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The Carnivores of Agate Fossil Beds National Monument University of Nebraska - Lincoln DigitalCommons@University of Nebraska - Lincoln Zea E-Books Zea E-Books 6-2018 The aC rnivores of Agate Fossil Beds National Monument: Miocene Dens and Waterhole in the Valley of a Dryland Paleoriver Robert M. Hunt Jr. University of Nebraska-Lincoln, [email protected] Robert Skolnick University of Nebraska State Museum, [email protected] Joshua Kaufman University of Nebraska State Museum Follow this and additional works at: https://digitalcommons.unl.edu/zeabook Part of the Geology Commons, Paleobiology Commons, Paleontology Commons, and the Sedimentology Commons Recommended Citation Hunt, Robert M. Jr.; Skolnick, Robert; and Kaufman, Joshua, "The aC rnivores of Agate Fossil Beds National Monument: Miocene Dens and Waterhole in the Valley of a Dryland Paleoriver" (2018). Zea E-Books. 74. https://digitalcommons.unl.edu/zeabook/74 This Book is brought to you for free and open access by the Zea E-Books at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Zea E-Books by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. The Carnivores of Agate Fossil Beds National Monument Miocene Dens and Waterhole in the Valley of a Dryland Paleoriver Robert M. Hunt, Jr. Robert Skolnick Joshua Kaufman In 1981 University of Nebraska paleontologists came upon an unexpected concentration of carnivore dens at Agate Fossil Beds National Monument in north west Nebraska. The discovery of bones of Miocene beardogs, mustelids, and canids in their burrows was unparalleled and marked an exceptional event in the fossil record. Survey and excavation (1981–1990) established that six species of carnivores had, over time, occupied the dens with traces of their prey: ju venile and adult oreodonts, camels, and a neonatal rhinoceros. At least nine individuals of the wolf-like beardog Daphoenodon superbus, the most common carnivore, were identified from remains of young, mature, and aged individuals that included in one den an adult female and her juvenile male offs pring. The carnivores found together in the dens represent a moment in time—the oldest carnivore den community yet discovered with remains of pred- ators, their prey, and their ecology in evidence. Dated at 22 to 23 Ma (million years), the den complex pro vided scientists with the oldest documented evidence of carnivore denning behavior. Front cover: Skull of Delotrochanter oryktes (UNSM 47800); hills from left to right, North Ridge (with Agate Ash), University Hill, Carnegie Hill, and Beardog Hill at far right, Agate Fossil Beds National Monument, with drawing of Daphoenodon superbus. Back cover: Entrance to Den 1, Quarry 3. ISBN 978-1-60962-143-8 doi 10.32873/unl.dc.zea.1070 The Carnivores of Agate Fossil Beds National Monument Miocene Dens and Waterhole in the Valley of a Dryland Paleoriver Robert M. Hunt, Jr. Robert Skolnick Joshua Kaufman Zea Books Lincoln, Nebraska 2018 Copyright © 2018 Robert M. Hunt, Jr. ISBN 978-1-60962-143-8 doi 10.32873/unl.dc.zea.1070 Composed in Chaparral types. Zea Books are published by the University of Nebraska–Lincoln Libraries Electronic (pdf) edition available online at https://digitalcommons.unl.edu/zeabook/ Print edition available from http://www.lulu.com/spotlight/unllib UNL does not discriminate based upon any protected status. Please go to http://www.unl.edu/equity/ notice-nondiscrimination Visitor Center diorama of the Miocene waterhole, Agate Fossil Beds National Monument Contents List of Illustrations . 6 Preface . 8 Introduction . 11 The Waterhole . 15 Carnegie Quarry 3 (Beardog Hill) . 22 Excavation and Survey Methods . 31 The Den Community . 35 The Dens of Quarry 3 . 41 1981–1982 Excavations . 42 1983 Survey and Mapping. 53 1984 Excavation. 53 1985 Excavation. 56 1989–1990 Excavations . 58 Dens of Living Carnivores . 60 Accumulation of Bones at Dens . 68 Bone-Processing by Carnivores . 72 Discovery of the “Bone Hash” Site . 74 The Carnivores and Their Dens: An Overview . 76 The Carnivore Species . 81 Beardogs (Family Amphicyonidae) . 81 Daphoenodon superbus . 81 Delotrochanter oryktes . 87 Mustelids (Family Mustelidae) . 89 Megalictis simplicidens . 89 Promartes olcotti . 93 Small Dogs (Family Canidae) . 94 Cormocyon, Phlaocyon 94 The Prey and Other Species . 98 Fossils of Merychyus from Quarry 3 . 98 Fossils of Small Camelids and Cervoids from Quarry 3 . 101 Fossils of Menoceras from Quarry 3 . 101 Fossils of Small Mammals from Quarry 3 . 103 Fossils of Lower Vertebrates from Quarry 3 . 103 Epilogue . 106 Key to Mapped Distribution of Taxa . 108 Map C . 108 Map D . 110 Map E . 110 Map F . 111 Map G . 112 4 CONTENTS Map H . 113 Map I . 114 Institutional abbreviations . 114 Introduction to the Tables . 115 Table 1. Fossils Collected in Carnegie Quarry 3 by Olaf A. Peterson and a Carnegie Museum Party (1904–1905) . 116 Table 2. Fossil Material Excavated in Carnegie Quarry 3, Beardog Hill, Agate Fossil Beds National Monument, Nebraska (1981–1990) . 117 Table 3. Postcranial Bones of Carnivores, Quarry 3 . 123 Table 4. Postcrania and Teeth of Ungulates, Quarry 3 . 127 Table 5. Dental Measurements (length × width in mm) of Lower Teeth of Daphoenodon superbus . 129 Table 6. Dental Measurements (length × width in mm) of Upper Teeth of Daphoenodon superbus . 129 Table 7. Dental Measurements (length × width in mm) of Upper Teeth of Delotrochanter oryktes, Megalictis simplicidens, Promartes olcotti, Phlaocyon annectens, and Cormocyon sp. 130 Table 8. Dental Measurements (length × width in mm) of Lower Teeth of Megalictis simplicidens, Promartes olcotti, Phlaocyon annectens, and Cormocyon sp. 130 Table 9. Measurements of Canines (length × width in mm) of Daphoenodon superbus, Megalictis simplicidens, and a Small Carnivore . 131 Table 10. Cranial and Mandibular Measurements of Carnivorans from the Dens . 131 Appendix A: Carnegie Hill Quarries—A Brief History of Excavations of the Bonebed . 133 Appendix B: Reopening of the Carnegie Hill Quarries by the University of Nebraska in 1986. 137 Appendix C: Sediments at Quarry 3 . 138 Appendix D: Analysis of Bone at Quarry 3 . 140 Acknowledgments . 142 References . 143 5 Illustrations Visitor Center diorama . 2-3 Excavation progressing meter by meter in North America in the Early Miocene . 8 the quarry . 32 Geologic Map of the Agate Fossil Beds Map A: The bones and partial skeletons at National Monument . 9 Quarry 3 (1905, 1981–1990) . 33 Carnegie and University Hills—Agate The Den 2 burrow as first exposed. 34 National Monument . 10-11 The carnivores of Quarry 3 . 36 Olaf Peterson . 12 Map B: Distribution of the bones of Erwin Barbour. 12 carnivores and their prey at Quarry 3 . 37 Southwest Excavation on Carnegie Hill Skull and jaw of the beardog Daphoenodon and the sands of the Agate paleoriver . 13 from Quarry 3 . 38 The Agate Bonebed . 13 Bones of prey found with a beardog . 39 Map of the Agate Spring Quarries . 14 Den 1 (burrow B)—unsurpassed among all burrows in its quality of preservation . 40-41 The abandoned stream channel and location of the fossil quarries . 15 Den 1 (burrow A)—demonstrated gradual filling of burrows with sediment . 43 Profile of the Agate waterhole at Carnegie Hill . 16 Discovery of the first evidence of carnivores Chronology of the waterhole . 17 remaining in the dens (1981) . 43 Regional drought in the valley of the The skull of the temnocyonine beardog paleoriver—Mark Marcuson Restoration . 18-19 Delotrochanter from Den 2 . 44 Foot bones of Eurasian immigrant beardog Evidence of the construction of Den 2 . 44 Ysengrinia at the waterhole . 20 Discovering Delotrochanter in the Molar tooth of immigrant Eurasian beardog preparation laboratory of the university . 44 Cynelos . 20 Appearance of Den 2 prior to its excavation The entelodont Dinohyus . 21 and two Daphoenodon foot bones . 45 The small rhinoceros Menoceras. 21 Excavation of the runway of Den 2 that Beardog Hill . 22-23 produced bones of Daphoenodon . 46 Initial trenching on west face of Beardog Hill . 24 Composition of sediment filling the dens . 46 Southern margin of the waterhole identified Layered sediment evident at the terminus in test trenches . 24 of Den 2 . 47 Discovery of Olaf Peterson’s Quarry 3 on Circular thin layers of sand at Den 2 . 47 Beardog Hill . 25 Profile of the face of Den 3 that exposed Josh Kaufman at cliff face with den, entrance bones of the aged Daphoenodon . 48 tunnel, and overlying fossil soil . 26 Excavation of the Den 3 Daphoenodon . 49 Location of Quarry 3 in 1977 . 27 From youth to old age: a male pup, young The providential limb bone: a tibia of female, and aged male at Quarry 3 . 50 Daphoenodon . 27 Recognition of Den 4 . 51 Discovery of the first burrow at Quarry .3 . 28 Excavation of Den 4 with suggestion of Quarry 3 in 1982—burrows begin to appear . 29 chambers dug by pups . 52 Probable location of Peterson’s beardogs in Spotted hyena (Crocuta crocuta) with cub Den 1. 29 at den (Kenya, Africa) . 53 Quarry 3 in 1989—the area in which Discovery of the unexcavated Den 6 in identified burrows occurred . 30-31 the north quarry . 54 Mapping the bones and burrows . 32 Evidence of digging within the burrow at Den 6 . 54 6 ILLUSTRATIONS The Den 5–Den 6 burrow complex during Jaw of Delotrochanter with teeth capable of excavation . 55 crushing bone. 88 Skulls of the mustelid Megalictis and the Bones of the predatory mustelid Megalictis wolverine Gulo . 56 found from 1981–1990 at Quarry 3 . 89 Jaw of Megalictis on floor of burrow in Damaged skull of young Megalictis from laminated sand . 57 Quarry 3 . 90 Comparison of the jaws of Megalictis Comparison of limb bones of Megalictis and Gulo. 57 and living Alaskan wolverine Gulo. 90 Scavenged foot bones of the small canid Claw marks of living badger (Taxidea taxus) Cormocyon . 59 in its burrow . 91 Temperature variation within a burrow on Map D: Distribution of the mustelid the East African plains.
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