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

Carbot-Chanona, Gerardo; Eng-Ponce, Joaquín; Gómez-Pérez, Luis Enrique Description of the Neochoerus specimens from the late Pleistocene (Rancholabrean) of Chiapas, and comments on the taxonomic identity of the fossil capybaras from other Mexican localities Boletín de la Sociedad Geológica Mexicana, vol. 72, no. 1, e021019, 2020 Sociedad Geológica Mexicana, A.C.

DOI: https://doi.org/10.18268/BSGM2020v72n1a021019

Available in: https://www.redalyc.org/articulo.oa?id=94366149003

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 This is an open access article under the CCBY-NC-SAThis Universidad Nacional Autónoma de México. Peer Review under the responsability of 18,2019 Manuscript April accepted: Corrected manuscript received: 10,2019 April Manuscript received: March 10,2019 BSGM2020v72n1a021019 72 (1), A021019.http://dx.doi.org/10.18268/ Boletín de la Sociedad Geológica Mexicana, capybaras from other Mexican localities: the fossil on the taxonomic identity of Chiapas, andcomments (Rancholabrean) of Neochoerus the Pérez, L.E., 2020,Descriptionof Carbot-Chanona, G., Eng-Ponce, J., Gómez- To citethisarticle: [email protected] * Corresponding author:(Carbot-Chanona) Michoacán, Mexico. s/n, Ciudad Universitaria, 58030, Morelia, de Hidalgo.Nicolás Av. FranciscoJ. Múgica Biología, Universidad Michoacana de San 2 Gutiérrez, Chiapas, Mexico. da de Los Hombres Ilustres s/n,29000, Tuxtla de MedioAmbiente e Historia Natural. Calza ilera”, Dirección dePaleontología, Secretaría 1 Gerardo comentarios sobrelataxonomíadeloscapibarasfósilesotraslocalidadesmexicanas Descripción deespecimenes Mexican localities Chiapas, andcommentsonthetaxonomicidentityoffossilcapybarasfromother Description ofthe (https://creativecommons.org/licenses/by-nc-sa/4.0/) Museo de Museo Paleontología “Eliseo Palacios Agu Laboratorio dePaleontología, Facultad de specimens from specimens the late Pleistocene Carbot-Chanona Neochoerus 1,* , Joaquín - - Neochoerus specimensfromthelatePleistocene(Rancholabrean)of Pleistocene, Mexico. rus Keywords: Capybara, its spacialrangeinNorthAmerica. thisspeciesbroadens identification of referred as and Puebla allowmaterial this to be examinedthe specimens fromJalisco characteristics and measurements of dibular and dental morphological respectively,reexamined. is man The Hydrochoerus fromJalisco andPuebla, referred as addition, the capybara material municipality, Chiapas, Mexico. In atlocality,Mangos Los Villaflores remains from the late Pleistocene najuato. Here, we describe Michoacán andGua the border of Jalisco; andLaCinta-Portalitos, on coya, Mexico State; Chapala lake, aesopi (late Pleistocene). In Mexico, only during the Rancholabrean/Lujanian North, Central, and South America extinct giant rodent that inhabited capybaraThe ABSTRACT Eng-Ponce Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín , has been reported has fromTlapa Hydrochoerus delPleistoceno(Rancholabreano)tardíodeChiapas,y sp. and N. aesopi 2 , LuisEnrique Neochoerus aesopi Neochoerus . The definitive. The , taxonomy,, Neochoerus Neochoe N. aesopi isan Gómez-Pérez sp., N. N. - - - - México, sólo Lujaniano (Pleistoceno tardío). En Sudamérica durante elRancholabreano/ que habitó Norte,gigante extinto Centro y El capibara RESUMEN onomía, Pleistoceno, México. choerus Palabras clave: Capibara, América delNorte. en esta especie amplía su rango geográfico N. referirlosPuebla también como permiten de Jaliscoespecímenes examinados y y dentales, así como las medidas de los características morfológicas mandibulares y y Puebla (referido como mos el material de capibaras de Jalisco Chiapas, México. También, reexamina Mangos,el Municipio en de Villaflores, ceno tardío que afloranla localidad en Los encontrados los sedimentos en del Pleisto Aquí describimos los restos de en los límites de Michoacán-Guanajuato. Jaliscode Chapala, y La Cinta-Portalitos, Estado de México;ellago para Tlapacoya,

Neochoerus aesopi 1 . La identificación definitiva de , /72(1)2020 Neochoerus sp., respectivamente). Las N. Hydrochoerus

aesopi ha sido reportado ha sido reportado

aesopi Hydrochoerus es un roedor N. , tax

Neo aesopi sp. 1 - - - -

Neochoerus in the late Pleistocene of Chiapas ABSTRACT Neochoerus in the late Pleistocene of Chiapas INTRODUCTION 1. Introduction 2 2 rmEsnda mdl litcn)o Tarija, from Ensenadian(middle Pleistocene) of dichroplax ranza-Castañeda, 2016), referred as Guanajuato (Car Blancan III(early Pliocene) of 2016); (ii) (Carranza-Castañeda, SonoraandJalisco Pliocene–earlyPleistocene) of occidentalis choerus SouthCarolina (Hay,Pleistocene of 1923). M3 (thirdbased on a single upper molar) from the species tocene, Only two genera are known from the late Pleis South America from the Miocene to the recent. erinae) are large rodents that lived in North and capybarasThe (Rodentia: Caviidae: Hydrocho 1991). North and South America (Mones, Pleistocene of record throughout the late Pliocene to the late Hydrochoerus Neochoerus et al. Grenada, Lesser Antilles island-arc (MacPhee of Grenadines Bank,island to early Pleistocene of (Mones, 1991),and theVilla Ballester,early PleistoceneArgentina of fossil record: two speciesof 2005) and doesnothave a fossil record. Only Venezuelaand western (Woods andKilpatrick, isthmius rus 1991; Kerber and Ribeiro, 2011) and Curaçao, Uruguay, Brazil,andArgentina (Mones, species hasalate Pleistocene fossilrecord from Argentina (Mones andOjasti, 1986). The ern Peru, to south and Bolivia,aswell as northeastern Ecuador, It also inhabitsthe Amazon basinof Brazil, Paraguay, Uruguay,Colombia. andeastern Today, two livingspecies: 1980). Anderson, / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Currently, sixspeciesare recognized: (i) , 2000). Neochoerus Hydrochoerus hydrochaeris Hydrochoerus pinckneyi Hydrochoerus by Vucetich inhabitsPanama, Colombia, western (Vucetich isanextinct genus closely related to H. ballesterensis Hydrochoerus was erected was byHay (1926) for the from Blancan-Irvingtonian (late Hydrochoerus H. hydrochaeris and H. gaylordi et al. et al. Neochoerus are represented in the (2015); (iii) from late Pliocene to , 2014), with a fossil , which he described inhabitsVenezuela, fromlate Pliocene is represented is by http://dx.doi.org/10.18268/BSGM2020v72n1a021019 and N. cordobai (Kurtén and Phugatherium H. isthmius N. tarijensis Hydrochoe from Neo ------/72(1)2020 .

locality. (1991), becausebothtaxa came fromthe same was considered a synonymy of Rufolo, 2016), but 1998;Carranza-Castañeda, it ( 1926) hasbeen considered valid by some authors N. pinckneyi ica (Mones, 1991;Lucas North,Central, and South Amer Pleistocene) of N. aesopi Brazil and Uruguay; and(vi) (late Pleistocene) of a Argentina (although Mones [1991] considered it Villa Ballester,Pleistocene BuenosAires,of Bolivia; (iv) localities inMexico. late Pleistocene capybara specimens from other the also comment on the taxonomic identity of Mexico.these rodents in south-southeastern We with locality first the adding state, Chiapas from mally describethe capybarasrecovered specimens thisworkto foris known.Therefore, the aim of capybaras inMexico ispoorly fossilThe record of (Lozano-Ramos nate hydrochoerids from LosTanques, Zacatecas 2016), and indetermi 2006; Carranza-Castañeda, dez-Junquera, 1977; Pichardo, 1997;Mead lake, Jalisco; andValsequillo, Puebla (Hernán Potosí;lagoon, SanLuis Térapa, Sonora; Chapala choerus referred2016). Inaddition,specimens to Shaw andMcDonald,1987;Carranza-Castañeda, (Carranza-Castañeda occidentalis and Miller,(Carranza-Castañeda 1988),and Guanajuato from the Blancan to Irvingtonian of the genus include Other records of pala lake, Jalisco (Lucas, 2008;Eng-Ponce, 2018). Michoacán andGuanajuato; andCha border of Álvarez, 1969);LaCinta-Portalitos locality, onthe Tlapacoya, Mexico State (referred toas cies, e.g. nomen dubium nomen The only North American RancholabreanThe spe , Álvarez, 1969;Kurtén1980; and Anderson, N. aesopi fo h aePesoeeo Media Luna from the late Pleistocene of from the Rancholabrean/Lujanian (late fo rigoino Jalisco and Sonora from Irvingtonian of from South Carolina, USA(Hay, 1923, N. fontanai , hasbeen described in Mexico from ); (v) et al. N. sulcidens , 2006)have been reported. from late Pliocene to early et al. et al. , 1981; Shaw, 1981; , 2008).Occasionally, from the Lujanian N. aesopi Artícle A021019 N. pinckneyi by Mones N. cordobai Hydro et al. N. N. ------, ;

Artícle A021019 2. Studyarea Figure 1 silt and reddish-brown fine sand. In this section, brown bedsof alternating m layerof composed above2014). Stratigraphically a 1.50 thisbed is nona, 2012;Carbot-Chanona and Gómez-Pérez, Bison eus virginianus in association with aesopi choerus light brownsilty clay where 0.50 mlayer of gravel.granular Over the base section, there is a tion is 1 coarse to composed m fineof of sand with the sec crops out there (Figure base of 1B). The 4mthicklacustrine sediments 1A). Asequence of (Figure 93°13ʹ28ʺW and 16°14ʹ25ʺN at laflores, fromMangos locality, the Los near downtownVil materialThe described in thiswork wascollected Location (A), and stratigraphic section (B)oftheLos Mangos, Location(A),andsection Villaflores, stratigraphic Chiapas,Mexico. sp. remains (Gómez-Pérez andCarbot-Cha material reported herecollected, was , Equus mexicanus Glyptotherium floridanum Glyptotherium , http://dx.doi.org/10.18268/BSGM2020v72n1a021019 E. conversidens Boletín de la Sociedad Geológica Mexicana Mexicana Geológica Sociedad la de Boletín , Odocoil , and Neo G. G. - - - - - 3.1. ANALYZED MATERIAL 3. Materialsandmethods with left and right p4-m3 tooth rows andboth M3; IHNFG-4866,analmost complete lower jaw, maxilla with complete right 4865, afragment of Specimens used in thisstudy include: IHNFG- (Bell nology, late Pleistocene) for the bearing deposits (North American LandMammalAgesbiochro the Rancholabrean NALMA sp. is indicative of matter.with organic soil presence The of of the sequence consists 1 upper part of m thick.The sand, fine and silt sandy of composed is layer floridanum et al. remainshave next beencollected. The , 2004). / 72(1)2020 Bison 3 3 -

STUDY AREA / MATERIALS AND Neochoerus in the late Pleistocene of Chiapas METHODS Neochoerus in the late Pleistocene of Chiapas MATERIALS AND METHODS toothrow length; IDA, internal distancebetweenthealveolioffossa p4;DMF,distance through ofthemasseteric thePr.I of them1. second prism;Pr.IIb-Pr.IIb’, posteriorplateofsecond prism. Measurements abbreviation:L, length; W,width;DL, diastemalength; TL, prism;anteriorofprism; Pr.Ib-Pr.Ib’,Pr.IIa-Pr.IIa’,p4; Pr.Ia-Pr.Ia’, prismofof first first posteriorplate anterior plate of anterior plate external fissure; Pr.I, first prism;Pr.II, second prism;Pr.V,fifth prism; Pr.XII, twelfth prism;Pr.XIII, thirteenthprism;Pr.s.a.,secondary internal fissure; h.p.i.,primaryinternal fissure; HFI, fundamental internal fissure; H2-12I, second-twelveth internalfissure; HPE,principal h.t.i., tertiaryinternal fissure; h.s.i.,secondary internalfissure; h.s.i.a.,anterior secondary internal fissure; h.s.i.p.,principal secondary fissure;fissure;externalfundamentalsecondaryh.f.e.,external h.s.e., column; internal fist-third c.1-3i., column;externalfirst-third fissure;external c.1-3e., first-third fissure;h.1-3e., internalfirst-fifth upperh.1-5i., third molar; molar; M3, third lower m3: molar,lower from Carranza-Castañeda (2016).Dentalabbreviations: i,lower incisor;p4,fourthlower molar;m2,second lower premolar;m1,first Figure 2 4 4 clay. Specimens are housed at Paleontological epoxy with filled were fissures the and water, to parts were joined with commercial insoluble glue bristle brushes to remove broken sediment. The and cleaned with needles, airscribe, andsoft the traditional vertebrate methods, paleontology pelvic girdle. Specimenswere recovered using the skull; andIHNFG-5761,a occipital part of dible with an incisive, p4 and m1;IHNFG-5760, incisives preserved; IHNFG-4873,left hemi-man / Nomenclatureof thelower(A)and upper teeth(B),modified from Mones(1991),and somemandible measurements (C)taken Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín http://dx.doi.org/10.18268/BSGM2020v72n1a021019 - /72(1)2020 3.2. NOMENCLATURE authors ( authors Mones (1975a, 1991), adapted andused by other capybaras (Figure 2) proposed by dentition of We use the nomenclature for upper and lower e HistoriaNatural, Chiapas, Mexico. Palacios Aguilera”, Secretaría de Medio Ambiente the Paleontological Museum “Eliseo Collection of e.g. Kerber and Ribeiro, 2011;Vucetich Artícle A021019 Artícle A021019 3.4. INSTITUTIONAL ABBREVIATIONS 3.3. MEASUREMENTSANDPHOTOGRAPHS hydrochaeris (mean), neyi in Figure measurements 2. The of (2016), as shown(1991) and Carranza-Castañeda caliper with 0.01mmaccuracyfollowing Mones measurementsThe were obtained with a digital the nomenclature are discussed. in modifications where work previous no is there the taxonomicname, and reason for the change of used the name and Miller, (2016) 1988).Carranza-Castañeda the original description (see Carranza-Castañeda cordobai choerus al. Liebich (2007), Bode ments the nomenclaturebased onKönig was and For the cranial, mandibular and postcranialele and lowercase letters, respectively ( uppercase by identified are dentition lower and et al. Arizona, USA; IHNFG, Instituto de Historia Ney York, USA;AWC, Arizona Western College, Natural History, American Museumof AMNH, final figures were made inCorelDraw X8. Adobe Illustrator to erase the background. The rect lighting. Later, the images were processed in camera Canon XS, using alightbox andindi Material).(Supplementary 1and2 Appendix using the measurements of were elaborated in JMPversion8 (SAS, 2008), comparative(Rasband, 2012).The bivariate plots ranza-Castañeda (2016) using ImageJ software Car in presented figures the from obtained were measurementsThe of and Ribeiro, 2016). 2011;Carranza-Castañeda, Miller, 1988;Mones, 1991;Sanders, 2002; Kerber from the literature and (Carranza-Castañeda (2018). In the text, we employ the name The photographs were photographs taken with adigital The , N. aesopi , 2015; Carranza-Castañeda, 2016).Upper , 2015;Carranza-Castañeda, N. cordobai usedforwere comparison obtained (mean), because it is the first name used in used name first the is it because N. cordobae , N. occidentalis N. sulcidens et al. N. occidentalis (2013) and Brombini and didnot explain the http://dx.doi.org/10.18268/BSGM2020v72n1a021019 (mean), , and Neochoerus pinck Neochoerus e.g. from Sonora Boletín de la Sociedad Geológica Mexicana Mexicana Geológica Sociedad la de Boletín Hydrochoerus , M1,m1). N. tarijensis Neo et - - - - -

4.1. SYSTEMATIC PALEONTOLOGY 4. Results cias da Pontifícia Universidade Católica do Rio Coleção de Paleovertebrados doMuseude Ciên Grande do Sul, Uruguaiana, Brazil; MCPU-PV, cias da Pontifícia Universidade Católica do Rio Sul, Uruguaiana, Brazil; MCPU,de Ciên Museu tifícia Universidade Católica doRioGrande Paleovertebrados do Museu de Ciências da Pon do Sul, Uruguaiana, Brazil; MCN-PV, Coleção de da Pontifícia Universidade Católica do RioGrande de Mexico, Mexico; MCN-D, Museude Ciências tuto Nacional de Antropología e Historia, Ciudad Historia Natural, Chiapas, Mexico); INAH,Insti the Secretaría de MedioAmbiente e Collection of the acronymtorically is for thePaleontological (his Mexico Chiapas, Geográfico, Fósil Natural, AMNH under thecatalog number FM485. South Carolina, USA (Leidy, 1853). Housed at the Prof. Holmesonthe Ashley River, Charleston, described as terminate), Holotype. Subfamily Hydrochoerinae (Gray, 1825)Gill,1872 Family Caviidae Fischer, 1817 at Austin, Texas, USA. TexasPaleontology Laboratory, University of The Geosciences, VertebrateTMM, JacksonSchool of de Paleontología Geociencias, Querétaro, Mexico; de La Plata, La Plata, Argentina; MPGJ, Museo Grande do Sul, Uruguaiana, Brazil; MLP, Museo cias da Pontifícia Universidade Católica do Rio Coleção de Mastozoologiade Ciên doMuseu Grande do Sul, Uruguaiana, Brazil; MCPU-M, Infraorder Hystricognathi Tullberg, 1899 Class MAMMALIALinnaeus, 1758 Orden RODENTIA Bowdich, 1821 Neochoerus aesopi Neochoerus Genus Famn fthe incisive (position inde Fragment of Neochoerus 1839 Oromys aesopi / 72(1)2020 (Leidy, 1853) Hay, 1926 sensu Waterhouse, , collected by 5 5 ------

MATERIALS AND METHODS / Neochoerus in the late Pleistocene of Chiapas RESULTS Neochoerus in the late Pleistocene of Chiapas RESULTS as inFigure 2. angular process;conpr, condylar corpr, coronoid mcr,massetericcrest; mfo,massetericfossa. process; Dentalabbreviations process; apr, andPr.IIb’.Pr.IIa’Abbreviations: union betweenthe unusual the indicates circle red the (C); view in occlusal toothrow left the of Detail Figure 3 6 6 / IHNFG-4866, nearlycompletemandible from Los Mangos (A)and locality, Villaflores, Chiapas,inocclusal left lateral(B)view. Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín http://dx.doi.org/10.18268/BSGM2020v72n1a021019 /72(1)2020 Artícle A021019 Artícle A021019 4.2. DESCRIPTION hydrochaeris is robust and more retracted than in masseteric crestits lingual portion is 34 mm. The the p4 (IDA) in nal distancebetween the alveoli of throw, 110mminthe right toothrow). inter The the toothrow length (TL: 100.5 mminthe left too tema length (DL) is shorter (nearly 90mm)than are dias complete and included the p4-m3. The are(Figures missing 3Aand3B).Bothtooth rows However, the right condylar and angular process IHNFG-4866 isanearly complete mandible. 1975b); 1952); (Hoffstteter, (Spillmann, 1948); (Spillmann, 1941); robustus magnus Neochoerus (Leidy, 1887); vaisand Ameghino, 1880); aesopi rus Synonymy. The Pr.s.a.The hasaYshapelikethe Pr.I, while Pr.II bridge. enamel narrow a leaving fissure) internal lingual portion by the h.s.i.p. inthe (principal secondary and portion labial the in fissure) external (first h.1e. by flanked Pr.IIPr.Iis and of junction the prism. The the total width of than 90% of and more to fissure) extends external present h.2e.(second the not is fissure) internal (fourth h.4i. and Pr.II are joined by a small enamel bridge. The p4 show the Pr.s.a. anterior prism), Pr.I (secondary Both m1. the of fissure) internal (secondary h.s.i. extends the incisors below the alveolusThe of with longitudinal groovesthe labial in portion. They are robust andhave an anteromedial canal enamel in their lingual portion (Hillson,2005). them1is97mm. through thePr.I of the masseteric fossa (DMF) and the distance of the m1 the mandible locatedis below the Pr.I of masseteric fossa p4.The in the two portions of of Like inall rodents, the lower lack incisors (Kraglievich, 1930); Neochoerus pinckneyi Neochoerus (Leidy, 1856); and begins below the Pr.I (first prism) Pr.I(first the below begins and

Oromys aesopi Hydrochoerus holmesi (Kraglievich, 1930); Protohydrochoerus schirasakae Neochoerus Neochoerus sirasakae Neochoerus Hydrochoerus magnus (Leidy, 1853); (Mones, 1991). Prohydrochoerus sirasakae ( Hydrochoerus robustus http://dx.doi.org/10.18268/BSGM2020v72n1a021019 Neochoerus (Simpson,1928); Boletín de la Sociedad Geológica Mexicana Mexicana Geológica Sociedad la de Boletín Hydrochoerus Hydrochoe (Mones, Neochoerus Neochoerus ) sirasakae (Ger [ sic ] - - - - -

the labial portion (Figure feature 3C). This has the enamel bridge in and maintain the union of the second prism) and Pr.IIb’ (posterior plate of the Pr.IIa the second prism) into Pr.IIa’ (anterior plate of divide not does fissure) internal tiary Pr.Ia and Pr.IIb. On the left molar the h.t.i. (ter the Pr.IIa are V shape.complete Both m2 possess arate Pr.I them. The andPr.IIb are Yshape, while but theprisms doesnotsep thewidthof 90% of exceeding deep, is fissure) internal (primary h.p.i. enamel; the the labial portionby a thinbridge of of the m2,thisfeaturecharacteristic is the length of greater the p4is than a Vshape.has length of The has V shape.has Pr.s.a. The similar tothe adult is shows the Pr.s.a. and Pr.I inYshapewhile Pr.II poorly marked longitudinal striations. p4 The sor, which presents the anteromedial groove and the inci m1.It retains most of below the Pr.II of masseteric crest andthemassetericfossalocatedis the specimen preservesThe the distal portion of 4C). and 4B (Figures fragment molar upper fied with incisor, p4,m1,andanassociated unidenti ajuvenile the left hemimandible of portion of their distalportion. mesial portion are robust andtend to thin outat in the prisms and the 20.67 mm.The mm width is 59.12 the M3 is enamel laminae. length of The and Pr.XVI is represented only by a fragment of which Pr.Iincomplete in itsmesialportion is of 16prisms the M3 (Figure 4A).Itiscomposedof have aVshape. themolar. complete prisms The thewidthof of total width andthe h.s.e. is deep, exceeding 90% the has acomplete Pr.I; the h.s.i. exceeds 90%of the h.s.e.widening making shorter. left m3 The divided into two plates; the Pr.IIb hasanabnormal that area right m3 hasthe Pr.I is damaged. The because the rightm2 1975a) andcannotbeseenin hydrochaeris been observedjuveniles in andyoung adultsof N. aesopi The m1presentsThe the Pr.IIa joinedtoPr.IIb in IHNFG-4873 corresponds to IHNFG-4873 corresponds the anterior fragment with IHNFG-4865 isaright maxillary , buta rare itis feature inadults(Mones, (Mones, 1991). / 72(1)2020 7 7 H. - - - -

Neochoerus in the late Pleistocene of Chiapas RESULTS Neochoerus in the late Pleistocene of Chiapas RESULTS rectus femoris.Dentalabbreviations asinFigurerectus 2. foramen magnum; bone;pcp,paracondylar stu,sacraltuberosity; mcr,masseteric crest;o,occipital tmrf,tuberosityforthem. process; condyle;fmg, notch;co, acetabular acetabulum;an, ac, Abbreviations: mm. 50 to equal cases in all scale Bar girdle (EandF). pelvic view(A);IHNFG-4873,hemimandible leftright M3inocclusal inlateral(B)and (C)views;IHNFG-5760, occlusal parietal (D);IHNFG-5761, Figure 4 8 8 / Dental,cranialand postcranialmaterialof Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín http://dx.doi.org/10.18268/BSGM2020v72n1a021019 Neochoerus aesopi /72(1)2020 from LosMangos locality, Villaflores,Chiapas.IHNFG-4865, Artícle A021019 Artícle A021019 5.1. TAXONOMIC ASSESSMENT 5. Discussion f h ih lu rsre h rxmlpr f the right ilium preserves the proximal part of of the body lay inasagittal plane. upper side of The butmissing is notorious that itis the ilium wing both ilium distal part of ilium is straight. The The are elongated and narrowshape.and withanY bara theextant capy served andresembles thepelvisof theoccipitalcondyle. theupperendof level of paracondylar processes are small andare at the num.foramen The magnum isoval in shape. The the foramen mag and border the lower part of the foramen magnum. condyles The are thin of side each in depression smooth a with flat, almost of (Figureskull similar tothe occipital 4D)andis Pr.IIb hasaYshape. specimen, the Pr.I andPr.IIa have aVshapeand be observedcan and, unliketheadult the m1 in the prisms and Pr.IIvery is thin.Onlythe shapeof marked and,like the adult, the union between Pr.I (fifth h.5i. the little deeper. is is which h.4i. fissure), The internal of exception the with specimen is wider; the supraoccipital is wider andmorewider; thesupraoccipitalis is bar and lacrimal are more robust; the skull roof rounded and oriented backward; the anterorbital the zygomatic arcmoreis the anterior portion of rus close relative The located intheposteriormargin (Figure 4E). circular inshape,acetabular withasmooth notch the pubis. acetabulum The and deep is branch of the not preservedis observedonly and is part as of the ilium is preserved. pubis The bic eminence of tuberosity for the a prominent the ilium bears direction. bodyof The the sacral tuberosity, which isthininlateral-medial isalmost twice as large; the rostrum islonger; IHNFG-5761 is a pelvicgirdleIHNFG-5761 is partially con the IHNFG-5760 isanoccipital portion of Hydrochoerus hydrochaeris Neochoerus H. hydrochaeris Hydrochoerus genus canbe distinguished from the m. rectus femoris . Inventral view, thehipbones bythe following: . The occipitalboneis . The http://dx.doi.org/10.18268/BSGM2020v72n1a021019 . Onlythe iliopu Boletín de la Sociedad Geológica Mexicana Mexicana Geológica Sociedad la de Boletín Neochoe - - - - - 5.2. HYDROCHOERUS dental morphology. Inthe oldest species, Neochoerus Chiapas tothegenus teristics allow usto assign the fossil material from of and IHNFG-4866fall within the observed range IHNFG-4865 In addition,themeasurements of arein the M3 IHNFG-486516prisms observed. and the coronoid processslightly elevated,is while ble IHNFG-4866, the masseteric crestretracted is 2008). In the mandi (Mones,prisms 1991;Flynn, 14 to17prisms,has while coronoidprocess greatlyis reduced; andtheM3 excavated; the massetericcrestretracted; is the Carranza-Castañeda, 2016). Carranza-Castañeda, quera, 1977; Pichardo, 1997;Mead to Potosí; andValsequillo, Puebla, have been referred Terapa, Sonora; Media Luna lagoon, San Luis capybaras from Chapala lake, Jalisco; Material of the rangeof the DL of longer 90.05mm, than IHNFG-4866 hasaDLof like laminas the on fissures external extraordinary the specimen IHNFG-4865, the M3doesnot have and 70mm,while in DL in dentalis not present intheM3of characteristic is dental morphological 2014). This choeropsis taxa the Pliocene because it is alsopresent in the M3 of is a primitive dental character in the capybaras, nal fissures (Mones, 1991).Apparently, this feature za-Castañeda, 2016), called extraordinary exter and Miller,(Carranza-Castañeda 1988; Carran M3 bear small invaginations inthe labial side bai and Neochoerus The North American speciesinthe genusThe Another difference between difference Another Hydrochoerus N. aesopi Phugatherium , and N. cordobai (Deschamps N. occidentalis can be differentiated by its size and size its by differentiated be can N. cordobai (Figures 5Aand5B). All these charac . Inthe same way, the mandible of N. aesopi N. aesopi (Álvarez, 1971; Hernández-Jun INMEXICO? and , “ and Chapalmatherium isobserved in the DL. The . h eta aia fthe , the central laminas of Neochoerus N. aesopi et al. N. occidentalis N. aesopi / 72(1)2020 N. occidentalis , 2013;Vucetich Hydrochoerus it is 70–90mm.In . N. cordobai . is between 60 ” and , falling into et al. has 12–13 has , N. cordo , 2006; N. occi Hydro et al. 9 9 ------,

Neochoerus in the late Pleistocene of Chiapas RESULTS / DISCUSSION Neochoerus in the late Pleistocene of Chiapas DISCUSSION genus occidentalis 1991); graydot: purple dot: dot: catalogued);black not City, Mexico (INAH, Jalisco Lake, Chapala from Figure 5 10 10 / Neochoerus Bivariateplotof M3and p4measurements. Black asterisks: Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín (Carranza-Castañeda, 2016);greentriangles: N. pinckneyi N. tarijensis ; greenlines:trend ofthegenus (= N. aesopi mean(Mones,1991);yellowdot: , Sanders, 2002); bluedots: http://dx.doi.org/10.18268/BSGM2020v72n1a021019 Hydrochoerus Hydrochoerus hydrochaeris /72(1)2020 . N. cordobai N. aesopi Neochoerus aesopi mean (Mones, 1991); pink dots: mean(Mones,1991);pink dots: N. aesopi (Carranza-Castañeda and Miller,1988);orange dots: (Kerber and Ribeiro,2011).Red lines:trend of the from Valsequillo, Puebla (INAH CDMX, “66-I;Y-I”); CDMX, Puebla(INAH Valsequillo,from from Villaflores,Chiapas;reddots: N. sulcidens mean(Mones, Artícle A021019 N. aesopi N. Artícle A021019 5.2.1. SPECIMENSFROM CHAPALA LAKE hydrochaeris Hydrochoerus = 102.6 mm, = 112mm, TL range is between 90 and112mm( Neochoerus in chaeris more evidentand theTL.In the DL in Neochoerus of differentiation the allows that (1991) Mones by Neochoerus (JEP) showedauthors thathave thesespecimens the the oratorio de Arqueozoología, INAH), byone of these specimens from Chapala (housed at the Lab choerus labeled with the numbers 78and126as of istics mentioned that some specimenshave character za-Castañeda (2016) reviewed thismaterial and Chapala lake as Álvarez (1971) referred the specimens from while in ies. In var prisms issimilar,morphology the number of Neochoerus oneyoung individual. are indicative of the specimen DLandTLmeasurementsThe of Pr.s.a, characteristics of the masseteric crest begins inthe posterior part of p4 length isgreater than the m2 length, and the within the range of 89.87mm,measurements 40.3mmanda TL of of Pr.s.a. Onthe other a DL hand, specimen126has Hydrochoerus the Pr.s.a andthe Pr.I (like in Additionally, the masseteric crest begins between measurementsthe rangeof within 99.26mm, and aTLof 71.97mm of a DL has mm) (Mones, 1991). Specimen 78 from Chapala H. isthmius In the same way, inthe The M3 also exhibits differences between differences exhibits also M3 The terneo the whileDL is35–52.1 mm, therange of sp. (fig. 9, Nevertheless, 310). p. a review of Neochoerus Hydrochoerus Hydrochoerus and theDLhasalongerrange(60–90mm). from mandibular characteristics mentioned =64–91.4mm, i eisi h neirpr fthe it begins inthe anterior part of the ( TL range 61.5–91.4 mm is it is34.6–41.7mm(Mones, 1991).In N. aesopi Hydrochoerus N. tarijensis , the number of prisms is 14 to 17, is prisms , the number of Hydrochoerus Hydrochoerus , andreferred the specimens it is 12 to13(Mones, itis 1991; =90–110mm, Hydrochoerus N. aesopi = 101 mm), while in . Although the general H. isthmius . Such differences are differences Such . Neochoerus Neochoerus . Recently, Carran Neochoerus http://dx.doi.org/10.18268/BSGM2020v72n1a021019 (Mones, 1991). . However, the =61.5–71.5 Boletín de la Sociedad Geológica Mexicana Mexicana Geológica Sociedad la de Boletín species, the N. sulcidens ), while in Neochoerus N. cordobai H. hydro Hydro H. ------.

5.2.2. that the size of this molar is more thismolar is informative that the size of not duetospecificdifferentiation. age, and the is result someindividuals of size of teristics (Vucetich charac morphological the significantly affect not age, which does capybaras isaconsequenceof of to of some specimensfrom Chapala falls into the range to specimens za-Castañeda, 2016), which allowsthe assigning (also mentioned by14 prisms the M3 has Carran from2008). Insomeindividuals Chapala, Flynn, men isfrom ajuvenile (Figure 5A). speci this that indicates M3 of size The fissures. laminae, andthesewithoutextraordinaryexternal rimal more robust andhigher, andthe M3 with15 the supraoccipital wider andmore excavated; lac iswider than skull roof M3. The the skull and the characteristics of morphological aesopi Y-I”) allow usto assignthis specimen to zoología, INAH, labeled with the number “66-I; Valsequillo (housed at Laboratorio de Arqueo assignment. the taxonomic they are not specific on the basis of presence of Pérez-Crespo these remains have not been illustrated. Later, Valsequillo, Puebla (Pichardo, 1997).However, Hydrochoerus ments withinthesamearea isnotpossible. or more specieswiththe same ecological require two rejected,lake is because thecoexistence of ically variable tooth(Álvarez, 1971). differentiation, although it is the most morpholog and abetter indicator for THECAPYBARA FROM VALSEQUILLO, PUEBLA Neochoerus Hydrochoerus h iait lto the p4 (Figure 5B) shows bivariateThe plot of However, the teeth the variation in the size of h xmnto fthe incomplete skull from examinationThe of Finally, the presence of . The taxonomic. The assignation is based on the remains have been reported from or Neochoerus , whereas the taxonomicassignation Neochoerus et al. Hydrochoerus et al. (2014, 2019) mentioned the sp. inValsequillo. However, , 2005).Therefore, the small h ieo the M3 in size of . The / 72(1)2020 Neochoerus Hydrochoerus isdifficult(Figure 5A). Hydrochoerus to inChapala Hydrochoerus Neochoerus Neochoerus , with 11 11 ------

Neochoerus in the late Pleistocene of Chiapas DISCUSSION Neochoerus in the late Pleistocene of Chiapas DISCUSSION / CONCLUSIONS 5.3. PALEOECOLOGICAL SIGNIFICANCE LUNA LAGOON 5.2.3. HYDROCHOERUS FROM TÉRAPA ANDMEDIA 12 12 described or illustrated. Unfortunately, the reposi al and Hydrochoeridae (Nunez indeterminate been referred to capybaras fromFossil Térapa have remains of Pleistocene, which with thelithology consistent is waterthe area bodiesin duringthelate permanent in LosMangoslocality indicates the existence of ported. Therefore, the presence of al. (Vucetich deposits swampy or lacustrine fluvial, grove swamps(MonesandOjasti,1986). riverbeds,former wetlands, brackish andman water,the proximityin likeforested riverbanks,of are semiaquaticand live lowlands within habitats extant species of baras have been inferred by comparisonwiththe the extinct capy paleoecological aspects of The Neochoerus as fied specimens from Chapala and Valsequillo, identi of fications of (1984) consideredtions: (1)Mones that the reports as for both faunas, it is possible to refer this material range (late Pleistocene, Rancholabrean) assigned are capybaras from Térapa and Media Luna lagoon pers. com.). Nevertheless, we do believe that the nowand is (Ferrusquía-Villafranca, missing 2018, inamuseum1977), wasnever deposited formally assigned as baras from Media Luna lagoon,San Luis Potosí, a revisionnot possible. is capy Further, the fossilof these fossilsis notmentioned and therefore of tory / ., 2010).However,material this never has been , 2013), so the lifestyle related to water is sup Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín N. aesopi Fossilcapybaras are recovered usually from twoassump upon is based affirmation Such Hydrochoerus Neochoerus Hydrochoerus . . Hydrochoerus Neochoerus , and, considering , and, considering the temporal inNorth America are misidenti Hydrochoerus Hydrochoerus , showed that theyare actually n 2 h eiwo the , and(2) the review of sp. (Hernández-Junquera, sp. (Mead . Recent capybaras http://dx.doi.org/10.18268/BSGM2020v72n1a021019 Neochoerus aesopi Neochoerus et al. , 2006) et et ------/72(1)2020 6. Conclusions locality fed on different types of grasses, reeds, grasses, and shrubbery. of supported byis assumption This types different on fed locality is possible to infer that (Barreto andQuintana,2013). Paratheria prostrata son, andfeeds on amplexicaulis that leaves (Ojasti, 1973). Recent observations show aquatic plants, andoccasionallybark, stems, and comparison withfeedinghabitsof habits of the outcrops inthe study locality.of feeding The areas is both in present species only the that confirms and housed inLaboratorio de Arqueozoología, INAH, the specimens rejected based uponthe review of Chapala lake, Jalisco, and Valsequillo, Puebla, is Pleistocene. increases the spatial range of most record, until now, for North America, and taxon inMexico. It also documentsthe southern as assignation taxonomy aesopi Neochoerus specific the usto confirm allows Chiapas, Villaflores, from remains the capybara fossil description of formal The C4 mixed feeder(MacFaden andShockey, 1997). tarijensis dataThese are similar to that obtained from 2014; Eng-Ponce wereindividuals (Pérez-Crespo C4grazers feeder diet based on C3/C4 plants and other and showsthat have someindividuals amixed Michoacán-Guanajuato and Valsequillo, Puebla, δ and the microwear and stable δ isotopes analyses of ae ntefeighbt fBased onthefeeding habits of h itrclpeec f historicalpresenceThe of Hydrochoerus hydrochaeris H. hydrochaeris 18 from Bolivia, species referred to as a C3/ O on Neochoerus aesopi Neochoerus Neochoerus (an aquatic grass) during the wet (anaquatic sea grass) N. aesopi , which addsa new locality for this (a grass) during the dry season duringthe dry (agrass) Eleocharis interstincta et al. haspreference for canalsobeinferred through from La Cinta-Portalitos, , 2017;Eng-Ponce, 2018). . Inthesameway, we refer N. aesopi consumes grasses, consumes Neochoerus from LosMangos H. hydrochaeris Hydrochoerus H. hydrochaeris Artícle A021019 (a reed) and Hymenachne inthe late et al. in 13 , it N. N. C . - - , Artícle A021019 References Acknowledgements they made, andFrancisco J. Vega for the editorial anonymousreviewer for the helpful suggestions manuscript. and Torrey the Nyborg for review the English of wikipaleo communityfor bibliography, sharing de Arqueozoología, INAH. We are to the grateful Valsequillo and Chapala housed at Laboratorio to review the capybarasfor the permission from of Government Chiapas state. We the thankJoaquín Arroyo-Cabrales by financed Chiapas”, de ción yresguardo del patrimonio paleontológico the project “Prospec lected with the supportof specimensdescribed in thisstudyThe were col Mexico. late Pleistoceneof capybarathe only was this present species the in Sonora to Chiapasallowsus tohypothesize that lagoon as the capybarasfrom Térapa and MediaLuna Barreto, G.R., Quintana, R.D., 2013,Foraging oclusal figura la de Variación T., 1971, Álvarez, Álvarez, T., 1969,Restos fósilesde mamíferosde support. Finally, weAlvaroto thank wish and an Mones Herrera, E.A., Macdonald,D.W. (eds.), in capybaras, strategies and feeding habits of e Historia,SerieInvestigaciones, 21,1–35. México: InstitutoNacionaldeAntropología (Rodentia: Hydrochoeridae) de Jalisco, fósiles del premolar inferior en carpinchos History, 51,93–112. 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Neochoerus in the late Pleistocene of Chiapas REFERENCES