Boletín de la Sociedad Geológica Mexicana ISSN: 1405-3322 Sociedad Geológica Mexicana, A.C.

Oliva, Cristian; Arregui, Mariano Mammalian Ichnopathology: a case study of Holartic Ungulates (Gomphotheriidae, Equidae, Camelidae) of the Late Pleistocene of South America. Ichnotaxomic implications Boletín de la Sociedad Geológica Mexicana, vol. 70, no. 2, 2018, May-August, pp. 417-447 Sociedad Geológica Mexicana, A.C.

DOI: https://doi.org/10.18268/BSGM2018v70n2a10

Available in: https://www.redalyc.org/articulo.oa?id=94362562011

How to cite Complete issue Scientific Information System Redalyc More information about this article Network of Scientific Journals from Latin America and the Caribbean, Spain and Journal's webpage in redalyc.org Portugal Project academic non-profit, developed under the open access initiative http://dx.doi.org/10.18268/BSGM2018v70n2a10 P. 417‒447 VOL. 70NO. 2 BOL. SOC.GEOL.MEX.2018 cas yTécnicas, CONICET. Científi Investigaciones de Nacional Consejo Provincia deBuenosAires, Argentina. no 900-1198, Berisso, CP 1923. BuenosAires, YPF Tecnología S.A. Av. del Petróleo Argenti Mariano Arregui Provincia deBuenosAires, Argentina. Aires.nos Calle 50 N°539.CP 1900. LaPlata, Secretaría de Cultura de la Provinciade Bue Preservacióny de Museos vincial Patrimonial, Paleontológicoy co (CRePAP), DirecciónPro Centro de Registro del Patrimonio Arqueológi [email protected] Cristian Oliva Cristian America. Ichnotaxomicimplications (Gomphotheriidae, Equidae, of Camelidae) ofSouth the LatePleistocene Mammalian Ichnopathology: study ofUngulates Holartic a case Manuscript accepted: September 26,2017. Manuscript September accepted: Corrected manuscript received: June 16,2017. Manuscript received: March 18,2017. Oliva , Mariano Arregui - - - - - Equidae. Gomphotheriidae, Camelidae, Late Pleistocene, South America, Keywords: Mammal ichnopathology, synonym of Bianco, 1987b),herein proposed asteratologic Lamaichnum tulipensis llamoid ichnospecies:a second to segregate (ichnopathological characters) are often used heteropody). intraspecificvariations These asecondary acquisitionof tracks, normal the ends], enlargement and wideningof [with ample separation between both digital interclavular gap digits, overexpansion of of prints) andoutward/divergent (lateral curving toepad digits, unevenof sizing of curving bilateral symmetry, medial subparallel (loss of overgrowth: horny inward/ tual evidence of and Manera de Bianco, 1987b displays even frog andbars). of andhypertrophy[inner and outer quarters], lateral walls showing hyper-lengthening of turkish-slipper foot (over-elongated hoofprint, anteriorly heel buttress), displacedexternal pillar [toe-quarter junction] and internal wall,withforwardlyprojected region andhoof (asymmetricalsole buttresses), distorted hoof the betweenspacing frog baseline and heel heel-bulb impressions andanenlargement of both onar region, with an approximation of tal hoof, narrowing and distal projection of to contracted heel (transversely compressed anomalies inthe digital impression,linked Hippipeda the sole (horizontal). (vertical) of and cracking prints, as fractures onthe hornwall interpreted 1987a) presents occasionalfissures onitshoof australis the group. by of livingmembers andalterationspedal affections experienced the main nopathologies) that some of reflect several (ichclinical signs the recognition of (Buenos Aires Province, Argentina), allows Pehuen CoandLagunadelMontetracksites the late Pleistocene (Lujanian Stage/Age) of to Holarcticcorresponding ungulates, from paleoichnites andtrackways of analysis The ABSTRACT (Aramayo andManera de Bianco, isp. exhibits a series of morphologic morphologic isp. exhibits aseriesof Lamaichnum guanicoe Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Lamaichnum guanicoe (Aramayo andManera de . Proboscipeda Aramayo - - - - permite elreconocimiento dediversos signos clínicospermite del Monte(Provincia de BuenosAires, Argentina), yacimientos paleoicnológicos de Pehuen CoyLaguna Pleistoceno tardío (piso/edad Lujanense) de los pondientes a ungulados holárticos, provenientes del El análisis de paleoicnitas yrastrilladas corres RESUMEN Camelidae, Equidae. Sudamérica, Gomphotheriidae, mamíferos, Pleistoceno tardío, Palabras clave: Icnopatología teratológico de de Bianco, 1987b), aquí propuesta comosinónimo pecie: icnoes unasegunda menudo utilizadas para segregar intraespecíficas (caracteres icnopatológicos) son a de una heteropodia secundaria). Estas variaciones adquisición y ensanchamiento de las huellas, normal raciónentre ambos extremos digitales], alargamiento del espacio interdigitalexpansión [conamplia sepa lateral de losdígitos, (curvamiento divergente/externo de las almohadillas digitales) y desigual desarrollo medial de los dígitos,simetría bilateral, curvamiento (pérdida de subparalelo/interno cimiento córneo: de sobrecre evidencias 1987b muestra eventuales Lamaichnum ehipertrofia de ranilla ybarras). yexterno interno mostrando hiper-alargamiento de los cuartos turcozapato (impresión del casco superelongada, de desplazado),forma pie en anteriormente externo proyectados hacia delante ycontrafuerteinternos solar ymuro asimétricos,nado (región conpilares la ranilla ylos ángulos del muro), casco distorsio del espacio entrey unalargamiento la línea base de con aproximación de ambas impresiones bulbares estrechamiento yproyección talonar, distal de la región talón contraído (casco transversalmente comprimido, morfológicas en la impresión digital, ligadas a un les). de la suela (horizonta y agrietamientos (verticales) pezuñas, interpretadas como fracturas del muro córneo presenta fisuras ocasionales enlas improntas de sus australis por los miembros vivientes del grupo. les afecciones yalteraciones pedales, experimentadas (icnopatologías) que reflejanalgunas de las principa Hippipeda Lamaichnum (Aramayo yManera de Bianco, 1987a)

guanicoe Lamaichnum isp. unaserie de anomalías exhibe

tulipensis Aramayo yManera de Bianco,

/2018 guanicoe (Aramayo yManera . Proboscipeda 417 ------

Ichnopathology of Holartic Ungulates; Late Pleistocene of South America ABSTRACT INTRODUCTION / MATERIALS AND Ichnopathology of Holartic Ungulates; Late Pleistocene of South America METHODS 418 418 1. Introduction in the last three decades,from mainly paleo Waldron, 2009), has experienced notable progress Tanke, 1992;Rothschild andMartin,1993; (Rothschild andShelton, 1992; Rothschild and Paleopathology, ancientdiseases the studyof Late Jurassic(Dantas Triassics. l.(Tucker andBurchette, 1977)and North America; Tanke andRothschild, 2015) of and Late Cretaceous (McCrea (Hitchcock,1844, 1858;Abel, 1935;Lull,1953) were described:Late Triassic–EarlyJurassic Hemisphere, Northern from the Mesozoic of anomalousrecords, wherein ascarce number of atively lesser, headed bysporadic contributions mid-nineteenth century, itsevolution was compar oped parallel to conventionalthe since ichnology In thisregard,devel although this specialtyhas Rothschild, 2002;McCrea (see Lockleyon dinosaurs focused specifically and restricted more much are the fossil-tracksbased on (paleopathology analysis) In contrast, the advances inichnopathology 2014). 2017b; Ferreira Barbosa Bell, 2010; Rega Druetta, 2009, 2012;Xing Farke andO’Connor, 2007;Lü Aramayo, 2006;Rothschild andMartin, 2006; Bennett, 2003; Rothschild andTanke, 2005; 2002; Tanke andRothschild, 2002,2010,2015; Tanke andCurrie, 2000;Molnar, 2001;Hanna, Rothschild andMolnar, 1988;Sullivan 2012; Henriques Ferigolo, 1992,2007;Rothschild points (Rothschild, 1988,1990,1997;Tanke, 1989; Bartosiewicz, 2008)andpaleontological view zooarqueological (Baker and Brothwell, 1980; and Malgosa,2003;Ortner, 2003a,2003b), Aufderheide and Rodriguez-Martin, 1998; Isidro anthropological (Ortner and Putschar, 1980; 08 f uoe ideJrsi fAfrica (Jenny Europe; Middle Jurassic of 2008) of / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín et al. , 2013,2014a,2014b, 2017a, et al et al. et al. ., 2013;BarbosaandLuna, , 1998; Marshall , 1998;Marshall et al. , 2010;Bell et al , 1994;Avanzini et al. et al. ., 1994;Tanke and , 2014,2015). et al. et al , 2009,2015; , 2014,2015; ., 1997,2001, et al. et al et al et al. ., 2011; , 2007; ., 1998; , 2000; et al. - - - - /2018 ,

2. Materialsandmethods camelids andgomphotherids), from the Late trackwaysrelated toHolarctic ungulates (equids, fossiltracks and the paleopathological studyof In thispaper we present the results obtained from speciesfor comparison. are nomodern 2015); made even worse inthe cases where there maker, (see McCrea taphonomicalfactors) the track track-bearing substrate, behaviour of several extrinsicagents ( of influence the reflect which trails, and footprints in signs pathological of that differentiation the difficulty imply high the to due probably is This al Asia(Currie Cretaceous of and Josen, 1986,1988) and Late 1982;Ishigaki, been madeindirectly, through of interpretation the ungual diseases has recognitionThe of infauna influence;seeMcCrea foreign objects, ical disturbance, presence of characters) ( (extramorphological factors linked to external phological changes) alterations and/or secondary (intramor variations intraspecific of types other ungulates); havingmodern previously for discarded known affections limb the of any to (related potential clinical signs based onthe detection of was footprints pathologic of differentiation The Buenos Aires Province, Argentina (Figure 1). Laguna del Monte (Partido de Guaminí) tracksites, Pehuen Co(Partido de Coronel Rosales) and of fromthe Late Pleistocene (Lujanian Stage/Age) and Manera de Bianco, 1987b); and Bianco, 1987b); Lamaichnum guanicoe australis imens assignable to the ichnospecies ichnopathologicalThe analysis wasmade on spec thesefindings. cal implications of the ichnopathologicaldiscuss and ichnotaxonomi Argentina (South America),and Pleistocene of ., 2015). (Aramayo andManera de Bianco, 1987a); e.g ,ntr fthe substrate, mechan ., nature of Lamaichnum tulipensis (Aramayo andManera de et al. e.g., et al. , 2003;McCrea ntr fthe nature of , 2015). Hippipeda (Aramayo Proboscipeda et al. isp.; et - - - - - , Monte (Guaminí);Buenos AiresProvince,Argentina. study material.(1)Pehuen Co (CoronelRosales);(2)Laguna del Figure 1 donkey); Equus africanus asinus ferus maximus Blumenbach, 1797(Africanelephant); the group: of livingmembers of cialized breeders) andcomparisonwithautopods specialists(rural and zoo veterinarian, spe tion of veterinaryliterature, and/ordirectly, byconsulta Geographic location of provenance tracksites ofthe Geographiclocation of provenance tracksites

caballus Linnaeus, 1758(Asianelephant); Equus quagga Linnaeus, 1758(domestic horse); Linnaeus, 1758 (domestic Boddaert, 1785 (plains Loxodonta africana Elephas Elephas Equus Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín - -

3. Podaldiseasesonlivingungulates (MG-Pic) and “Carlos(Punta Darwin” Alta) the Museums “Marcelino Freyre” (Guaminí) of resideimens inthe Paleoichnological Collections Pehuen CoTracksite), while the remaining spec PCT: Tracksite; del Monte Laguna (LDM: field the study material was examined inthe Most of ological) subtypes. palmar/talonar; interdigital; and phalangeal (oste according intoclavular (ungular); solar; bulbar/ grouped and classified were disorders podal the Following veterinary tendencies, the modern 1758 (alpaca). Molina, 1782(vicugna); Lama glama zebra); Rhinocerotidae Gray, 1821; Tapiridae Burnett, Perissodactyla Owen, 1848(Equidae Gray, 1821; Owen, Gray, 1848 (Elephantidae 1821); ied for severalProboscidea hoofedmammals: have been recognized andsystematically stud foot disorders In thisregard, a great number of digital pads, foot pad)regions. (ungular/solar) and palmar/talonar (heel bulbs, cracking, overgrowth, infections) inthe clavular (osteopathies, bonelesions) andcomplications ( affections phalangeal/osteological of variety wide cutting wounds, contusions, foreign bodies) to a lesions andtrauma ( podal diseases, from simple to a broad range of nature, ungulates are particularly susceptible their unguligrade/subunguligrade Because of Aires Province, Argentina. (Pehuen Co, Partido de Coronel Rosales); Buenos Partido de Guaminí). PCT: Pehuen Co Tracksite Freyre; Colección Paleoicnológica (Guaminí, Rosales). MG-Pic:MuseoMunicipalMarcelino Paleoicnológico (Punta Alta, Partido de Coronel MD-YPI: MuseoDarwin;ColecciónYacimiento Monte Tracksite (Guaminí, Partido de Guaminí). abbreviations.Institutional (MD-YPI). Lama guanicoe Linnaeus, 1758(llama); Müller, 1776 (guanaco); e.g. Vicugna pacos , penetrating injuries, LDM: Laguna del LDM: /2018 Vicugna vicugna Linnaeus, 419 419 e.g. - - - ,

MATERIALS AND METHODS / PODAL Ichnopathology of Holartic Ungulates; Late Pleistocene of South America DISEASES ON LIVING UNGULATES PODAL DISEASES ON LIVING Ichnopathology of Holartic Ungulates; Late Pleistocene of South America UNGULATES 420 420 Among the main clinical conditions detected on 1978, 1980). 1897), either on wild or captive animals (Fowler, 1821; Suidae Gray, Gray, 1821;Tayassuidae Palmer, Hippopotamidae 1821; Gray, Giraffidae 1821; Cervidae Gray, 1821;Bobidae Gray, 1821; 1829); Artiodactyla Owen, 1848 (Camelidae Gray, Fgr A.Te r loafce yasre f of series a by affected also are They 2A). (Figure 1980, 2001;von Houwald, 2001;Benz,2005) (Kuntze, theheel cushion 1972, 1980;Fowler,of chronicapododermatitis suppurativa orfoot rot) traumatica or abscess; infection (pododermatitis susceptible to the overgrowth, cracking, and rhinoceros, hippopotamus), are particularly as other foot-paddedelephant), ungulates (tapirs, proboscidean, neospecies of The from onegroup toanother. incidenceandgravity varynotably of and degrees However, their etiology, evolution, clinical pattern, cited pathologiesThe are common to all ungulates. luxations, osseousfractures andfissures. arthritis, heterotopic ossification, subluxations and hyperplasia; and(5) matitis, interdigital phlegmon, necrobacillosis, and (4) white line, gravel, seedy toe); hypertrophyand abscessation of (inflammation, diseases horn), white-line overgrown trauma; and fissures, cracks, split toenails), hoofwallalterations ( nary phlegmon, flexor-zone abscess, exungulation, ulceration, digital and verrucous coro dermatitis, nous); (3) matitis (purulent, suppurative, necrotic, gangre orpododermatitis circumpscripta), infectiousorbacterial pododer ulceration solar diffusa; tica asep (laminitis orpododermatitis pododermatitis dropped sole, solarcallus, solarpressure, aseptic sole, penetrating wounds, solarhemorrhage), (2) necrosis,heel, heel, crushed sheared greasy heel; heel wart (talonar hyperplasia), heel abscess, heel (1) these mammalian lineages can be mentioned: (African elephant) and (African elephant) / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín solar interdigital talonar : solarinjuriesandtrauma ( clavular / palmar : postdigital and interdigital der : proximal ( dermic osteological / bulbar Elephas maximus Elephas : heel erosion, Loxodonta africana : osteomyelitis, e.g., e.g., punctured hoofwall (Asian e.g., toe - - - - - /2018

their natural (forwardlyposition directed, aligned the toenails, which from depart and deviation) of and consequentalteration (elongation, curving, distinguished bycondition is This the hypertrophy ( Lama glama genera the SouthAmerican lids, suchof the species as came digital pads) inmodern of pododermatitis frequent limb disease(along with theulcerative overdevelopment Horny constitutes the most West, 2001;Benz,2005)(Figure 2B–D). Kuntze, 1980;Fowler, 1993;Mikota illnesses(see severe (laminitis, exungulation) hoof moderate (overgrown cracking) to horn-wall horn, first case may potentially collapse, causing the capsule, and outer wall andhoof causing separation of collapse, potentially may case first highandunstableThe hooves generated in the (Figure 4C) appearance. a “turkishslipper” (Figure 4B) and/or “helical” thehooves, whichacquire strong of deformation (Figure 4A), thisdisorder can evolve towards a the hoofwall vation, thickening,of andcracking (plains zebra). Initially manifested as anover-ele africanus asinus equids: overdevelopment aggravates inunguligrade disturbance inthe cuticular keratogenesis, horny wear and a Linked with an inadequate corneous theseanimals. tion andlateral stabilityof locomo the long-term at affect does it laminitis, generates chronicproblem llamoids,in except if overgrowth doesnotconstitute necessarily a the individuals. For thisreason, while horny of remain crucial during the traction and propulsion the load. However,support most of healthy nails mammals,and unguligrade inwhich the hooves on the toe cushions, unlike semi-unguligrade weight bearing, withthe entire body weight falling camelids are non- unguals in digitigrade The oriented. (Figure 3C),oroutwardly (divergent) (Figure 3D) (subparallel) (Figure 3B), medially (convergent) with thedigital axis) (Figure 3A)tobeinwardly Vicugna vicugna Lama Equus ferus caballus Equus ferus [domestic])and Cuvier, 1800 ( [wild]); (domesticdonkey); Vicugna pacos (domestic horse); (domestichorse); Lama guanicoe Vicugna [domestic]). Miller, 1924 Equus quagga et al. [wild]; , 1994; Equus - - -

(schematic). (A)Pododermatitis; (B)hornwall(schematic). cracking; (C)overgrownhorn; (D)exungulation. BasedonBenz(2005). Figure 2 xre ytecroa egto thehorse. exerted by weight of the corporal the force the foot, by action of coverthe horny of thedistalphalanxinto rotation,of and sinking laminar corion,withaconsequent detachment, inflammatory the sive degeneration, separation, andnecrosisof an constitutes hoof-membranes diseasethatthe progres causes laminitis The thebars. rant overdevelopment of the white line and anaber a frontal stretching of hypergrowth), the hoofwall (corneous bending of nitis, comprisesthe anterior projection and dorsal secondstate,The typicallyassociated with lami tious necrosis. their infec a grey crumbly residue as a result of of cavities between bothstructures andthe formation minorslitsto extensive with the development of the white line, the hoofwall andwideningof of wall separation, is characterized by a horadation dition, known asseedy toe, white line disease, or infections andabscesses.con terial/fungal This bac the hoofwall, besidesthe occurrence of of fissuration and weakening the facilitating thereby Principal podaldiseasesinliving elephants (Proboscidea,Elephantidae, Elephantinae); lateraland palmar/plantarviews Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín ------

accompanied by important behavioral changes: In bothclinical phases, the physicalaresigns versal grooves alongthe wall. trans thick coronarybands, withemergence of of developmenthoofwall (helical foot); (4) abnormal the slipper foot) and/or spiralingovergrowth of the toe (turkish with antero-dorsal projection of the hoof, white line; (3)distaloverdevelopment of the bone; (2)hypertrophyand coronal tearing of the ungual rotation anddescentof severeof cases the same (puncture sole) in with perforation of the sole (dropped sole), thinning, andweakening of the autopodium: (1) drop,alterations flattening,of significant causes hoof, the within phalanx distal the laminar supportthatthe positions of the loss Conversely, achronicinvolving clinical pattern, deformation. toe and coronet, do not result into podal ness of although generating intense painandfeverish circulatorydeep changes intothehooves, which, and processes inflammatory of series a embraces thispathology of acute An clinicalpattern /2018 421 421 - -

PODAL DISEASES ON LIVING Ichnopathology of Holartic Ungulates; Late Pleistocene of South America UNGULATES PODAL DISEASES ON LIVING Ichnopathology of Holartic Ungulates; Late Pleistocene of South America UNGULATES 422 422 overgrowths. outward/divergent,toenail (D) medial/convergent; inward/subparallel; (C) (B) growth; natural showing:(A) Footprints (schematic). Figure 3 expansion and steepness of the hornwall) and the the hornwall) expansionof and steepness (dorsoventral hoof the in modifications of series tain a bilateral symmetry, they experience a Although autopodes withcontracted heel main dissipater for thehoof. structure, acting as circulatory pumpandshock soleandfrog; thislatter with apassive contactof hoofwall exertspressure on the ground surface, In these unaltered feet, only the weight-bearing line andheelbulbs (Figure 5A). approximately at the same level as the frog base transversely aligned heel buttresses, disposed autopodes) areand present almostbisymmetric orbalanced equine hooves normal The (unaltered (forwardly directed limbs). frequent limping march); and (3) antalgic posture ( gait abnormal (1) recurrent recline and resistance to walking; (2) / Principalclavularalterationsinliving and llamoids(Artiodactyla,Camelidae,Lamini);left/lateral palmar/plantarviews Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín e.g ., measure, shortandfast steps, - - /2018 in the foot,in well as the support an imbalance in as medial/lateral balance of anomaly implies aloss hoof heel-buttresses (Figure type of 5C). This junctions), andanteriorly displaced external forwardly(toe-quarter pillars projected internal and lateral (short and upright) walls into the hoof, flared) and (long medial of development uneven hoof) displaya strong pedal asymmetry, with the ungular region (distorted bilateral distortion of On the other hand, feet experiencing (Figure 5B). apex andhornwall the spacing between frog sole, withanincrease of the bya lateral of compressionand deformation tresses). changes These are usually accompanied distance between frog baseline and heel but of the frog andenlargement heel bulbs, atrophy of both the talonar area, withanapproximation of heel (narrowing, elevation anddistal projection of -

4. Systematicichnology overgrowth). elevated hoof (incipientovergrowth);(B)turkish-slipper hoof (acutehypertrophic overgrowth); (C)helicalhoof (chronichypertrophic Figure 4 concussion; and (3)solefrog:concussion; solarpressure, (bruising, injuries, lacerations, traumas), coronet ungular: quarter cracks, coronary bandlesions (2) foot gangrene; heel (infected abscess corn), sheared heels, heels (mud greasy rash,mud fever), living in equids. include(1)bulbar: These crushed heels, identified are diseases hoof numerous afflictions, and alterations described the Besides a greater strainthantheouterquarter). the bodyweight (exerting on the inner quarter of Proboscipeda australis Type Ichnospecies. Gomphotheriipedidae Remeika,Morphofamily 1995). (pythiosis),canker, thrush,keratoma (see Pollitt, ing at the white linezone, sarcoid,phycomycosis sole/frog,traumas), underrun stain haemoglobin (injuries,lesions puncture/penetrating wounds, Ichnogenus Ichnorder Proboscipedida Remeika Types of horny overdevelopment in living equines (Perissodactyla; Equidae; Equini); lateral views (schematic). (A) over- (A) (schematic). views Equidae;Equini);lateral (Perissodactyla; equinesliving in overdevelopment horny of Types Ichnodivision Vertebratichnia Vialov, 1966 Ichnoclass MammalipediaVialov, 1966 Manera deBianco, 1987a) Proboscipeda (Table 1;Figure 6) and Avram, 1962.

2001 Proboscipeda Panin and Avram, 1962 (Aramayo and

enigmatica et al. , 1995 Panin Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín - Stegomastodonichnum australis Stegomastodonichnum digital impressions, along tangentially arranged five to Three 1). ≥ ratio length/width with wide, larger than pes (isodiametric or longer than than long, withlength/width ratio < 1) slightly Family. Moderate heteropody, withmanus(wider theGomphotheriidae um-sized proboscideanof type, semi-plantigrade related to medi mm), of prints (length: 230 –270mm;width:300 et al. Extended diagnosis. (excepting MPA-87-56-I-E). and Arregui, 2016); all specimens remain imprints (Figure 6H) (Oliva four overlapped forefoot and rearfoot trackway of incomplete paleoichnites (Figure 6F); LDM-S (Figure 6E);LDM-S overstepped forefoot and hindfoot imprints of forefoot imprint(Figure 6D);LDM-S 2015). three paleoichnites (Aramayo trackway of forefoot imprint (Figure 6G) corresponding to a de Bianco, 1987a; Aramayo the holotype (Figure 6C) (Aramayo andManera footprint belonging to 87-56-I-E: plaster cast of seven footprints (holotype) (Figure 6A–B); MPA-of Study material. et al. de Bianco Bianco, 1987a, 2000; Lucas de Bianco, 1987a(Aramayo andManera de , 2015). , 2015). Large subcircular to subovoidal foot Laguna del Monte Laguna del et al. , 2010; Oliva

Pehuen Co 1 T 6 (modified from Aramayo from (modified ata rcwyo four : partial trackwayof Aramayo andManera ; LDM-S et al et al. et al. et al. ; PCT-I: trackway ., 2013; Aramayo , 2015); PCT-II: , 2007;Manera /2018 , 2013;Oliva 2 I 1 22 S : isolated 7 : couple in situ et al. 423 423 1 T 7 - - : ,

PODAL DISEASES ON LIVING UNGULATES Ichnopathology of Holartic Ungulates; Late Pleistocene of South America / SYSTEMATIC ICHNOLOGY Ichnopathology of Holartic Ungulates; Late Pleistocene of South America SYSTEMATIC ICHNOLOGY 424 424 hoof (B)unbalanced (healthyfoot); hoof, withcompressed heel;(C)distortedhoof, withbilateralasymmetry. Figure 5 defines the ichnogenus the defines track that elephantine of archetypal morphology Description. Cabrera, 1929;Anancinae Hay, 1922). sis platensis Possible trackmaker. et al. Bianco, 1987a, 2000; Oliva Province, Argentina (Aramayo andManera de tribution. dis and chronostratigraphic Geographic preserved, respectively. wrinkled), according thetrace beingpoorlyto well (pockmarked textured,ornamented cracked, and theichnite, either or irregularlysmooth part of most the encompasses that surface flattened wide rounded and a metapodial (footpad cushion print), conforming Broad hoofprints. differentiated slightundulations and/ormore or less series of thetrack,appearing a as the anterior margin of / Principal hoofin modern equinesBalanced Principal alterations (Perissodactyla; (A) Equidae;(schematic). Equini);views palmar/plantar ) (Proboscidea Owen, 1848; Gomphotheriidae Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín , 2015). (Ameghino, 1888) (= aePesoeeo Buenos Aires Late Pleistocene of

Proboscipeda australis Proboscipeda et al. Notiomastodon platen

, 2013; Aramayo ; characterized exhibits the Stegomastodon Stegomastodon - - /2018

whose surfacecan be pockmarked (cracked and ated by athickmetapodial-cushion (footpad-print), as the imprint talonar gener region, interpreted These footprints possess a broad flattened palmar/ width ratio: 1.00–1.10)(Table 1). isodiametric and/or longer than wide (length/ (length: 230 –270mm;width:250mm), ratio: 0.80 –0.96),isinvariably larger than the pes mm), typically wider than long(length/width manus(length:width: 250 – 300 240–290mm; foot impressions are nonexistent. Nevertheless the Anatomical differences between forefoot and hind rim) (Figure 6A–F). rounded bya noticeable annular ridge (marginal metrical, almost isodiametrical and usuallysur cular/ovoidal outline, which are bilaterally sym roughly cir imprinted paleoichnitesdeeply of Unaltered footprints. et al. structural (see simplicity Lucas by itsgreat size, hemispherical constitution and , 2007;Aramayo et al., Constitute large and 2015). et al. , 2007;McNeil - - - - - (2015). manus (forefoot); p:pes(hindfoot);manus (forefoot); r:rim(marginalsedimentary ridge);II toIV: hoofprints. A–Cand G,modified from Aramayo displaying evidenceof hornwall fractures and solefissuresarrows ).Abbreviations:d:undifferentiated (white digitalimpressions;m: (unhealthy (G)forefoot feet): imprint (belonging toapartialtrackway)and (H)couple ofoverlappedforefoot and rearfoot tracks;both footprintAltered trackway. partial a footprintpreservedof better (F) tracks; hindfootforefootandoversteppedcoupleof (E) imprint; forefoot isolated (D) (C); for cast a as served that paleoichnite the indicates arrow white the in (B) holotypespecimen; (A–C) feet): Figure 6 DF H ado mr o ls differentiated, less or more and/or H) 6D–F, can appear as slight undulations (Figure Similar in size and shape, these toe imprints pad print). transverse tangentially to the foot axisarranging the track (with their longer anterior margin of disposed inclosehemicircular along the pattern smalldigital impressions, appreciate a series of On better-preserved specimens, itispossible to state. (Figure on the preservational 6D–F) depending wrinkled) (Figure 6A–C) orfeatureless (smooth) Gomphotherid[ paleoichnites Proboscipeda australis Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín (Aramayoand Manera deBianco,1987a)].Unaltered footprints(healthy -

(see Ramsay andHenry, 2001;Benz,2005). proboscidean pes, speciesof such asinmodern and manus in either five, to three indistinctly be hooves in the number of to whatContrary Aramayo divergent). lateral digits (antero-laterally directed; somewhat forwardmost projected than the slightly delayed (anteriorlypointed; subparallely disposed) are In bothcases, the nearly aligned central toes (Figure 6B–C,G). subcircular to subrectangular shaped, hoofprints Proboscipeda australis et al. (2015) observed, /2018 may et al. 425 425

Ichnopathology of Holartic Ungulates; Late Pleistocene of South America SYSTEMATIC ICHNOLOGY Ichnopathology of Holartic Ungulates; Late Pleistocene of South America SYSTEMATIC ICHNOLOGY 426 426 **Extended from:Aramayoand Manera deBianco,1987a;Aramayo from*Compiled Aramayo and Manera deBianco,1987a,2010;Aramayo Manera de Bianco,1987a).L: tracklength; W:trackwidth;L/W: length/width ratio. proboscidean-tracks: Pleistocene, LateSouth American,measurementsof Footprint 1. Table Morphofamily PecoripedidaeMorphofamily Remeika overgrown laminitis, exungulation). horn, the group ( documented for of living members fossilelephants, unlike the numerous pathologies other clavular alterations in the occurrence of of paleoichnological record doesnotprovide evidence Leaving aside these scarce examples, the available 6G–H). inward, prolong towards the sole region (solar cracking) (Figure that hornwall) the on fissures as verticalridges on itshoofprints(interpreted of However, somewell preserved isolated specimens ungualdisorders. the identification of digital impressions(hoofprints), greatly hindering Additionally, these footprints rarely preserve the palmar/plantar diseases. notably the correct recognition and diagnosisof (both extinctboscidean tracks asextant), precludes pro the footpad print of texture, characteristic of Pathological footprints. / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Proboscipeda australis Order Artiodactypedida Vialov, 1966 LDM-S LDM-S PCT-I (MPA-87-56-I-E)* PCT-II* Ichnospecifical Range** Designation 1 1 LDM-S LDM-S (acronym) T S 7 7 exhibit inboundperipheral 2 1 I T 22 6 The irregular skin The I I I I I I I 3 2 1 2 1 2 1 et al. Referred idot270 230 230 Hindfoot >240 Hindfoot Hindfoot idot20–2020–201.0–1.10 210–240 230–270 240 Hindfoot Hindfoot idot235 Hindfoot oeot240 240 Forefoot Forefoot - - Forefoot Forefoot oeot290 Forefoot oeot20–2020–300.8–0.97 230–300 240–290 Forefoot Imprint , 1995 e.g ., - /2018 et al. et al , 2015. L ofrig eea takas noye (figures (neotype) trackways several conforming Bianco, 1987b); PCT-IV: twenty-one footprints, footprints (holotype) (Aramayo andManera de icoe Study material. Bianco, 1987b) nov. sin.(teratological synonym). tulipensis Megalamaichnum de Bianco, 1987b) Aramayo 1999; Reynolds, 1999; 1999; Lamaichnum obliquiclavum alfi Lamaichnum Lamaichnum guanicoe Aramayo andManeradeBianco, 1987b. Type ichnospecies. Ichnogenus ., 2015. ). Measurements Pehuen Co Lamaichnum tulipensis Lamaichnum etoromorphum Manera deBianco, 1987b Lamaichnum 8 0.85 300 280 250 250 5 1.08 0.97 1.08 250 300 250 240 3 1.02 230 (tables 2–3;figures 7–8) W Sarjeant and Reynolds, Sarjeant 1999; Proboscipeda australis Lamaichnum borregoensis Bianco, 1987b C-I:taka feleven ; PCT-III: trackway of (referred to (mm) Aramayo andManera de Aramayo andManera de Sarjeant and Reynolds, Sarjeant

(Aramayo andManera L/W Lamaichnum guanicoe 0.8 Aramayo and - 1 - - Lamaichnum guan et al. Sarjeant and Sarjeant (Aramayo and (Aramayo , 2015(= Remeika, -

(2013). (forefoot); A–B,D,and p:pes(hindfoot). F,modified from Aramayo footprints (healthy and feet) D–H:altered footprints (unhealthy showing feet), signsoftoenailovergrowths.Abbreviations:m:manus Figure 7 LDM-S LDM-S LDM-S LDM-S (Figure 8C); LDM-S LDM-S 8A) 7G, (figures (unpublished). footprints seven of trackway bioturbed area (Aramayo this MD-YPI-15-01: silicone rubber cast of chonicus mammals ( atrample bird ( [sharing surface withother tracks of 7B) (Figure footprints fifty the anterior specimen; PCT-V:cast of around 7A, 7C,8E);MD-YPI-05-15: siliconerubber Llamoid fossiltrackways( Aramayo andManera de Bianco, 1987a)]; 1 1 1 1 1 I I I I I 134 94 73 43 25 ; LDM-S ; LDM-S ; LDM-S ; LDM-S ; LDM-S Canipeda Laguna del Monte Laguna del isp.; 1 I 1 1 1 1 1 I I I 127 I I 78 44 40 178 176 ; LDM-S ; LDM-S ; LDM-S ; LDM-S (Figure 8F); LDM-S (Figure 8B); LDM-S Eumacrauchenichnus pata et al Lamaichnum guanicoe ., 2015); PCT-VI: 1 Gruipeda I 1 1 1 I I I 128 79 45 41 ; LDM-S ; LDM-S ; LDM-S ; LDM-S ; LDM-S isp.) and 1 1 1 I 1 1 1 1 Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín I I I I I I Aramayoand Manera deBianco,1987a)composed byA–C:unaltered 132 179 177 82 52 42 24 - ; ; ; ; ;

et al. (2015);Cand E,modified from Manera deBiancoand Aramayo Laguna del Monte specimen (Aramayo this ber cast of (figures 7E, 8M) and MD-YPI-06-10: silicone rub thirteen footprints 2015); PCT-IX: trackwayof block (paratype)sedimentary in (Aramayo et al. four footprints (neotype) (Figure 7F) (Aramayo of Manera de Bianco, 1987b);PCT-VIII: trackway footprints (holotype)(Figure 7D)(Aramayo and Co Pehuen Study material. prints (Oliva (Figure 8H);MG-Pic17(Figure 8D): isolated foot LDM-S (Figure 8K);LDM-S (Figure 8G); LDM-S , 2015); MD-YPI-06-04: isolated footprint, 2 I 12 ; LDM-S C-I:taka ftwenty-one ; PCT-VII: trackwayof et al ., 2013;Oliva andArregui, 2016). (referred to 2 ; LDM-S 1 I I 2 16 180 I (Figure 8L);MG-Pic10 5 ; LDM-S (Figure 8I); LDM-S Lamaichnum tulipensis 1 T 1 andLDM-S /2018 2 I 6 (Figure 8J); et al ., 2015). et al 427 427 1 1 I T 181 ., ). - - 2

Ichnopathology of Holartic Ungulates; Late Pleistocene of South America SYSTEMATIC ICHNOLOGY Ichnopathology of Holartic Ungulates; Late Pleistocene of South America SYSTEMATIC ICHNOLOGY 428 428 subparallel and M–P:outward/divergent,toenail overgrowths. ScalebarinB,F, G,I,K,and N equals 10 cm. andshallowest/lesser-preserved E–H:deepest/better-preserved, specimens.Altered footprints (unhealthy evidencing feet): I–L: inward/ Figure 8 / Llamoid paleoichnites ( Llamoid paleoichnites Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Lamaichnum guanicoe Aramayo and Manera deBianco,1987a).Unaltered footprints A–D: (healthyfeet): /2018 ratio. de Bianco,1987b= Table 2.Footprintmeasurements of SouthAmerican, Late Pleistocene,camelid-trackways: (Aramayo andManerade (Aramayo andManerade Lamaichnum tulipensis Lamaichnum guanicoe Bianco, 1987b) Bianco, 1987b) cntxnIchnopathology Ichnotaxon both are closely resembling inshapeandsize, digitsIII–IV); generated byof the toe cushions ceolate, digital impressions(attributed to the prints curved lateral walls. Two oblong, acicular to lan bilobed/biconvex posterior border and softly higher than 1),withbicuspidate anterior margin, ichnite is typicallyelongated (length/width ratio teratologicwith a secondary heteropody). Each foot and hindfoot imprints (strong homopody, to subrectangular-subtrapezoidal outlined, fore Family. Almostindistinguishable, heart-shaped theCamelidae small to great-sized artiodactylsof type, digitigrade related to didactyl footprints, of and Manera de Bianco, 1987b).Paraxonic Extended diagnosis. et al. MG-Pic 16 (Figure 8P): isolated footprints (Oliva LDM-S footprints, respectively; LDM-S nineandfour (Figure 7H):trackwaysof composed , 2013;Oliva andArregui, 2016). 2 I 9 (Figure 8N);MG-Pic11(Figure 8O); Lamaichnum tulipensis Unaltered Footrpints Outward /divergent Altered Footprints toenail overgrowth Neither disease (healthy foot) (modified from Aramayo from (modified (MD-YPI-15-01) Hypodigm Neotype (MD-YPI-05-15) (Aramayo and Manera deBianco,1987b).L: tracklength; W:trackwidth; L/W: length/width (MD-YPI-06-10) 1 I 72 LDM-S LDM-S Hypodigm Material Holotype Holotype Neotype ; LDM-S 1 1 T T 2 1 IUdtrie 0–156 511 .49 1.14–1.64 15 60–85 1.14–1.54 80–115 65–80 Undetermined 85–100 VI Undetermined IV I neemnd9 55 011 .26 1.13–1.72 55–80 90–95 Undetermined III neemnd8 56 511 .83 1.13–1.58 60–85 85–95 Undetermined V IUdtrie 0–9 5–8 .9–13 4 1.19–1.39 65–80 90–95 Undetermined II neemnd9 2 5–8 .3–16 12 1.13–1.69 65–80 90–120 Undetermined I Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín 2 I 7 - - ; cnseiia 0–107 0 0.90–1.57 70–105 90–120 1.25–1.45 Ichnospecifical 80–100 100–145 Ichnospecifical neemnd10–108 0– 85–90 110–120 Undetermined – 70–75 90–100 Undetermined neemnd10–157 0 .0–16 4 1.20–1.67 75–100 120–125 Undetermined 21 0.94–1.88 46–87 75–116 Undetermined otrn L/W W L Footprint Referred idot158 01.27–1.34 85–90 115 1.05–1.28 70–95 Hindfoot 90–110 1.25–1.33 Hindfoot 80–92.5 100–120 Hindfoot oeot109 0 1.14 90–105 120 0.90–1.57 70–105 Forefoot 1.35–1.45 95–120 100 Forefoot 135–145 Forefoot and Hunt,2007;Cabral-Perdomo tribution. dis and chronostratigraphic Geographic reduced orabsent. trough andmedial pocket Interphalangeal depth. variable amplitude and parabolic heel notch, of during toenail overgrowth) and rearward ina short gence reached by the digits; distally overexpanded the separation and diver open, infunctionof interclavular (more gap orlessextended and state), expanding forward inalong V-shaped ing development onthe preservational (depending contrast overdevelopment). Interdigital of septum and/or outwardly, horny inpathological cases of from their natural position, either curving inwardly with thedigital axes (stretching and departing rectilinear, anteriorly directed, and coincident nailprints;these latter are commonly small sharp acuminate anterior tips,ending in andusually having broad, rounded posterior edges, narrow Measurement Range(mm) Noeeo North America (Lucas Neogene of Lamaichnum guanicoe °Reference N° 21 11 13 9 4 Aramayoand Manera /2018 Aramayo Aramayo Aramayo andManera Aramayo andManera Aramayo et al. New StudyMaterial de Bianco,1987b de Bianco,1987a tab. 3 tab.4 , 2018). et al. et al. et al. , 2015; , 2015; , 2015 429 429 - - -

Ichnopathology of Holartic Ungulates; Late Pleistocene of South America SYSTEMATIC ICHNOLOGY Ichnopathology of Holartic Ungulates; Late Pleistocene of South America SYSTEMATIC ICHNOLOGY 430 430 *Extended from:Aramayo and Manera deBianco,1987b,1996; Aramayo Bianco, 1987b= Table 3.Footprintmeasurements of SouthAmerican,Late Pleistocene,camelid-tracks: / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Lamaichnum tulipensis Unaltered Footprints (Aramayo andManeradeBianco, guanicoe ihoeu n cnseis (acronym) (ichnogenus andichnospecies) Feet) Lamaichnum tulipensis Footprints associatedto Footprints associatedto Outward /Divergent Inward /Subparallel Toenail Overgrowth Toenail Overgrowth de Bianco,1987b) Ichnotaxon Designation (Aramayo andManera 1987b Lamaichnum Ichnospecifical Range* ) (Aramayoand Manera deBianco,1987b).L:tracklength; W:trackwidth;L/W:length/width ratio. (Health y ouainRne6 1 0–8 1.03–1.57 50–85 65–110 Population Range ouainRne7 2 0–101.13–1.6 60–110 75–125 Population Range ouainRne6 3 0–8 1.07–1.67 60–80 65–130 Population Range LDM-S LDM-S LDM-S LDM-S LDM-S LDM-S LDM-S LDM-S LDM-S MG-Pic 17 MG-Pic 16 MG-Pic 11 MG-Pic 10 LDM-S LDM-S LDM-S LDM-S LDM-S LDM-S LDM-S LDM-S LDM-S LDM-S LDM-S LDM-S LDM-S LDM-S LDM-S LDM-S LDM-S LDM-S LDM-S LDM-S LDM-S 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 2 2 1 1 1 2 2 2 I I 2 I I I I I I I I I I I I I I I I I I I I I I I I /2018 I I I 181 180 I 179 178 176 134 132 128 127 78 73 52 45 44 43 42 41 40 25 24 72 16 12 94 82 79 9 7 6 5 et al 0–105 1 1.03–1.67 50–110 40 –150 ., 2015;Lucas and Hunt,2007. 100 110 110 100 100 100 130 125 125 120 100 105 100 100 65 75 75 75 85 95 80 65 75 75 85 75 80 80 85 95 95 75 90 65 L Measurements Lamaichnum guanicoe 1 1.13 110 65 60 70 60 55 82 75 65 60 65 65 75 60 70 65 55 65 75 65 70 50 70 75 75 75 70 80 75 75 65 75 75 85 W (mm) L/W 1.53 1.08 1.07 1.25 1.36 1.03 1.27 1.23 1.08 1.15 1.15 1.13 1.25 1.14 1.23 1.54 1.46 1.27 1.15 1.57 1.47 1.33 1.33 1.43 1.63 1.67 1.54 1.33 1.17 1.2 1.3 1.6 1.4 Aramayoand Manera de intraspecific and/or interespecific/intergeneric and/or intraspecific interaction (taphonomical variations) insteadof Hunt, 2007),which onautopode-substrate depend variations extramorphological (see Lucas and of extremely changeable; they display a broad range tracks, their appearance is Similar to modern larger extinct forms. existence of mm (see Lucas andHunt,2007), revealing the exhibiting ahigher measurement range: 40 –150 generated by livingllamoid species, although indistinguishable frommorphostructurally those South American camelids( Description. 1893; LaminiWebb, 1965). 1848; Camelidae Gray, 1821;Camelinae Zittel, Ameghino, 1880(great size) (Artiodactyla Owen, 1884) and† (medium size); † 1880) (smallsize); 1782) (= Possible trackmakers. 2007; Oliva Manera de Bianco, 1987b, 1996;Tassara South America (Aramayo and Late Pleistocene of served trackways (Figure 7A–C). virtually differentiation impossible (strong homopody), even inwell-pre their making sionally, and dimen nearly identical, both morphologically In thissense, forefoot andhindfoot tracks are width varyingfrom 45to90mm (tables 2,3). 1), with length ranging from 65to 120 mm and longer than wide (length/width ratio higher than byhealthy feet aretypically almostbisymmetric, Unaltered footprints. microbial mat, etc. of content and water saturation level, development solidation, compactionandcementation, moisture con granulometry,as the composition, of degrees sediment plasticity, such closelylinkedto factors andbodying tothe gait mass, of andthe grade the ground surface, whose intensityvaries accord respond to the pressure that the foot exerts over Consequently, inthese would thetrackmorphology ungulates). (insignificant differences autopodial Lama gracilis et al. The paleoichnites The associated with Hemiauchenia paradoxa , 2013;Aramayo Eulamaops parallelus Lama guanicoe Gervais and Ameghino, The imprintsproduced The

Vicugna vicugna Lamaichnum guanicoe (Müller, 1776) et al. Gervais and (Ameghino, , 2015). (Molina, et al ) are Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín ., - - - - pointed and posteriorly rounded lanceolate con ichnites, imprintsrepresent thetoepad anteriorly Meanwhile, andbetter-preserved inthe deepest typically related tocompactsediments. the trackmaker, is palmar/plantar anatomyof the reflect faithfully not does whichsubtype, This (interdigitalmedian sulcus (Figure septum) 8A–D). thatsions remain disconnectedthrough athick constitute narrow imprints toepad acicular depres In shallowest and less-preserved specimens, these toe cushions. smooth,antero-posteriorly elongated, presence of concave,undivided and uniformly indicating the parallel, digital impressions, which are invariably two acuminate, sub footprintsof These consist gent toenail overgrowth (Figure 8N–P) depart paleotracksassociated The with outward/diver narrower, thanouterdigit(IV). anddeeper prints, withinner toe (III) comparatively shorter, toepad and/or cracking); and (3) uneven sizing of nails (usuallyaccompanied withdistaltruncation toe elongation andantero-medial projection of digital axes and digital impressions;(2) curving of featuresary related to this condition: (1) inward second bilateral andacquire symmetry a seriesof and interclavularital septum spacing), lose their interdig toenail-prints; regular development of and toepad- width; parallelism and alignment of footprints ( features the intrinsic observed inthehealthy of overgrowth (Figure 8I–L), while retaining many attributedThose to an inward/subparallel clavular 7D–H), are relatively frequent inthefossil record. overdevelopment horny (Figure exhibitingof signs P digital axes. wardly directed andcoincidentwiththe rectilinear truncated appendices, which are straight, for appearing either as thinlineations and/orsharp/ In bothcases, toenail prints canbe present, theproducer organism. plantar physiognomy of deposits, isthe one that best manifests the palmar/ subtype, This generally linked touncompacted (which may beabsent) occasionally (Figure 8E–H). cavities separated by areduced mediumridge athological footprints. e.g. , similartracklength and track Llamoid paleoichnites, /2018 431 431 ------

Ichnopathology of Holartic Ungulates; Late Pleistocene of South America SYSTEMATIC ICHNOLOGY Ichnopathology of Holartic Ungulates; Late Pleistocene of South America SYSTEMATIC ICHNOLOGY 432 432 rlbnigo the toenail prints; (2) distal overex eral bending of imprints,toepad experienced hypertrophyand lat the development of despite maintaining a normal outwardlyfacing, digital impressions, which, divergent, forwardly (1) modifications: anatomical a bilateral symmetry, although undergoing deep notably from their natural morphology, conserving forefoot and hindfoot imprints: the first ones first the imprints: hindfoot and well-differentiated forefoot with ichnospecies, latter the between comb., baseduponthe heteropody exhibited by distinction specific Lamaichnum guanicoe a retained they generic synonymy by suggested these authors, Despite Aramayo with consequently the ichnogenera proposingto synonymize bothichnospeciesand such features aspotential ichnotaxobases, of Lucas and Hunt (2007) doubted the usefulness thelatter.divergence of mainly due to the greater and digital dimensions level: ichnogeneric and Lamaichnumguanicoe ichnospecific at both initially these ichnotaxa,1996) segregate Aramayo and Manera de Bianco (1987b, sis and South America: Pleistocene of llamoid ichnospecies are recognized for the Late Ichnotaxonomic remarks. the overdeveloped nails. tered footprints due to the anterior projection of development,to be shorter than inunal appears interdigital although retaining septum, a regular specially being notable this conditioninthe forefeet; and (6) the limb, affected the by exercised imprinted footprints, because the higher pressure deeper (5) ones; first the in greater toenails, pedal and manual of upgrowth differential a to owing heteropody, asecondary acquisitionof (4) normal the overgrownnails; antero-lateral projection of the tracks, due to the and transversal widening of the axial borders; (3) longitudinal enlargement of separation between both digital ends andcurving the interclavular gap, withanomalous of pansion / ) (Aramayo Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Lamaichnum tulipensis Lamaichnum et al . ., 2015). and and et al. (= Lamaichnum tulipensis Megalamaichnum tulipensis Megalamaichnum (2015) accepting the (2015)accepting Megalamaichnum tulipen Megalamaichnum Traditionally, two Lamaichnum guanicoe Megalamaichnum nov. - - - - /2018 ;

(see Aramayo andManera de Bianco, 1987b, guanicoe entiate ichnotaxobasesdiffer the thatto establishedcates More precisely, the new available evidence indi analysed andconfirmed). the toenails (hypothesis herein of deformation by overdevelopment horny and the consequent tulipensis of pattern same morphological Recently, Oliva andArregui (2016) recognize the directed, digitaltips. ones culminatingsecond insubparallel, forwardly pointed anterior ends facing outward, while the slightly larger impressed, and deeper showing and hindfootLDM-S tracks; superimposed forefoot (Figure 9A): couples of del Monte four footprints (Figure 9E). trackway of Aramayo, 2003;Aramayo undertracks (Figure 9C)(Manera de Biancoand four footprints, includingtwo tracksandtwo of Study material. 1966. Type ichnospecies. and Hunt(2007). Megalamaichnum sis Lamaichnum guanicoe Given the exposed results, we synonymize ichnotaxonomical order. of differences as interpreted anuniquespecies)thatnot be should variation of intraspecific of range the within (falling llamoids outward/divergent toenail overgrowth in living 1996; Aramayo aswell as the ichnogenera Ichnorder Perissodactypedida Vialov, 1966 Morphofamily HippipedidaeVialov,Morphofamily 1966 (and the ichnogenera that they typify) Megalamaichnum tulipensis Megalamaichnum Ichnogenus in footprints of modern camelids affected modern in footprints of ; LDM-S efimn h rtrao Lucas of criteria the reaffirming , (Table 4;Figure 9) et al Hippipeda Pehuen Co Pehuen and Hippipeda 1 S ,21)rpeetsgso ., 2015) represent of signs 15

Hippipeda aurelianis (Figure 9I) andLDM-S Megalamaichnum tulipen Megalamaichnum isp. et al 2 Vialov, 1966 S ; PCT-X: trackway 10 ., 2015);PCT-XI: from : three associated Lamaichnum Megalamaichnum Megalamaichnum Lamaichnum Laguna Vialov, and 2 S - - - 6

hindfoot tracks.Altered footprints (unbalanced (E–F)rearfoot hooves): impressionsshowing potentialsignsof contracted-heelhooves; Figure 9 Taken fromManera-de Bianco and Aramayo(2013). (G–H) hindfoot (I) imprints evidencing asymmetricaldistortion of forefoot thehooves; trackassociated withaturkish-slipper foot. C, Equid ( paleoichnites Hippipeda isp.). Unalteredfootprints (balancedhoofwall hooves):(A–B)forefoot imprints;(C–D) natural Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín /2018 433 433

Ichnopathology of Holartic Ungulates; Late Pleistocene of South America SYSTEMATIC ICHNOLOGY Ichnopathology of Holartic Ungulates; Late Pleistocene of South America SYSTEMATIC ICHNOLOGY 434 434 hoofwall thickness;L/W:length/width ratio. measurementsTable 4.Footprint ofSouthAmerican, Late Pleistocene,horse-tracks: Unbalanced remain 2013; Oliva and Arregui, 2016); all specimens footprints, preserved in loose slabs (Oliva 7 (Figure 9F) andMG-Pic 8 (Figure 9H): isolated three footprints); MG-Pic (eachof onecomposed and LDM-S footprints (Figure 9G); LDM-S distinguished basicallythrough their contrasting to the trace generated by a central digit III); lipsoidal to hemiovoidal toe impression (attributed tracks hindfoot hemiel a single (slight heteropody) of consisting and forefoot undifferentiated the Equidae Family.hoofed mammals of Almost type, unguligrade related to moderate-sized of Diagnosis. / Altered = Hooves Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Unaltered =Balanced Hoof Typology in situ Hooves Monodactylmesaxonic footprints, 2 (excepting MG-Pic7and8). T Contracted Heel Turkish Slipper Distorted Hoof 5 (Figure 9B): two related trackways Hoof PCT-X* Dimensional Range LDM-S LDM-S LDM-S LDM-S LDM-S 2 Designation T (acronym) MG-Pic 8 MG-Pic 7 MG-Pic 6 4 (Figure 9D) 2 2 2 1 2 T T S S S 6 5 4 15 10 I I 4 2 I I I I I I I I et al. I I I I I 3 2 1 3 2 1 2 1 2 1 3 2 1 neemnd156 01.62 10 65 105 Undetermined - /2018 , Footprint Referred idot1759 1.75 1.23 - 1.25 - 90 10 85 157.5 80 Hindfoot 105 100 Hindfoot Hindfoot idot8 5-1.3 - 65 1.3 1.2 85 - 8 50–100 Hindfoot 100 60–130 1.26 1.27 50 Hindfoot 130 8 8 Hindfoot 60 95 Hindfoot 90 120 115 Hindfoot Hindfoot oeot1258 2.03 - 1.25 80 - 162.5 100 Forefoot 125 Forefoot oeot1010-1.25 1.14 - 1.14 8 70–120 120 8 80–150 70 Forefoot 1.25 150 - 70 Forefoot 80 - 1.25 10 80 1.24 Forefoot 120 10 115 Forefoot 10 150 120 - 85 Forefoot 150 Forefoot 105 Forefoot Forefoot (Lund, 1846)and geus Possible trackmakers. a mediantalonarnotch. two mirrored, heel-bulb impressionsseparated by limited posterolaterallyprint usuallyappears; by thetrack,asubtriangular inner frog of rior half Surrounded bythe latter structures, ontheposte forward andinward inwedge-shaped bar marks. the wall bend on each side, the posterior ends of downward concave solar region, can be observed; curved hoofwall print, enclosing a more elevated, served specimens, anexcavated and strongly smaller and narrower than manus). Onbetter-pre and proportionsdimensions (pescomparatively Lund,1840(medium size); L/W H W L Hippipeda Measurements isp.L:tracklength; W:trackwidth;H: Hippidion devillei

Equus (mm) Hippidion principale ( (Gervais, 1855) 1.20 –1.30 1.14 –1.25 Amerhippus ) neo - - -

impressions, which canbe evenly(compact sunk anced hoof),constituting elongate hemi-ellipsoidal the producer autopode (bal anatomy of a normal Unaltered footprints. frog area. a thickhoofwall print) and aforwardly projected with an expanded ungular region (encircled by exhibited by equids,conformation the modern They display, onthe other hand, the autopodial genus the ichno distinctivemonodactyl track-pattern of Laguna del Monte (Oliva (Aramayo 2015), Monte Hermoso de Bianco andAramayo, 2003;Aramayo American equids, from Pehuen Co (Manera Description. et al Aramayo, 2003;Aramayo Province, Argentina (Manera deBiancoand tribution. dis and chronostratigraphic Geographic 1821). Gray, 1821;Equinae Gray, 1821;EquiniGray, (small size) (Perissodactyla Owen, 1848; Equidae bends and sharps forwardly andinwardlybends and sharps (heel buttress)postero-external angle andthen a V-shaped defines structure this side, each On thetrack. ference of the circum (8 – 10mm)thatthickness determines convex frontal wall (toe); and a nearly constant lateral respectively; walls (quarters), auniformly toU-shaped (Figure 9B) outline, withcurvilinear and rectilinear 9A) (Figure epicycloidal of flange, perimetral hoofwallprintconstitutesadeep The 1.20 –1.30)(Table 4). mm; width: 50 – 100 mm; length/width ratio: portionally longer than the pes (length: 60 –130 ratio:comparatively 1.14–1.25)is larger and pro 80 –150mm;width:70120length/width heteropody). Inthisregard, the manus(length: differentiating throughand proportions their dimensions (slight only pattern, morphostructural Forefoot andrearfoot imprintsexhibit the same sediment). substrate) forward and/or deeper (uncompacted ., 2013). Hippipeda aePesoeeo Buenos Aires Late Pleistocene of The paleoichnites The related to South . Those equid tracks reflect et al. et al ., 2013)exhibit the , 2007,2015;Oliva et al ., 2007)and et al. Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín - - - - - , Pathological footprints. (Figure 9C–D). the frogas baseline and theheel-bulb impressions tresses, disposedapproximately at the same level metric and having transversely aligned heel but tracks) are characterized by being almost bisym Invariably, the balanced hoofprints (unaltered axis) may bepresent sometimes. the track (symmetrically disposedonboth sides of and two postero-external heel-bulb impressions subtriangular inner frog print (axially positioned), cent-shaped sole region (radially distributed), a Enclosed insidethe hoofwall, amediumcres separated from itsoppositeby thefrog apex. (antero-medially) a wedge-shapedin bar mark per feet” (Figure notably 9I)depart from their Finally, the fossil tracksattributed to “turkish-slip asymmetry). sole area (solar development and distribution of heel buttress;displaced external and(5) uneven pillar (toe-quarter junction); (4) anteriorlyinternal nal wall (outer quarter); (3) forwardly projected longer and straighter than the exter abnormally frontal wall (toe); (2) medial wall (inner quarter) lateral, eccentric (sagitally disproportionate), thisanomaly:(1) unequi features characteristic of distinctive the solar region, through a series of or print hoofwall the into either reflected clearly distinguished immediately by their asymmetry, are 4), (Table 1.30) ≤ ratio (length/width tracks ilar andproportionsdimensions to the unaltered (Figure 9G–H),while maintainingsim the hoof footprintsThe indicating bilateral distortion of buttresses. space existent between the frog baselineandheel the heel-bulb impressionsand anenlargement of both talonar region, withanapproximation of the iting anarrowing anddistalprojection of width ratio >1.30)(Tablewell 4),as byas exhib elongation andtransversal compression(length/ from specifically healthy footprints by their greater antero-posterior differs These area). (ungular bilateral andanunaltered symmetry hoofprint with contracted heel (Figure 9E–F), conserve the “unbalancedhooves” evidencing potential of signs Equidpaleoichnites, /2018 435 435 ------

Ichnopathology of Holartic Ungulates; Late Pleistocene of South America SYSTEMATIC ICHNOLOGY Ichnopathology of Holartic Ungulates; Late Pleistocene of South America SYSTEMATIC ICHNOLOGY / DISCUSSION 436 436 secondary characteristics secondary linked tothe hypertro Likewise, these paleoichnites display several thebilateral symmetry. of thehoofprint,andaneventual loss projection) of (Table 4) with a considerable lengthening (distal 1.75) ≥ ratio (length/width enlargement gitudinal naturalmorphology, lon significant a manifesting Santucci Reynolds, 1999;SantucciandNyborg, 1999; 1984; Scrivner and Bottjer, and 1986;Sarjeant fossilIn tracks, besidesthehoofwallshape(Scrivner, as themostreliable diagnosticcharacters. unbalanced hooves). So, herein they are proposed or in balanced (either individuals conspecific on lateral remain walls (quarters) relatively constant of curvature and profile hoofwall the Regardless, unaltered hoofprints(non-pathological tracks). Therefore, it is advisable to restrict its useonlyto of measurements (length, width, length/width ratio). variations intraspecific teratologic order, to aswell as the overall footprint subject also are these features previouslyAs most of discussed, transversal widening). development (antero-posterior elongation and frog (3) and concavity/flattening); of degree and (profundity the wall); (2) solesagging of conicity ( of differentiation specific hooves andhoofprints:(1)hoofwall morphology the in determinant that three factors are primary recognition of comparative equids allows the study onmodern Ichnotaxonomicremarks. hooves). forward inanacute apex, inhealthy asoccurs thefrog print(whichdo notsharpen expansionof fected feet); and (3)overdevelopment and anterior unaf marks(unlike those medially convergent of postero-medially in overexpanded subparallel bar andouter prolongingquarters) longitudinally and ending (inner walls lateral (2) hoof; affected the rearward due theantero-dorsal bendingof plantar imprint (toe impression) unusually deeper the ungular region: (1) phy of anddeformation / e.g., Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín height, contour, thickness, inclination, and et al ., 2011), the length/width ratio, Acomprehensive - - - /2018 5. Discussion moreequid ichnospe thanoneSouthAmerican material preliminarily suggest the presence of and quarter convexities observed in the study profiles hoofwall contrasting the context, this In cific level. Arregui, 2016), have not been assigned to a spe et al (Manera de BiancoandAramayo, 2003;Aramayo Argentina Pampean Region of Pleistocene of austral recordsThe of ichnospecies. 2007) are particularly relevant in the distinction of bar marks(see Reynolds, 2006;McNeil of frog print, and morphology/extension absence of hoofprint area, hoofwall thickness, presence/ possible, through comparative analysiswith traces intrataxonomic alterations inpaleotracks, asfar as to identify thistype of it is necessary Then anomalies, necessarilylack taxonomic validity. pathological and/or congenital dimorphism, variations”such asontogenetic changes, sexual intramorphological or “intraspecifical the while theautopodes, the structurefunctionality of and characterictics are those intrinsicallyrelated to useful most the regard, this In criteria. differential or trackway are applicable as diagnostic and/or atrack Nevertheless, notall physicalfeatures of foot (Lucas, 2001,2005). producer the of characteristics anatomical reflect choosingcarefully ichnotaxobases which only mended to work with the best-preserved footprints, the pedal physiognomy). So, it is alwaysrecom deviations secondary unlinkedto of ing all kinds criteria (exclud based strictlyonmorphostructural theichnotaxa are imperative that thediagnosesof to vertebrateWhen itcomes ichnotaxonomy, itis them. define correctly each oneof differentiateordernotaxonomicto in analysis and cies. the ich to deepen Although, it is necessary ., 2007,2015;Oliva Hippipeda et al ., 2013;Oliva and , fromthe Late et al ., - - - - - h slr rcig n hrwl fissuration, hornwall and herein described for proboscidean paleoichnites cracking solar The differentiallyand/or all affecting (not extremities). rather erratic distribution, isolatedly occurring a show fractures), and fissures, trauma, injuries, as exungulation, ( lesions and horny digit, orosseous missing such affections, limb Certain et al McCrea 2006; Martin, and (Rothschild tification thatichnological allow patterns their correct iden the pathologies develop particular andrepetitive that so condition, this reflect that trackways and shape oroperation, producing anomaloustracks podal disease somehow its alters affected by sometype of limb a that expected is it Additionally, specifically important. see Rothschild andMartin, 1993, 2006) becomes (a standard procedure used inpaleopathology, the treated groups of parison withlivingmembers pathologic footprints. Consequently,of the com vertebrates) the recognition modern that permit in known affections limb to (related signs clinical pathological viewpoint, it is fundamental to detect species. Accordingly, modern from anichnoof mrns h cnptooi eerho llamoid imprints. ichnopathologic The research of alteredto distinguish successfully from unaltered the resulting trackways,of it is much more difficult In these cases, characterized by a greater regularity separately fore- and hind-feet). the opposite autopodes (at least considering of the mirroredand the specularanatomy symmetry portionately to the limbs,preserving commonly pro more affect to tend laminitis) deformation, overdevelopment, horny hooves distortionand In contrast, other podal disorders ( et al. ( feet) (unaffected elements unaltered anichnotaxon to the the systematic diagnosis of and unhealthyfootprints, crucial torestrict itis Hence, insetsandtrackwayshealthycombining maladies. ( Proboscipeda australis ., 2014,2015). , 2014). ,cnttt xmlso these ), constitute examples of e.g ., polydactyly, e.g ., McCrea e.g Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín ., - - - - only determined by the podal anatomy,determined only but also not is and the substrate. trackmorphology The a complex interaction between the autopode anichnite is the result of of formation The group. described diseases would be caused by the fourth (4) sicknesses, and(5) age-related problems. The biomechanical injuries, (3)brainorspinallesions, (2) ananimal: (1) congenital abnormalities, of foot mor the firstly phology, andultimatelyfunction thelocomotory affecting of potential the pathologies, with causative factors primary of McCrea sil tracksexpound clearly thisproblematic. ( Within ichnotaxa thismaincategory,of asubclass istics are called“phantomtaxa” (Haubold, 1996). while the ichnotaxa erected from these character characters”(Peabody, “extramorphological 1948), the trackmaker receive the denomination of tion of constitu podal the reflect not does that alteration any ichnologicalThe features that are productsof theichnotaxon. from thediagnosisof properly recognized and systematically excluded appearance and trackwayquality, andthat must be al (McCrea effects post-impressional and pre- of and Brokx,1999; Schulp, 2002),aswell as aseries factors, (Schulp either or post-depositional syn- external Therefore, therea great is number of McCrea 2007; Falkingham, 2014; Falkingham microbial avegetal cover) (Minter mat, development of superficial of presence rates, erosion energy, weathering intensity, flow sedimentation and environment, subaquatic ( and/or interface subaerial fluid/sediment the of properties consistency, infaunal activity), and of degree ( the track-bearer deposits type), the naturegait of maker behavior ( onextrinsicdepends agents such asthe track Lamaichnum guanicoe e.g. ., 2015), that can actively influence the footprint the activelyinfluence can that 2015), ., , composition, granulometry,, composition, moisture content, et al et al ., 2015). (05 pooe fv ctgre of categories five proposed (2015) . e.g ., foot pressure, speed, motion, ) andequid ( Hippipeda /2018 et al isp.) fos ., 2014; et al 437 437 e.g et ., ., - - - - -

Ichnopathology of Holartic Ungulates; Late Pleistocene of South America DISCUSSION DISCUSSION / CONCLUSIONS / Ichnopathology of Holartic Ungulates; Late Pleistocene of South America ACKNOWLEDGEMENTS 438 438 6. Conclusions Based on the exposed, we define the intraspecific the define we exposed, the on Based through non-anatomical ichnotaxobases. be considered invalid, becausetheyare diagnosed Necessarily, phantomtaxa and taphotaxa should 2001). on taphonomicalinducedcharacters (Lucas, “taphotaxa” is distinguished,based strictly termed • • thegroup: of living members by experienced alterations and affections pedal main the of some reflect that signs severalclinical Province, Argentina), allows the recognition of and Laguna del Monte tracksites (Buenos Aires Pehuen Co Pleistocene (Lujanian Stage/Age) of responding toHolarctic ungulates from the Late paleoichnites andtrackways, cor of analysis The previously known ichnospecies. acters, that constituting “teratologic synonyms”of those ichnotaxa basedonichnopathological char On the other hand, we denominate “teratotaxa” to physiognomy. factors, that from usuallydepart the real foot from due to extrinsic thoseextramorphological, differentiating autopodes), unhealthy case this (in anatomy podal the faithfully reflect that features secondary which constitute intramorphological trackmaker as “ichnopathological characters”, the variationslinked toanunhealthy condition of / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín f opooia nmle fthe digital anomalies of morphological of Equid footprints ( ichnotaxonomic implications. direct evidence, but paleopathology simultaneouslyof lacks significant represent which thesole), wall andtransversalof cracking as vertical(interpreted fractures onthe horn hoofprints the on fissures occasional present Proboscidean tracks( variations intraspecific will be essential to achieve a correct these of study and recognition future The horses). podal disorders inmodern hoof, andturkish-slipper foot (three habitual impression,linked tocontracted heel, distorted Hippipeda Proboscipeda australis isp.) exhibit a series - - - /2018 ) Acknowledgements • Manera, whose pioneer studies inpaleoichnology contribution to Ph. Silvia Aramayo and Dr. Teresa substantially this paper. We specially dedicate this Chávez andTom Musselman),who improved Dr. BruceRothschild),(Elizabeth andtheeditors made by(Dr. the arbiters Richard McCrea and thank the reviews, commentsandsuggestions tive paleoichnologic collections. We alsowantto Museums, for allowing usto access their respec Freyre” “Marcelino of (Guaminí) and “Carlos Darwin”(Punta Alta) staff the and directives Y-Tec/CONICET. We are to the grateful Cultura de la Provinciade Buenos Aires) and y Preservación Patrimonial (Secretaría de Guaminí, Dirección Provincial de Museos work This supported bywas de Municipio tional) variations. phological ( extramor and pathologic) dimorfic, ontogenetic, ( excluding intramorphological secondary characters, morphostructural based on only to establish ichnotaxonomic schemes intetrapods evidenceThe exposed inthisstudyclarify the need fichnotaxonomical order. Therefore,of we differences as species interpreted be aunique not should and of variation intraspecific changes clearlythe range fall withinof et al Manera de Bianco, 1987b, 1996;Aramayo tulipensis Megalamaichnum ichnospecies:second lids), which are often utilized a to segregate most frequent claw disorder in extant came overgrown(the horn playstrong evidence of Lamoid footprints ( ichnogeneric level). and ichnospecific at (both ichnotaxon the of definition and interpretation ichnosystematic h rtrao LucasandHunt(2007). the criteriaof typify, reaffirming these that ichnogenera the propose tosynonymize bothichnospecies, and ., 2015).However, these morphological e.g ., taphonomic, postural, locomo Lamaichnum guanicoe Lamaichnum tulipensis ) (see Aramayo and ) dis (= e.g. - - - - - ,

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