the natural history of HOEWENEGG A SKELETON, BERNOR ET AL. 1997 HORSES

RAY BERNOR,COLLEGE OF MEDICINE, DEPARTMENT OF ANATOMY, HOWARD UNIVERSITY, WASHINGTON D.C. the natural history of HORSES •BRIEF HISTORY OF EQUID RESEARCH •FIELD-BASED RESEARCH BEARING ON EVOLUTIONARY HISTORY OF O.W. HIPPARIONS •BIOGEOGRAPHIC PROVINCIALITY AND ITS BEARING ON EQUID EVOLUTION •RECONSTRUCTING PALEODIET •EVOLUTIONARY STUDIES OF THE GENUS EQUUS •ANATOMICAL AND BIOLOGICAL BASES OF RACE HORSE FOOT INJURIES • MOLECULAR AND PALEONTOLOGICAL ON EQUUS CONTRARY TO MIVART, Darwin DID not BELIEVE THAT THE EXTINCT THREE-TOED HORSE Hipparion evolved suddenly (DARWIN, 1872: 201) Gaudry 1866 T.H. Huxley 1876

VISITS YALE PEABODY MUSEUM TO DISCUSS HORSE EVOLUTION WITH O.C. MARSH, 1876 GAUDRY EARLY STUDENTS OF EQUID EVOLUTION

MATTHEW

MARSH

OSBORN

STIRTON Orthogenetic Evolution

MATTHEW, 1903, 1926 STIRTON, 1940 George G. Simpson MacFadden, 2005 Science Vol. 307. no. 5716, pp. 1728 - 1730 Maragheh, N.W. Iran, 1974 Maragheh Field Team, 1976 Maragheh Geochronology 1980-1996

7.4 7.67.8

8.2 8.28.6 9.0 Maragheh Hipparion Evolution

Cr. matthewi

7.9 aff. H. brachypus Cr. moldavicum

8.2 8.6

Taxa in red after Bernor et al. 2016

9.0 Old World Later Neogene Provinciality: Cluster Analyses on Genus Faunal Resemblances

BERNOR, BERNOR AND PAVLAKIS, 1978 1987 BERNOR, 1983

EASTERN AFRICA

PROVINCIALITY AND ZOOGEOGRAPHIC EVOLUTION OF THE OLD WORLD LATER NEOGENE BIOME 16-5.2 MA The Late Miocene Pikermian Chronofauna: The Samos family portrait

Nikos 1993 Samos Horse Lineages: 10 taxa 8-6.7 Ma

6.7 Ma

Cr. mediterraneum

Hippotherium sp. Cr. matthewi

H. dietrichi By: G. Koufos and D. Vlachou 8.2 Ma Cr. proboscideum Raup-Crick Genus Faunal Resemblance Index Values as Compared to Pikermi (MN 11, ca. 8.0 Ma)

MN 7+8 MN 9 MN 12 MIO-PLIOCENE BOUNDARY = MN13/14 MN 14 (5.3-4.2) MN 13 (6.6-5.3) MN 12 (8.0-6.6) MN 11 (8.7-8.0) MN 10 (9.7-8.7) MN 9 (11.1-9.7) MN 13 MN 14 MN 7+8 (13.5-11.1)

ERONEN, MIRZAIE, KARME, MICHEELS, BERNOR AND FORTELIUS,2009, PNAS Hegau District, S.W. Germany HEGAU DISTRICT, SOUTHERN GERMANY

GRABUNG HOEWENEGG Hoewenegg Excavations in the 1950s and 60s Ho E Skeleton With Fetus in Situs Utero RENEWED RESEARCH

1986

1991

1992

2001-2016 TOBIEN REVISIT IN 1986 1986 TRENCH 1991 DISCOVERY OF MIOTRAGOCERUS SKULL Section Skeletons Reopened

in 2003 Hoewenegg 2003 Miotragocerus with 2 fetuses in situs utero 2004 Extension of the Site to the NW 2005 - Extending the Excavation to the West Hoewenegg West Quarry Biotic Elements Collected 2004-2006 Field Seasons Exclusive of Gastropods

38 4 21 Plants

18 Crustacea 2 Vertebrata indet.

Pices

Amphibia

81 Reptilia 266 Aves

Mammalia 2 Figure 11 - Proportions of Biotic Contents by Stratigraphic Level

100%

Mammalia 80% Aves

Reptilia

60% Amphibia

Pices

Vert. indet. 40% Insecta

Crustacea 20% Gastropoda Plants

0% 5 7 9 Sol 11 13 15 17 19 Proportion of Hoewenegg Mammal Species Skeletons

5% 2% 2% Hippotherium 7%

Miotragocerus 37%

Rhinos

Deinotherium

Dorcatherium

Muntiak 47% Stratigraphic Provenance of the Höwenegg Skeletons – The Jorg and Tobien Stratigraphic Numbering System

HORIZO Hippotherium Miotragocerus Rhinos Deinotherium Dorcatherium Muntjak N 18 1 20 6 (2) 9 (6, f) 1 1 (1) 1 22a 25 4 (f) 2 1 1 27 1 3 32 3 2 34 2 42 1 Hoewenegg Lageplan 2012 Central European Vallesian Habitats 11-9.7 Ma

BERNOR ET AL., 1988 VEGETATION RECONSTRUCTION BY JOHANNA EDER

GYRAULUS KLEINI ULMUS

Populus TROPIDOMPHALUS (P.) ZELLI HOEWENEGG FLORA & INVERTEBRATES Hoewenegg Hippotherium primigenium 10.3 Ma.

POF-LACRIMAL SPINOUS PROCESSES SACRUM

TEETH

LIMBS COMPOSITE CHRONOLOGY OF THE SINAP FORMATION FOSSIL LOCALITIES, TURKEY 11-9 MA

DETAIL OF SINAP TEPE "Hipparion" uzunagizli 9.97 Cormohipparion "Hipparion" sp. sinapensis cursorial form 10.692 10.08 "Hipparion" kecigibi "Hipparion" Old World 9.92 sp. large Hipparions 9.59

Cormohipparion sp.(Woodburne, 2007&2009) 11 Ma

40 11.2-6MA

11.2 10.08-9.59 11.2 10.7

10.5 10.7

10.5 A Systematic, Biostratigraphic, and Paleogeographic Reevaluation of the Siwalik Hipparion Horse Assemblage from the Potwar Plateau, Northern Pakistan

- an application of paleontological and paleoecological methodology to the assessment of a regional population of hipparionine horses

Wolf, 2012. Siwalik Taxonomy – Sivalhippus perimense

Wolf, Bernor and Hussain, 2013) Sivalhippus theobaldi

Bernor et al. (2010)

AMNH 98728

GSI C153; holotype

MacFadden and Woodburne (1982) Stratigraphic Distribution of Siwalik Hipparionines

Wolf, Bernor and Hussain, 2013 MAJOR OLD WORLD LATE MIOCENE CLADES 1996 Plio-Pleistocene Lineages: Eurygnathohippus, Plesiohipparion, Proboscidipparion DIACHRONEITY OF CROWN HEIGHT SPIKES IN OLD WORLD HIPPARIONS 2013

11.4 Ma = 50 MM 2 MA > 90 MM

6.7 MA = 10.5 = 85 MM 60 MM

8.5 MA = 70 MM

1.2 MA = 90 MM

5 MA = 80 MM Horses Eat Grass MESOWEAR ANALYSIS Fortelius and Solounias, 2000, Mihlbachler et al., 2011, Wolf et al., 2012

0 0 Sharp CUSPS & HIGH RELIEF 0.51 2 1 Rounded CUSPS & HIGH RELIEF 1.53

2 4 Rounded CUPS & LOW RELIEF 2.55 Blunt CUSPS & LOW 6 RELIEF 3

VARIABLES: RELIEF RELIEF AND SHAPE Old World Hipparionine Diets

EURYGNATHOHIPPUS CORNELIANUS DINOTHERIENSANDE 1.4 MA 10 MA

CLUSTER ANALYSIS OF MESOWEAR PARAMETER FREQUENCIES IN 27 EXTANT UNGULATE SPECIES AND 10 HIPPARIONINE POPULATIONS (Methodology Following Fortelius & Solounias, 2000; Bernor & Kaiser 2006 a & b; Bernor et al., 2004)

Bison bison CERATOTHERIUM SIMUM EQUUS BURCHELLI EQUUS GREVYI DAMALISCUS LUNATUS GRAZER Eurygnathohippus (LANGEBAANWEG & SAHABI) CONNOCHAETES TAURINUS Alcelaphus buselaphus HIPPOTRAGUS NIGER Redunca redunca HIPPOTRAGUS EQUINUS KOBUS ELLIPSIPRYMNUS Cormohipparion goorisi Cormohipparion quinni Merychippus insignis (BALTAVAR) AEPYCEROS MELAMPUS CAPRICORNIS SUMATRAENSIS CERVUS CANADENSIS Cormohipparion sp. (CHORORA) MIXED TAUROTRAGUS ORYX TRAGELAPHUS SCRIPTUS FEEDER Hippotherium prim/intrans (DD & RUD) GAZELLA GRANTI GAZELLA THOMSONI OVIBOS MOSCHATUS GIRAFFA CAMELOPARDALIS ODOCOILEUS HEMIONUS Hippotherium primigenium (HOEWENEGG) DICERORHINUS SUMATRENSIS OKAPIA JOHNSTONI ODOCOILEUS VIRGINIANUS DICEROS BICORNIS BROWSER ALCES ALCES RHINOCEROS SONDAICUS

NED 0 10 20 30 40 50 60 70 80 90 100 SCRATCHES

PITS LIGHT MICROSCOPE MICROWEAR

Courtesy of Gina Semprebon (SCALE BAR = 0.4 MM) Phylogeny of Equus Azzaroli, 2003 Equus asinus

OLD WORLD GEOGRAPHY OF LIVING EQUUS SPECIES

Mitochondrial Research 2005 Leonard, Rohland, Glaberman, Fleischer, Caccone and Hofreiter Barbaro and 8 Bells

BARBARO AT THE DERBY 8 BELLS AT THE DERBY

Jockey Gabriel Saez "heard the worst sound possible - a pop” [the Huffington Post, October 6, 2008 BARBARO BREAK AND THERAPY ! DR. CHRIS KAWCAK DVM, COLORADO STATE VETERINARY SCHOOL ANATOMY OF RACE HORSE FOOT FRACTURE

DR. KAWCAK’S FACT-LIST • 97 % OF ALL FRACTURES ARE TO THE LATERAL CONDYLE OF MPIII [MORE FORELIMB THAN HINDLIMB] FOLLOWED BY FRACTURES OF THE SESAMOID AND THE PASTERN [1ST + 2ND PHALANGES]

• FRACTURES ARE FOUND IN HORSES WITH LONG MPIII’S

THERE IS REDUCED VOLUME [=SIZE] OF LATERAL CONDYLES THAT BREAK

• A “GOOD HORSE” HAS SHORT METAPODIALS Do Race Horses Need 3 Toes?

A. The Horse has MPII & IV splints with fibrous connection to MPIII

B. Hipparions have fully functional MPIII/MPII & IV joints and complete digits II and IV

C. MP IV is more stoutly developed than MPII giving greater support laterally than medially Diagrammatic expression maps for 50 genes of the Hox A (top) and Hox D (bottom) clusters during early, middle and late stages of forelimb outgrowth. Reno et al., 2007 Fig. 2 Anatomical Maps of the Autopod (wrist and hand) Reno et al., 2007 Fig. 3 CONSERVE THE SOMALIAN ASS

THANK YOU PSW, 02-15-2017 END