarbor Caves Madura Cave (fossil) Observed Observed Variate N range Mean N range Mean M»L — — — 3 3.48-4.09 3.78 AW — — — 3 2.49-2.68 2.59 PW — — — 3 2.77-3.20 2.96 M2L 4 2.96-3.01 2.97 2 2.68-2.87 2.78 AW 4 2.82-3.15 2.94 2 3.15-3.24 3.20 PW 4 2.77-3.01 2.89 2 3.06-3.24 3.15 M3L — — — 2 2.68-2.87 2.78 AW — — — 2 3.20-3.34 3.27 PW — — — 2 2.91-3.06 2.99 M4L — — — 1 — 1.88 W — — — 1 — 2.54 M,L — — — 1 — 3.24 AW — — — 1 — 1.41 PW — — — 1 — 2.44 M2L — — — 2 3.15-3.20 3.18 AW — — — 2 2.21-2.44 2.33 PW — — — 2 2.30-2.59 2.45 M2 „ 3 L 6 3.24-3.52 3.35 — — — AW 7 2.12-2.30 2.19 — — — PW 7 2.16-2.35 2.27 — — — M3L — — — 4 3.06-3.24 3.15 AW — — — 4 2.12-2.30 2.24 PW — — — 4 2.07-2.26 2.18 M4L — — — 5 3.38-3.57 3.44 AW — — — 5 1.74-1.93 1.82 PW — — — 5 1.27-1.41 1.37 of the metacone where it climbs partway up the labial face of the metacone. In some teeth (TMM 41106-732; WAM 75.1.140, fig. 16C), a short ridge extends from the lingual end of the mesostyle to the premetacrista. The labial ends of the V's are higher than the central areas, resulting in the formation of basins. Mandible and lower molars. — The one mandibular fragment (TMM 41106-2763; fig. 16D) is slender and deep and has the alveoli of all the premolars. It is referred to Chaeropus on the basis of the distinctive form of the ramus and upon the presence of a prominent diastema in front of and behind the first premolar (compare fig. 16D with figs. 6D and 10E). Other peramelids have much less space between the first two premolars, Pi and P3, than does Chaeropus and the ramus is shallower. No Mis or M4s were recovered, and M2 and M3 are so similar in size (table 6; fig. 17B) and form that we cannot be certain about a means of distinguishing them when they are found as isolated teeth. The lower molars of Chaeropus have the basic structure seen in most peramelids (figs. 15B, E; 16E, F). There is a high degree of the same sort of asymmetrical crown hypsodonty as in the upper teeth, particularly in the area underlying the protoconid and hypoconid. The trigonid is well defined in an unworn tooth, but its three cusps are close to- gether, and the trigonid basin is small. The protoconid lies at the apex of an equilateral triangle formed by the anterior half of the eocristid and is joined to LUNDELIUS & TURNBULL: MADURA CAVE 51 LENGTH B 1 M2\ \ M3 vl/y/ M2or3 V r7 * * A • • ] M, Fig. 17. Bivariate graphs showing plots of length vs. anterior width measures for each of the molar teeth of sub-Recent specimens of Chaeropus ecaudatus from several Nullarbor Caves. Stars designate those from the Madura Cave sample. Straight lines connect the midpoints of each tooth cluster of measures to provide the characteristic pattern for the species (compare with figs. 8C, D; 12A, B; 13A, C; 21A, B). Note extensive overlap of M2s and M3s and of M2s and Mas. the paraconid and metaconid by the paralophid (paracristid, I') and metalophid (epicristid II', II"), respectively. Each of these ridges has an open V close to its midpoint, but there is no carnassial notch. There is no connection between the paraconid and metaconid, except where their bases abut one another. The talonid is lower than the trigonid and is also V-shaped. The hypoconid lies at the apex of the V. A cristid obliqua (premetacristid I'"b) joins it to the entocristid (V) at the anterior edge of the entoconid. The postmetacristid (I") 52 FIELDIANA: GEOLOGY joins the hypoconid to the small hypoconulid at the posterolingual corner of the tooth. The entoconid is circular in cross section and is located at the lingual edge of the tooth in the V formed by the posterior half of the eocristid. A weak crest (entocristid V) joins the anterior edge of the entoconid to the adjacent cristid obliqua. Because the cristid obliqua does not join the protolophid, the hypoflexid extends all the way to the entocristid. A cingulum is present at the base of the tooth between the protoconid and hypoconid. It may or may not bear a small cuspule. A large procingulum ex- tends upward from the base of the protoconid to the base of the paraconid and incorporates a parastylid. It forms a prominent elongate cingular basin. Discussion This species has been recorded as a living animal in two disjunct areas. One area includes southwestern New South Wales, much of South Australia, and the southern part of the Northern Territory. The other is a small area near Northam in the inland district of Western Australia. The Madura Cave specimens and notes by Merrilees (1968) and Archer (1972) demonstrate its former presence in an intermediate area. Its Recent occurrence in arid areas in South Australia suggests that it may have been widespread in the arid parts of southern Aus- tralia until very recently and that the disjunct nature of its known Recent dis- tribution is the result of either a collecting bias or recent historic restrictions in its distribution. This is supported by the presence of its remains in surficial deposits in other caves of the Nullarbor Plain (Merrilees, 1968; Archer, 1972). Its presence throughout the sequence at Madura Cave indicates that it has had a long residence in this region and that it was one of the species able to tolerate the drying of the climate that took place at the end of the Pleistocene. Macrotis Reid, 1837 Macrotis lagotis (Reid, 1837) Material Trench 3, Unit 1, Level 1 TMM 41106-32, edentulous right ramus fragment, juvenile Trench 3, Unit 2 TMM 41106-73, left M2 or 3 TMM 41106-56, right ramus fragment with M2-3 and M4 removed from crypt (fig. 20A) TMM 41106-57, edentulous left ramus fragment of juvenile with alveoli of P4-M4 TMM 41106-58, edentulous right ramus fragment with alveoli of M2-4 TMM 41106-59, edentulous right ramus fragment with alveoli of M24? PM 26336, left M1 (fig. 19D) PM 26337, right M2 or 3 PM 26338, right M2 or 3 PM 26339, right M2 or 3 PM 26340, left Mi (fig. 20B) PM 26341, left M2 or 3 PM 26342, edentulous left maxillary with alveoli of C-M1 (fig. 19 A) PM 26343, left P3 (fig. 20C) LUNDELIUS & TURNBULL: MADURA CAVE 53 Trench 3, Unit 2 (continued) PM 26344, right P1 or P3 (fig. 19B) PM 26345, broken left M2 or 3 PM 26346, right Mi PM 26347, broken right M2 or 3 PM 26348, left Pa (or Pi) Trench 3, Unit 2, Level 2 TMM 41106-53, left mandibular fragment with alveoli of two molars TMM 41106-54, right M2 or 3 PM 26349, right M2 or 3 PM 26350, right M2 or 3 (fig. 19E) PM 26351, left M1 PM 26352, right M2 or 3 PM 26353, left M2 or 3 PM 26354, left Ma (or M2) PM 26355, right M2 or 3 PM 26356, right M1 PM 26357, left P3 (or P1) Trench 4, Unit 1, Level 1 TMM 41106-476, right M2 or 3 TMM 41106-573, right M1 TMM 41106-575, left M1 (fig. 19C) TMM 41106-584, left Ma (or M2) TMM 41106-611, left M1 TMM 41106-612, right Ma (or M2) TMM 41106-676, left Mi PM 26358, right M3 (or M2) (fig. 19G) PM 26359, right premolar, probably P4 PM 26360, anterior half of a left P3 or > PM 26361, right M3 Trench 4, Unit 2, Level 1 WAM 75.1.128, left P3? PM 26363, parastylar area of right M2 or 3 PM 26364, lingual side upper molar PM 26365, right Ma (or M2) PM 26366, right M2 or 3 PM 26367, right M2 or 3 PM 26368, left M2 or 3 PM 26369, left M4 (fig. 19H) PM 26370, left Mi PM 26921, right M2 or 3 PM 26922, right M2 or 3 PM 26923, left M2 or 3 Trench 4, Unit 2, Level 2 PM 26924, left M1 PM 26925, right M2 or 3 PM 26926, right Ma PM 26927, left M1 54 FIELDIANA: GEOLOGY Trench 4, Unit 2, Level 2 (continued) PM 26928, right M2 or 3 PM 26929, left Mi PM 26930, right Mi Trench 4, Units 4-5 PM 26931, edentulous right ramus with alveoli of P4-M4 (fig. 20D) PM 26932, left M1 PM 26933, left M1 PM 26934, right M2, 3, or 4 PM 26935, left M2 or 3 Trench 4, Units 4-5 (continued) PM 26936, left Ms (or M2) PM 26937, right M1 PM 26938, left M2 or 3 PM 26939, right M1 PM 26940, left Ms (or Ma) PM 26941, right M4 PM 26942, edentulous left ramus with alveoli of P1-M4 (fig. 20E) PM 26943, edentulous left ramus with alveoli of Mi -4 PM 26944, left Ms (or M2) PM 26945, right M2 or 3 (fig. 19F) PM 26946, anterior half, left M2 or 3 PM 26947, right M2 or 3 PM 26948, left M1 PM 26949, anterolabial corner of left M2 PM 26950, trigonid of right M2 or 3 PM 26951, left upper molar PM 26952, trigonid of right lower molar PM 26953, right Mi PM 26954, trigon of right upper molar PM 26955, right M3 (or M2) PM 26956, left M2 or 3 PM 26957, left M2 or 3 PM 26958, right upper molar fragment, probably M2 or 3 PM 26959, left P1 or P3 PM 26960, left premolar, probably a P4 PM 26961, right premolar, probably a P4 PM 30549, right Mi PM 30550, left Mi Trench 4, Unit 7 PM 26963, left M2 (or M3) Trench 4, Unit 7, Level 1 PM 26964, left M2 (or M3) PM 26965, left M3 (or M2) PM 26966, right Mi PM 34417, edentulous ramus fragment Trench 4, Unit 7, Level 2 WAM 75.1.129, edentulous right maxillary fragment with alveoli of P3"4 PM 26967, right ramus with alveoli of P4-M3 LUNDELIUS & TURNBULL: MADURA CAVE 55 Trench 4, Unit 7, Level 2 (continued) PM 26968, right M2 PM 26969, right Ma (or Ma) PM 26970, left M2 (or M3) PM 26971, left Ma (or M2) PM 26972, right M1 PM 26973, right M1 PM 26974, left Mi PM 26975, right M2 or 3 PM 26976, right M2 Trench 5, Unit 5 PM 26335, right ramus fragment with alveoli of P1-M2 (fig. 20F) PM 26977, left M2 (or Ma) PM 26978, left M2 (or Ma) PM 26979, left Ma (or M2) PM 26980, right M3 (or M2) PM 26981, left M1 fragment PM 26982, left M1 fragment PM 26983, right M2 or 3 PM 26984, left M2 or 3 PM 26985, partial right M2 or 3 Trench 5, Unit 5 or 6? PM 26986, left Mi PM 26987, left Pa (or Pi; "1 ... . . .,..,, PM 26988, right Pa (or Pl)| Possibly from the same individual PM 26989, left Pa Descriptions Skull. — Very little cranial material of this species is represented in the Madura collection. Most of the specimens represent immature individuals, probably indicating that owls were the predators responsible for bringing the material to the cave. An adult Macrotis is probably too large to be regularly taken by owls. One edentulous left maxilla of a juvenile (PM 26342, fig. 19A) is the only specimen that represents the skull. It has alveoli for the canine, the first two upper premolars, and dP4, and the M1. The developing alveolus for the un- erupted P4 is located lingual to that of the dP4. The infraorbital foramen is located above the anterior edge of the alveolus of M1. It is elliptical in cross section and opens into a broad rounded groove on the lateral face of the maxilla. A comparison of the Madura Cave specimen with the skull of a Recent speci- men (FM 35331; fig. 18) reveals no differences that are not the result of the difference in ontogenetic age. The Madura Cave specimen is slightly younger, and the anterior part of the maxillary that carries the premolars is relatively shorter than in the Recent specimen. Upper premolars. — Several upper premolars have been recognized. In one of them (PM 26344), a worn right P1 or P3, the form agrees well with the modern comparative specimen. The tooth is double rooted, and the anterior root has a distinctive, markedly curved appearance. There is a single, large, central cusp with rounded crests. The anterior crest is very weak, and there is no anterior 56 57 58 FIELDIANA: GEOLOGY cingular cuspule. The posterior crest runs to a posterior cingular cuspule that shows wear lingually. The crown is slightly narrower than that of the modern specimen when viewed occlusally. Another specimen (WAM 75.1.128), a left P3 (or P1), is also worn. It is wider in the area of the posterior root than is PM 26344 and the recent specimen, and the anterior crest of the central cusp is more distinct as in the recent specimen. In PM 26959, a left P1 or P3, there is a minute but distinct anterior cingular cuspule. None of the other teeth shows this fea- ture, nor does the modern specimen. In PM 26357, a left P3 or P1, the profile is unusual in that instead of being convexly curved from crown tip to root tip, here there is a slight concavity between the tip of the cusp and the base of the crown. Upper molars. — The upper molars, with the exception of M4, are roughly rectangular in occlusal view when unworn. When worn, they become more square. The M1"3 have achieved this shape by an enlargement and lingual movement of the metacone rather than by the addition of a hypocone as in Isoodon and Chaeropus. The M4 has the shape of a rounded triangle. All have massive rounded cusps and tooth base hypsodonty that is more developed on the lingual half of the tooth. The M1 is characterized by the presence of a well-developed parastyle with a medially directed crest that joins the paracone so as to delimit a tiny basin and the crowding together of the subequal-sized paracone and stylocone. The metacone is the largest cusp. It has the shape of an asymmetrical crescent in occlusal view, with the short anterior horn extending to the base of stylar cusp 1. The longer posterior horn joins the metastyle. The protocone is the lowest major cusp on the tooth. It is elongated anteroposteriorly and reaches from the an- terolingual corner of the tooth to the metacone. There is no connection with the parastyle. Stylar cusp 1 is the tallest cusp on the tooth. It is rounded anteriorly where it is separated from the stylocone by a sharp valley. It is compressed posteriorly into a ridge that approaches but does not join the metastyle in the unworn state. With wear they become joined. A slight bulge on this ridge might represent stylar cusp m (PM 26336, PM 26351). Specimens TMM 41106-573 and 41106-575 (fig. 19C), although little worn, do not show this feature. Wear results in a joining of the metacone and stylar cusp 1 around the posterior part of the tooth and at a later stage in the joining of the paracone and stylocone, which in still later stages are joined to the protocone. The M2 and M3 are so similar that they will be treated together. The stylocone and stylar cusp 1 are nearly equal in size and height and are the two highest cusps on the tooth. In the M2, the anterior face of stylar cusp 1 usually has a ridge (PM 26350 lacks this ridge: fig 19E), which is absent on the M3. The labial face of the stylocone of the M2 bulges more than in M3, and the base of the enamel swings sharply toward the parastyle. In the M3, the labial surfaces of these two cusps bulge more nearly equally, and the labial enamel line does not turn toward the parastyle. The posterior face of the stylocone is ridged, usually better developed in the M2. The metacone is the largest cusp. It is like that of the M1, with a long posterior part that extends around the posterior part of the tooth and joins the metastyle. The anterior arm swings labially to the base of stylar cusp 1. In the M3, the crescents of metacone and paracone are not as disproportionate in size as in the M2, mostly because the crescent of the metacone is tighter in M3 than it is in the M2. Fig. 19. Macrotis lagotis specimens from Madura Cave. A, PM 26342, an edentulous left maxillary fragment with the alveoli of C, P,, P3, dP4/ M,.2, shown in labial and palatal views. Note the characteristic, large molar alveolus for the root of the protocone (and hypocone) which can be seen in the palatal view. Note too that the dP4 had its two roots fused. B, PM 26344, a right P1 or P3 shown in lingual and crown views. C, TMM 41106-575, a left M1 shown in labial, mesial, lingual, and crown views. D, PM 26336, a left M1 shown in labial and crown views. E, PM 26350, a right M2 or M3 shown in labial, mesial, lingual, distal, and crown views. F, PM 26945, a right M2 or M3 shown in labial and crown views. G, PM 26358, a right M3 or M2 shown in labial and crown views. H, PM 26369, a left M4 shown in labial and crown views. 59 60 FIELDIANA: GEOLOGY The protocone is the lowest major cusp. It extends from the base of the metacone around the anterolingual corner of the tooth and joins an anterior cingulum that continues to the parastyle. The paracone is V-shaped, with the anterior arm joining the base of the stylocone at the notch between the stylocone and parastyle. In unworn or little-worn teeth, M2 can be distinguished from M3 by its nar- rower anterior absolute (but not relative) width (fig. 18B). This is demonstrated by the Recent specimens that have both teeth present in the maxillary. In the plot of the Madura Cave sample, we interpret the group with the narrower anterior width measure to be the M2s; that with the wider anterior width to be the M3s (fig. 21). At advanced wear stages in which all cusp, crest, and valley features of the crown have been worn away, the anterior width measures of M2 and M3 have become reversed. This reflects the nature of the crown hypsodonty in which the taper causes the anterior width vs. transverse proportions to shift at the different heights (wear stages). The M4 is a three-rooted tooth with the shape of a rounded triangle in cross section (fig. 19H). The posterior part of the tooth is reduced. The paracone is the largest cusp on the tooth. It is crescentic, with the anterior arm joining the base of a reduced stylocone. The posterior arm ends against a small cuspule on the labial edge of the tooth. A well-developed anterior cingulum extends from the parastyle to the base of the paracone or a fused paracone-protocone. The latter alternative is suggested by the shape of the worn crescent. The posterior edge of the tooth is marked by a small cuspule. Wear removes these features early in the life of the tooth. The result is an enamel ring surrounding the dentine core of the tooth. Mandible.— Three specimens (fig. 20A, F, D), TMM 41106-156, PM 26335, and PM 26931, provide the basis for the description of the mandible of M. lagotis from Madura Cave. The horizontal ramus is characterized by its rapid increase in width at Mi to accommodate the wide molars. The entire ramus is twisted so as to turn the anterior alveoli outward. A mental foramen is located under the Pi. The posterior end of the symphysis is located between Pi and P2. The anterior edge of the ascending ramus makes a high angle (more than 45°) with the tooth row. The masseteric fossa is broad and bordered by ridges that become better defined with age. The ventral ridge of the masseteric fossa is enlarged above the angular process and forms a tubercle. A flattened rugose ridge extends from this tubercle onto the lateral edge of the angular process. The mandibular foramen, which is located in a broad, posteriorly open groove, opens near the anterior edge of the base of the angular process. The condyle is located closer to the top of the coronoid process than to the angular process. Its articular surface is almost circular. It is slightly concave transversely and convex anteroposteriorly. Lower premolars. — Eight lower premolars have been recognized. Five of them (PM 26343, 26348, and PM 26987-9) are very much alike and are comparable to Pi and P3 of the modern form. They consistently differ from the modern form in having a wider posterior half of the crown base. They all show traces of two subsidiary cusps associated with the crests of the primary cusp, one on each crest. In the modern specimen, only one of the four teeth bears one of these features. On some of the fossils, there is a slight bulge where the anterior cingular cusp might be expected, but none has a cuspule. LUNDELIUS & TURNBULL: MADURA CAVE 61 The P4S (PM 26359 and PM 26960-1) are shorter anteroposteriorly than the other premolars, and the crown base is proportionately wider. The crown has a central cusp as its main feature, and this is laterally compressed and has anterior and posterior crests. The cingulae show variation. There is a weak anterior cingulum enveloping the anterior quarter of the crown base. On two (PM 26960-1), there is a small cusp present, and on one (PM 26359), this cusp is missing, and instead on the anterolingual side there are two minute cuspules tight against the base of the central cusp just where the cingulum rises slightly before it merges into the base of the central cusp. All have a distinct posterior cingular cusp at the rear end of the posterior crest, usually somewhat lingually situated. In PM 26960 and 26961, short cingulae extend forward from this cusp, one on each side along the crown base margin for up to a fourth of the length of the tooth. In PM 26359, from the rear side of this posterior cingular cusp or heel, a weak cingulum runs down and labially across the midaxis of the tooth and then rises and runs forward for about one fourth the length of the tooth to merge with the base of the main cusp. It bears two minute, freestanding cuspules. There is no lingual cingulum going off from this posterior cusp as in the other teeth. Lower molars. — The Mi of M. lagotis is elongate and tapers toward the anterior end. There is considerable tooth base hypsodonty beneath the protoconid as compared with the lingual side of the tooth where there is little or none and a much greater development of tooth base hypsodonty beneath the hypoconid similar to that of the labial side of the rest of the molars. The trigonid is narrow and small and variable in its development. It is slightly raised above the talonid, which is much wider. The protoconid and metaconid are subequal in size and height and are connected by a weakly developed epicristid (II' and II") that runs across the valley between the cusps to connect to their back edges. The pro- toconid part of the epicristid lies just ahead of a better developed postparacris- tid (I'"a). The paraconid is the smallest and lowest trigonid cusp, but it is better devel- oped than on the Recent comparative specimen (FMNH 35331; fig. 18D). It lies ahead of the metaconid, appressed against its base. There usually is a connec- tion to the protoconid by a paracristid (I'). In PM 26930, PM 26953, and PM 26966, this is a weak connection, but in PM 26370, PM 26974, and PM 30549, it is a stronger one, and in PM 26340 (fig. 20B), the ridges from each cusp are strong yet they fail to meet in the valley. In PM 26346, PM 26929, and PM 26986, there are not even any ridges suggestive of a connecting crest. There is always a parastylid that is low and lies anterior to the paraconid, near the crown base at the anterolingual corner of the tooth. It too varies in size and in the extent to which it tends to be extended labially into a short cingular ridge. In most, it is nearly a small cone, whereas in PM 26346, PM 26370, PM 26974, and PM 30549, there is an expressed tendency for the cingular crest to form. In the talonid, the entoconid and hypoconid stand about equally high in an unworn tooth, but the latter, with its extreme crown base hypsodonty, is far larger. The entoconid is rounded posteriorly and ridged from its crest both anteriorly and anterolabially. The anterior crest runs toward the transverse cen- tral valley and bends labially without entering the bottom of the valley. The anterolabially directed crest also fails to reach the base of the cusp. The hypoconid has a crescentic form with an anterior crest, the premetacristid Fig. 20. Macrotis lagotis from Madura Cave. A, TMM 41106-156, a right mandibular fragment with half of the horizontal ramus, most of the coronoid and angular processes, but lacking the condyle. M2 and M3 are in place, and the unerupted M4 has been removed from its crypt. The jaw is shown in medial and dorsal views, the M4 in lingual, labial, and crown views. B, PM 26340, a left M, shown in crown, lingual, mesial, labial, and distal views. C, PM 26343, a left P3 shown in labial, lingual, and crown views. Opposite: Fig. 20. D, PM 26931, an edentulous partial right mandible of a juvenile individual shown in lingual and dorsal views. Preserved is the ascending ramus and the posterior portion of the horizontal ramus, with part of the alveolus of dP4, part of the crypt of P4, the alveoli of M,.3, and the crypt for P4. E, PM 26942, an edentulous left ramus with the alveoli of P, and P3, dP4, and P4 at an early developmental stage within its crypt (the tip of the main cusp is broken off), the alveoli of M,.3/ and the crypt for M4. F, PM 26335, an edentulous right ramus fragment of a juvenile individual with alveoli of P,- M2 including that for dP4 and the crypt of P4, shown in labial and crown views. 62 63 64 FIELDIANA: GEOLOGY or cristid obliqua (I'"b), which runs anterolingually to join the postparacristid behind the protoconid above the transverse valley floor. The posterior crest of the hypoconid, the postmetacristid (I"), runs labially to join the hypoconulid at the posterolingual corner of the tooth. The hypoconulid is a rounded projection with a weakly developed crest connecting it to the base of the entoconid. In four specimens there is a spur from the postmetacristid to the base of the entoconid (PM 26340, PM 26370, PM 26986, and PM 30550). Occasionally there may be a small cingular cuspule between the protoconid and hypoconid (PM 26370 and PM 26986), but usually there is no trace of this element. Comparisons with the Mi of two subfossil specimens (PM 4979 and PM 4980) from Murraelellevan Cave on the Nullarbor are very close. They have the crested parastylid, but it is even more expanded and complex than in the Madura Cave specimens. Also, each has a spur from its postmetacristid to the base of the entoconid, and one (PM 4979) has the paracristid developed, the other does not, and neither has the small cingular cusplet between the protoconid and hypoconid as do a few of the Madura Cave specimens. The major difference between the fossils and the Recent specimen is the much more reduced paraconid in the latter as well as a strong cingular cusp between protoconid and hypoconid. The M2 and M3 are roundly rectangular teeth in crown view, and they are so similar that one description fits them both almost completely. The metaconid and entoconid are the highest cusps, and the former reaches slightly higher than the latter. The protoconid and hypoconid are smaller and lower, and in the unworn state the protoconid is slightly higher, but the hypoconid is a larger cusp with more open crescentic form than the protoconid — especially in M2. There is no paraconid on M2: the paracristid simply runs from the protoconid to the anterolabial base of the metaconid. In M3 there is a small bulge at the union of crest and cusp that may be the homologue of the paraconid, and in one specimen, PM 26337, there is a weak crest from this "paracristid" to the para- stylid. The epicristid connects the back sides of protoconid and metaconid. The trigonid of M2 is distinctly narrower than the talonid, whereas in M3 both are about equal in width or the trigonid may be slightly wider. There is a heavy procumbent anterior cingulum that commences in a cuspule one-half to two- thirds of the way up the anterolingual side of the protoconid. The cingulum runs forward and swings labially to form the stout parastylid labial to the mid-axis of the tooth. It thereby cuts off a deep, curved depression between its crest and the anterolabial wall of the protoconid. There is a weak postparacristid that joins the cristid obliqua above the floor of the transverse valley. Usually the hypoconid in its hypsodonty suddenly flares forward, occluding the labial side of the transverse valley so that it abuts the back of the protoconid for the ventral one-half to two-thirds of the height of the crown beneath these cusps. An exception to this is seen on PM 26337 where the flare is gentle and the pro- toconid also flares backward a bit. In another specimen (PM 26365), neither cusp flares, they just gradually expand. The posterior arm of the hypoconid crescent, leading to the hypoconulid, sends off a spur to the base of the en- toconid. The hypoconulid of M2 is slightly larger than that of M3, and either tooth may have minute cingular cuspules associated with it. In one specimen (PM 26337), an M3, there is no hypoconulid, and the postmetacristid just fades away without reaching the posterolingual corner of the tooth. The M2S and Mss are similar in size and morphology but can be separated on LUNDELIUS & TURNBULL: MADURA CAVE 65 bivariate scatter diagrams of tooth length vs. anterior and posterior widths. In each case, M2S are shorter than Mas (fig. 21B). The scatter diagram of L vs. AW shows the M2S to be narrower, too, so that there is a clear separation into two discrete groups. In M3, the two width measures do not show a significant difference, and although there is a complete separation along the length scale, there is no gap between the clouds of points. Larger samples might be expected to show some overlap. The M4 has a well-developed trigonid and a reduced and posteriorly tapered talonid (fig. 20A). The protoconid and metaconid lie close to one another, about as in the Mi, and they are nearly equal in size. Both are somewhat crescentic. The posterior arm of each crescent descends, and they meet in the center along the back edge of the trigonid. The anterior arm of the metaconid is enveloped by that of the protoconid. This results from the more open crescentic form of the protoconid. The anterior cingulum is pronounced and is similar to those of M2 and M3. The two basins in the trigonid region, a central trigonid basin and an anterior cingular basin, are joined. The central basin that lies between the facing crescents of protoconid and metaconid is small and has an anterolingu- ally directed valley leading from it. The anterior cingular basin lies transverse to the tooth axis, and at its lingual end it joins the lingual end of the valley of the central basin. There may or may not be a common exit to the lingual side of the tooth. A parastylid cusp is incorporated into the lingual terminus of the cin- gulum. The trigonid is raised above the talonid. The entoconid is the largest talonid cusp, and it is weakly crescentic. The hypoconid is very low and not well defined, and there is only the faintest trace of a hypoconulid. The talonid basin is triangular and deeply clefted labially. Comparisons. — The M*s from Madura Cave and those of young individuals from Murraelellevan Cave (PM 4872) have a well-developed parastyle. The more complete specimens in the Murraelellevan sample show that this feature is soon reduced, then eliminated by wear. One Recent specimen from Woyaline Wells (FMNH 35331; fig. 18A, B) has no parastyle on its M1. Its absence cannot be attributed to wear because this specimen is a juvenile with M3 still incompletely erupted, with P4 and M4 within their respective crypts. On M1, the Madura Cave specimens have no anterior cingulum connecting protocone to parastyle as is seen in unworn specimens from Murraelellevan Cave. Such a cingulum is present on all the other molars of the Madura Cave materials and in unworn or little worn M2-M4 of all of our comparative speci- mens. On M2 of the Murraelellevan Cave specimen, the anterior ridge of the paracone also goes to the parastyle; in the Madura Cave specimens of M2 and in M2 of the comparative materials and in all M3s, this ridge goes toward the notch between the parastyle and stylar cusp 1. Last molars are rare in spite of a relatively large number of specimens of Macrotis in the Madura Cave fauna. This correlates with the young age stages of most of the other teeth. The M4s and M4S had not fully formed at the time of death. The other lower molars compare well in their main features to the avail- able Recent and fossil materials. Discussion Macrotis lagotis is known as a living animal from widely scattered areas in the dryer parts of Australia, including two localities, Ooldea and Rawlinna, on the 2.50 2.00 B ^—^ (• M3 • / o / M2 \ ° \ o o / \ \ • \V I ° ° 1 > o/ o o / v^ % \ ^ / • / • / • M, \ • • J LENGTH Fig. 21. 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