http://dx.doi.org/10.18268/BSGM2018v70n2a5 P. 351‒359 VOL. 70NO. 2 BOL. SOC.GEOL.MEX.2018 Mexico. Escondido –Oaxaca,C.P. 71985,Oaxaca, versidad delMar, Km2.5,Carretera Puerto Recursos, campus PuertoEscondido, Uni Laboratorio dePaleobiología, Institutode Rosalía Guerrero-Arenas [email protected] Eduardo Jiménez-Hidalgo Eduardo The OldestCamelFootprintsfromMexico Manuscript accepted: September 29,2017. Manuscript September accepted: Corrected manuscript received: July 15,2017. Manuscript received: March 31,2017. Jiménez-Hidalgo , Rosal - í a Guerrero-Arenas ewrs Camelidae, Mixteca. Oaxaca, Tezoatlán, Oligocene, guanicoe Lamaichnum Keywords: Eocene-early late Oligocene. the by evolved already had foot camel peculiar the of sug characteristics some camels, that gesting extant with features track foot anatomical some the shared makers that indicate Oaxacan the footprints, of imprints nail and Pseudolabis makers include track Potential imprints connected. digit have that footprints and anteriorly converge not do that impressions digit with ends, posterior rounded and ends anterior pointed guanicoe of features diagnostic the display footprints These late Formation. the Huajuapan Oligocene Eocene-early of sandstone tuffaceous fine-grained in epirelief concave as preserved are footprints camel The Mexico. southern Oaxaca, western north Luna, y Segura of Tezoatlánde from outskirts the in footprints located Mexico, camel oldest the describe we Here, known. is less much record Paleogene their however Mexico, including America, North pre well fromstrata Neogene served recordin a have footprints Camel ABSTRACT , such as bidigital tracks with tracks bidigital as such , Paratylopus . The concavesurface The axial . Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín , Paralabis Eocene- , Lamaichnum and - - - menos conocido.menos artículo describimos este En mucho es Paleógeno del registro sin el embargo, preservado; bien camélidos de huellas de registro un existe México, a incluyendo norteamericano, Neógeno del estratos los En RESUMEN Mixteca. Oaxaca, Tezoatlán, Oligoceno, Camelidae, Lamaichnum clave: Palabras dío-Oligoceno temprano. tar Eoceno el en ya evolucionado habían recientes camellos los de pie del anatómicas características algunas que sugiere anterior, actuales.Lo camellos los con podiales micas anató características algunas productores compartían los que huellas indican las oaxaqueñas, de uñas de impresiones las y y dígitos las huellas incluyen a de potenciales de productores Los impresiones conectadas. e anteriormente convergen no que dígitos de impresiones dos, redondea posteriores bordes y acuminados anteriores bordes con bidigitígradas incluyen:huellas cuales los presentan, que rasgos identificaron como se huellas Las Huajuapan. Formación la de temprano tardío-Oligoceno Eoceno del tobáceas areniscas en cóncavos epirelieves como preservadas están huellas de impresiones Las México. de sur de Oaxaca, noroeste Luna, y Segura de de Tezoatlán alrededores de los de antiguas procedentes más México, camello de huellas las Pseudolabis Lamaichnum L sprii ail cóncava axial superficie La .
guanicoe
guanicoe Paratylopus /2018 dbd a los a debido , , Eoceno- , Paralabis 351 - - -
The Oldest Camel Footprints from Mexico ABSTRACT The Oldest Camel Footprints from Mexico INTRODUCTION 352 352 Eocene-early of Tezoatlán, Oaxaca. Oligoceneage intheoutskirts de Rodríguez, Plio-Pleistocene of Puebla; 4. Tehuacán, Plio-Pleistocene of Puebla. The red stars indicate the ichnofossil localities of late Tepexi 3. Jalisco; of Plio-Pleistocene Lagos, los de Juan San 2. Durango; of Miocene Salto, El 1. Mexico. southern Oaxaca, Luna, y Segura Figure 1 1. Introduction ee eod s cre pbihd eod include records published scarce; is record gene Paleo the contrast, In 2007). Hunt, and (Lucas Mexico central to Canada western from ranging geographically strata, Neogene American North in recorded well relatively are footprints Camel 1 1 and (Lockley tracks turkey-like a with associated Salto), (El Durango of Miocene the from ported re formally been have ichnites camel Mexico, In 19 3 199 . 19 and Dakota South of Oligocene the from those / Map of Mexico showing the states with camel footprint records and a hypsometric map of the study area in Tezoatlán de Tezoatlán in area study the of map hypsometric a and records footprint camel with states the showing Mexico of Map Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín - - /2018 (Figure 1). dr (Dávalos-Álvarez ichnites canid with associated are footprints (Tehu camel where acán), Puebla southeastern of Plio-Pleistocene 2013; Jiménez-Hidalgo (Cabral-Perdomo, footprints ruminant small and proboscidean footprints, felid of kinds two prints, foot bird with associated are footprints camel the (TepexiRodrPuebla de southern of Plio-Pleistocene the from 2012); mán-Gutiérrez, (Rodr ichnites felid and proboscidean bird, with los associated de Lagos), Juan (San Jalisco of Plio-Pleistocene the í guez-de la Rosala guez-de 2012) Guzmán-Gutiérrez, and í uzd l Rs ad Guz and Rosa la guez-de et al. , 2015a); and from the t al et í guez), where guez), , 07 Ro 2007; ., - - - - is incm. with one footprint; 2. MGRN-02, isolated slab with two footprints (A and B); 3. MGRN-03, isolated slab with five footprints (A to E). Scale Figure 2 about theirpaleobiological implications. nites from Tezoatlán and to make some comments this paper is to describe the The aim camel of ich 1), where somecamel ichnites were discovered. (Figure Mexico southern Oaxaca, northwestern in Luna, y Segura Tezoatlánde near are calities lo these of Two localities. fossil Oligocene and Eocene new several study to us allowed has aca, Oax northwestern from Paleogenesediments the in 2009 since Paleontologicalout research carried Late Eocene-early Oligocene camel impressions from the surroundings of Juquila de León, Oaxaca. 1. MGRN-01, isolated slab isolated MGRN-01, 1. Oaxaca. León, de Juquila of surroundings the from impressions camel Oligocene Eocene-early Late Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín - - - 2. Geologicalcontext lower unit of Martiny of unit lower the with correlated be can beds fossiliferous The and R 19 9 breccia, conglomerate,sandstone, of consists that P Oaxaca (Figure 1), which the is Huajuapan part of Tezoatláneast of de Segura y Luna, northwestern í os-Mart í nez, 2009). et al. et (2000), which consists which (2000), /2018 353 353 -
INTRODUCTION / The Oldest Camel Footprints from Mexico GEOLOGICAL CONTEXT MATERIALS AND METHODS / The Oldest Camel Footprints from Mexico SYSTEMATIC ICHNOLOGY 354 354 4. Systematicichnology 3. MaterialsandMethods cene agefor theichfossiliferous beds. Oligo Eocene-early late a suggest which 2000), (Martinyavailablelowerunit are the Ma for 1. 3 . 1. 33. . el rsre fopit. he fopit pre footprints Three footprints. preserved well 3 with slab isolated MGRN-02, footprint; one with material. Referred Ichnogenus PecoripedidaeMorphofamily Remeika 199 . were derived fromProthero the data(1996) and of producers potential of estimations mass body the and (1996) Prothero of measurements on estimatedbased was length limb producer’s Potential and Hunt(2007). Lucas of ichnotaxonomy camel the followedWe length and widthare expressed inmm. maximum as taken were Measurements Naranjo. El to road the near bed sandstone a discoveredin were ichnites additional Three Nuevo. Rosario of town the in located (MGRN), Nuevo Rosario from collected slabs three foot in preserved are Eight prints 3). and 2 (Figures epirelief concave in footprints 11 of consists material studied The / Lamaichnum guanicoe Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Order ArtiodactipedidaVialov, 1966 Class MammalipediaVialov, 1966 Lamaichnum Aramayo andBianco, 1987 GN0, sltd slab isolated MGRN-01, 1987 rmy ad Bianco, and Aramayo et al. , 1995 et al. et - - - - /2018 , the pushup of sediment intherearof the print.Scaleisincm. note H3; Footprint 3. underprint; an represent to appears which H2, Footprint 2. H1; Footprint 1. Oaxaca. Naranjo, El to road the Figure 3 ae oeeery lgcn cml mrsin from impressions camel Oligocene Eocene-early Late * Standard* deviation. the lateEocene-early OligoceneofTezoatlán, Oaxaca. Table 1. Measurements of GN0. 50 65 MGRN-03.D 45 MGRN-03.A MGRN-02.A GN0. 45 60 MGRN-03.C MGRN-03.B MGRN-02.B GN0. 55 MGRN-03.E MGRN-01 pcmnLnt m)Wdh(m Depth(mm) Width(mm) Length(mm) Specimen Mean SD* . by separated are and divergent slightly or parallel and impressions are digit impressions digit ends.The roundedposterior two with an outline, with ovoid 1), (Table 1.21) = ratio (length/width Description. to ElNaranjo. road the near located bed sandstone a in served Lamaichnum guanicoe . (Lucas and Hunt, 2007). Thus, the Oaxacan prints of range length reported the mm (Table 1), which falls within the lower limit of The studied footprints have53.3 a mean length of imprints connected(LucasandHunt,2007). digit have that footprints and anteriorlyconverge not do that impressions digit with ends, posterior roundedand ends anterior pointed with tracks tal of features diagnostic Discussion. it probably (Figure 3.2). represents anunderprint so, others, the than shallower and smaller is print footOne (Figure3.1). cleft posterior absent an or such as a continuous interdigital sulcus (Figure 3.1) Some ichnites show extramorphological variation, are (Table 10to26mmindepth 1). . . the of most in evident are impressions nail Two H2 H1 H3 11.6 53.3 The ichnitesfrom showthe The Oaxaca 31 65 64 otrns r lne ta wide than longer are Footprints Lamaichnum guanicoe does not imply that we assume Lamaichnum 61.3 43.8 9.6 42 39 40 40 38 42 33 59 Lamaichnum guanicoe Lamaichnum , such as: bidigi as: such , footprints from 16.4 5.8 13 21 24 26 16 10 12 12 14 Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín - -
Oaxacan footprints 5. Paleobiologicalimplicationsofthe 1998), aswillbediscussedbelow. (Honey Oligocene Eocene-early late the members be should makers track Potential ichnofossiliferous beds. the of age estimated the than al. et (Honey age in Miocene early is camels lamine of record oldest The erected. was ichnospecies the when suggested (1987) Bianco and Aramayo as was maker track potential the that Tezoatlán deSegura yLuna. of southeast Yolomécatl.the to arerecords These in taxon camelid another of presence the dicate in specimens postcranial new collected recently (Jiménez-Hidalgo reported localities where Oaxaca, Yolomécatl, Eocene of late the from from comes camels Mexico of record osteological oldest The tomical pedalfeatures withextant camels. ana some shared Oaxaca, of Eocene-Oligocene the from imprints the produced that camelids the . 1 hoovescloven not and camels, extant of those to makers,similar track the footof the in nails terior (Thomson camels extant of of surface axial foot’sconcave anatomy. The the producer of some indicate do Oaxaca zoatlán, Te from footprints camel the of Characteristics 19 3 199 . 19 the amongst are and country the in record gene P The 1). and western northwestern, central Mexico (Figure includes Miocene and Plio-Pleistocene localities in ichnites camel of record Mexican published The 199 1 1 199 Lamaichnum guanicoe Lamaichnum footprints from Tezofrom footprints et al. et t al. et , 2007). The nail The 2007). , Poebrotherium /2018 21b. Also, 2015b). , Lama guanicoe Lama t al. et was 355 355 ------, ,
SYSTEMATIC ICHNOLOGY / The Oldest Camel Footprints from Mexico PALEOBIOLOGICAL IMPLICATIONS The Oldest Camel Footprints from Mexico PALEOBIOLOGICAL IMPLICATIONS 356 356 otrns n te rsne f n neir cleft/interclavular of anterior gap. Scaleisincm. an concavity of the presence the impressions, and footprints digit the from differentiated imprints nail the Note MGRN-03.C. 3. MGRN-03.B; 2. MGRN-03.A; Figure 4 / Close-up of some footprints of slab MGRN-03. 1. Ichnite Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín /2018 h camel The maker. Poebrotherium the of characters 19 Honey characteriz that cushion the lacked foot its and deer,and pronghorn resemblingpointed, of those and long,high are phalanges ungual the minants: of my anatoHowever,pedal Oaxaca. fromthe ichnites the of makers track potential the be would taxon this of individuals some that possible be could it of record logical geochrono Oligocene Eocene-late late the Given iat ad aie aeis (Janis camelids lamine and minants ru of that between intermediate is morphology bis somewhat divergent at their distal end, in In camelid living smallest the to comparable length limb and mass body ed estimat an with camels, medium-sized were ers mak track potential These footprints. Oaxacan the produced taxa these of individuals that sible Tezoatlán; therefore, pos is fossiliferousof it beds within the temporal range estimated for the ichno (Honey America the early Oligocene to North the early Miocene of and abis Oligocene, early to Eocene late from range otaaoia etrso the peculiar camel foot soft anatomical some features of that suggests footprints camel studied of the imprints nail and surface axial concave The the largest imprints. for least maker,at track potential as out ruled be of masses Pseudolabis Footichnites.of Oaxacan structure the of ducers abis of length limb Paratylopus and mass body estimated The (Prothero, 1996). of skeleton postcranial The the metapodials are partially fused and its foot its and fused partially are metapodials the Paratylopus has been recorded from the early Oligocene, early the from recorded been has make them good candidates as potential pro potential as candidates good them make Pseudolabis Poebrotherium t al. et adteetmtdbd aso of mass body estimated the and and the observed wide range of body of range wide observed the and Vicugna can be excluded as a potential track potential a as excluded be can 19) Hne gvn te observed the giving Hence, 1998). , and Paratylopus has a geochronological range fromgeochronological range a has Poebrotherium L. guanicoe L. , suggest that suggest , t al. et Pseudolabis is similar to that of typical ru typical of that to similar is 19) Tee eea are genera These 1998). , a a geochronological a has iun vicugna Vicugna imprints herestudied, imprints the metapodials are metapodials the Paralabis (Honey Pseudolabis is unknown is t al. et et al. et Tbe 2). (Table Pseudola 2002). , cannot , 1998), , Paral Paral ------Acknowledgements 6. Conclusions Data from: a)Quispe Eocene-early OligoceneofTezoatlán, Oaxacaand theextantlaminecamel of trackmakers potential of length limb estimated and estimation mass Body 2. Table n n f h letrcrso camel imprints in the oldest records of and one of Mexico in ichnospecies this of record oldest the icoe The late Eocene-early Oligocene based on osteological evidence by Janis determined independently also was this gocene; Oli Eocene-early late evolvedthe byalready had aul rv Cea hle t ipoe this Renter improveJiménez Jorge to grateful paper.Weare to helped Cuevas Bravo Manuel Victor and Lucas G. Spencer of comments number. The special this in participate to us invited for Barroso Quiroz Sara acknowledge kindly We thedistinctive camel foot. tures of gocene some camel taxa had some anatomical fea Oli Eocene-early late the bythat indicates which trackmakers, the in camels extant of features foot impres studied the of morphology footprint The as trackmakergiven itsfoot morphology. genus The Oaxaca. or of thatindividuals some probableis geochronologicalrange,it their Given North America. Paratylopus footprints from Tezoatlán, Oaxaca, represent Tezoatlán,Oaxaca, from footprints Limb length Body mass produced the camel impressions from et al (mm) (kg) . (2010);b)ChristiansenWalker (1964). (2002);c) Poebrotherium Paratylopus Paralabis labiatus 682 25 can be excluded be can , Lamaichnum guan Pseudolabis et al. Paratylopus primaevus 26.7 (2002). -- and/ Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín í a - - - - - Paralabis cedrensis References 32.6 -- Vicugna vicugna Nuevo. . Rosario to and Luna y Segura de atlán Tezo to visits our during attentions kind his for rmy, .. Bac, .. 18, Hallazgo 1987, T.M., Bianco, S.A., Aramayo, . fosil idades “Asociaciones255883 Nº project 2015 Básica cia Cien CONACYT by funded partially was study Torrey G. Nyborg for improving the This English. . P 3 13 . . P from tracks Mammalian 1976, P.R., Bjork, Mexicana, 3(1), 51–58. Rodr P TardCenozoico del terrestres vertebrados 55 15 15 . Dakota: South Academy Dakota South the of of Proceedings Formation Brule the P 53 5 . Santa Paleontologia: de Latinoamericano Carnivora, Congreso IV Suárez-Soruco,R., M., Riglos, II. (resumen), Aves y Parte Artiodactyla Argentina. Aires, Buenos de Provincia Rosales), Coronel de Tard (Pleistoceno continental icnofauna una de Pseudolabis dakotensis Lamaichnum guanicoe Lamaichnum í í o) en la localidad de Pehuencode (Partidolocalidad la en o) 45.57 guex, Estado de Puebla: Paleontolog Puebla: de Estado guex, í 875 feras continentales del Eoceno tard Eoceno del continentales feras . 760 –860 (shoulder Vicugna vicugna 35 –65 22 –54 height) 668.34 44.97 footprints from the late the from footprints c b c b a /2018 in Suárez- 357 357 í í í o o a - - -
CONCLUSIONS / The Oldest Camel Footprints from Mexico AKNOWLEDGEMENTS / REFERENCES The Oldest Camel Footprints from Mexico REFERENCES 358 358 hitasn P, 02 Lcmto i terrestrial in Locomotion P., 2002, Christiansen, . . 19 3 iée-iag, . Cro-hnn, G., Carbot-Chanona, E., Jiménez-Hidalgo, 2002, B., Boisvert, J.M., Theodor, C.M., Janis, cranial of Correlation 1990, C.M., Janis, D.R., Prothero, J.A., Harrison, J.G., Honey, R C., Fitz-Bravo, D / á Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín valos- egh n bn pootos n speed: on proportions bone and length of Sciences Natural Philadelphia, 116,1–13. of Academy the Naturae of Notulae Oligocene: River White P . . P P . 15 Vertebrate Paleontology, 22,110–121. a revisited: camels in evolution Locomotor Cambridge University Press, 255–299. U.S.A., York, New implications: biological and in estimation size Paleobiology: Body mammalian (eds.), B.J. MacFadden, in J., size macropodoids, body and with ungulates variables dental and 39 . Press, University Cambridge U.S.A.,York, New Mammals: Ungulatelike and Ungulates Carnivores, Terrestrial 1: Volume (eds.), North America, of L.L. Mammals Tertiary Jacobs, of Evolution K.M., Scott, C.M., Camelidae, 1998, M.S., Stevens, 1 mapa. P 1 5 1 Ciencias de Mexicana Revista Oaxaca: Estratigrafía 2007, E., Ramírez-Arriaga, E., Alaniz- 13 5 1 . Society, Linnean the of Journal Zoological Á vrz OG, it-aaig, A.F., Nieto-Samaniego, O.G., lvarez, 19 15. Á vrz SA, Martínez-Hern S.A., lvarez, í os-Mart í e, . 20, Carta 2009, D., nez, in in Damith, in Janis, á ndez, /2018 .. . 1 1 . . . Guerrero- K.T., Smith, E., Jiménez-Hidalgo, Quispe, E.C., Ramos, H., Mayhua, P., Alfonso, L., Camelidae, 1996, D.R., Prothero, Mart B., Martiny, S.G., Lucas,A.P., Hunt, Ichnotaxonomyof 2007, Lockley,Tracking2007, M.G.,an C.R., Delgado, eio uem f Ntrl itr & History Natural of Museum Mexico New America: North Western Footprintsof in activity mammal Mexico, Durango, near site and Miocene a from avian of evidence 5 39 . Sciences, Earth American South of Journal America: North Tropical from Assemblage Faunal Continental 2015b, Eocene Late J., First The Alvarado-Ortega, R., Arenas, P 1 1 1 . Biolog de Escuela México, Zaragoza, de Puebla Puebla: de estado del 93 . mensalis vicugna 1 609–651. Press, University Cambridge U.S.A.,York, Oligocene transition in North America: New D.R., Emry, R. (eds.), The terrestrial Eocene- Mexico: Tectonophysics, 318,71–98. magmatic rockswestern Oaxaca, of southern geochemistry Stratigraphy, 2000, Ayuso, R.A., C., Macías-Romo, D.J., Zenteno, 155–168. Museum Mexico New traces: and tracks J.A., Lockley, M.G. (eds.), Cenozoic vertebrate footprints, camel 67–72. Museum Mexico New traces: and tracks vertebrate Cenozoic (eds.), M.G. Lockley, Durango, Mexico, near from ichnofauna Miocene ancient turkey: a preliminary report on a new Science Bulletin,62,501–507. oke, .. Lcs SG (d., Fossil (eds.), S.G. Lucas, M.G., Lockley, in ua, .. Silan J.A., Spielmann, S.G., Lucas, í : ml Rmnn Research, Ruminant Small ): e-ern, .. Morán- R.G., nez-Serrano, in Lucas, S.G.,Spielmann, Lucas, í , Benemérita a, in Prothero, Vicugna . . . . . 199 19 9 .P. Rodr P. . . 1995 . . atn ruh eahet Anza-Borrego Detachment, Trough Salton Western the of Geology and Paleontology California, Anza-Borrego Desert State Park and vicinity, Basins, Felipe Borrego-San and Creek Fish Vallecito-the for list faunal Fossilvertebrate 36, 1–86. Bulletin, Museum Memorial Texas Texas: Trans-Pecos of Eocene) (Late Chadronian the from traces invertebrate and footprints Petroleros, 1,79–156. Bolet Oaxaca: de sedimentaria Cuenca 1 15 . P mam y aves de Huellas 2012, J.R., Field Trip Guidebook,I,82–93. Diego, Geologists, U.S.A., AssociationDiego San of San California: Park, State Desert í guez-de la Rosa, R.A., Guzmán-Gutiérrez, R.A., Rosa, la guez-de in eek, . Suz A (eds.), A. Sturz, P., Remeika, í feros del feros Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín í n ilv OS, 96 Sey zhiznedeyatelnosti Sledy 1966, O.S., Vialov, G.S., Morgan, R.S.jr., White, M.E., Thomson, . . 19 . . P 9 19 .P. ak kanky S Isiu Gooi I Geologii Akademiya Institut SSR Ukrainskoy paleontologicheskoe Ukraine, Nauk Kiev, ikh znacheniye: i organizmov 3 9 31 . & History Natural of Museum Mexico New traces: and vertebratetracks Cenozoic(eds.), Lockley,J.A.,M.G. Spielmann, S.G., Lucas, trackways?, fossil from determined be gait speed medium can trot: versus Pace 2007, P 3 3 . American the of Transactions Artiodactyla: the White River Oligocene: Partfauna of IV. P 5 . Baltimore,U.S.A.,II: volume JohnsHopkins Inskopayemykh, Naukova Dumka, 219p. Goryuchikh Geokhimii /2018 359 359 in
The Oldest Camel Footprints from Mexico REFERENCES