Hipparion tracks and horses’ toes: the evolution of the equid single hoof.
Abstract: The traditional story of the evolution of the horse (family Equidae) has been in large part about the evolution of their feet. How did modern horses come to have a single toe (digit III), with the hoof bearing a characteristic V-shaped keratinous frog on the sole, and what happened to the other digits? While it has long been known that the proximal portions of digits II and IV are retained as the splint bones, a recent hypothesis suggested that the distal portion of these digits have also been retained as part of the frog, drawing upon the famous Laetoli footprints of the tridactyl (three-toed) equid as part of the evidence. We show here that, while there is good anatomical and embryological evidence for the proximal portions of all the accessory digits (i.e. I and V, as well as II and IV) being retained in the feet of modern horses, evidence is lacking for the retention of any distal portions of these digits. There is also good ichnological evidence that many tridactyl equids possessed a frog, and that the frog has been part of the equid foot for much of equid evolutionary history.
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Publication Date: 2023-06-21 PubMed ID: 37351494PubMed Central: PMC10282582DOI: 10.1098/rsos.230358Google Scholar: Lookup
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Summary
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The research article discusses the evolution of the horse’s foot, mainly focusing on how the modern horse has transitioned to a single-toe structure, with a characteristic V-shaped keratinous frog part on the sole, from a three-toe structure. The researchers further investigate an hypothesis suggesting that the remains of the distal portions of the vanished digits form part of the frog.
Introduction
- The research provides a detailed insight into the evolution of the horse’s foot, coming from the family Equidae.
- It mainly focuses on how horses have evolved from having three toes (digits II, III, and IV) to just one toe (digit III) with a unique V-shaped keratinous component on the sole known as the ‘frog’.
- Then, it conducts a deep dive into an hypothesis that suggests that the frog on a horse’s foot could contain remnants of the distal portions of the lost digits.
The Evolution of the Horse Foot
- The researchers indicate that horse’s fee evolution has traditionally been a major fascination in the realm of biological and evolutionary research.
- They focus on how modern horses, instead of having three separate toes like their ancestors, now possess a single toe (digit III).
- A probing question is what happened to the missing digits. It has been established that the proximal parts of the absent digits are retained as splint bones.
The Hypothesis on Digit Retainment
- The researchers examine a recent hypothesis suggesting that not only the proximal parts, but also the distal parts of the disappeared digits might be retained as a part of the frog.
- To back up this hypothesis, they refer to the renowned Laetoli footprints of the tridactyl (three-toed) equine species.
The Findings
- The researchers find compelling anatomical and embryological evidence supporting the retainment of the proximal parts of all the accessory digits (i.e. I and V, as well as II and IV) in the feet of modern horses.
- The evidence regarding the distal retainment of these discontinued digits in the form of the ‘frog’ is, however, inadequate.
- They also establish that many three-toed equines had a frog, implying that the frog has been a part of the equine foot throughout most of its evolutionary history.
Cite This Article
APA
Vincelette AR, Renders E, Scott KM, Falkingham PL, Janis CM.
(2023).
Hipparion tracks and horses’ toes: the evolution of the equid single hoof.
R Soc Open Sci, 10(6), 230358.
https://doi.org/10.1098/rsos.230358 Publication
Researcher Affiliations
- Department of Pathology, St. John's Seminary, Camarillo, 93012, CA, USA.
- Department of Functional Morphology, Faculty of Veterinary Medicine, Utrecht University (Ret.), Utrecht, 3584 CM, The Netherlands.
- Department of Cell Biology and Neuroscience, Rutgers University, New Brunswick, 08854, NJ, USA.
- School of Biological and Environmental Sciences, Liverpool John Moores University, Liverpool, L3 3AF, UK.
- Bristol Palaeobiology Group, School of Earth Sciences, University of Bristol, Bristol, BS8 1RJ, UK.
- Department of Ecology and Evolutionary Biology, Brown University, Providence, 02912, RI, USA.
Conflict of Interest Statement
We declare we have no competing interests.
References
This article includes 140 references
- Robb RC. A study of mutations in evolution, III, The evolution of the equine foot.. J. Genet. 33, 267-273.
- Forsten A. Adaptive evolution of the equine foot.. Z. Säugetierkd. 40, 304-308.
- Hussain ST. Evolutionary and functional anatomy of the pelvic limb in fossil and recent Equidae (Perissodactyla, Mammalia).. Zentralbl Veterinarmed C 1975 Sep;4(3):193-222 concl.
- MacFadden BJ. Fossil horses, systematics, paleobiology, and evolution of the family Equidae.. New York, NY: Cambridge University Press.
- De Vos J, Sondaar PY, Reumer JWF. The evolution of hominid bipedalism.. Anthropologie 36, 5-16.
- Floyd AE. Evolution of the equine digit and its relevance to the modern horse.. In Equine podiatry (eds Floyd AE, Mansmann RA), pp. 102-111. St. Louis, MO: Saunders.
- Wood AR, Bebej RM, Manz CL, Begun DL, Gingerich PD. Postcranial functional morphology of Hyracotherium (Equidae, Perissodactyla) and locomotion in the earliest horses.. J. Mamm. Evol. 18, 1-32.
- MacLaren JA, Nauwelaerts S. Interspecific variation in the tetradactyl manus of modern tapirs (Perissodactyla: Tapirus) exposed using geometric morphometrics.. J Morphol 2017 Nov;278(11):1517-1535.
- MacLaren JA, Nauwelaerts S. Modern tapirs as morphofunctional analogues for locomotion in endemic Eocene European perissodactyls.. J. Mamm. Evol. 27, 245-263.
- Marsh OC. Fossil horses in America.. Am. Nat. 8, 288-294.
- Janis CM. Asymmetry of evolutionary patterns between New World and Old World equids and among New World equine tribes.. In Evolution of Cenozoic land mammal faunas and ecosystems: 25 years of the NOW database of fossil mammals (eds Casanovas I, Janis CM, van der Hoek Ostend LW, Saarinen J), pp. 143-164. New York, NY: Springer.
- Singh B. Dyce, Sack, and Wensing's textbook of veterinary anatomy, 5th edn.. St. Louis, MO: Elsevier.
- Dzierzęcka M, Purzyc H, Charuta A, Barszcz K, Komosa M, Hecold M, Kłosińska D. Evaluation of distal phalanx formation and association with front hoof conformation in coldblooded horses.. Biologia 71, 337-342.
- Carlson N. Foreign body penetration of the hoof.. In Comparative veterinary anatomy (eds Orsini J, Greneger N, de Lahunta A), pp. 918-924. St. Louis, MO: Elsevier.
- Bhattacharjee S. The osseous fusion patterns in an equine limb: A theoretical deconstruction of the evolutionary mechanisms.. Biosystems 2023 Mar;225:104871.
- Simpson GG. Horses: the story of the horse family in the modern world and through sixty million years.. Oxford, UK: Oxford University Press.
- Janis CM, Bernor RL. The evolution of equid monodactyly: a review including a new hypothesis.. Front. Ecol. Evol. 12, 1-19.
- Solounias N, Danowitz M, Stachtiaris E, Khurana A, Araim M, Sayegh M, Natale J. The evolution and anatomy of the horse manus with an emphasis on digit reduction.. R Soc Open Sci 2018 Jan;5(1):171782.
- Falkingham P. A standard protocol for documenting modern and fossil ichnological data.. Front. Palaeontol. 61, 469-480.
- Camp CL, Smith N. Phylogeny and function of the digital ligaments of the horse.. Mem. Univ. Calif. 13, 64-124.
- Sondaar PY. The osteology of the manus of fossil and recent Equidae with special reference to phylogeny and function.. Verh. Konik. Ned. Akad. Wet. 25, 1-76.
- Thomason JJ. The relationship of trabecular architecture to inferred loading patterns in the third metacarpals of the extinct equids Merychippus and Mesohippus.. Paleobiology 11, 323-335.
- Thomason JJ. The functional morphology of the manus in the tridactyl equids Merychippus and Mesohippus: paleontological inferences from neontological models.. J. Vertebr. Paleontol. 6, 143-161.
- Tobien H. Über die Funktion der Seitenzehen tridactyler Equiden.. Neues Jahrb. Geol. Paleontol. 96, 137-172.
- Biewener AA. Muscle-tendon stresses and elastic energy storage during locomotion in the horse.. Comp Biochem Physiol B Biochem Mol Biol 1998 May;120(1):73-87.
- Rooney JR, Quddus MA, Kingsbury HB. A laboratory investigation of the function of the stay apparatus of the equine foreleg.. J. Equine Med. Surg. 2, 173-180.
- Pollitt CC. The anatomy and physiology of the suspensory apparatus of the distal phalanx.. Vet Clin North Am Equine Pract 2010 Apr;26(1):29-49.
- Pollitt CC, Collins SN. The suspensory apparatus of the distal phalanx in normal horses.. Equine Vet J 2016 Jul;48(4):496-501.
- Abel O. Die Stamme der Wirbeltiere.. Berlin, Germany: Walter de Gruyter & Co.
- Janis CM, Franklin E, Baird CN, Tyler J. The Miocene browsing horses: another way to be a successful large equid.. In The equids, a successful group of species (eds Prins HHT, Gordon IJ). New York, NY: Springer.
- O'Sullivan JA. Evolution of the proximal third phalanx in Oligocene and Miocene equids, and the utility of phalangeal indices in phylogeny reconstruction.. In Mammalian evolutionary morphology, a tribute to Fred Szalay (eds Sargis EJ, Dagosto M), pp. 159-165. New York, NY: Springer.
- McHorse BK, Biewener AA, Pierce SE. Mechanics of evolutionary digit reduction in fossil horses (Equidae).. Proc Biol Sci 2017 Aug 30;284(1861).
- Ewart JC. The Development of the Skeleton of the Limbs of the Horse, with Observations on Polydactyly.. J Anat Physiol 1894 Jan;28(Pt 2):236-56.
- Kavanagh KD, Bailey CS, Sears KE. Evidence of five digits in embryonic horses and developmental stabilization of tetrapod digit number.. Proc Biol Sci 2020 Feb 12;287(1920):20192756.
- Tucker ST, Otto RE, Joeckel RM, Voorhies MR. The geology and paleontology of Ashfall Fossil Beds: a late Miocene (Clarendonian) mass-death assemblage, Antelope County and adjacent Knox County, Nebraska, USA.. In Geologic field trips along the boundary between the central lowlands and great plains, 2014 meeting of the GSA north-central section (eds Koris JT), pp. 1-22. Boulder, CO: Geological Society of America.
- Renders E. The gait of Hipparion sp. from fossil footprints in Laetoli, Tanzania.. Nature 1984 Mar 8-14;308(5955):179-81.
- Renders E, Sondaar PY. Animal prints and trails, Hipparion.. In Laetoli: a Pliocene site in northern Tanzania (eds Leakey MD, Harris JM), pp. 471-481. Oxford, UK: Clarendon Press.
- McHorse BK, Biewener AA, Pierce SE. The Evolution of a Single Toe in Horses: Causes, Consequences, and the Way Forward.. Integr Comp Biol 2019 Sep 1;59(3):638-655.
- Lull RS. The evolution of the horse family as illustrated in the Yale collections.. Am. J. Sci. 23, 161-182.
- Osborn HF. Equidae of the Oligocene, Miocene, and Pliocene of North America, iconographic type revision.. Mem. Am. Mus. Nat. Hist. 2, 1-217.
- Stock C. Neohipparion leptode (Merriam) from the Pliocene of northwestern Nevada.. Am. J. Sci. 249, 430-438.
- MacFadden BJ. Systematics and phylogeny of the Hipparion, Neohipparion, Nannippus, and Cormohipparion (Mammalia, Equidae) from the Miocene and Pliocene of the New World.. B. Am. Mus. Nat. Hist. 179, 1-195.
- Bernor RL, Tobien H, Hayek LAC, Mittmann HL. Hippotherium primigenium (Equidae, Mammalia) from the late Miocene of Höwenegg (Hegao, Germany).. Andrias 10, 1-230.
- Lazaridis G, Tsoukala E. Hipparion phlegrae, sp. nov. (Mammalia, Perissodactyla): a new species from the Turolian locality of Kryopigi (Kassandra, Chalkidiki, Greece).. J. Vertebr. Paleontol. 34, 164-178.
- Li Y, Deng T, Hua H, Sun B, Zhang Y. Locomotor adaptations of 7.4 Ma hipparionine fossils from the middle reaches of the Yellow River and their palaeoecological significance.. Hist. Biol. 33, 927-940.
- Walther K. Der Bewegungsablauf der freien Gliedmaßen des Pferdes im Schritt, Trab und Galopp.. Arch. Wiss. Prak. Tierheilk. 53, 316-352.
- Krüger W. Über die Arbeit der vier Gliedmaßen des Pferdes beim Galopp.. Tierarztl. Rundsch 45, 250-255.
- Ratzlaff MH, Wilson PD, Hyde ML, Balch OK, Grant BD. Relationship between locomotor forces, hoof position and joint motion during the support phase of the stride of galloping horses.. Acta Anat (Basel) 1993;146(2-3):200-4.
- Clayton HM, Singleton W, Lanovaz JL, Prades M. Pastern joint motion in trotting horses.. P. Am. Assoc. Equine Practit. 46, 217-219.
- Clayton HM, Sha D, Stick J, Elvin N. 3D kinematics of the equine metacarpophalangeal joint at walk and trot.. Vet Comp Orthop Traumatol 2007;20(2):86-91.
- Butcher MT, Ashley-Ross MA. Fetlock joint kinematics differ with age in Thoroughbred [was thoroughbred] racehorses.. J Biomech 2002 May;35(5):563-71.
- Setterbo J, Garcia TC, Campbell I, Kim S, Hubbard M, Stover SM, Estivalet M, Brisson P. Forelimb kinematics of galloping thoroughbred racehorses measured on dirt, synthetic, and turf track surfaces (P235).. In The engineering of sport 7, pp. 437-446. Paris, France: Springer.
- Crevier-Denoix N, Falala S, Holden-Douilly L, Camus M, Martino J, Ravary-Plumioen B, Vergari C, Desquilbet L, Denoix JM, Chateau H, Pourcelot P. Comparative kinematic analysis of the leading and trailing forelimbs of horses cantering on a turf and a synthetic surface.. Equine Vet J Suppl 2013 Dec;(45):54-61.
- Solé M, Gómez MD, Martínez Galisteo A, Santos R, Valera M. Kinematic characterization of the Menorca horse at the walk and the trot: influence of hind limb pastern angle.. J. Equine Vet. Sci. 33, 726-732.
- Symons JE, Garcia TC, Stover SM. Distal hindlimb kinematics of galloping Thoroughbred racehorses on dirt and synthetic racetrack surfaces.. Equine Vet J 2014 Mar;46(2):227-32.
- Schade J, de Souza AF, Vincensi LC, Fonteque JH. The influence of the metacarpophalangeal joint angle on the transversal area and mean echogenicity of the superficial digital flexor tendon and suspensory ligament in gaited horses.. J Equine Sci 2021 Dec;32(4):135-141.
- Pagliara E, Marenchino M, Antenucci L, Costantini M, Zoppi G, Giacobini MDL, Bullone M, Riccio B, Bertuglia A. Fetlock Joint Angle Pattern and Range of Motion Quantification Using Two Synchronized Wearable Inertial Sensors per Limb in Sound Horses and Horses with Single Limb Naturally Occurring Lameness.. Vet Sci 2022 Aug 25;9(9).
- Turner ML, Falkingham PL, Gatesy SM. What is Stance Phase On Deformable Substrates?. Integr Comp Biol 2022 Mar 23;.
- Herrero C, Herrero E, Martín-Chivelet J, Pérez-Lorente F. Vertebrate ichnofauna from Sierra de las Cabras tracksite (Late Miocene, Jumilla, SE Spain): mammalian ichnofauna.. J. Iber. Geol. 48, 241-279.
- Lallensack JN, Farlow JO, Falkingham PL. A new solution to an old riddle, elongate dinosaur tracks explained as deep penetration of the foot, not plantigrade locomotion.. Palaeontology 64, e12584.
- Schamhardt HC, Merkens HW. Objective determination of ground contact of equine limbs at the walk and trot: comparison between ground reaction forces, accelerometer data and kinematics.. Equine Vet. J. 26, 75-79.
- Masao FT, Ichumbaki EB, Cherin M, Barili A, Boschian G, Iurino DA, Menconero S, Moggi-Cecchi J, Manzi G. New footprints from Laetoli (Tanzania) provide evidence for marked body size variation in early hominins.. Elife 2016 Dec 14;5.
- Robbins LM. Hominid footprints from Site G.. In Laetoli: a Pliocene site in northern Tanzania (eds Leakey MD, Harris JM), pp. 497-502. Oxford, UK: Clarendon Press.
- Moussa MT. Fossil tracks from the Green River Formation (Eocene) near Soldier Summit, Utah.. J. Paleontol. 42, 1433-1483.
- Lockley MG, Ritts BD, Leonardi G. Mammal track assemblages from the early Tertiary of China, Peru, Europe, and North America.. Palaios 14, 398-404.
- Mustoe GE, Hopkins DQ. Mammal and bird tracks from the Eocene Puget Group, Northwest Washington, USA.. Ichnos 20, 36-42.
- Shumer A, Wilkens H, Vollmerhaus B, Habermehl KH. The circulatory system, the skin, and the cutaneous organs of the domestic mammals.. Berlin, Germany: Springer.
- Onar V, Alpak H, Pazvant G, Armutak A, Chrószcz A. Byzantine horse skeletons of Theodosius harbour: 1. Paleopathology.. Revue Méd. Vét. 163, 139-146.
- Rooney JR, Scott E. Sidebone in a fossil horse.. J. Equine Vet. Sci. 18, 519-520.
- Quse V, Fernandes-Santos RC. Tapir veterinary manual, 2nd edn.. Belize City, Belize: IUCN/SSC Tapir Specialist Group (TSG).
- Scrivner PJ, Bottjer DJ. Neogene avian and mammalian tracks from Death Valley National Monument, California.. Palaeogeogr. Palaeocl. Palaeoecol. 57, 285-331.
- Sarjeant WAS, Reynolds RE. Camel and horse footprints from the Miocene of California and Nevada.. San Bernadin. Co. Mus. Assoc. Q. 46, 3-20.
- Reynolds RE. Horse hoof prints in the fossil record.. In Making tracks across the Southwest: the 2006 desert symposium (eds Reynolds RE), pp. 25-28. Redlands, CA: California State University Desert Studies Consortium.
- Morgan GS, Williamson TE. Middle Miocene (Late Barstovian) mammal and bird tracks from the Benevidez Ranch local fauna.. New Mex. Mus. Nat. Hist. Sci. Bull. 42, 319-330.
- Nieminen M. Hoof and foot loads for reindeer (Rangifer tarandus).. Rangifer 10, 249-254.
- Getty R. Sisson and Grossman's the anatomy of the domestic animals, 5th edn.. Philadelphia, PN: W.B. Saunders Company.
- Bragulla H. Fetal development of the segment-specific papillary body in the equine hoof.. J Morphol 2003 Nov;258(2):207-24.
- Miyaki H, Ohnishi T, Yamamoto T, Kameya T. Measurement of water contents of hoof wall, sole, and frog in horses.. Exp. Rep. Equine Heal. Lab. 11, 15-20.
- Bragulla HH, Reese S, Mulling C. Histochemical and immunohistochemical studies of the horn quality of equine hoof.. Anat. Histol. Embryol. 23, 44-45.
- Hinterhofer C, Stanek C, Binder K. Elastic modulus of equine hoof horn, tested in wall samples, sole samples and frog samples at varying levels of moisture.. Berl Munch Tierarztl Wochenschr 1998 Jun;111(6):217-21.
- Chard R. The wet dry cycle.. Equine Heal. 21, e14.
- Coleman E. Observations on the formation and uses of the natural frog of the horse, with a description of a patent artificial frog.. London, UK: J. Crowder.
- Lungwitz A. The changes in the form of the horse's hoof under the action of the body-weight.. J. Comp. Pathol. Therap. 4, 191-211.
- Barrey E. Investigation of the vertical hoof force distribution in the equine forelimb with an instrumented horseboot.. Equine Vet J Suppl 1990 Jun;(9):35-8.
- König HE, Macher R, Polsterer-Heindl E, Sora C, Hinterhofer C, Helmreich M, Böck P. Stoßbrechende Einrichtungen am Zehenendorgan des Pferdes.. Wien. Tierarztl. Monat. 90, 267-273.
- Hinterhofer C, Weißbacher N, Buchner HHF, Peham C, Stanek C. Motion analysis of hoof wall, sole and frog under cyclic load in vitro: deformation of the equine hoof shod with regular horse shoe, straight bar shoe and bare hoof.. Pferdeheilkunde 22, 314-319.
- Back W, Pille F. The role of the hoof and shoeing.. In Equine locomotion (eds Back W, Clayton HM), pp. 147-174, 2nd edn. St. Louis, MO: Saunders.
- Davies Z. Equine science, 3rd edn.. Hoboken, NJ: Wiley-Blackwell.
- Douglas JE, Biddick TL, Thomason JJ, Jofriet JC. Stress/strain behaviour of the equine laminar junction.. J Exp Biol 1998 Aug;201(Pt 15):2287-97.
- Douglas JE, Mittal C, Thomason JJ, Jofriet JC. The modulus of elasticity of equine hoof wall: implications for the mechanical function of the hoof.. J Exp Biol 1996 Aug;199(Pt 8):1829-36.
- Douglas JE, Thomason JJ. Shape, orientation and spacing of the primary epidermal laminae in the hooves of neonatal and adult horses (Equus caballus).. Cells Tissues Organs 2000;166(3):304-18.
- Bragulla H, Hirschberg RM. Horse hooves and bird feathers: Two model systems for studying the structure and development of highly adapted integumentary accessory organs--the role of the dermo-epidermal interface for the micro-architecture of complex epidermal structures.. J Exp Zool B Mol Dev Evol 2003 Aug 15;298(1):140-51.
- Bragulla HH, Homberger DG. Zur Funktion der spezifischen Profilaggrin entgaltenden Keratohyalingranula in der Hufepidermis des Pferdefötus.. Pferdeheilkunde 23, 5-20.
- Chmielewski Z. Identification of horse hoof impressions.. J. Forensic Identificat. 50, 563-571.
- Aramayo SA, de Bianco T M, Bastianelli NV, Melchor RN. Pehuen Co, Updated taxonomic review of a late Pleistocene ichnological site in Argentina.. Palaeogeogr. Palaeocl. Palaeoecol. 439, 144-165.
- Oliva C, Arregui M. Mammalian ichnopathology: a case study of Holartic Ungulates (Gomphotheriidae, Equidae, Camelidae) of the Late Pleistocene of South America, ichnotaxomic implications.. B. Soc. Geol. Mex. 70, 417-447.
- Remeika P. Fossil footprints of the Anza-Borrego.. In Fossil treasures of the Anza-Borrego Desert (eds Jefferson GT, Lindsay L), pp. 311-327. San Diego, CA: Sunbelt Publications.
- Reynolds RE. The 2006 Desert Symposium field trip (guide), day 1.. In Making tracks across the Southwest: the 2006 desert symposium (eds Reynolds RE), pp. 3-5. Redlands, CA: California State University Desert Studies Consortium.
- Burns A. The prehistoric footprints at Formby.. Merseyside, UK: Ainsdale Discovery Center.
- Pillola GL, Palombo MR, Panarello A, Zoboli D. The Pleistocene non-hominid vertebrate ichnofossil record of Italy.. J. Mediterr. Earth Sci. 12, 193-212.
- McNeil PE, Hills LV, Kooyman B, Tolman S. Significance of latest Pleistocene tracks, trackways, and trample grounds from Southern Alberta, Canada.. New Mex. Mus. Nat. Hist. Sci. Bull. 42, 209-223.
- Kulchytzky YO, Kulchytzky AY. Finding of the maxilla of [a] hollow-horned and the traces of vertebrates in the Miocene molasses of the Precarpathian and Transcarpathian Foredeeps.. Paleontol. Sbornik 17, 69-74.
- Pérez-Lorente F, Serrano F, Rodríguez-Estrella T, Mancheño MA, Romero M. Pisadas fósiles de mamíferos en el Mioceno superior de La Hoya de la Sima (Jumilla, Murcia. España).. Rev. Espan. Paleontol. 14, 257-267.
- Sauqué V, García-González R, Cuenca-Bescós G. A Late Pleistocene (MIS3) ungulate mammal assemblage (Los Rincones, Zaragoza, Spain) in the Eurosiberian–Mediterranean boundary.. Hist. Biol. 28, 1-26.
- Hulbert RC Jr, MacFadden BJ. Morphological transformation and cladogenesis at the base of the adaptive radiation of Miocene hypsodont horses.. Am. Mus. Nov. 3000, 1-61.
- Kelly TS. New middle Miocene equid crania from California and their implications for the phylogeny of the Equini.. Nat. Hist. Mus. L.A. Co. Contrib. Sci. 473, 1-44.
- Cantalapiedra JL, Prado JL, Hernández Fernández M, Alberdi MT. Decoupled ecomorphological evolution and diversification in Neogene-Quaternary horses.. Science 2017 Feb 10;355(6325):627-630.
- Eisenmann V. Sur quelques caractères adaptatifs du squelette d’Equus et leurs implications paléoécologiques.. B. Mus. Natl Hist. Paris 6, 185-195.
- Bernor RL, Kaya F, Kaakinen A, Fortelius M. Old world hipparion evolution, biogeography, climatology, and ecology.. Earth Sci. Rev. 221, e103784.
- Heintzman PD, Zazula GD, MacPhee R, Scott E, Cahill JA, McHorse BK, Kapp JD, Stiller M, Wooller MJ, Orlando L, Southon J, Froese DG, Shapiro B. A new genus of horse from Pleistocene North America.. Elife 2017 Nov 28;6.
- Jimenez-Hidalgo E, Diaz-Sibaja R. Was Equus cedralensis a non-stilt legged horse? Taxonomical implications for the Mexican Pleistocene horses.. Ameghiniana 57, 284-288.
- Oliveira EV, Bélo PS, Fambrini GL, Siala AN, Silva AKB, Barreto AMF. A new late Pleistocene ichnological site with mammal footprints from Brazil.. J. S. Am. Earth Sci. 94, e102216.
- Boulbes N, Van Asperen EN. Biostratigraphy and palaeoecology of European Equus.. Front. Ecol. Evol. 7, 1-30.
- Uzunidis A. Middle Pleistocene variations in the diet of Equus in the South of France and its morphometric adaptations to local environments.. Quaternary 4, 1-28.
- Cooper KL, Sears KE, Uygur A, Maier J, Baczkowski KS, Brosnahan M, Antczak D, Skidmore JA, Tabin CJ. Patterning and post-patterning modes of evolutionary digit loss in mammals.. Nature 2014 Jul 3;511(7507):41-5.
- Ewart JC. The second and fourth digits in the horse: their development and subsequent degeneration.. Proc. R. Soc. Edin. B 20, 185-191.
- Franciolli AL, Cordeiro BM, da Fonseca ET, Rodrigues MN, Sarmento CA, Ambrosio CE, de Carvalho AF, Miglino MA, Silva LA. Characteristics of the equine embryo and fetus from days 15 to 107 of pregnancy.. Theriogenology 2011 Sep 15;76(5):819-32.
- Barreto Rda S, Rodrigues MN, Carvalho RC, De Oliveira E Silva FM, Rigoglio NN, Jacob JC, Gastal EL, Miglino MA. Organogenesis of the Musculoskeletal System in Horse Embryos and Early Fetuses.. Anat Rec (Hoboken) 2016 Jun;299(6):722-9.
- Bragulla H, Budras KD, Reilly JD. Fetal development of the white line (Zona alba) of the equine hoof.. Equine Vet J Suppl 1998 Sep;(26):22-6.
- Bidwell LA, Bowker RM. Evaluation of changes in architecture of the stratum internum of the hoof wall from fetal, newborn, and yearling horses.. Am J Vet Res 2006 Dec;67(12):1947-55.
- Bragulla H. [The deciduous hoof capsule (Capsula ungulae decidua) of the equine fetus and newborn foal].. Anat Histol Embryol 1991 Mar;20(1):66-74.
- Bragulla HH, Budras KD. Die Homologisierung des foetal-perinatalen Eponychiums' als hinfällige Hufkapsel des Pferdes (Capsula ungulae decidua).. Anat. Anzeiger 170, 539-540.
- Smith S, Yohe LR, Solounias N. The bony cap and its distinction from the distal phalanx in humans, cats, and horses.. PeerJ 2023;11:e14352.
- Bragulla HH, Reese S, Mulling C, Sachtleben M. Die hinfällige Klauenkapsel des Rindes.. Anat. Histol. Embryol. 26, 62.
- Hamrick MW. Development and evolution of the mammalian limb: adaptive diversification of nails, hooves, and claws.. Evol Dev 2001 Sep-Oct;3(5):355-63.
- Homberger DG, Ham K, Ogunbakin T, Bonin JA, Hopkins BA, Osborn ML, Hossain I, Barnett HA, Matthews KL 2nd, Butler LG, Bragulla HH. The structure of the cornified claw sheath in the domesticated cat (Felis catus): implications for the claw-shedding mechanism and the evolution of cornified digital end organs.. J Anat 2009 Apr;214(4):620-43.
- Bragulla H, Ernsberger S, Budras KD. On the development of the papillary body in the feline claw.. Anat Histol Embryol 2001 Aug;30(4):211-7.
- Thomson TJ. The functional morphology of vertebrate claws and the morphological relationships between unguals and sheaths applied to predicting claw function in Archaeopteryx.. Doctoral dissertation, University of California, Davis.
- Thomson TJ, Motani R. Functional morphology of vertebrate claws investigated using functionally based categories and multiple morphological metrics.. J Morphol 2021 Mar;282(3):449-471.
- Hedrick BP, Cordero SA, Zanno LE, Noto C, Dodson P. Quantifying shape and ecology in avian pedal claws: The relationship between the bony core and keratinous sheath.. Ecol Evol 2019 Oct;9(20):11545-11556.
- Alibardi L. Development, comparative morphology and cornification of reptilian claws in relation to claws evolution in tetrapods.. Contrib. Zool. 78, 25-42.
- Sato F, Otsuka N, Kuwano A. Toe white line separation associated with the notch (crena marginalis) on the distal phalanges in Thoroughbred yearling horses.. J Vet Med Sci 2021 Nov 16;83(11):1745-1749.
- Coombs MC. Large mammalian clawed herbivores: a comparative study.. T. Am. Phil. Soc. 73, 1-96.
- Marsh OC. Recent polydactyle horses.. Am. J. Sci. 43, 339-355.
- Carstanjen B, Abitbol M, Desbois C. Bilateral Polydactyly in a foal.. J Vet Sci 2007 Jun;8(2):201-3.
- Wolff K, Lühken G, Litzkel LF. Polydaktylie, in Gedenken an das Urpferdchen: ein Fallbericht.. Pferdeheilkunde 25, 586-589.
- Beste KJ, Whitfield-Cargile CM. Considerations for surgical correction of polydactyly in horses.. Equine Vet. Educ. 31, 468-471.
- Gould SJ. Hen's teeth and horse's toes: further reflections in natural history.. New York, NY: Norton.
- Stanek C, Hantak E. Bilateral atavistic polydactyly in a colt and its dam.. Equine Vet J 1986 Jan;18(1):76-9.
- Vincelette AR, Renders E, Scott KM, Falkingham PL, Janis CM. Hipparion tracks and horses’ toes: the evolution of the equid single hoof.. Figshare .
Citations
This article has been cited 1 times.- Vincelette A. The Characteristics, Distribution, Function, and Origin of Alternative Lateral Horse Gaits.. Animals (Basel) 2023 Aug 8;13(16).
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