UNIVERSITY of CALIFORNIA Los Angeles Carnivory

Total Page:16

File Type:pdf, Size:1020Kb

UNIVERSITY of CALIFORNIA Los Angeles Carnivory UNIVERSITY OF CALIFORNIA Los Angeles Carnivory in the Oligo-Miocene: Resource Specialization, Competition, and Coexistence Among North American Fossil Canids A dissertation submitted in partial satisfaction of the requirements for the degree Doctor of Philosophy in Biology by Mairin Francesca Aragones Balisi 2018 © Copyright by Mairin Francesca Aragones Balisi 2018 ABSTRACT OF THE DISSERTATION Carnivory in the Oligo-Miocene: Resource Specialization, Competition, and Coexistence Among North American Fossil Canids by Mairin Francesca Aragones Balisi Doctor of Philosophy in Biology University of California, Los Angeles 2018 Professor Blaire Van Valkenburgh, Chair Competition likely shaped the evolution of the mammalian family Canidae (dogs and their ancestors; order Carnivora). From their origin 40 million years ago (Ma), early canids lived alongside potential competitors that may have monopolized the large-carnivore niche—such as bears, bear-dogs, and nimravid saber-toothed cats—before becoming large and hypercarnivorous (diet of >70% meat) later in their history. This ecomorphological context, along with high phylogenetic resolution and an outstanding fossil record, makes North American fossil canids an ideal system for investigating how biotic interactions (e.g. competition) and ecological specialization (e.g. hypercarnivory) might influence clade evolution. Chapter 1 investigates whether ecological generalization enables species to have longer durations and broader geographic range. I developed and applied a carnivory index to 100+ ii species, processing 3708 occurrences through a new duration-estimation method accounting for varying fossil preservation. A non-linear relationship between duration and carnivory emerged: both hyper- and hypocarnivores have shorter durations than mesocarnivores. Chapter 2 tests the cost of hypercarnivory, quantified as elevated extinction risk. Large hypercarnivorous canids experienced extinction rates on par with other canids. However, in each canid subfamily, extinction rates rose after the first occurrence of large hypercarnivores, supporting the idea of hypercarnivory as a “macroevolutionary ratchet” for individual clades. Large hypercarnivores originated just over 10 Ma after the origin of Canidae, exhibiting constant diversification rates and peaking in richness around 12 Ma. Relationships emerge between diversification rates of canid subfamilies and temperature, suggesting future hypotheses to be investigated. Chapter 3 quantifies functional morphology in 114 fossil canids and non-canid competitors from 40 to 15 Ma, testing for ecological congruence between continental and regional scales and whether ecomorphological disparity reflects taxonomic diversity. Disparity appears inversely related to diversity. All four geographic regions examined preserved a narrower range of ecomorphology than at the continental scale. These results suggest that partitioning dietary as well as geographic resources, despite constraints to size, permitted canids to minimize competition and diversify. These macroecological and macroevolutionary studies of early canids and their potential competitors improve understanding of the resilience of the carnivore niche over long timescales, including periods of accelerated global change. iii The dissertation of Mairin Francesca Aragones Balisi is approved. Michael Edward Alfaro Paul Jeffrey Brantingham Blaire Van Valkenburgh, Committee Chair University of California, Los Angeles 2018 iv I dedicated my master’s thesis to the memory of my mother, Fe Aragones Balisi, who gave me my first biology lesson, and to the memory of my grandfather, Benjamin Aragones, who left me his National Geographic collection. I now dedicate my Ph.D. dissertation to my father, Mario Luczon Balisi, and my brother, John Paul Aragones Balisi, who have supported me in the pursuit of this degree even when memory faltered. v TABLE OF CONTENTS LIST OF TABLES .................................................................................................................. viii LIST OF FIGURES ................................................................................................................... ix ACKNOWLEDGEMENTS ........................................................................................................ x VITA ....................................................................................................................................... xiv I. DIETARY SPECIALIZATION IS LINKED TO REDUCED SPECIES DURATIONS IN NORTH AMERICAN FOSSIL CANIDS ............................................................................... 1 Abstract................................................................................................................................. 2 Background .......................................................................................................................... 3 Methods ................................................................................................................................ 7 Results ................................................................................................................................ 14 Discussion .......................................................................................................................... 18 Conclusion.......................................................................................................................... 23 Tables and Figures ............................................................................................................ 24 Supplementary Information ............................................................................................. 31 II. LARGE-BODIED HYPERCARNIVORY AS CONVEYOR BELT TO EXTINCTION? DIVERSIFICATION DYNAMICS OF NORTH AMERICAN FOSSIL CANIDS ................ 41 Abstract............................................................................................................................... 42 Background ........................................................................................................................ 43 Methods .............................................................................................................................. 46 Results ................................................................................................................................ 51 Discussion .......................................................................................................................... 55 Conclusions ........................................................................................................................ 59 Tables and Figures ............................................................................................................ 61 Supplementary Information ............................................................................................. 67 III. FUNCTIONAL DIVERGENCE IN BODY SIZE AND DIETARY ADAPTATIONS WITHIN NORTH AMERICAN FOSSIL CARNIVORAN COMMUNITIES ....................... 71 Abstract............................................................................................................................... 72 Background ........................................................................................................................ 73 Methods .............................................................................................................................. 77 Results ................................................................................................................................ 80 Discussion .......................................................................................................................... 82 Conclusions ........................................................................................................................ 86 vi Tables and Figures ............................................................................................................ 87 Supplementary Information ............................................................................................. 92 REFERENCES ......................................................................................................................... 93 vii LIST OF TABLES Table 1.1. Summary statistics and measures of phylogenetic signal.......................................... 24 Table 1.2. Quantile regression results at four levels.................................................................... 24 Table 1.3. Results of nested linear regressions........................................................................... 25 Table 2.1. Median posterior estimates of the parameters al and aµ........................................... 61 Table 2.2. Median posterior estimates of the parameters gl and gµ............................................. 62 Table 3.1. Subdivisions of North American Land Mammal Ages.............................................. 87 Table 3.2. Diversity metrics for each region............................................................................... 87 viii LIST OF FIGURES Figure 1.1. First two axes of principal component analysis......................................................... 26 Figure 1.2. Bivariate plots of emergent properties against intrinsic traits.................................... 27 Figure 1.3. Relationship between species duration and specialization........................................ 28 Figure 1.4. Bivariate plots of species duration as a function of the
Recommended publications
  • The Impact of Large Terrestrial Carnivores on Pleistocene Ecosystems Blaire Van Valkenburgh, Matthew W
    The impact of large terrestrial carnivores on SPECIAL FEATURE Pleistocene ecosystems Blaire Van Valkenburgha,1, Matthew W. Haywardb,c,d, William J. Ripplee, Carlo Melorof, and V. Louise Rothg aDepartment of Ecology and Evolutionary Biology, University of California, Los Angeles, CA 90095; bCollege of Natural Sciences, Bangor University, Bangor, Gwynedd LL57 2UW, United Kingdom; cCentre for African Conservation Ecology, Nelson Mandela Metropolitan University, Port Elizabeth, South Africa; dCentre for Wildlife Management, University of Pretoria, Pretoria, South Africa; eTrophic Cascades Program, Department of Forest Ecosystems and Society, Oregon State University, Corvallis, OR 97331; fResearch Centre in Evolutionary Anthropology and Palaeoecology, School of Natural Sciences and Psychology, Liverpool John Moores University, Liverpool L3 3AF, United Kingdom; and gDepartment of Biology, Duke University, Durham, NC 27708-0338 Edited by Yadvinder Malhi, Oxford University, Oxford, United Kingdom, and accepted by the Editorial Board August 6, 2015 (received for review February 28, 2015) Large mammalian terrestrial herbivores, such as elephants, have analogs, making their prey preferences a matter of inference, dramatic effects on the ecosystems they inhabit and at high rather than observation. population densities their environmental impacts can be devas- In this article, we estimate the predatory impact of large (>21 tating. Pleistocene terrestrial ecosystems included a much greater kg, ref. 11) Pleistocene carnivores using a variety of data from diversity of megaherbivores (e.g., mammoths, mastodons, giant the fossil record, including species richness within guilds, pop- ground sloths) and thus a greater potential for widespread habitat ulation density inferences based on tooth wear, and dietary in- degradation if population sizes were not limited.
    [Show full text]
  • Shape Evolution and Sexual Dimorphism in the Mandible of the Dire Wolf, Canis Dirus, at Rancho La Brea Alexandria L
    Marshall University Marshall Digital Scholar Theses, Dissertations and Capstones 2014 Shape evolution and sexual dimorphism in the mandible of the dire wolf, Canis Dirus, at Rancho la Brea Alexandria L. Brannick [email protected] Follow this and additional works at: http://mds.marshall.edu/etd Part of the Animal Sciences Commons, and the Paleontology Commons Recommended Citation Brannick, Alexandria L., "Shape evolution and sexual dimorphism in the mandible of the dire wolf, Canis Dirus, at Rancho la Brea" (2014). Theses, Dissertations and Capstones. Paper 804. This Thesis is brought to you for free and open access by Marshall Digital Scholar. It has been accepted for inclusion in Theses, Dissertations and Capstones by an authorized administrator of Marshall Digital Scholar. For more information, please contact [email protected]. SHAPE EVOLUTION AND SEXUAL DIMORPHISM IN THE MANDIBLE OF THE DIRE WOLF, CANIS DIRUS, AT RANCHO LA BREA A thesis submitted to the Graduate College of Marshall University In partial fulfillment of the requirements for the degree of Master of Science in Biological Sciences by Alexandria L. Brannick Approved by Dr. F. Robin O’Keefe, Committee Chairperson Dr. Julie Meachen Dr. Paul Constantino Marshall University May 2014 ©2014 Alexandria L. Brannick ALL RIGHTS RESERVED ii ACKNOWLEDGEMENTS I thank my advisor, Dr. F. Robin O’Keefe, for all of his help with this project, the many scientific opportunities he has given me, and his guidance throughout my graduate education. I thank Dr. Julie Meachen for her help with collecting data from the Page Museum, her insight and advice, as well as her support. I learned so much from Dr.
    [Show full text]
  • Mammalia, Felidae, Canidae, and Mustelidae) from the Earliest Hemphillian Screw Bean Local Fauna, Big Bend National Park, Brewster County, Texas
    Chapter 9 Carnivora (Mammalia, Felidae, Canidae, and Mustelidae) From the Earliest Hemphillian Screw Bean Local Fauna, Big Bend National Park, Brewster County, Texas MARGARET SKEELS STEVENS1 AND JAMES BOWIE STEVENS2 ABSTRACT The Screw Bean Local Fauna is the earliest Hemphillian fauna of the southwestern United States. The fossil remains occur in all parts of the informal Banta Shut-in formation, nowhere very fossiliferous. The formation is informally subdivided on the basis of stepwise ®ning and slowing deposition into Lower (least fossiliferous), Middle, and Red clay members, succeeded by the valley-®lling, Bench member (most fossiliferous). Identi®ed Carnivora include: cf. Pseudaelurus sp. and cf. Nimravides catocopis, medium and large extinct cats; Epicyon haydeni, large borophagine dog; Vulpes sp., small fox; cf. Eucyon sp., extinct primitive canine; Buisnictis chisoensis, n. sp., extinct skunk; and Martes sp., marten. B. chisoensis may be allied with Spilogale on the basis of mastoid specialization. Some of the Screw Bean taxa are late survivors of the Clarendonian Chronofauna, which extended through most or all of the early Hemphillian. The early early Hemphillian, late Miocene age attributed to the fauna is based on the Screw Bean assemblage postdating or- eodont and predating North American edentate occurrences, on lack of de®ning Hemphillian taxa, and on stage of evolution. INTRODUCTION southwestern North America, and ®ll a pa- leobiogeographic gap. In Trans-Pecos Texas NAMING AND IMPORTANCE OF THE SCREW and adjacent Chihuahua and Coahuila, Mex- BEAN LOCAL FAUNA: The name ``Screw Bean ico, they provide an age determination for Local Fauna,'' Banta Shut-in formation, postvolcanic (,18±20 Ma; Henry et al., Trans-Pecos Texas (®g.
    [Show full text]
  • Mammalia: Carnivora) in the Americas: Past to Present
    Journal of Mammalian Evolution https://doi.org/10.1007/s10914-020-09496-8 ORIGINAL PAPER Environmental Drivers and Distribution Patterns of Carnivoran Assemblages (Mammalia: Carnivora) in the Americas: Past to Present Andrés Arias-Alzate1,2 & José F. González-Maya3 & Joaquín Arroyo-Cabrales4 & Rodrigo A. Medellín5 & Enrique Martínez-Meyer2 # Springer Science+Business Media, LLC, part of Springer Nature 2020 Abstract Understanding species distributions and the variation of assemblage structure in time and space are fundamental goals of biogeography and ecology. Here, we use an ecological niche modeling and macroecological approach in order to assess whether constraints patterns in carnivoran richness and composition structures in replicated assemblages through time and space should reflect environmental filtering through ecological niche constraints from the Last Inter-glacial (LIG), Last Glacial Maximum (LGM) to the present (C) time. Our results suggest a diverse distribution of carnivoran co-occurrence patterns at the continental scale as a result of spatial climatic variation as an important driver constrained by the ecological niches of the species. This influence was an important factor restructuring assemblages (more directly on richness than composition patterns) not only at the continental level, but also from regional and local scales and this influence was geographically different throughout the space in the continent. These climatic restrictions and disruption of the niche during the environmental changes at the LIG-LGM-C transition show a considerable shift in assemblage richness and composition across the Americas, which suggests an environ- mental filtering mainly during the LGM, explaining between 30 and 75% of these variations through space and time, with more accentuated changes in North than South America.
    [Show full text]
  • Photographic Evidence of a Jaguar (Panthera Onca) Killing an Ocelot (Leopardus Pardalis)
    Received: 12 May 2020 | Revised: 14 October 2020 | Accepted: 15 November 2020 DOI: 10.1111/btp.12916 NATURAL HISTORY FIELD NOTES When waterholes get busy, rare interactions thrive: Photographic evidence of a jaguar (Panthera onca) killing an ocelot (Leopardus pardalis) Lucy Perera-Romero1 | Rony Garcia-Anleu2 | Roan Balas McNab2 | Daniel H. Thornton1 1School of the Environment, Washington State University, Pullman, WA, USA Abstract 2Wildlife Conservation Society – During a camera trap survey conducted in Guatemala in the 2019 dry season, we doc- Guatemala Program, Petén, Guatemala umented a jaguar killing an ocelot at a waterhole with high mammal activity. During Correspondence severe droughts, the probability of aggressive interactions between carnivores might Lucy Perera-Romero, School of the Environment, Washington State increase when fixed, valuable resources such as water cannot be easily partitioned. University, Pullman, WA, 99163, USA. Email: [email protected] KEYWORDS activity overlap, activity patterns, carnivores, interspecific killing, drought, climate change, Funding information Maya forest, Guatemala Coypu Foundation; Rufford Foundation Associate Editor: Eleanor Slade Handling Editor: Kim McConkey 1 | INTRODUCTION and Johnson 2009). Interspecific killing has been documented in many different pairs of carnivores and is more likely when the larger Interference competition is an important process working to shape species is 2–5.4 times the mass of the victim species, or when the mammalian carnivore communities (Palomares and Caro 1999; larger species is a hypercarnivore (Donadio and Buskirk 2006; de Donadio and Buskirk 2006). Dominance in these interactions is Oliveria and Pereira 2014). Carnivores may reduce the likelihood often asymmetric based on body size (Palomares and Caro 1999; de of these types of encounters through the partitioning of habitat or Oliviera and Pereira 2014), and the threat of intraguild strife from temporal activity.
    [Show full text]
  • Carnivore-Specific Stable Isotope Variables and Variation in The
    458 Carnivore-specific stable isotope variables and variation in the foraging ecology of modern and ancient wolf populations: case studies from Isle Royale, Minnesota, and La Brea K. Fox-Dobbs, J.K. Bump, R.O. Peterson, D.L. Fox, and P.L. Koch Abstract: We use carbon and nitrogen isotope data collected from two North American gray wolf (Canis lupus L., 1758) populations (Isle Royale and northern Minnesota) to both calculate carnivore-specific isotopic variables and investigate wolf foraging ecology. The isotopic enrichments of 13C and 15N that occur between mammalian carnivores and their prey have not been well defined in modern populations. We use bone collagen from the Isle Royale National Park wolf, moose (Alces alces (L., 1758)), and beaver (Castor canadensis Kuhl, 1820) populations to determine trophic enrichment factors of 1.3% ± 0.6% for d13C and 4.6% ± 0.7% for d15N. We apply these carnivore-specific fractionation factors to a case study from the fossil record, and reconstruct the diets of late-Pleistocene dire wolves (Canis dirus (Leidy, 1858)) from the La Brea tar pits. We use the Minnesota wolf tissue (collagen, hair, muscle) isotopic data to estimate carnivore population subsample sizes needed to replicate the mean values of the whole population within one standard deviation. Finally, we compare the Isle Royale and Minnesota collagen and hair isotopic data to published d13C and d15N values for North Amer- ican gray wolf populations. We find that interpopulation differences in isotope variances provide insight into wolf foraging ecology. Re´sume´ : Nous utilisons des donne´es sur les isotopes de carbone et d’azote re´colte´es dans deux populations (Isle Royale et nord du Minnesota) de loups gris (Canis lupus L., 1758) pour calculer les variables isotopiques spe´cifiques aux carni- vores et aussi pour e´tudier l’e´cologie alimentaire des loups.
    [Show full text]
  • Paleoecological Comparison Between Late Miocene Localities of China and Greece Based on Hipparion Faunas
    Paleoecological comparison between late Miocene localities of China and Greece based on Hipparion faunas Tao DENG Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing 100044 (China) [email protected] Deng T. 2006. — Paleoecological comparison between late Miocene localities of China and Greece based on Hipparion faunas. Geodiversitas 28 (3) : 499-516. ABSTRACT Both China and Greece have abundant fossils of the late Miocene Hipparion fauna. Th e habitat of the Hipparion fauna in Greece was a sclerophyllous ever- green woodland. Th e Chinese late Miocene Hipparion fauna is represented respectively in the Guonigou fauna (MN 9), the Dashengou fauna (MN 10), and the Yangjiashan fauna (MN 11) from Linxia, Gansu, and the Baode fauna (MN 12) from Baode, Shanxi. According to the evidence from lithology, carbon isotopes, paleobotany, taxonomic framework, mammalian diversity and faunal similarity, the paleoenvironment of the Hipparion fauna in China was a subarid open steppe, which is diff erent from that of Greece. Th e red clay bearing the Hipparion fauna in China is windblown in origin, i.e. eolian deposits. Stable carbon isotopes from tooth enamel and paleosols indicate that C3 plants domi- nated the vegetation during the late Miocene in China. Pollens of xerophilous and sub-xerophilous grasses show a signal of steppe or dry grassland. Forest mammals, such as primates and chalicotheres, are absent or scarce, but grass- land mammals, such as horses and rhinoceroses, are abundant in the Chinese Hipparion fauna. Th e species richness of China and Greece exhibits a similar KEY WORDS trend with a clear increase from MN 9 to MN 12, but the two regions have Hipparion fauna, low similarities at the species level.
    [Show full text]
  • Quaternary Records of the Dire Wolf, Canis Dirus, in North and South America
    Quaternary records of the dire wolf, Canis dirus, in North and South America ROBERT G. DUNDAS Dundas, R. G. 1999 (September): Quaternary records of the dire wolf, Canis dirus, in North and South Ameri- ca. Boreas, Vol. 28, pp. 375–385. Oslo. ISSN 0300-9483. The dire wolf was an important large, late Pleistocene predator in North and South America, well adapted to preying on megaherbivores. Geographically widespread, Canis dirus is reported from 136 localities in North America from Alberta, Canada, southward and from three localities in South America (Muaco, Venezuela; Ta- lara, Peru; and Tarija, Bolivia). The species lived in a variety of environments, from forested mountains to open grasslands and plains ranging in elevation from sea level to 2255 m (7400 feet). Canis dirus is assigned to the Rancholabrean land mammal age of North America and the Lujanian land mammal age of South Amer- ica and was among the many large carnivores and megaherbivores that became extinct in North and South America near the end of the Pleistocene Epoch. Robert G. Dundas, Department of Geology, California State University, Fresno, California 93740-8031, USA. E-mail: [email protected]; received 20th May 1998, accepted 23rd March 1999 Because of the large number of Canis dirus localities Rancho La Brea, comparing them with Canis lupus and and individuals recovered from the fossil record, the dire wolf specimens from other localities. Although dire wolf is the most commonly occurring large knowledge of the animal’s biology had greatly predator in the Pleistocene of North America. By increased by 1912, little was known about its strati- contrast, the species is rare in South America.
    [Show full text]
  • University of Florida Thesis Or Dissertation Formatting
    UNDERSTANDING CARNIVORAN ECOMORPHOLOGY THROUGH DEEP TIME, WITH A CASE STUDY DURING THE CAT-GAP OF FLORIDA By SHARON ELIZABETH HOLTE A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA 2018 © 2018 Sharon Elizabeth Holte To Dr. Larry, thank you ACKNOWLEDGMENTS I would like to thank my family for encouraging me to pursue my interests. They have always believed in me and never doubted that I would reach my goals. I am eternally grateful to my mentors, Dr. Jim Mead and the late Dr. Larry Agenbroad, who have shaped me as a paleontologist and have provided me to the strength and knowledge to continue to grow as a scientist. I would like to thank my colleagues from the Florida Museum of Natural History who provided insight and open discussion on my research. In particular, I would like to thank Dr. Aldo Rincon for his help in researching procyonids. I am so grateful to Dr. Anne-Claire Fabre; without her understanding of R and knowledge of 3D morphometrics this project would have been an immense struggle. I would also to thank Rachel Short for the late-night work sessions and discussions. I am extremely grateful to my advisor Dr. David Steadman for his comments, feedback, and guidance through my time here at the University of Florida. I also thank my committee, Dr. Bruce MacFadden, Dr. Jon Bloch, Dr. Elizabeth Screaton, for their feedback and encouragement. I am grateful to the geosciences department at East Tennessee State University, the American Museum of Natural History, and the Museum of Comparative Zoology at Harvard for the loans of specimens.
    [Show full text]
  • La Brea and Beyond: the Paleontology of Asphalt-Preserved Biotas
    La Brea and Beyond: The Paleontology of Asphalt-Preserved Biotas Edited by John M. Harris Natural History Museum of Los Angeles County Science Series 42 September 15, 2015 Cover Illustration: Pit 91 in 1915 An asphaltic bone mass in Pit 91 was discovered and exposed by the Los Angeles County Museum of History, Science and Art in the summer of 1915. The Los Angeles County Museum of Natural History resumed excavation at this site in 1969. Retrieval of the “microfossils” from the asphaltic matrix has yielded a wealth of insect, mollusk, and plant remains, more than doubling the number of species recovered by earlier excavations. Today, the current excavation site is 900 square feet in extent, yielding fossils that range in age from about 15,000 to about 42,000 radiocarbon years. Natural History Museum of Los Angeles County Archives, RLB 347. LA BREA AND BEYOND: THE PALEONTOLOGY OF ASPHALT-PRESERVED BIOTAS Edited By John M. Harris NO. 42 SCIENCE SERIES NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY SCIENTIFIC PUBLICATIONS COMMITTEE Luis M. Chiappe, Vice President for Research and Collections John M. Harris, Committee Chairman Joel W. Martin Gregory Pauly Christine Thacker Xiaoming Wang K. Victoria Brown, Managing Editor Go Online to www.nhm.org/scholarlypublications for open access to volumes of Science Series and Contributions in Science. Natural History Museum of Los Angeles County Los Angeles, California 90007 ISSN 1-891276-27-1 Published on September 15, 2015 Printed at Allen Press, Inc., Lawrence, Kansas PREFACE Rancho La Brea was a Mexican land grant Basin during the Late Pleistocene—sagebrush located to the west of El Pueblo de Nuestra scrub dotted with groves of oak and juniper with Sen˜ora la Reina de los A´ ngeles del Rı´ode riparian woodland along the major stream courses Porciu´ncula, now better known as downtown and with chaparral vegetation on the surrounding Los Angeles.
    [Show full text]
  • Download PDF File
    1.08 1.19 1.46 Nimravus brachyops Nandinia binotata Neofelis nebulosa 115 Panthera onca 111 114 Panthera atrox 113 Uncia uncia 116 Panthera leo 112 Panthera pardus Panthera tigris Lynx issiodorensis 220 Lynx rufus 221 Lynx pardinus 222 223 Lynx canadensis Lynx lynx 119 Acinonyx jubatus 110 225 226 Puma concolor Puma yagouaroundi 224 Felis nigripes 228 Felis chaus 229 Felis margarita 118 330 227 331Felis catus Felis silvestris 332 Otocolobus manul Prionailurus bengalensis Felis rexroadensis 99 117 334 335 Leopardus pardalis 44 333 Leopardus wiedii 336 Leopardus geoffroyi Leopardus tigrinus 337 Pardofelis marmorata Pardofelis temminckii 440 Pseudaelurus intrepidus Pseudaelurus stouti 88 339 Nimravides pedionomus 442 443 Nimravides galiani 22 338 441 Nimravides thinobates Pseudaelurus marshi Pseudaelurus validus 446 Machairodus alberdiae 77 Machairodus coloradensis 445 Homotherium serum 447 444 448 Smilodon fatalis Smilodon gracilis 66 Pseudaelurus quadridentatus Barbourofelis morrisi 449 Barbourofelis whitfordi 550 551 Barbourofelis fricki Barbourofelis loveorum Stenogale Hemigalus derbyanus 554 555 Arctictis binturong 55 Paradoxurus hermaphroditus Genetta victoriae 553 558 Genetta maculata 559 557 660 Genetta genetta Genetta servalina Poiana richardsonii 556 Civettictis civetta 662 Viverra tangalunga 661 663 552 Viverra zibetha Viverricula indica Crocuta crocuta 666 667 Hyaena brunnea 665 Hyaena hyaena Proteles cristata Fossa fossana 664 669 770 Cryptoprocta ferox Salanoia concolor 668 772 Crossarchus alexandri 33 Suricata suricatta 775
    [Show full text]
  • PHYLOGENETIC SYSTEMATICS of the BOROPHAGINAE (CARNIVORA: CANIDAE) Xiaoming Wang
    University of Nebraska - Lincoln DigitalCommons@University of Nebraska - Lincoln Mammalogy Papers: University of Nebraska State Museum, University of Nebraska State Museum 1999 PHYLOGENETIC SYSTEMATICS OF THE BOROPHAGINAE (CARNIVORA: CANIDAE) Xiaoming Wang Richard H. Tedford Beryl E. Taylor Follow this and additional works at: http://digitalcommons.unl.edu/museummammalogy This Article is brought to you for free and open access by the Museum, University of Nebraska State at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Mammalogy Papers: University of Nebraska State Museum by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. PHYLOGENETIC SYSTEMATICS OF THE BOROPHAGINAE (CARNIVORA: CANIDAE) XIAOMING WANG Research Associate, Division of Paleontology American Museum of Natural History and Department of Biology, Long Island University, C. W. Post Campus, 720 Northern Blvd., Brookville, New York 11548 -1300 RICHARD H. TEDFORD Curator, Division of Paleontology American Museum of Natural History BERYL E. TAYLOR Curator Emeritus, Division of Paleontology American Museum of Natural History BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY Number 243, 391 pages, 147 figures, 2 tables, 3 appendices Issued November 17, 1999 Price: $32.00 a copy Copyright O American Museum of Natural History 1999 ISSN 0003-0090 AMNH BULLETIN Monday Oct 04 02:19 PM 1999 amnb 99111 Mp 2 Allen Press • DTPro System File # 01acc (large individual, composite figure, based Epicyon haydeni ´n. (small individual, based on AMNH 8305) and Epicyon saevus Reconstruction of on specimens from JackNorth Swayze America. Quarry). Illustration These by two Mauricio species Anto co-occur extensively during the late Clarendonian and early Hemphillian of western 2 1999 WANG ET AL.: SYSTEMATICS OF BOROPHAGINAE 3 CONTENTS Abstract ....................................................................
    [Show full text]