This is anopenaccess article undertheCCBY-NC-SAThis Universidad Nacional Autónoma de México. Peer Reviewing under the responsibility of Manuscript November accepted: 5,2019 Corrected manuscript received: October20,2019 Manuscript received: August 25,2019 org/10.18268/BSGM2021v73n1a100720 Mexicana, 73(1),A100720.http://dx.doi. Mexico:Boletín de laSociedad Geológica in its paleobiogeography andpaleoecology Valsequillo, Puebla,on withcomments of Mylodontidae) from the late Pleistocene harlani E., 2021,Anewrecord of E.,Jiménez-Moreno,F.J.,Benítez-Gálvez, Carbot-Chanona,G.,Jiménez-Hidalgo, How tocitethisarticle: na) * Corresponding author:(G. Carbot-Chano Puebla, Mexico. 4 Colonia Azcarate, 72000,Puebla, Puebla. 3 Escondido, 71985, Oaxaca,Mexico. sidad del Mar. Ciudad Universitaria Puerto Recursos, Campus Puerto Escondido, Univer 2 rez, Chiapas, Mexico. da de Las PersonasIlustres Tuxtla s/n, Gutiér de MedioAmbiente e Historia Natural. Calza ilera”, Dirección dePaleontología, Secretaría 1 Enrique Gerardo tardío deValsequillo,Puebla,concomentariossobresupaleogeografíaypaleoecologíaenMéxico Nuevo registrode paleobiogeography andpaleoecologyinMexico from thelatePleistoceneofValsequillo,Puebla,withcommentsonits A newrecordof license(https://creativecommons.org/licenses/by-nc-sa/4.0/) Museo de Museo Paleontología “Eliseo Palacios Agu Independent researcher.Independent Puebla deZaragoza, Universidad Angelópolis, Calle 3 Ote. 1603, Laboratorio de Paleobiología, Instituto de [email protected] (Owen 1840) (Xenarthra, Pilosa, Benítez-Gálvez Carbot-Chanona Paramylodon harlani Paramylodon harlani Paramylodon 4 1,* Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín , Eduardo - - - - - habitat. corridors,biogeographic Quaternary, late Pleistocene, Keywords: Mexico. north tothesoutheastof allowed it to extend its range from the wooded areas, andthis adaptation ronments, fromto more grasslands P the United States.indicates that This for this speciesbased on localities in with the preferred habitat proposed dominated by grasslands, andagree showed heterogenous vegetation Mexico where matic inference in somelocalities of Paleoenvironmental located between 1500 to 2000 m.a.s.l. the country. Mostlocalities are of part in the north and southeastern central Mexico, withsomerecords in theTrans-Mexican Volcanic Belt, Mexico, adding anewrecord for Mexico. In to identify it as andmeasurements allowedogy us Basin, Puebla state. Itsmorphol a left tibiafromthe Valsequillo illustrated. In thiswork, we describe few specimenshave been described or just mentionthe taxon and inpassing these records localities, but most of knownis from several late Pleistocene from Canada to Mexico. InMexico, it sloth recorded acrossNorth America, Paramylodon ABSTRACT . (Owen1840)(Xenarthra,Pilosa,Mylodontidae)delPleistoceno Jiménez-Hidalgo harlani couldinhabitdifferent envi (Owen1840)(Xenarthra,Pilosa,Mylodontidae) P .
harlani harlani P was a large was ground . Paramylodon occurred mainly Paramylodon
harlani and paleocli 2 , Francisco J. occurred,
harlani , - - - , Jiménez-Moreno /73(1)A1007202021 varias localidades del Pleistoceno tardío, hasta México. EnMéxicoseconoce en vés de América del Norte, desde Canadá tamaño distribuido a tra degran terrestre Paramylodon harlani RESUMEN hábitat. dío, Corredoresbiogeográficos, Pleistoceno tar Cuaternario, Palabras clave: desde elnortealsureste deMéxico. extenderse le hayaesa adaptación permitido hasta zonas más arboladas, y es posible que habitar diferentes ambientes, desde pastizales de EUA. Esto indica que propuestos para la especie en localidades pastizales, lo que concuerda con los hábitats heterogénea dominada por vegetación existió donde se encuentra paleoclimáticas de algunas de las localidades msnm. Las inferencias paleoambientales y localidades se ubica entre los 1500 a 2000 el norteysureste.en Elmayor número de el centro deMéxico, conalgunos registros en la Faja Volcánica Transmexicana, en P. harlani México. Nuestros resultados muestranque para registro con loqueseañade un nuevo referirla a permiten Valsequillo, Puebla; la morfología y medidas tibia izquierda provenientede la Cuenca de o ilustrado. Enestetrabajo describimos una menciones y poco material ha sido descrito pero la mayoría son sólo de esos reportes seencontraba principalmente P. harlani 3 , Paramylodon harlani fueunperezoso Paramylodon P. harlani , muestran que podía 1 , - - ,
Paramylodon harlani from the late Pleistocene of Puebla ABSTRACT Paramylodon harlani from the late Pleistocene of Puebla INTRODUCTION 1. Introduction 2 2 Croft, 2015). Extinct taxa were more diverse and the Cenozoic (McKenna range in thissubcontinent throughoutgeographic late Eocene-early Oligocene and they hada broad Groundoriginatedsloths the South Americain in the North Americanmylodontidsbelonged to the don synonymized and tooth, upper first the th isvariable, due to the presence or absence of upper tee Stock (1917)notedthat the number of don don nebraskensis material from HaySpring, Nebraskaas Kentucky, USA. Later, Brown (1903) described species th America.Inthe same paper, Owen erected the sed on a complete mandible and teeth from Sou genus lani McDonald andNaples, 2008). NorthAmerica (Kurténin 1980; Anderson, and several Irvingtonian and Rancholabrean localities 2017; 2018);thelater wellspecies is knownfrom 2017; Stinnesbeck and Carranza-Castañeda, 1985; McDonaldand De Iuliis, 2008;McDonald as mylodont X jeffersonii throtheriops there the four families, the mega comprised taxa of NorthAmerica sloths inthe late Pleistocene of ground1988; Webb, record 1989,2006). The of the Great American Biotic Interchange (Marshall ne-Pleistocene interval, being an important part of dontidae and Nothrotheriidae during the Plioce late Miocene, andlater Megatheriidae, Mylo Megalonychidae and Mylodontidae during the first, events; migration principal two in America day. had abroader spatial distribution range than to / / . Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín microcaninus, with upper teeth. However, basedonthenumber of h oecaua itr f nomenclatural of The history ground slothsreached North Four families of is problematic. Owen First, (1840) erected the Eremotherium Mylodon, M , Mylodon Meizonyx .
shastensis harlani Paramylodon , which he differentiatedwhich fromhe , and with the type species . Kraglievich (1928) argued that for material fromBig BoneLick,
; the megalonychids salvadorensis laurillardi Nohochichak
harlani et al , the nothrothere ., 2006;Gaudinand , (Webb andPerrigo,
xibalbahkah, Xibalbaonyx http://dx.doi.org/10.18268/BSGM2021v73n1a100720 Paramylodon M . darwinii Megalonyx Megalonyx Paramylo Paramylo
as well oviceps Mylo et
ba har
No al ., /73(1)A1007202021 ------/73(1)A1007202021 , separation between support for the and morphometric morphologic recommended. Finally, McAfee(2009) provided Rancholabrean specimens from North America is use of mylodonts, the and South American lineages of North isolationphic andseparate evolution of Donald (1995) mentioned that due to the geogra 1980;Polaco-Ramos,and Anderson, 1981).Mc name instead of ones, andthey shouldbe referred as generically separated fromthe SouthAmerican are not tioned that the North American forms genus (Arroyo-Cabrales and Polaco, 2003), Santiago co (Polaco-Ramos, 1981), Actun Spukil, Yucatán món andPérez-Rodríguez, 2017); Teapa, Tabas Morro delaMancha, Veracruz(Barrañón-Sal Valsequillo, Puebla (Pérez-Crespo Cruz Aquiahuac, Tlaxcala (Polaco-Ramos, 1981); lán, Morelos (Arroyo-Cabrales lleried, 1934); Cueva EncantadadeChimalacat Montellano-Ballesteros, 2008);MexicoCity (Mu berg, 1921;Mones, 1971;Cristín-Ponciano and and Valle de Bravo, EstadodeMexico (Freuden and Dalquest, 1975); Tequixquiac, Tlalnepantla 1981); Arroyo Cedazo, Aguascalientes (Mooser 2018); ElCedral, PotosíSan Luis (Polaco-Ramos, ta-Portalitos, Michoacán-Guanajuato (Eng-Ponce, Jalisco (Polaco-Ramos, 1981;Lucas, 2008);LaCin León (McDonald,2002);ZacoalcoandChapala, Bustamante Cave, Baja California; thern Nuevo CaboColnett nor in Rancholabrean localities of local fauna, Sonora (McDonald, 2002), and the ElGolfo reported viously from theIrvingtonian of Iuliis,Mexico, 2008).In burne, 1969;Churcher,and De 1984;McDonald subcontinent, fromCanada to Guatemala (Wood distribution extended acrossthe North American (Lambert and Holling, 1998;Smith reached abodybetweenmass 1,587to1,990kg Paramylodon Pleistocene specimensfrom NorthAmerica. as thevalidwith the second genusfor the Late Glossotherium Paramylodon Paramylodon Mylodon
harlani
. Later, Simpson(1945) men harlani
, andinsubsequentworksthe harlani Glossotherium was a large was groundsloth that for the Irvingtonian and P. wasused(
harlani et al and has been pre has ., 2004);Santa et al et al. Glossotherium Paramylodon e.g ., 2003); its . Kurtén , 2014); ------, , 2. Studyareaandgeologicalsetting Figure 1 sediments (Armenta Camacho, 1978; Pichardo,sediments (Armenta humanactivityhave been recovered inthese of specimens,and megafauna aswell as evidence 38.9 ka(Gonzalez that has been radiocarbon dated between 9.15 to the Valsequillo Gravelsexposure30 mthick of et al Gravels crop out in the Valsequillo Basin (Metcalfe namedValsequillo alluvial deposits A sequence of and paleobiogeography inMexico. Basin, Puebla Stateits paleoecology and to discuss to describeatibia of thiswork is described or illustrated. goal of The these specimens have been fully passing and few of these records are only mentioned in less, much of record of Gómez-Pérez, 2014). and (Carbot-Chanona Chiapas Villaflores, and Chazumba, Oaxaca (Viñas-Vallverdú Location of ElBalneariosite,BarrancaCaulapan,Valsequillo Basin,Puebla, Mexico. ., 2016). In the Barranca Caulapan there., 2016).Inthe Barranca a is P . harlani inMexico isbroad. Neverthe et al P. As demonstrated above, the
harlani ., 2006).Several mollusks Boletín de la Sociedad Geológica Mexicana Mexicana Mexicana Geológica Geológica Sociedad Sociedad la la de de Boletín Boletín , fromthe Valsequillo http://dx.doi.org/10.18268/BSGM2021v73n1a100720 et al ., 2017), - North American LandMammalAge. presence of The sand. fine-grained of m 0.5 by constituted is the section sandy-silt layer ispresent. top of The clay withorganic matter. layer Above, of 2mof sandy-silt., which iscovered by a0.6mthick m of sloth tibia described here. Overlying therea 1.4 is recoveredhorses in association with the ground mammoths, wolfs, bison, camels and bones of claywith organic matter contain fossil 0.4 mof 1.30mthickispresent. Above sandy-silt layer of Oversand. medium-grained this layer a layer of 1.1minthickness. Above follows a 0.7 mthick of constituted is bygravel sand and coarse-grained the section in progress).this localityis baseof The descriptionof sediments (the detailed stratigraphic at the El Balneario site consists 8 of meters fluvial of sequence (Figure general city stratigraphic 1).The Puebla deZaragoza m.a.s.l.); which liessouthof the Valsequillo Basin(18°57’N, 98°8’W, 2100 Caulapan, CerroBarranca LaMesaGrande, in described herefrom comes the ElBalneario site, 1997; Stevens /73(1)A1007202021 / 73(1)A1007202021 Bison et al indicates aRancholabrean ., 2012). The specimen ., 2012).The 3 3
STUDY AREA AND GEOLOGICAL Paramylodon harlani from the late Pleistocene of Puebla SETTING Paramylodon harlani from the late Pleistocene of Puebla MATERIAL AND METHODS COMPARISON 3.2. MEASUREMENTSANDMATERIAL FOR 3.1. STUDIEDMATERIAL 3. Materialandmethods Figure 2 4 4 vertebrate paleontology, andishousedinthe Mu left tibia collected with the traditional method of acomplete of specimendescribed consists The rudsoh rmteLt litcn fNorth ground slothsfrom the Late Pleistocene of large-sized taxa of comparedwas tothe tibia of order to taxonomically identify the specimen, it were madewithameasuringtape (Figure 2). In measurementsThe follow McAfee(2007) and der thecode3117P.F.10. Instituto Nacional deAntropología e Historia,un the and Archaeological and HistoricalZones)of Monuments Public Registry of (Unique System of MonumentosArqueológicasZonas y eHistóricas red in the Sistema Únicode Registro Público de under the numberregiste IHNFF-180,anditis seo dePaleontología “Eliseo Palacios Aguilera”, / / Schematic drawing Schematic showing the taken measurements. Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín http://dx.doi.org/10.18268/BSGM2021v73n1a100720 /73(1)A1007202021 - - - /73(1)A1007202021 3.2.1. MEASUREMENTSABBREVIATIONS it fthe diaphysisin the midshaft (MnW); 3, width of 1, Greatest length (GL); 2,mediolateral minimum number AMNH2780. Naturalunder the History of American Museum nebraskensis compared it with the type specimen South America. Additionally, we35-III-10-1) of (specimens MLP3-114, MLP 3-128, and MLP G with Stock, 1925;DeIuliis, 1996).We alsocompared it jeffersonii America, such as surface (AWDAS). distalarticular (DW); 10,anteroposterior width of distalend width (ICW); 9, mediolateral width of end acrossthe condyles (PW); 8, intercondylar proximal dyle (AWLC); 7, mediolateral width of lateral con (MWLC); 6, anteroposterior width of lateral condyle (AWMC); 5, mediolateral width of medial condyle 4, anteroposterior width of medialcondyle (MWMC); mediolateral width of . phoenesis Glossotherium , and (= (Cartelle Paramylodon harlani Nothrotheriops shastensis
Eremotherium tropicorum tropicorum et al ., 2019),and (De Iuliis
laurillardi ) housedinthe (Leidy, 1855; et al Paramylodon G , Megalonyx . ., 2017), robustum -
Figure 3 3.3. INSTITUTIONAL ABBREVIATIONS Ontario Museum,Toronto, Canada. de La Plata, La Plata, Argentina; CatólicaGerais, deMinas Brazil; de Ciências Naturais da Pontifícia Universidade Tuxtla Gutiérrez, Chiapas, México; the of de PaleontologíaMuseo Palacios“Eliseo Aguilera”, Collection Paleontological Geográfico, IHNFG Aguilera”, Tuxtla Gutiérrez, Chiapas, México; the Museo de Paleontología “Eliseo Palacios of Natural, Fósil Foráneo, Paleontological Collection New York, USA; AMNH TibiaIHNFF-180, mrcnMsu fNatural History, Museum of , American , Instituto de Historia Natural, Fósil Paramylodon harlani IHNFF Boletín de la Sociedad Geológica Mexicana Mexicana Mexicana Geológica Geológica Sociedad Sociedad la la de de Boletín Boletín , Instituto de Historia http://dx.doi.org/10.18268/BSGM2021v73n1a100720 , incranial (A),caudal (B),proximal (C)and distal(D)view. MLP MCL ROM , Museo , Museo , Royal
4.1. SYSTEMATIC PALEONTOLOGY 4. Results Genus Paramylodon harlani Subfamily MylodontinaeGill, 1872 Family Gill, Mylodontidae 1872 Davies inForster, 1795 Infraorder Tardigrada Latham and Order Pilosa Flower, 1883 Superorder Xenarthra Cope, 1889 Paramylodon /73(1)A1007202021 / 73(1)A1007202021 Figure 3,Table 1 Brown, 1903 (Owen, 1840) 5 5
Paramylodon harlani from the late Pleistocene of Puebla RESULTS
Paramylodon harlani from the late Pleistocene of Puebla RESULTS / DISCUSSION Valsequillo Basin,Puebla. Table 1.Measurements (inmm)of thetibiaIHNFF-180, from 4.2. DESCRIPTION 6 6 the medialmalleolus. tendinal groove is located in the posterior side of andsturdy. medial short malleolus is The One two bears large foramina inthe proximal surface. lateral3D). The tibial condyle isprominent and concavethe surfaceis shape and moon (Figure concave.its surfaceis facet ahalf- has discoid The wide andcoversalmost the entire distalarea and isanteromedially the astragalus positioned, it is of ally. articular facet for The the odontoid process later lies and flat shape, in semicircular is fibula (Figure 3C). Distally, the articular facet for the separated by asulcusinthe intercondylar area the lateral articular facet. Both facetsare size of medial-lateralin directionalmost twiceits andis shape andconcave, withthe longaxisoriented circular, while the medial articular facet is oval in almost and flat is condyle femoral the for facet lar (Figure 3A and3B). Proximally, the lateral articu straight, but thelateral bordermedially curved is the diaphysisalmost is medial borderThe of proximal and distal ends lateromedially expanded. tibia is longer than wide,individual. The withthe fused, indicating thatan adult the specimen was at dragging both ends. epiphyses The are nals of tibia IHNFF-180 iscomplete,The it shows sig / / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín AWDAS DW ICW P AWLC MWLC AWMC MWMC MnW GL W
1
106 136.1 29.2 166.1 71 69.3 97.1 94.5 96.2 259 http://dx.doi.org/10.18268/BSGM2021v73n1a100720 /73(1)A1007202021 - - - /73(1)A1007202021 5.1. COMPARISON ANDTAXONOMIC ASSESSMENT 5. Discussion phoenesis uccasamamensis thebe seenin SouthAmericantaxa mylodontine mylodonts, ascan to be typical of Paramylodon M. laurillardi the diaphysis islessconcave in medial border of shastensis and thininthe midshaft,like that of (Table 2,Figure tibiaof 4).The laurillardi of the tibia as in size, difference significant a literature (DeIullis, 1996)withIHNFF-180,show IHNFG-2715 and specimens reported on the A comparisonbetween eoa odl fIHNFF-180is wider in the femoral condyle of In proximal view, the articular medial facet for G lateral tibial condyleoriented is more upward in than the tibiae of moreis prominent andextends more upwardly the lateral epicondyle inthe tibia IHNFF-180 et al. IHNFF-180 and XIV, figs. 1,1a,1b, 2,3). (Leidy, 1885, plate XII, figs. 1-3; Stock, 1925, plate and the articular surface forwider is the astragalus less prominent; the medial malleolus is very short Tomiya grooves are present in thisregion (Stock, 1925; malleolus, while in the medial tendinal groove inthe posterior side of N. shastensis 2017; Cartelle the diaphysiswider thanin is of 1925). In IHNFF-180the mediolateral midshaft M. distinguished from the tibiae of .
The tibia IHNFF-180 isalso easily The We noted a major difference between difference major a noted We of those from differs IHNFF-180, tibia, The phoenesis jeffersoni jeffersonii , 2019)and (Saint-Andre and et al thaninIHNFF-180. the studiedspecimen twice the issize of than inIHNFF-180 and
u thstesm rprin f, but it has the same proportions of harlani ., 2001). The lateral., 2001).The tibial condyle is and reported in theliterature (Stock, Megalonyx , et al Glossotherium . jeffersonii M. (Tablerobustness 2).This seems G G ., 2019). Glossotherium N. shastensis . . phoenesis robustum et al Eremotherium
jeffersonii ., 2010;DeIuliis
and in that in one itbears . Inposterior view, and tropicorum tropicorum
E. . Additionally, the phoenesis N. G
N. laurillardi M. jeffersonii
laurillardi .
robustum shastensis shastensis Nothrotheriops G Simomylodon (Cartelle . robustum and is long is tibiae . The . The et al and and two G E. E. ., . .
South America. Table 2.ComparativemeasurementsofIHNFF-180 andPleistocene large-sized selectedLate ground slothtaxafrom North America and width of thediaphysis;PW, proximalwidth; RLB, Rancho La Brea. PWandMnW. Abbreviations: DW;C)DWvs D)GLvs PW;B)GL length;America. DW,distalwidth;GL, vs A)GLvs greatest MnW,minimal Figure 4 Bivariate plots of themeasurements Bivariateplots ofselected Late Pleistocenelarge-sized ground slothspeciesfrom North and South
Note: *Themeasurementsrepresentthemean;**estimated.
(MCL4303) Glossotherium Glossotherium laurillardi Eremotherium (n=6)* Megalonyxjeffersonii (n= shastensis Nothrotheriops Paramylodon (n=32)* Paramylodon AMNH specimen 2780) harlani Paramylodon (IHNFF harlani Paramylodon 2 )*
-
180) Taxon
harlani harlani
phoenesis tropicorum Boletín de la Sociedad Geológica Mexicana Mexicana Mexicana Geológica Geológica Sociedad Sociedad la la de de Boletín Boletín
(n=4) (type
*
* http://dx.doi.org/10.18268/BSGM2021v73n1a100720
182** 251.4 302.1 312.5 270.3 247.3 555 287 259 GL
138.4 182.6 184.4 206.6 185.2 180.7 PW 168 315 136
152.5 142.5 149.2 166.1 114 D 213 144 136 125 * W
*
1
MnW 106.8 75.6 61.7 93.8 94.5 96.2 78.7 70.7 110
MnW Ratio 0. 0.25 0.21 0.39 0.38 0.32 0.37 0.32 0.38 19
/GL
Hay Spring, Nebraska Spring, Hay Toca dos Ossos, Ouro Ossos, dos Toca pantla, Mexico pantla, Tlalne Corralito, Ecuador Several localities Several localities RanchoLa Brea, RanchoLa Brea, Califor Califor Puebla State Bahia, Brazil Valse Locality Branco, USA , Mexico , , nia , Mexico , /73(1)A1007202021 n quillo,
/ 73(1)A1007202021 ia ia
USA USA
, Ponciano, 2003 Ponciano, De Iuliis De Iuliis 1996 Cartelle Stock, 1925 Stock, Fields 2010 Fields Stock 1925 Stock This work This work This Cristín Source 2019 2017
et al et et al et -
.,
.,
7 7
Paramylodon harlani from the late Pleistocene of Puebla DISCUSSION Paramylodon harlani from the late Pleistocene of Puebla DISCUSSION epicondyle. condyle; genera.Abbreviations: between both proximal (I, I’),distal (J,J’),cranial(K,K’)and caudalto scale.Thearrowsshow (L, theprincipalmorphological L’)views.Not differences proximal (E,E’),distal(F,F’),cranial (G,G’)and caudal (H,H’)views;andphotograph anddraws of thetibia schematic of in proximal(A,A’),distal(B,B’), cranial(C,C’)and caudal (D,D’)views;photographdraws ofthetibia and schematic Cartelle MCL 4303, Figure 5 8 8 process of the astragalus isoval the astragalus in IHNFF-180 process of robustum anteroromedialThe border in IHNFF-180 thanin in the contrary, the lateral articular facet is smaller cranial side than in described not possible,and IHNFF-180was becauseboth between the tibiae of G in than IHNFF-180 in wider fibula is the for facet while in / / . Comparison oftherighttibiae Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín amf Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín phoenesis In distal view, the articular facet for the odontoid On the other hand, anobjective comparison , articularmedial facet for femoralcondyle; ismore oblique thaninIHNFF-180 G
. tibiae, left and right (ROM 4237and and phoenesis et al., G. robustum 2019)and and G .
phoenesis G G . . G. robustum phoenesis . robustum Glossotherium afo , articular facet for the odontoid process of astragalus; of process odontoid the for facetarticular , and Paramylodon harlani G http://dx.doi.org/10.18268/BSGM2021v73n1a100720 . and is circular. is The phoenesis G (specimen MLP 35-III-10-1). (specimenMLP Photographthe tibiaof draws of and schematic . robustum G
. tropicorum robustum and ff , facetforthefibula; le,lateralepicondyle; ; on G /73(1)A1007202021 /73(1)A1007202021 . . (typespecimenAMNH 2780),
nebraskensis the type specimen the tibiae of the ontogeneticdisparitybetween thespecimens. of consequence a be may differences minor these However, we donotdiscardthat the possibility the proximalwider than inIHNFF-180. endis the diaphysismoreis inwardly curved and of with IHNFF-180.InROM 4154the medial side 8K) itispossible to left tibiae (ROM 4154)(DeIuliis Iuliis specimen andthe epiphyses are (De missing ROM 4154,respectively) belong to ajuvenile The comparison between comparison The IHNFF-180 and et al (= ., 2017).However, intheillustrated Paramylodon harlani Glossotherium phoenesis alf ltc
, articular lateral facet for femoral for facet lateral articular , note some minor differences minor some note , lateraltibialcondyle; ) didnotshow any et al (typespecimen G. phoenesis G. robustum ., 2017, fig. 2017, ., Paramylodon me P. harlani , medial in in
Veracruz; 17. Teapa, Tabasco; 18. Actun Spukil, Yucatán; 19.Santiago Chazumba,Tabasco; 18.ActunSpukil, Veracruz; 17.Teapa, Oaxaca;20.Villaflores,Chiapas. Majorbiogeographic Mancha, la de Morro Puebla;Valsequillo,16. 15. Cruz Aquiahuac,Tlaxcala; Santa 14. Morelos; Chimalacatlán, 13. City; Mexico 12. State; Potosí; 8.ArroyoCedazo, Aguascalientes;9.Tequixquiac, EstadodeMexico;10.Tlalnepantla, MexicoState;11.ValledeBravo, Mexico Bustamante Cave,Nuevo León;4.Zocoalco, Jalisco;5.Chapala,6.LaCinta-Portalitos,Michoacán-Guanajuato; 7.ElCedral, SanLuis Black squareIrvingtonian Rancholabrean localities;blackdots localities.Localities: 1.El Golfo,Sonora; 2.CaboColnett,BajaCalifornia; 3. Figure 6 Tamaulipas –Central America Gulf Lowlands. Madre Oriental, 4.TransvolcanicBelt–Sierra Madre delSur.Tropicalcorridors: 5.Sonora –Central AmericaPacificlowlands and 6. Sierra – 3. Eastern US Madre Occidental, Sierra 2. RockyMountains – California, Baja – Western US corridors: 1. corridors: temperate h yeseie fthe type specimen of phoenesis tibiae IHNFF-180 andthe tibiae of same morphological differences noted between the Glossotherium size the in differences size. We with McAfee (2007) that agree there are tibiae of the between differences significant any exist not indistinguishable, assigned to almost are therefore, our studied specimen can be confidently and difference characteristic ormeasurements morphological Localities with McAfee (2007) mentioned that there does , G. robustum Paramylodon Paramylodon (Figure 4, Table 2). However, the Paramylodon harlani and and
harlani G between Boletín de la Sociedad Geológica Mexicana Mexicana Mexicana Geológica Geológica Sociedad Sociedad la la de de Boletín Boletín . Glossotherium Paramylodon tropicorum . http://dx.doi.org/10.18268/BSGM2021v73n1a100720 specimensinMexico(black numbers) and mainbiogeographiccorridors (blue numbers). Paramylodon exist between and these Glossotherium , except the and
AND PALEOECOLOGY 5.2. COMMENTSONTHEPALEOBIOGEOGRAPHY (Figure 6); thiscontrast with the much larger Rancholabrean localities are recorded in Mexico (Chiapas). One Irvingtonian and19 the country part of and Sonora) to the southeastern California distribution,geographic from the north (Baja In Mexico, (Figure 5). the species usingthe tibiae morphology between differentiate to possible is it and genera, both between differences morphological are there Glossotherium species. Therefore, we consider that /73(1)A1007202021 / 73(1)A1007202021 Paramylodon
harlani hada wide 9 9
Paramylodon harlani from the late Pleistocene of Puebla DISCUSSION Paramylodon harlani from the late Pleistocene of Puebla DISCUSSION Table 3.Altitudeofthelocalitieswithrecords of 10 10 (Stock, 1925; Naples, 1989);even,been ithas (Dalquest and Schultz, 1992) or mixed feeder (Brown,a grazer 1903; Webb, 1978), browser Historically, tropical lowlands Pacific ( corridors America Central – Lowlands, and Sonora Central America Gulf del Sur temperate andTamaulipascorridors, – Occidental, Transvolcanic Madre Belt – Sierra RockyCalifornia, MadreMountains –Sierra MadreUS –Sierra Oriental, Western US–Baja to move through Mexicoinclude the Eastern (Figure 6)thatcorridors altitudes (0 to1000m.a.s.l.). biogeographic The Mexico (Figurethree 6), at in southeastern lower Only three localities are located in the north, and Colorado (2,330 m.a.s.l.) (McDonald m.a.s.l.) and at the Magna Site, Saguache County, Utah(1,952 Silverin as Creek localfaunaof has been recorded also at high elevations, such Basin is at 2,100 m.a.s.l. In the USA, the species for the distribution of m.a.s.l. (Table 3), soaltitude is notalimiting factor MexicanVolcanic Belt, between 1500to2200 thelocalities arethe Trans- in 2012). Mostof localities from USA(see McDonald, number of / / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín sensu
Ceballos 20 19 18 17 16 15 14 13 12 11 10 9 8 7 6 5 4 3 2 1 Id
P
. harlani Villaflores,Chiapas OaxacaChazumba, Santiago YucatánActun Spukil, Teapa,Tabasco Veracruz Mancha, la de Morro Valsequillo,Puebla Tlaxcala Aquiahuac, Cruz Santa Chimalacatlán,Morelos City Mexico ValleBravo,Mexicode Tlalnepantla,Mexico Tequixquiac,Mexico ArroyoCedazo,Aguascalientes ElLuisCedral,San Potosí LaCinta Chapala,Jalisco Zocoalco,Jalisco BustamanteCave,Nuevo León CaboColnett,Baja California ElGolfo,Sonora Locality P. harlani. et al P has been considered has . ., 2010). harlani
- Portalitos, Michoacán http://dx.doi.org/10.18268/BSGM2021v73n1a100720
The ValsequilloThe probably used
Paramylodon harlan
et al
State State
., 2004).
State
- /73(1)A1007202021
/73(1)A1007202021 Guanajuato
i inMexico. 1
Marín-Leyva dominated by (Pérez-Crespo grasslands lived in areas withheterogenous vegetation with records of localities Villaflores, and Cinta-Portalitos some herbivorous mammalsfrom El Cedral, La tomore woodedgrasslands areas. environments, from different inhabit could it that of flexibility dietary The and showed thata grazer. thisindividualwas specimen of Pérez-Crespo a grazer.are InMexico, near to values indicative of inferred a mixed diet, but the values found (-4‰) from fauna, SanPatricio Ingleside County, Texas, fgrazer, browser andmixedherbivores feeders of In thesameway, inthe Valsequillo a mixturebasin δ using (2005) Rues showedBrea, California, that they were browsers. (δ nitrogen (δ carbon of isotopes stable using (2004) harlani 2006). Recent work hasinferred the diet of by andsedges (McDonald and Pelikan, grasses referred as granivore with a diet dominated
h eosrcino tedeayhbt f habits of the dietary reconstructionThe of basedonstable isotopes. Coltrain Paramylodon 15 Altitude (m.a.s.l.) P N) of specimens from specimens Rancho La N) of et al. et al . harlani 21)ifre h ito one . (2014) inferred the diet of , 2016;Díaz-Sibaja 13 from Valsequillofrom δ using rmteho specimens C from teeth of , indicate that these species 2100 1750 1358 1500 - - - -
2100 2220 2250 1850 2240 2600 1890 1750 2350 1540 1500 2200 1710 Paramylodon 915 560 45 38 72 8 7
et al. et al. indicates 13 , 2018). C) and , 2015; et al 13 P C . . 6. Conclusions 2004; Wilson etal.2005;McDonald2012). River ValleyMontana ( in Creek in Iowa, andDoeden gravel pit, Yellowstone and Springerville localities in Arizona, West Tarkio proposed to in Mexico with the habitats agree previously conditions reconstructed for some localities conditions. have had ageneralist diet to be adapted to these vegetation. Therefore, the influenced turn in which time, through Basin shows highclimatic variability in the Valsequillo moisture (Stevens raised again conditions increased, and later, after 20,000 years, one. Between 35,000and20,000yearswet Puebla was very similarto the present region of showsthat 35,000 yearsago, the climate thatin freshwater gastropods from Caulapan Barranca δ Valsequillo δ basin using stable isotopes of the the area. paleoclimatic The reconstruction of heterogenous vegetation hasbeen proposed for been reportedhas (Pérez-Crespo HF-8,tetp pcmno IHNFF-180, the type specimen of Mexico. the southeastof adaptation allowed it to extend from the north to fromto more wooded grasslands areas, andthis P indicates localities in the thatUSA. This northern proposed habitats for thespecies foundmorein dominated by withthe grasslands, andagrees harlani some localities in Mexico where inference of the speciesfor theValsequillo Basin. although thereunpublished isasecond record of described for thisarea, specimen the first formally and meristic characteristics. is morphological This us to identify it as the tibia IHNFF-180allows examinationThe of 13 . xrce rmtesel fterrestrial and C extracted fromof the shells The paleoclimaticThe andpaleoenvironmental The comparison between comparison The thetibiae paleoenvironmentalThe and paleoclimatic harlani occurred, indicate heterogenous vegetation could inhabit different environments, different inhabit could P . harlani Paramylodon from USA,such asShonto Paramylodon Boletín de la Sociedad Geológica Mexicana Mexicana Mexicana Geológica Geológica Sociedad Sociedad la la de de Boletín Boletín e.g.
harlani http://dx.doi.org/10.18268/BSGM2021v73n1a100720 McDonald et al et al.
, based on the ., 2012). This ., 2012). This harlani P . , 2014).So, nebraskensis 18 should O and et al. P , . 7. Acknowledgements References Paramylodon the type specimenof of for the photograph raphy, aswell as Miguel Díaz de León Muñoz the wikipaleo community for sharingbibliog and Eng fieldwork; Ponce Joaquín the in port We thank AbrahamLópezRojas for the sup both genera. between differentiation morphological the about and between differences morphological G Barrañón-Salmón, A.E., Pérez-Rodríguez,Barrañón-Salmón, V.J., Arroyo-Cabrales, J., Polaco, O.J., 2003,Caves Arroyo-Cabrales, J., Corona-M., E.,Polaco, O.J., Camacho, J.,Armenta 1978,Vestigios delabor ments that improved themanuscript. ald and one anonymousreviewer for the com G Alberto providedBoscaini of the photographs to used it; and Castor Cartelle the permission of vided the photographs . . tropicorum robustum Glossotherium 2017, Identificación de perezoso fósil del fósil Morro de la Mancha, Veracruz (abstract), perezoso de Identificación 2017, Indiana University Press, 273–291. America, North America:UnitedStates of Graham, R.W. (eds), Ice Age cave faunas of Mexico, inSchubert, B.W.,of Mead,J.I., and the Pleistocene vertebrate paleontology 21, 9–11. México: Current Researchthe Pleistocene,in Cueva Encantada, Chimalacatlán, Morelos, Basante, O., 2004,Recent excavations inthe Cruz Silva, J.A., Córdova, M.,Canto, G., Puebla, Mexico. del Estado dePuebla. Editorial del Gobierno de Valsequillo, Puebla,Consejo México: humana en huesos de animales extintos . We thanktoH.GregoryMcDon /73(1)A1007202021 ,
/ 73(1)A1007202021 G nebraskensis . robustum , contributing to the discussion and . FrançoisPujos pro Glossotherium G . phoenesis Paramylodon , revealed
phoenesis 11 11 - - - - -
CONCLUSIONS / Paramylodon harlani from the late Pleistocene of Puebla ACKNOWLEDGEMENTS Paramylodon harlani from the late Pleistocene of Puebla REFERENCES 12 12 rw,B,10,Anwgnso ground sloth Brown, B., 1903, A newgenus of Cristín-Ponciano,2003. Variación A., North Cope, E.D., Edentata 1889,The of Coltrain, J.B., Harris, J.M., Cerling, T.E., Churcher, C.S., 1984,Faunal correlations of Ceballos, G., Arroyo-Cabrales, J., Ponce, E.,2010, Cartelle, C.,DeIuliis, G., Boscaini, A.,Pujos, F., Carbot-Chanona, G., Gómez-Pérez, L.E.,2014, / / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín 569–583. Natural History, 19, of American Museum Nebraska:Bulletin from the Pleistocene of Special 2,12. Potosí,San Luis Paleontología Mexicana, Nacional de Paleontología:in XVCongreso morfométrica in https://doi.org/10.1086/274985 America: AmericanNaturalist, 23,657–664. palaeo.2003.12.008 199– 219.https://doi.org/10.1016/j. Palaeoclimatology, Palaeoecology, 205, Palaeogeography, California: southern late Pleistocene, coastal of palaeoecology biogeochemistryand itsimplications for the Allen, J., 2004,RanchoLa Brea stable isotope Ehleringer, J.R., Dearing, M.-D., Ward, J., GeoBooks,England, 145–158. Chronologies: Norwich, Quaternary in Mahaney, W.C. (ed.), Correlation of Canada, inwestern Pleistocene deposits doi.org/10.1016/j.yqres.2010.02.006 Research, Quaternary 73,464–473.https:// Mexico: the mammalian faunaof of on the distribution andcommunity structure Pleistocene environmentalEffects changes of doi.org/10.1080/14772019.2019.1574406 Palaeontology, anew 17(23),1957-1988.https:// Systematic intertropical Brazil: Journal of of of affinities mylodontine slothfromthe late Pleistocene phylogenetic and 2019, Anatomy, possible sexual dimorphism, Universidad delMar, 24. de México:Puerto Escondido, Oaxaca, II Simposiode Paleontología en el sureste Pleistoceno tardío(abstract), deChiapas in al registroAdición de megamamíferos del Paramylodon http://dx.doi.org/10.18268/BSGM2021v73n1a100720
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harlani harlani Glossotherium (Mammalia: ) (Xenarthra: (Mammalia: and 13 13
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