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LOCOMOTOR ADAPTATIONS and ECOMORPHOLOGY of SHORT-FACED BEARS (Arctodus Simus) in EASTERN BERINGIA

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LOCOMOTOR ADAPTATIONS and ECOMORPHOLOGY of SHORT-FACED BEARS (Arctodus Simus) in EASTERN BERINGIA Palaeontology Program Government of the Yukon Occasional Papers in Earth Sciences No. 7 LOCOMOTOR ADAPTATIONS AND ECOMORPHOLOGY OF SHORT-FACED BEARS (Arctodus simus) IN EASTERN BERINGIA Paul E. Matheus The Alaska Quaternary Center, Department of Geology and Geophysics and The Institute of Arctic Biology University of Alaska Fairbanks YUKON Palaeontology Program Department of Tourism and Culture Elaine Taylor, Minister 2003 Publication Note: This monograph was originally written as as Matheus 1995 and 2001. The dissertation also Chapters 2, 3, and 4 of the Ph.D. dissertation contained an appendix with stable isotope data on entitled, “Paleoecology and Ecomorphology of the modern and Pleistocene carnivores along with a Giant Short-Faced Bear in Eastern Beringia,” manual for extracting and purifying collagen from completed by the author in 1997 at The University bone. The present monograph may be cited of Alaska Fairbanks. The content is essentially directly, but if the citation is used to establish unchanged, except for minor editing, typographic when the ideas herein were established or data corrections, and re-formatting. The complete herein made public, then the dissertation or dissertation contained two additional papers Matheus (1995) take precedence. (Chapters 1 and 5) that are cited in this monograph YUKON Palaeontology Program Department of Tourism and Culture Elaine Taylor, Minister 2003 LOCOMOTOR ADAPTATIONS AND ECOMORPHOLOGY OF SHORT-FACED BEARS (Arctodus simus) IN EASTERN BERINGIA Paul E. Matheus TABLE OF CONTENTS Frontispiece Arctodus simus skeletal reconstruction by author .................................................. i Publication Note .......................................................................................................................... ii Abstract ....................................................................................................................................... v Part I: General Effects of Body Size and Limb Morphology in Large Quadrupedal Mammals— Designs for Speed, Acceleration, and Endurance Introduction .................................................................................................... 1 Current Ecological Models and the Hypotheses Being Tested ......... 1 Primer on Arctodus’ Morphology .................................................... 2 Limbs are Compromises .................................................................. 5 The Comparative Approach and Layout of This Paper ................... 5 Definitions ..................................................................................................... 6 Confusions Regarding Cursorialism .............................................................. 7 Quadrupedal Gaits ......................................................................................... 8 Preliminary Remarks on Scaling and Allometry ............................................ 11 Allometric Functions ....................................................................... 11 Similarity Hypotheses and Scaling Theory ...................................... 12 Reservations About Scaling Applications ........................................ 13 Functional Interpretations Using Allometry .................................... 14 Energetics, Metabolism, and the Cost of Locomotion ................................... 17 Setting the Stage .............................................................................. 18 The Cost of Transport ...................................................................... 18 Mechanics of Walking and Running ................................................. 20 Locomotor Efficiency and the Role of Stored Elastic Strain Energy— Advantages of Being Big ................................................. 21 Optimal Speeds Within a Gait and the Effect of Limb Length— Reconsidering the Cost of Transport ................................ 22 Scaling of Muscle Force ................................................................................. 24 Bones and Skeletal Stresses ............................................................................ 28 Predicted vs. Actual Bone Stress ...................................................... 28 How do Large Animals Reduce Realized Bone Stress? ................... 29 What Triggers Gait Changes— More Clues into the Role of Body Size ...................................................................................... 31 An Upper Size Limit for Predators ................................................... 33 Summarizing the Relationships Between Force Generation, Bone Stress, and Body Size .............................................................. 35 Limbs as Levers .............................................................................................. 37 Lever Systems ................................................................................... 37 The Effects of Mechanical Advantage on Locomotor Performance ..................................................................................... 37 Low Gear and High Gear Muscle Systems ....................................... 39 Kinematics of Limbs ....................................................................................... 41 Summary of Relationships Between Morphology and Locomotor Performance .................................................................................................... 44 Good Accelerators ........................................................................... 44 Efficient Runners ............................................................................. 44 iii PART II: Specific Morphological Features of Arctodus Introduction ................................................................................................... 46 Was Arctodus Cursorially Adapted ? ............................................................ 47 Proximal-Distal Limb Proportions .................................................. 47 Limb Mass and Allometry ............................................................... 50 podials ............................................................................... 50 metapodials ........................................................................ 50 long bones .......................................................................... 50 humerus ............................................................... 51 radius/ulna ........................................................... 51 femur ................................................................... 61 tibia .................................................................... 61 overall allometry and functional interpretations ................ 61 Limb Posture and Conformational Features .................................... 66 Front Limbs ....................................................................... 66 Hip and Femur ................................................................... 68 summary on cursorialism .................................................................. 73 Gait Selection and Speed: What Kind of Cursor was Arctodus? ................... 75 Gait Selection ................................................................................... 75 Acceleration ..................................................................................... 76 The Hyena Analogy– pros and cons ..................................................... 77 Speed of Travel ................................................................................ 79 Postlogue and Preview of the Part III ............................................................. 81 Part III: Foraging Energetics, Carcass Productivity, and the Evolution of Arctodus’ Scavenging Niche Introduction .................................................................................................... 83 Forage Requirements and Population Parameters .......................................... 84 Forage Requirements ....................................................................... 84 Carcass Requirements ...................................................................... 85 Population Density and Home Range Size ....................................... 87 Minimum Viable Population ............................................................ 89 Modeling Carcass Abundance ........................................................................ 90 Carcass Production from Predation .................................................. 90 Carcass Production from Natural Mortality ..................................... 95 Modeling Herbivore Populations in Eastern Beringia ....... 95 Redmann’s Model ............................................... 95 Bliss and Richard’s Model .................................. 98 Combined Model ................................................. 99 Non-Predatory Mortality in Eastern Beringia .................... 99 Corollaries to the Model .................................................... 102 Short-Faced Bear Population Levels ..... 102 Predator-Prey Ratios ............................. 102 Relying on Carcasses as a Continuous Food Source and the Evolution of Arctodus’ Scavenging Niche .......................................................................... 104 Feasibility of a Pure Scavenging Niche ............................................ 104 A theory for the Evolution of Scavenging in Short-Faced Bears ....... 106 Extinction .......................................................................................... 111 Literature Cited ........................................................................................................................
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