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Functional Morphology of Marsupial Moles ( Marsupialia, N Otoryctidae) Contents Verh. naturwiss. Ver. Hamburg (NF) 42 39-149 Hamburg 2006 Functional morphology of marsupial moles ( Marsupialia, N otoryctidae) By NATALIE MARINA WARBURTON, Nedlands (Western Australia*) With 22 Figures Abstract: Marsupial moles (Notoryctes) are the most highly specialised burrowing marsupials. The specialisa­ tions of the appendicular musculo-skeletal system of the marsupial moles are extensive and widespread; the ma­ jor alterations are concentrated in, but not restricted to, the forelimb. Many of the derived features of the mus­ cular system appear to be adaptations for improving the mechanical advantage of the limbs for burrowing. A number of the specialisations of the muscular system of the marsupial moles are convergent with those pre­ viously documented in other fossorial mammals, including golden moles ( Chrysochloris), rodents (Spalacidae) and armadillos (Dasypodidae: Chlamyphorus). There are, however, a number of unique specialisations of the musculo-skeletal system of Notoryctes. The functional morphology of the locomotor apparatus of marsupial moles is interpreted on the basis of the descriptions of the anatomy of the skeletal and muscular systems. The burrowing technique of the marsupial moles is a modified form of the parasagittal digging method that is used by other fossorial mammals, such as golden moles, armadillos and some rodents including pocket gophers (Geomyidae). Differences in the functional morphology of the hindlimb between marsupial moles and other fossorial mammals are a reflection of the fact that marsupial moles do not construct permanent open burrow systems, but instead constantly dig through loose soil, backfilling as they progress. The functional morphology of the tail is uniquely specialised in the marsupial moles to function as the fifth limb during the pentapedal bur­ rowing locomotion. Phylogenetic relationships of marsupial moles within the Marsupialia have long been enig­ matic. While specialisation of the musculo-skeletal system have been so widespread as to obscure almost any phylogenetically relevant patterns, there is evidence to support a sister group relationship of notoryctids and peramelid bandicoots (Peramelidae). Interspecific differences between the two species of marsupial moles, No­ toryctes typhlops and N caurinus, are minor. Contents A. Introduction . 41 I. Marsupial radiation . 41 II. Marsupial moles . 42 III. Fossil record of the Notoryctidae . 46 IV. Phylogenetic affinities of the Notoryctidae . 46 V. Other fossorial mammals . 48 * Author's address: Dr. N. WARBURTON, School of Animal Biology M092, The University of Western Australia, 35 Stirling Highway, Crawley WA 6009, Australia.- e-mail: [email protected] 39 1. Insectivora ......................................................... 49 3. Other forearm muscles . 105 2. Rodentia ........................................................... 49 III. Summary: Pectoral girdle and limb musculature . 106 3. Xenarthra .......................................................... 50 IV. Muscles of the pelvic girdle and thigh . 106 VI. Aim of study .......................................................... 50 1. Iliacus group . 107 Acknowledgements ..................................................... 52 2. Gluteal group . 107 3. Quadriceps femoris group . 108 B. Materials and methods ....................................................... 52 4. Adductor group . 109 I. Osteology ............................................................ 52 5. Ischiotrochanteric group . 111 II. Myology ............................................................. 53 6. Hamstring group . 112 C. Skeleton .................................................................. 53 V. Muscles of the lower leg . 113 I. Skull ................................................................ 54 1. Flexor group . 113 II. Vertebral column ....................................................... 56 2. Deep flexors . 114 1. Results ............................................................ 56 3. Tibial extensor group . 115 2. Review ............................................................ 57 4. Peroneal group . 116 III. Thorax ............................................................... 60 VI. Summary: Pelvic girdle and hindlimb musculature . 116 1. Ribs .............................................................. 60 VII. Musculature of the tail . 117 2. Sternum ........................................................... 60 VIII. Summary . 118 3. Review ............................................................ 60 1. Interspecific and intraspecific variation in Notoryctes . 119 IV. Pectoral girdle ......................................................... 61 2. Marsupial moles: Comparison with other marsupials . 119 1. Scapula ............................................................ 61 3. Marsupial moles as subterranean mammals . 121 2. Clavicle ............................................................ 64 E. Functional Morphology . 122 3. Review ............................................................ 65 I. Movements of the pectoral limb . 123 V. Forelimb ............................................................. 67 1. Rotation of scapula . 123 1. Humerus ........................................................... 67 2. Mobility of pectoral girdle . 124 2. illna and radius ...................................................... 69 3. Movements about shoulder joint . 126 3. Manus ............................................................ 70 4. Movements about elbow joint . 129 4. Review ............................................................ 70 5. Movements about wrist joint . 130 VI. Pelvic girdle ........................................................... 72 6. Summary . 131 1. Results ............................................................ 73 II. Movements ofthe pelviclimb . 132 2. Review 74 1. Mobility of pelvic girdle . 133 VII. Hindlimb 76 2. Movements about hip joint . 133 1. Femur ............................................................. 76 3. Movements about knee joint . 136 2. Patella ............................................................ 76 4. Movements about tarsal joint ............................................. 137 3. Tibia and Fibula ..................................................... 79 5. Summary . 138 4. Pes ............................................................... 79 III. Movements of the tail . 139 5. Review ............................................................ 79 IV. Summary . 141 VIII. Discussion ............................................................ 81 V. Evolutionary convergence . 142 IX. Conclusions ........................................................... 82 F. Conclusions . 143 D. Muscular system . 83 G. Bibliography . 145 I. Pectoral girdle and brachium . 85 1. Accessory field . 85 2. Suprazonal matrix . 89 3. Infrazonal matrix . 91 4. Deltoid group . 91 A. Introduction 5. Subscapular group . 92 6. Latissimus dorsi group . 95 7. Triceps . 96 I. Marsupial radiation 8. Pectoral group . 97 The current classification of Australian marsupials is based on the hypothesis of a mono­ 9. Supracoracoid group . ..
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