the diversity of glyptodontidae (xenarthra, cingulata) in the tarija valley (bolivia): systematic,...

DOI 1011270077-774920090251-0225 0077-7749090251-0225 $ 325copy2009 E Schweizerbartrsquosche Verlagsbuchhandlung D-70176 Stuttgart

N Jb Geol Palaumlont Abh2009 vol 2512 p 225ndash237 Stuttgart February 2009 published online 2009

The diversity of Glyptodontidae (Xenarthra Cingulata) in the TarijaValley (Bolivia) systematic biostratigraphic and paleobiogeographicaspects of a particular assemblage

Alfredo Eduardo Zurita Aacutengel Ramoacuten Mintildeo-Boilini Corrientes Esteban SoibelzonAlfredo Armando Carlini La Plata and Freddy Paredes Riacuteos Tarija

With 3 figures and 1 table

ZURITA A E MINtildeO-BOILINI Aacute R SOIBELZON E CARLINI A A amp PAREDES RIacuteOS F (2009)The diversity of Glyptodontidae (Xenarthra Cingulata) in the Tarija Valley (Bolivia) systematicbiostratigraphic and paleobiogeographic aspects of a particular assemblage ndash N Jb Geol PalaumlontAbh 251 225ndash237 Stuttgart

Abstract A revaluation on the main systematic biostratigraphic and paleobiogeographic aspectsof the Glyptodontidae fauna from the Pleistocene of Tarija Valley Bolivia are presented TheGlyptodontidae assemblage of this area is unique with respect to faunas known for other areas ofSouth America Some noteworthy features are the abundance of remains assignable to genusGlyptodon OWEN and the scarcity or absence of others (Panochthus BURMEISTER and Neosclero-calyptus PAULA COUTO) that are very frequent in the fossil record of the Pampean and north-centralregions of Argentina The validity of Hoplophorus echazui HOFFSTETTER as well as the presence ofH euphractus LUND and P tuberculatus (OWEN) are questioned pending discovery of more completematerials All specimens referred to Neothoracophorus AMEGHINO very probably correspond tosubadult Glyptodon specimens Biostratigraphically all but one of the Glyptodon specimens resemblethose from the Middle Pleistocene ndash Early Holocene (Bonaerian ndash Lujanian) of the Pampean regionin Argentina However one of the specimens studied (MNPA-v 006118) from the locality Armadoscorresponds to the species G munizi AMEGHINO restricted to the Ensenadan (Early Middle Pleisto-cene) in the Pampean region

Key words South America Bolivia Pleistocene biostratigraphy Glyptodontidae Glyptodonpalaeobiogeography systematics

1 Introduction

The Tarija Valley (21ordm 31rsquo S and 64ordm 43rsquoW) is locatedapproximately 1000 km southeast of La Paz (BoliviaFig 1) Tarija city is situated within it The valleyextends for approx 4500 km2 at nearly 2000 m abovesea level (MACFADDEN amp SHOCKEY 1997 COLTORTI etal 2007)The paleontological importance of the Tarija Valley

has been evident since the times of AMEGHINO (1902)and BOULE amp THEacuteVENIN (1920) although the first

fossil findings in the area date back to 1602 (seeBOULE amp THEacuteVENIN 1920 1 MARSHALL amp SEMPERE1991) In this context one of the main problemsthat affects most of the collections of Pleistocenemammals from the area is the lack of precise strati-graphic and geographic data for exhumed materials(HOFFSTETTER 1963 MACFADDEN 2000 TONNI et alin press) Nevertheless despite the uncertain pro-venance and specific identification of the collectedmaterials these have been and are still being usedto propose correlations and to establish biochrono-

eschweizerbartxxx ingenta

226 A E Zurita et al

logical schemes always in comparison to the schemefor the Pampean region of Argentina (eg CIONE ampTONNI 1995 1999 COLTORTI et al 2007 CIONE et al2007)Thus from a historical perspective the sedi-

mentary sequences that crop out in Tarija Valley havebeen successively assigned to different ages andstages ranging from the Lower to the Upper Pleisto-cene (DE CARLES 1888 AMEGHINO 1902 ROVERETO1914 BOULE amp THEacuteVENIN 1920 KRAGLIEVICH 1934OPPENHEIM 1943 PATTERSON amp PASCUAL 1972TAKAI et al 1982 1984 MACFADDEN et al 1983MACFADDEN amp SHOCKEY 1997 MACFADDEN 2000COLTORTI et al 2007 MARSHALL et al 1984 33)More recently and on the basis of a preliminaryrevision SOIBELZON et al (2007) and TONNI et al (inpress) have indicated that from a biostratigraphicviewpoint Tarija includes taxa characteristic of theEnsenadan Bonaerian and Lujanian interval (Early-Middle Pliocene ndash Early Holocene) included in thePampean chronological standard (see CIONE amp TONNI2005)From a geological and stratigraphical perspective

the Tarija Valley is part of a Quaternary sedimentationbasin filled with fluvio-lacustrine sediments thatdiscordantly overlie a Paleozoic basement (SUAacuteREZ-

MONTERO 1996) The most outstanding feature of thearea is the characteristic badlands landscape withirregular relief produced by differential sedimenterosion (OPPENHEIN 1943 SUAacuteREZ-MONTERO 1996)In this context COLTORTI et al (2007) included

the entire Pleistocene sequence within the TolomosaFormation subdivided into two major Units AncoacutenGrande Unit (AG) and San Jacinto Unit (SJ) Thetransition from AG Unit to SJ Unit is evident alongmost of the badlands The upper part (SJ) is yel-lowish-gray and fine-grained The lower part (AG)is reddish and coarser-grained (see more details inTONNI et al in press)The fossil mammal fauna exhumed from the Tarija

Valley comprises a wide taxonomical diversity (ca 55species HOFFSTETTER 1963 MACFADDEN 2000)including taxa characteristic for both forested or sub-forested relatively warm and humid environments(eg Tapirus BRUumlNICH Myocastor KERR NeochoerusHAY) as well as others adapted to open arid orsemiarid and cold areas where grasslands are domi-nant (Equus LINNAEUS Camelidae Glyptodontidae)(MACFADDEN amp SHOCKEY 1997) Some palynologicalstudies (TAKAI et al 1982) suggest that during most ofthe Pleistocene the Valley was dominated by an opensemiarid environment occupied by grasslands with

Fig 1 Location map showing the Tarija Valley (Bolivia)


scattered trees and shrubs that occurred along watercoursesThe Xenarthra (Cingulata and Tardigrada) are some

of the most frequent faunal elements approximately12 genera have been recognized (see HOFFSTETTER1963 TAKAI et al 1982 1984 COLTORTI et al 2007)4 of which correspond to Glyptodontidae Neothora-cophorus AMEGHINO Panochthus BURMEISTER Hop-lophorus LUND and Glyptodon OWENHere we discuss the taxonomical identifications

of Xenarthra Glyptodontidae that allegedly occur insediments from the Tarija Valley (Bolivia) their mainpaleobiogeographical features and their biostratigra-phic implications

Abbreviations GCF Grupo Conservacionista de FoacutesilesMuseo Paleontoloacutegico Fray Manuel de Torres (San PedroBuenos Aires Argentina) MACN Seccioacuten PaleontologiacuteaVertebrados Museo Argentino de Ciencias NaturalesldquoBernardino Rivadaviardquo (Buenos Aires Argentina) MCAMuseo de Ciencias Naturales ldquoCarlos Ameghinordquo (Mer-cedes Buenos Aires Argentina) MFCA Museo Univer-sitario ldquoFlorentino y Carlos Ameghinordquo UniversidadNacional de Rosario (ex Instituto de Fisiografiacutea y GeologiacutealdquoAlfredo Castellanosrdquo) (Rosario Argentina) MLP Colec-cioacuten Paleontologiacutea de Vertebrados Museo de La Plata (LaPlata Argentina) MMP Museo Municipal de CienciasNaturales del Mar del Plata ldquoLorenzo Scagliardquo (Mar delPlata Argentina) MNHNP Museacuteum National drsquoHistoireNaturelle Paris (France) MNK-PAL Coleccioacuten Paleonto-logiacutea del Museo de Historia Natural Noel Kempf Mercado(Santa Cruz de la Sierra Bolivia) MNPA-V MuseoNacional Paleontoloacutegico ndash Arqueoloacutegico Vertebrados(Tarija Bolivia)

2 Taxonomic biostratigraphic andpaleobiogeographic context of theCingulata Glyptodontidae from theTarija Valley Bolivia

Cingulata Glyptodontidae are among the most fre-quent elements of the megafauna known for theTarija Valley (HOFFSTETTER 1963 WERDELIN 1991COLTORTI et al 2007) and were first described byAMEGHINO (1902) and BOULE amp THEacuteVENIN (1920)although the greatest contributions correspond toHOFFSTETTER (1963 1964) TAKAI et al (1982 1984)and WERDELIN (1991)From a taxonomical perspective and taking into

account the existing associations and the frequency ofrecords the ensemble of Cingulata Glyptodontidaeexhumed from the Tarija Valley shows some signi-ficant differences with respect to the associationsknown for the Pleistocene of the Pampean region

central-northern Argentina Paraguay western Uru-guay and even with the locality Ntildeuapua (20ordm 52rsquo S and63ordm 04rsquoW) in southeastern Bolivia In these latterareas the dominant taxa are the genera GlyptodonNeosclerocalyptus Panochthus and to a lesser extentDoedicurus BURMEISTER (AMEGHINO 1889 HOFF-STETTER 1968 1978 MARSHALL et al 1984 SCILLATO-YANEacute et al 1995 CARLINI amp SCILLATO-YANEacute 1999CARLINI amp TONNI 2000 BAacuteEZ-PRESSER et al 2004UBILLA 2004 ZURITA et al 2004 ZUrita 2007)The composition of the glyptodont association is

remarkably different in Tarija Valley where more than90 of the records correspond to the genus Glypto-don In this sense most of the materials have beentraditionally assigned to the species G reticulatusOWEN (AMEGHINO 1902 HOFFSTETTER 1963 TAKAIet al 1982 1984 WERDELIN 1991 MARSHALL ampSEMPERE 1991 COLTORTI et al 2007) and G clavipesOWEN (BOULE amp THEacuteVENIN 1920) However andcontrasting with the case of the North AmericanGlyptodontinae that have been recently reviewedusing modern systematic criteria (GILLETTE amp RAY1981) thus far no similar updated study has beenmade for the South American taxa (SOIBELZON et al2006) In the South American case several speciesincluded in the genus Glyptodon were recognizedby numerous authors (eg OWEN 1839 1845 BUR-MEISTER 1866 AMEGHINO 1881 1883 1889) mainlyon the basis of dorsal carapace fragments This clearlytypological species concept (see MAYR 1996) charac-teristic of the 19th century (GIRAUDO 1997 TINAUT ampRUANO 2000) has indubitably led to remarkable over-estimation of the specific diversity of this genus(DUARTE 1997) a phenomenon that had already beennoted by AMEGHINO (1889) himself One of the directconsequences of this peculiar situation is the lack ofmorphological support for many species assigna-tions Despite this evident constraint the numerousmaterials (mainly skulls mandibles and dorsal cara-paces) referable to Glyptodon deposited in the collec-tions of MNPA-V (eg MNPA-V 006084 006088006119 006102 006077) (Fig 1A) in addition to thespecimens illustrated by TAKAI et al (1982 26 f 27figs 20-22) show morphological similarities to thespecimens collected from the Middle Pleistocene ndashEarly Holocene of the Pampean region (Bonaerianand Lujanian Stages see BURMEISTER 1870-1874pls 25-28 AMEGHINO 1889 pl 52 fig 1 LYDEKKER1894 pls 4-5 MLP 18 MCA 2015 1086) (Fig 3Table 1) Although most specimens lack precisestratigraphic and geographical provenance such data

The diversity of Glyptodontidae in the Tarija Valley (Bolivia) 227



Fig 2 (Legend see p 229)


are available for at least some of them [MNPA-V006088 from the locality San Pedro Tarija Valley andthe materials illustrated by TAKAI et al (1982 27 fig21) from the upper member of ldquoTarijardquo Formation(= Tolomosa Formation see SUAacuteREZ SORUCO amp DIacuteAZMARTIacuteNEZ 1996) (sic TAKAI et al 1982 11)Recently SOIBELZON et al (2006) have provided an

enhanced morphological and stratigraphical charac-terization of G munizi AMEGHINO which is at presentthe only species of Glyptodon that occurs with cer-tainty in the Ensenadan Stage (Early-Middle Pleisto-cene) outcropping at the ldquoToscasrdquo del Riacuteo de LaPlata (MACN 8706) San Pedro (GCF 10) and Mardel Plata (MMP 3985) in the Pampean region ofArgentina) Furthermore a specimen assignable to G

munizi (MNPA-V 006118) exhumed from the localityArmados 40 km South from Tarija has been foundin the collections of MNPA (see TAKAI et al 1984 55)(Fig 2B) This species is characterized by a robustskull with great dorso-ventral diameter in additionthe skull is more elongated than that of MiddlePleistocene ndash Late Pleistocene forms with markedlynarrow region between the postorbital apophyses andthe supraoccipital crest clearly developed antero-inferior margin of the orbital notch and lower thirdof descending processes of maxillaries and firstmolariform with more primitive morphology (seeSOIBELZON et al 2006) (Table 1) The presence of thisglyptodontine in the Tarija Valley represents an im-portant biostratigraphical observation because it im-


Fig 2 Skull in lateral view A ndash Glyptodon sp (MNPA-V 006084) B ndash Glyptodon munizi (MNPA-V 006118) C ndash Glypto-don cf elongatus dorsal carapace of a juvenile specimen (MCA 2017) D ndash Detail of lateral osteoderms (MCA 2017)E ndash Glyptodon sp (originally classified as Neothoracophorus) lateral osteoderms of a juvenile specimen (MNPA-V005423) F ndash Panochthus sp proximal fragments of a caudal tube (MNPA-V 006598) G ndash Hoplophorus echazui distalportion of a caudal tube (MNPA-V 142)

Table 1 Comparativemeasurements (in mm)of the Glyptodontinaespecies present in theTarija Valley



plies the presence of some levels of Ensenadan ageat least at the locality Armados (Fig 3) Anotherremarkable aspect is that the fossilization type of thisspecimen is quite different from that of most materialsreferred to Tarija Valley as it is heavily mineralized(in contrast to the very poor mineralization of othermaterials)The other taxon that occurs exclusively in the

Ensenadan of the Pampean region and has been foundin Tarija Valley is the Ursidae Arctotherium angusti-dens GERVAIS amp AMEGHINO regrettably the materiallacks well-defined geographical or stratigraphical pro-venance (SOIBELZON 2004 SOIBELZON et al 2005) Itis noteworthy that its fossilization type is the same asthat of the G munizi specimenFrom a palaeobiogeographical perspective the

species of Glyptodon were the most frequent Pleisto-cene Glyptodontidae from approximately 22ordm S to 4ordm N(BOMBIN 1981 MARSHALL et al 1984 MARSHALL

amp SALINAS 1991 PUJOS amp SALAS 2004 CARLINI etal 2008a b) probably following ldquopara-Andeanrdquocorridors (CARLINI amp ZURITA 2007 CARLINI et al2008a) In the region of the Cusco Valley Peruremains of Glyptodon cf G clavipes have been citedat an altitude of 3350 m above sea level (HOFFSTETTER1970 PUJOS amp SALAS 2004) while MARSHALL ampSALINAS (1991) have reported the presence of thisgenus in Ulloma Formation Bolivia at over 3880 mabove sea level In this context BENJAMIN et al (1987)estimate that the uplift of Eastern Cordillera in Boliviaand Peru accelerate at the beginning of the Quater-nary reaching a uplift rate of 07 mmyr (see alsoREGUERO et al 2007) In Venezuela the Glypto-dontidae from the Late Pleistocene that had tradi-tionally been assigned to Glyptodon (see BOCQUENTINVILLANUEVA 1982 AGUILERA 2006 RINCOacuteN et al2008) actually correspond to the genus GlyptotheriumOsborn (see CIONE et al 2007 CARLINI et al 2008a)

Glyptodon is also one of the most frequent taxarecorded on the ldquoAtlanticrdquo eastern slope in southernBrazil but in this area it is associated with otherGlyptodontidae particularly Panochthus as far as 5ordm S(PAULA COUTO 1979 PORPINO amp BERGQVIST 2002PORPINO et al 2004 DANTAS amp ZUCON 2005 DASILVA et al 2006) The presence of the Equidae Equus(A) neogeus LUND associated with Glyptodon at theselatitudes allows the existence of open savanna-typeenvironments to be inferred during the Late Pleisto-cene of that region (see ALBERDI et al 2003) Thegenus has also been recorded in Bahia (Brazil)synchronically associated with typical ldquopampeanrdquo

forms (eg Toxodon platensis OWEN Morenelaphussp) as well as intertropical taxa (eg Trigonodopslopesi Xenorhinotheriun bahiense) (CARTELLE amp DEIULIIS 1995) An association of Panochthus ndash Glypto-don has also been observed in the south-westernAmazon region in sediments corresponding to theLast Glacial Maximum (RANCY 1992 LATRUBESSE ampFRANZINELLI 1995 RANZI 2000) The remarkableabsence of Cingulata Glyptodontidae in the area of theAmazon Basin could be related to the existence of acold-arid to warm-humid climatic and environmentalgradient with a north-western orientation (MARSHALLet al 1984CARTELLE amp LESSA1988 OLIVEIRA 1996)Nevertheless the evidence suggests that during thelatest Pleistocene the dominant habitats in Brazilwere characterized by open or semi-forested environ-ments (MARSHALL et al 1984 PASCUAL amp ORTIZ-JAU-REGUIZAR 1990 PENNINGTON et al 2000) althoughfluvial systems were present that could have acted asbiogeographical barriers Southwards (Rio Grande doSul State western Uruguay and Argentine Meso-potamia) this association also includes the genusNeuryurus AMEGHINO and in much lower proportionDoedicurus (CARLINI et al 2004 2008 NORIEGA etal 2004 UBILLA 2004 UBILLA et al 2004 RIBEIROet al 2007)Another interesting feature of the Tarija Valley

is the absence of genera such as Neosclerocalyptus(see ZURITA 2007) and the evident scarcity of otherssuch as Panochthus Remarkably Neosclerocalyptuswhich is one of the most common forms in the Pam-pean region and north-central Argentina (ZURITA etal 2005 ZURITA 2007) is recorded again furthernorthwards at the locality of Santa Cruz de la Sierra(17ordm 47rsquo S and 63ordm 11rsquoW) and at much lesser altitude(ca 439 m) this material deposited in MNK-PALalso represents the northernmost record of thisgenus

Panochthus is very rare in the Tarija Valley thisgenus is represented only by one isolated osteodermfrom the mid-central region of the dorsal carapace andfragments of a caudal armor (MNPA-V 006598) (Fig2F) A new analysis of these specimens indicates thatthe original assignation made by HOFFSTETTER (1964131-132) as Panochthus cf P tuberculatus has nomorphological support and that it can only besustained at the genus level (Fig 3) Particularly thereticular ornamentation pattern of the dorsal carapaceosteoderm and fragments of caudal armor is acharacter useful for generic identification only (seeCASTELLANOS 1941 PORPINO amp BERGQVIST 2002


among others) Future findings of more completematerials may allow a more precise determination

Hoplophorus echazui HOFFSTETTER is a speciesthat thus far has only been recorded in this area (HOFF-STETTER 1964) it is known only from the holotype(MNPA-V 142 distal portion of a caudal armour)several dorsal carapace fragments and one caudal ringosteoderm (MNPA-V141) (Fig 2G) Among othersthe diagnostic characters provided by its author thatdistinguish this taxon from the other recognizedspecies (H euphractus) an intertropical form fromthe late Pleistocene of Minas Gerais Brazil (LUND1839 PAULA COUTO 1957 CARTELLE amp DE IULIIS1995 RANZI 2000) include the following a) lateralmargins of distal end of caudal armor more parallel toeach other than in H euphractus b) sheath tip morerounded with fewer peripheral figures on its dorsalsurface and c) central figures that tend to be moreclearly circular than in H euphractus It is worthnoting that these morphological characters show highintraspecific variability in the Hoplophorinae Hoplo-phorini (ZURITA 2007) In addition the ornamentationof the dorsal carapace osteoderms does not showany diagnostic features and these plates are virtuallyidentical to those of H euphractus as recognized bythe author himself (HOFFSTETTER 1963 128) Eachosteoderm is characterized by a central figure en-circled by a row of peripheral figures creating therosette-type pattern typical of the Hoplophorinae

Hoplophorini with parabolic transversal section ofboth main and radial sulci (ZURITA 2007) To sum upthe extremely fragmentary nature of this materialtogether with the lack of clear diagnostic characterssuggest that the validity of this species should beprovisional pending the discovery of more completematerials HOFFSTETTER himself (1964 131) acknow-ledged this limitation when he stated ldquoIl conviendracependant de reacuteviser cette interpreacutetation provisoirelorsquacute on connaicirctra plusieurs tubes caudaux dechacune de ces formes et quacuteon sera en mesure depreacuteciser leurs variations respectivesrdquo Consequentlyalthough this material undoubtedly corresponds to thegenus Hoplophorus (Fig 3) the validity of the speciesH echazui is uncertainLastly TAKAI et al (1984 pl 20 figs 50-52) as

part of a paleofaunal survey have reported and illust-rated H euphractus in the Pleistocene of Tarija repre-sented by approximately six isolated dorsal carapaceosteoderms As in the case of H echazui these osteo-derms show only the typical rosette-type ornamenta-tion The absence of diagnostic features combinedwith the scarcity and poor preservation of thematerial preclude any specific assignation As in theprevious case referral to the genus Hoplophorus issupported by the rugosity of the exposed osteodermsurface greater than that of Neosclerocalyptus (PAULACOUTO 1957 ZURITA 2007)


Fig 3 Chronological distribution in the Pampean region of the Cingulata Glyptodontidae (except Hoplophorus) present inthe Tarija Valley 1 ndash Glyptodon sp 2 ndash Glyptodon munizi 3 ndash Panochthus 4 ndash Hoplophorus 5 ndash Neothoracophorus



Another Glyptodontidae recognized for the TarijaValley is Neothoracophorus AMEGHINO The occur-rence of this taxon has been mentioned by severalauthors including HOFFSTETTER (1963 1964)MARSHALL et al (1984) and COLTORTI et al (2007)although none of these workers have justified thisassignation or illustrated the materials in questionThe dorsal carapace of this genus is formed by smallbut extremely thick osteoderms with practically novisible lateral sutures Each osteoderm comprises arelatively large and elevated central figure surroundedby a series of large foramina (AMEGHINO 1889CASTELLANOS 1951) Currently this Pleistocenegenus includes one species N elevatus (NODOT) (Fig3) which is only known from isolated dorsal carapaceosteoderms The holotype of this species (MNHNPBRD 20 see MONES 1994) consists of a fragmentarydorsal carapace in relatively poor preservation statethe latter circumstance according to AMEGHINO(1889 791) precluded detailed study of the specimenHowever the characters provided originally by NODOT(1857) and later by AMEGHINO (1889) imply that thiscould actually be a juvenile specimen (eg small-sized osteoderms large foramina and lax sutures) Inaddition its stratigraphic provenance is not precisealthough it has been tentatively assigned to the Plei-stocene (MONES 1986)As a consequence of these fac-tors this is certainly the least known and worst charac-terized of the Pleistocene Glyptodontidae (PAULACOUTO 1979) For the Tarija Valley only COLTORTI etal (2007 7 fig 3a) have illustrated the material con-sisting of 9 osteoderms from the lateral region of adorsal carapace However this specimen cannot bereferred to Neothoracophorus because the exposedsurface of the osteoderms is flat and smooth and theyare larger with no evident foramina This combinationof characters refutes the assignation made by theseauthors In this context the remarkable morphologicalsimilarity between the dorsal carapace osteoderms ofjuvenile Glyptodon specimens (see LYDEKKER 1894RINDERKNECHT 2000) and those that have been refer-red by different authors (eg CASTELLANOS 1951) toNeothoracophorus together with the evident scarcityof records suggest that this latter genus is probably asynonym of Glyptodon something that had alreadybeen noted by certain authors (see CASTELLANOS1951 74-75) Furthermore the type species wasoriginally assigned to genus Glyptodon (G elevatusNODOT) and later transferred to Thoracophorus byGERVAIS amp AMEGHINO (1880) Finally AMEGHINO(1889) replaced the genus name (since it was pre-

occupied) and erected Neothoracophorus (seeCASTELLANOS 1951)The examination of the materials deposited in the

Museo Nacional Paleontoloacutegico ndash Arqueoloacutegico deTarija (Bolivia) that have been referred to Neothoraco-phorus does not reveal significant differences whencompared to dorsal carapace osteoderms from juve-nile Glyptodon specimens (eg MCFA 760 MCA2017 LYDEKKER 1894 pl 2) (Fig 2C-D) In particu-lar the dorsal carapace osteoderms of these juvenilesare characterized by a) relatively small but very thickosteoderms mostly pentagonal or hexagonal b) pre-sence of a protruding central figure surrounded by arow of poorly defined peripheral figures c) evidentforamina at the intersection of the main and radialsulci d) poorly co-ossified sutures between osteo-derms (Fig 2D) In a word this morphology isnot much different from the diagnostic features ofNeothoracophorus Furthermore in the localityRugero (situated about 40 km south of Tarija) one ofthe richest fossiliferous sites of the Tarija Valley (seeTAKAI et al 1982 COLTORTI et al 2007) large frag-ments of Glyptodon carapace have been observed inclose spatial and stratigraphic association with smallerfragments with similar morphology to that of thematerials assigned to Neothoracophorus (Fig 2E)To sum up most of the evidence strongly suggests

that at least in Tarija Valley the records assigned toNeothoracophorus actually correspond to juvenileGlyptodon specimens Along these lines it cannot beruled out that Neothoracophorus is a synonym ofGlyptodon although testing this hypothesis wouldrequire a more complete revision

3 Discussion of results

As previously discussed the association of CingulataGlyptodontidae exhumed from the Tarija Valley(Bolivia) is substantially different from the assem-blages known for other regions of South America (egChacoan-Pampean region continued in the easternldquoAtlanticrdquo sector of south Brazil and subandeanareas) and permits the paleofaunal and paleobio-geographical characterization of this particular regionsituated between the Andean Altiplano the Amazonregion and the Chacoan-Pampean area (MOURGUIARTet al 1997) As stated by COLTORTI et al (2007) themammalian faunal assemblage exhumed from TarijaValley includes taxa characteristic of both flatlandsand higher altitude environments (HippocamelusLEUCKART Cuvieronius hyodon FISCHER)


In this context the almost exclusively dominantglyptodonts in Tarija Valley belong to genus Glypto-don eastwards and northwards and in parallel tosubandean areas this taxon is practically the onlyglyptodont occurring up to 4deg N (BOMBIN 1981MARSHALL amp SALINAS 1991 PUJOS amp SALAS 2004CARLINI et al 2008a)In contrast some genera that are very frequent

in the Chaco-Pampean plains such as Neosclero-calyptus are absent fromTarijarsquos fossil record Otherssuch as Panochthus which have a good record in thelateral eastern portion of South America between 5deg Sand 22deg S are quite poorly represented in the TarijaValley In addition the validity of H echazui andthe presence of an intertropical taxon such as Heuphractus (see RANZI 2000 CARTELLE amp DE IULIIS1995) in Tarija Valley are unconfirmed until new andmore complete specimens are found The currentavailable evidence only allows confirmation of thepresence of the genus HoplophorusApart from this the presence and validity of the

poorly known genus Neothoracophorus is highlyuncertain This is due on one hand to the fact thatthe published records were never illustrated or theirassignations justified and on the other hand theanalysis of those materials deposited in the collectionsof MNPA-V strongly suggests that these correspond tojuvenile Glyptodon specimensFrom a biostratigraphical perspective the Glypto-

dontidae (Glyptodon) are important for the resolutionof the problematic chrono-stratigraphy of this area Onseveral occasions MACFADDEN and collaboratorshave supported an age of 11 to 07 Ma for the entiresequence (see MACFADDEN amp SHOCKEY 1997MACFADDEN et al 1983 MACFADDEN 2000) How-ever a possible Lujanian age had already been sug-gested by some authors (eg CASAMIQUELA 1969MARSHALL et al 1984 HOFFSTETTER 1986) for atleast some areas of the Tarija Valley More recentlyCOLTORTI et al (2007) adopting a completely dif-ferent perspective have proposed a much younger ageof between 44 and 21 ka Likewise TONNI et al (inpress) remark that the paleofaunal association ofTarija Valley comprises taxa whose biochrons in thePampean region fall within the Ensenadan-Lujanianinterval (Early-Middle Pliocene ndash Eearly Holocene)(see CIONE amp TONNI 1995 1999 2005) and that atpresent it is not possible to assume that the entiresequence necessarily corresponds to only one of thoseintervals

It is also important to remark that although thespecies of Glyptodon are in need of urgent revisionthe morphology of all the exhumed specimens but oneis similar to that of the specimens known from theMiddle Pleistocene (Bonaerian) and Late Pleistocenendash Early Holocene (Lujanian) of the Pampean regionHowever the exceptional specimen deposited in thecollection MNPA-V from the localityArmados corre-sponds to the species G munizi whose stratigraphicdistribution in the Pampean region is limitedto the Ensenadan (AMEGHINO 1881 SOIBELZON et al2006) This specimen in addition to being the firstrecord of the taxon outside the Pampean region wouldbe the only Ensenadan taxon with precise geo-graphical provenance

4 Conclusions

From a taxonomic perspective the Glyptodontidaepresent in the Tarija Valley (Bolivia) are representedwith certainty by the genera Glyptodon Panochthusand HoplophorusThe presence of the speciesH echazui H euphrac-

tus and P tuberculatus is conditional pending dis-covery of more complete materialsThe records of Neothoracophorus correspond with

high probability to juvenile Glyptodon specimensFrom a taxonomical-stratigraphic viewpoint the

great majority of Glyptodon specimens are morpho-logically similar to those exhumed from the MiddlePleistocene ndash Eearly Holocene of the Pampean regionin Argentina (Bonaerian-Lujanian stages)Nevertheless one of the specimens (from the

locality Armados) is assignable to the species Gmunizi whose stratigraphic distribution is restricted tothe Ensenadan Stage (Early ndash Middle Pleistocene) inthe Pampean region of Argentina and this is the firstrecord of this taxon outside this regionThe assemblage of Cingulata Glyptodontidae from

the Tarija Valley differs from those known for theChacoan-Pampean region southern Brazil andwestern Uruguay In this context the remarkablefrequency of Glyptodon records and the scarcity ofPanochthus as well as the absence of Neosclero-calyptus one of the commonest taxa in the chacoan-pampean plains are noteworthy

Acknowledgements

The authors thank the staff at Museo Nacional Paleonto-loacutegico ndash Arqueoloacutegico Vertebrados (Tarija Bolivia) forallowing study of the materials presented here Dr D CROFT




and an anonymous reviewer are also thanked for theirthorough reviews and helpful suggestions This work wasfunded by project grant PICTO-UNNE (2007-00164) andPI (UNNE-06805)

References

AGUILERA O (2006) Tesoros paleontoloacutegicos de Vene-zuela El Cuaternario del Nordeste del estado Falcoacuten ndashEditorial Arte Caracas 120 pp Caracas (EditorialArte)

ALBERDI M T CARTELLE C amp PRADO J L (2003) Elregistro pleistoceno de Equus (Amerhippus) e Hippidion(Mammalia Perisodactyla) de Brasil Consideracionespaleoecoloacutegicas y biogeograacuteficas ndash Ameghiniana 40(2) 173-196

AMEGHINO F (1881) La antiguumledad del hombre en elPlata Vol 2 ndash 557 pp Paris amp Buenos Aires (Massonamp Igon Hermanos)

ndash (1883) Sobre la necesidad de borrar el geacutenero Schisto-pleurum y sobre la clasificacioacuten y sinonimia de losGliptodontes en general ndash Boletiacuten de la AcademiaNacional de Ciencias Coacuterdoba 5 1-34

ndash (1889) Contribucioacuten al conocimiento de los mamiacuteferosfoacutesiles de la Repuacuteblica Argentina ndash Academia Nacionalde Ciencias de la Repuacuteblica Argentina (Coacuterdoba)Buenos Aires 6 1-1027

ndash (1902) Notas sobre algunos mamiacuteferos foacutesiles nuevos opoco conocidos del Valle de Tarija ndash Anales MuseoNacional Buenos Aires 8 225-261

BAacuteEZ-PRESSER J L BUONGERMINI E AMAacuteBILE V O FCROSSA V F BAacuteEZ-ALMADA A B ZARZA-LIMA P Ramp MIGONE O O (2004) Algunos antecedentes paleon-toloacutegicos del Paraguay ndash Boletiacuten del Museo de HistoriaNatural del Paraguay 15 (1-2) 95-110

BENJAMIN M T JOHNSON N M amp NAESER CW (1987)Recent rapid uplift in the Bolivian Andes evidencesfrom fission-track dating ndash Geology 15 680-683

BOCQUENTIN-VILLANUEVA J (1982) Notas sobre la faunadel Pleistoceno superior de Taima-Taima depositada enel Museo del Hombre de Coro Estado Falcoacuten Vene-zuela ndash Acta Cientiacutefica Venezolana 33 479-487

BOMBIN M (1981) Ocurrencia de Glyptodon clavipes onColombia ndash Revista CIAF Bogotaacute 6 (1-3) 17-18

BOULE M ampTHEVENIN A (1920) Mammifegraveres fossiles deTarija Vol 4 ndash 256 pp Paris (Imprimerie National)

BURMEISTER H (1866) Einige Bemerkungen uumlber die imMuseum zu Buenos Aires befindlichen Glyptodonartenndash Zeitschrift fuumlr die gesammten Naturwissenschaften28 138-142

ndash (1870-1874) Monografiacutea de los Glyptodontes en elMuseo Puacuteblico de Buenos Aires ndash Anales del MuseoPuacuteblico de Buenos Aires 2 367-377

CARLINI A A amp SCILLATO-YANEacute G J (1999) Evolutionof Quaternary Xenarthrans (Mammalia) of Argentina ndashIn RABASSA J amp SALEMME M (Eds) Quaternary ofSouth America and Antarctic Peninsula 12 149-175Rotterdam (AA Balkema)

CARLINI A A amp TONNI E P (2000) Mamiacuteferos Foacutesilesdel Paraguay ndash 108 pp Buenos Aires (CooperacioacutenTeacutecnica Paraguayo-Alemana Proyecto SistemaAmbien-tal del Chaco-Proyecto Sistema Ambiental RegioacutenOriental)

CARLINI A A amp ZURITA A E (2007) Evolucioacuten y paleo-biogeografiacutea de los Glyptodontidae Glyptodontinae(Mammalia Xenarthra) una nueva interpretacioacuten 3degCongreso de Mastozoologiacutea en Bolivia Resuacutemenes 24Santa Cruz de la Sierra Bolivia

CARLINI A A ZURITA A E amp AGUILERA O (2008a)NorthAmerican Glyptodontines (Xenarthra Mammalia)in the upper Pleistocene of northern South AmericaPalaeontologische Zeitschrift 82 (2) 125-138

CARLINIAAZURITA AE GASPARINI GM amp NORIEGAJ I (2004) Los mamiacuteferos del Pleistoceno de la Meso-potamia argentina y su relacioacuten tanto con aquellos delCentro-Norte de la Argentina Paraguay sur de Boliviacomo con los del sur de Brasil y oeste de Uruguaypaleobiogeografiacutea y paleoambientes ndash INSUGEOMiscelaacuteneas 12 5-12

CARLINI A A ZURITA A E amp MINO BOILINI A R (Inpress) Resentildea paleobiogeograacutefica de los Xenarthra(Mammalia) del Pleistoceno tardiacuteo de la regioacuten Meso-potaacutemica (Argentina) ndash INSUGEO Miscelaacuteneas 17259-270

CARLINI A A ZURITA A E SCILLATO-YANEacute G JSANCHEZ R amp AGUILERA O (2008b) A new Glypto-dont species from Codore Formation (Pliocene) EstadoFalcoacuten (Venezuela) and the lsquoAsterostemmarsquo problem ndashPalaumlontologische Zeitschrift 82 (2) 139-152

CARTELLE C amp DE IULIIS G (1995) Eremotherium lauril-lardi The Panamerican Late Pleistocene MegatheriidSloth ndash Journal of Vertebrate Paleontology 15 (4) 830-841

CARTELLE C amp LESSA G (1988) Presenccedila de Myocatorcoypus (Molina 1782) Rodentia Myocatoridae doPleistoceno final-Holoceno no centro-oeste da BahiaBrasil 11ordm Congresso Brasilero de Paleontologiacutea Actas1 583-591

CASAMIQUELA R (1969) Enumeracioacuten criacutetica de algunosvertebrados foacutesiles continentales pleistoceacutenicos deChile ndash Rehue 2 143-172

CASTELLANOS A (1941) A propoacutesito de los geacutenerosPlohophorus Nopachthus y Panochthus ndash Publicacio-nes del Instituto de Fisiografiacutea y Geologiacutea 11 417-583

ndash (1951) Acotaciones al geacutenero NeothocoraphorusAmeghino ndash Revista de la Asociacioacuten Geoloacutegica Argen-tina 6 (1) 63-82

CIONE A L amp TONNI E P (1995) Bioestratigrafiacutea ycronologiacutea del Cenozoico de la regioacuten Pampeana ndash InALBERDI M T LEONE G amp TONNI E P (Eds)Evolucioacuten bioloacutegica y climaacutetica de la regioacuten Pampeanadurante los uacuteltimos cinco millones de antildeos Un ensayode correlacioacuten con el Mediterraacuteneo occidental 1247-74 Madrid

ndash (1999) Biostratigraphy and chronological scale ofupper-most Cenozoic in the Pampean Area Argentina ndashIn RABASSA J amp SALEMME M (Eds) Quaternary ofSouth America and Antarctic Peninsula 12 23-51Rotterdam (Balkema)


CIONE A L amp TONNI E P (2005) Bioestratigrafiacutea basadaen mamiacuteferos del Cenozoico superior de la provincia deBuenos Aires Argentina ndash In DE BARRIO R EETCHEVERRY R O CABALLEacute M F amp LLAMBIacuteAS E(Eds) Relatorio del 16ordm Congreso GeoloacutegicoArgentino183-200 La Plata

CIONE A L TONNI E P BARGO S BOND M CANDELAA M CARLINI A A DESCHAMPS C M DOZO M TESTEBAN G GOIN F J MONTALVO C I NASIF NNORIEGA J I ORTIZ JAUREGUIZAR E PASCUAL RPRADO J L REGUERO M A SCILLATO-YANEacute G JSOIBELZAN L VERZI D H VIEYTES C E VIZCAINOS F amp VUCETICH M G (2007) Mamiacuteferos continen-tales del Mioceno tardiacuteo a la actualidad en la Argentinacincuenta antildeos de estudios ndash Asociacioacuten PaleontoloacutegicaArgentina Publicacioacuten Especial 11 257-278

COLTORTI M ABBAZZI L FERRETI M LACUMIC PPAREDES RIacuteOS F PELLEGRINI M PIERUCCINI PRUSTIONI M TITO G amp ROOK L (2007) Last GlacialMammals in South America a new scenario from theTarija Basin (Bolivia) ndash Naturwissenschaften 94 288-299

DANTAS M A T amp ZUCON M H (2005) Sobre a ocor-recircncia de dois taxa Pleistocecircnicos na Fazenda TytoyaPoccedilo Redondo Sergipe ndash Scientia Plena 1 (4) 92-97

DA SILA F M DA SILVA ALVES R FRANCA BERRETO AM BEZERRA DE SAacute F amp BORGES LINS E SILVA A C(2006) A megafauna Pleistocecircnica do Estado dePernambuco ndash Estudos Geoloacutegicos 16 (2) 55-66

DANTAS M A T amp ZUCON M H (2005) Sobre a ocor-recircncia de dois taxa Pleistocecircnicos na Fazenda TytoyaPoccedilo Redondo Sergipe Scientia Plena 1 (4) 92-97

DE CARLES E (1888) Noticias sobre un viaje a Tarija(Bolivia) ndash Boletiacuten del Instituto Geograacutefico Argentino9 (1) 35-40

DUARTE R G (1997) Gliptodontes del Pleistoceno tardiacuteode Aguas de las Palomas Campo de Pucaraacute CatamarcaArgentina Variaciones morfoloacutegicas del caparazoacuten deGlyptodon reticulatus OWEN 1845 ndash Ameghiniana 34345-355

GERVAIS H ampAMEGHINO F (1880) Los mamiacuteferos foacutesilesde la Ameacuterica del Sur ndash 225 pp Paris amp Buenos Aires(Sabih amp Igon)

GILLETTE D D amp RAY C E (1981) Glyptodonts of NorthAmerica ndash Smithsonian Contributions to Palaeobiology40 1-251

GIRAUDO A (1997) El Concepto de especie Parte I ndashNatura Neotropicalis 28 (2) 161-169

HOFFSTETTER R (1963) La faune Pleacuteistocegravene de Tarija(Bolivie) Nota preacuteliminaire ndash Bulletin du MuseacuteumdacuteHistoire Naturelle 35 (2)194-203

ndash (1964) Les Glyptodontes du Pleacuteistocene de Tarija(Bolivie) I Genres Hoplophorus et Panochthus ndashBulletin de la Socieacuteteacute Geacuteologique de France 7 (5) 126-133

ndash (1968) Ntildeuapua un gisement de verteacutebreacutes pleacuteistocegravenesdans le Chaco Bolivien ndash Bulletin du Museacuteum NationaldrsquoHistoire Naturelle 2deg Seacuterie 40 (4) 823-836

ndash (1970) Vertebrados cenozoicos y mamiacuteferos cretaacutecicosdel Peruacute 4ordm Congreso Latinoamericano de ZoologiacuteaActas 2 971-983 Caracas

HOFFSTETTER R (1978) Une faune de Mammifegraveres pleacutei-stocegravenes au Paraguay ndash Comptes Rendus Sommairesdes Seacuteances de la Socieacuteteacute Geacuteologique de France 132-33

KRAGLIEVICH L (1934) La antiguumledad Pliocena de lasfaunas de Monte Hermoso y Chapadmalal deducidas desu comparacioacuten con las que le precedieron y sucedieronndash 136 pp Montevideo (Imprenta El Siglo Ilustrado)

LATRUBESSE E M amp FRANZINELLII E (1995) Cambiosclimaacuteticos en Amazonia durante el Pleistoceno tardiacuteo-Holoceno ndash In ARGOLLO J amp MOURGUIART PH(Eds) Cambios Cuaternarios en Ameacuterica del SurORSTOM ndash Institut Franccedilais de Recherche Scientifiquepour le Deacuteveloppement en Coopeacuteration La Paz Boliviandash 1-344 pp

LYDEKKER R (1894) The extinct edentates of Argentina ndashAnales del Museo de La Plata 3 1-118

LUND P W (1839) Blik paa Brasiliens dyreverden foumlr sid-ste jordomvaeltning Anden afhandling Pattedyrene(Lagoa Santa d 16111837) ndash Det kongelige DanskeVidenskabernes Selskabs naturvidenskabelige og math-ematiske Afhandlinger 8 61-144

MACFADDEN B J (2000) Middle Pleistocene ClimateChange Recorded in Fossil Mammal Teeth from TarijaBolivia and Upper Limit of the Ensenadan Land-Mammal Age ndash Quaternary Research 54 (1) 121-131

MACFADDEN B J amp SHOCKEY B J (1997) Ancientfeeding ecology and niche differentiation of Pleistocenemammalian herbivores from Tarija Bolivia Morpho-logical and isotopic evidence ndash Paleobiology 23 (1) 77-100

MACFADDEN B J SILES O ZEITLER P JOHNSON N Mamp CAMPBELL Jr K E (1983) Magnetic polarity strati-graphy of the middle Pleistocene (Ensenadan) TarijaFormation of southern Bolivia ndash Quaternary Research19 (2) 172-187

MARSHALL L G BERTA A HOFFSTETTER R PASCUALR REIG O A BOMBIN M amp MONES A (1984)Mammals and stratigraphy geochronology of the con-tinental mammal-bearing quaternary of South Americandash Palaeovertebrata (Meacutemoire Extraordinaire) 1-76

MARSHALL L G amp SALINAS P Z (1991) The LorenzoSundt Collection of Pleistocene Mammals from UllomaBolivia in the Museum Nacional de Historia NaturalSantiago Chile ndash In SUAREZ-SORUCO R (Ed) Foacutesilesy Facies de Bolivia (I) Vertebrados 12 (3-4) 685-692

MARSHALL L G amp SEMPERE T (1991) The EocenePleistocene vertebrales of Bolivia and their stratigraphiccontext a review ndash In SUAREZ-SORUCO R (Ed)Foacutesiles y Facies de Bolivia (I) Vertebrados 12 (3-4)631-652

MAYR E (1996) What is species and what is not ndash Philo-sophy of Science 63 262-277

MONES A (1986) Palaeovertebrata Sudamericana Cataacute-logo sistemaacutetico de los vertebrados foacutesiles de Ameacutericadel Sur Parte I Lista preliminar y bibliografiacutea ndashCourier Forschungsinstitut Senckenberg 82 1-625

ndash (1994) Las vicisitudes del geacutenero Panochthus BUR-MEISTER 1866 (Mammalia Cingulata Glyptodontidae)ndash Comunicaciones Paleontoloacutegicas del Museo deHistoria Natural de Montevideo 2 (27) 79-86




MOURGUIART P ARGOLLO J MARTIacuteN L MONTENEGROM E SIFEDDINE A amp WIRRMAN D (1997) Change-ments limnologiques et climatologiques dans le basin dulac Titicaca (Bolivia) depuis 30000 ans ndash ComptesRendu de lrsquoAcadeacutemie de Science de la terre et desplanegravetes 325 139-146

NODOT L (1857) Description drsquo un nouveaux genredrsquoeacutedenteacute fossile renfermant plusieurs espegraveces voisinesdu Glyptodon etc ndash Meacutemoires de lrsquoAcadeacutemie Impeacuterialdes Sciences Arts et Belles-Lettres de Dijon (2) 5 1-170

NORIEGA J I CARLINII A A ampTONNI E P (2004) Verte-brados del Pleistoceno tardiacuteo de la cuenca del arroyoEnsenada (Departamento Diamante provincia de EntreRiacuteos Argentina) ndash INSUGEO Miscelaacuteneas 12 71-76

OLIVEIRA E V (1992) Mamiacuteferos foacutesseis do Quaternariodo Estado do Rio Grande do Sul Brasil ndash Dissertaccedilatildeode Mestrado (Universidade Federal do Riacuteo Grande doSul) 118 pp

ndash (1996) Mamiacuteferos Xenarthra (Edentata) do Quaternariodo Estado do Rio Grande do Sul Brasil ndash Ameghiniana33 65-75

OPPENHEIM V (1943) The fossiliferous basin of TarijaBolivia ndash Journal of Geology 51 (8) 548-555

OWEN R (1839) Description of a tooth and part of theskeleton of the Glyptodon a large quadruped of theedentate order to which belongs the tessellated bonyarmour figured by Mr CLIFT in his memoir on theremains of the Megatherium brought to England by SirWOODBINE PARISH FGS ndash Proceedings of the Geo-logical Society of London 3 108-113

ndash (1845) Descriptive and illustrated catalogue of the fossilorganic remains of Mammalia and Aves contained in theMuseum of the Royal College of Surgeons of LondonEngland ndash 391 pp

PASCUAL R amp ORTIZ JAUREGUIZAR E O (1990) Evolvingclimates and mammal faunas in Cenozoic SouthAmerica ndash Journal of Human Evolution 19 23-60

PATTERSON B amp PASCUAL R (1972) The fossil mammalfauna of South America ndash In KEAST A ERK F ampGLASS B (Eds) Evolution mammals and southerncontinents 247-309

PAULA COUTO J C (1957) Socircbre um gliptodonte do Brasilndash Boletim Divisatildeo de Geologia e Mineralogia 165 1-37

ndash (1979) Tratado de Paleomastozoologiacutea ndash 590 pp Riode Janeiro (Academia Brasileira de Ciecircncias)

PENNINGTON R T PRADO D E amp PENDRY C A (2000)Neotropical seasonally dry forest and Quaternary vege-tation changes ndash Journal of Biogeography 27 261-273

PORPINO K O amp BERGQVIST L P (2002) Novos achadosde Panochthus (Mammalia Cingulata Glyptodontoidea)no Nordeste do Brasil ndash Revista Brasileira de Paleonto-logia 4 51-62

PORPINO O K DOS SANTOS M F C F amp BERGQVIST L P(2004) Registros de mamiacuteferos foacutesseis no Lajedo deSoledade Apodi Rio Grande do Norte Brasil ndash RevistaBrasileira de Paleontologia 7 349-358

PUJOS F amp SALAS R (2004) A systematic reassessmentand paleogeographic review of fossil Xenarthra fromPeru ndash Bulletin de lrsquoInstitute Franccedilais drsquoeacutetudes Andine33 331-377

RANZY A (1992) Western Amazon Paleomammals andthe forest refugia model ndash Resumos e ContribuiccedilotildeesCientificas Simp Int Cuat DaAmaz UFAM ManausBrasil 45-48

ndash (2000) Paleoecologia da Amazocircnia Megafauna doPleistoceno ndash 101 pp Brasil (Editorial de la Uni-versidade Federal de Santa Catarina)

REGUERO MA CANDELA A M ampALONSO R N (2007)Biochronology and biostratigraphy of the Uquiacutea For-mation (Pliocene-early Pleistocene NW Argentina) andits significance in the Great American Biotic Inter-change ndash Journal of SouthAmerican Earth Sciences 231-16

RIBEIRO A N SCHERER C D amp PITANA V G (2007)Mamiacuteferos do Pleistoceno do Rio Grande do Sul BrasilEstado atual do conhecimento ndash Quaternaacuterio do RSIntegrando Conhecimentos resumos 25

RINCOacuteN A D WHITE R S amp MCDONALD H G (2008)Late Pleistocene Cingulates (Mammalia Xenarthra)from Mene de Inciarte Tar Pits Sierra de Perijaacute WesternVenezuela ndash Journal of Vertebrate Paleontology 28 (1)197-207

RINDERKNECHT A (2000) Estudios sobre la familia Glypt-dontidae Gray 1869 II Variacioacuten morfoloacutegica en lacoraza de Glyptodon sp juvenil (Mammalia Cingulata)ndash Revista de la Sociedad Uruguaya de Geologiacutea 3 (7)32-35

ROVERETO C (1914) Los estratos araucanos y sus foacutesiles ndashAnales del Museo Nacional de Historia Natural deBuenos Aires 25 1-247

SCILLATO-YANEacute G J CARLINI A A VIZCAIacuteNO S F ampORTIZ JAUREGUIZAR E (1995) Los Xenarthros ndash InALBERDI M T LEONE G amp TONNI E P (Eds) Evo-lucioacuten bioloacutegica y climaacutetica de la regioacuten Pampeanadurante los uacuteltimos cinco millones de antildeos Un ensayode correlacioacuten con el Mediterraacuteneo occidental 12 183-209 Madrid

SOIBELZON L H (2004) Revisioacuten sistemaacutetica de losTremactinae (Carnivora Ursidae) foacutesiles de Ameacutericadel Sur ndash Revista del Museo Argentino de CienciasNaturales 6 (3) 107-133

SOIBELZON L H TONNI E P amp BIDEGAIN J C (2008)Cronologiacutea magnetoestratigrafiacutea y caracterizacioacuten bio-estratigraacutefica del Ensenadense (Pleistoceno inferior-medio) en la ciudad de Buenos Aires ndash Revista de laAsociacioacuten Geoloacutegica Argentina 63 (3) 421-429

SOIBELZON L H TONNI E P amp BOND M (2005) Thefossil record of South American short-faced bears(Ursidae Tremarctinae) ndash Journal of South AmericanEarth Sciences 20 105-113

SOIBELZON E ZURITA A E amp CARLINI A A (2006)Glyptodon munizi AMEGHINO (Mammalia CingulataGlyptodontidae) redescripcioacuten y anatomiacutea ndash Ameghi-niana 43 377-384

SOIBELZON E ZURITA A E MINO BOILINI A R TONNIE P amp PAREDES RIacuteOS F (2007) Los mamiacuteferos pleisto-cenos del Valle de Tarija (Bolivia) ndash Reunioacuten Anual deComunicaciones de laAsociacioacuten PaleontoloacutegicaArgen-tina Corrientes Resuacutemenes 37

SUAacuteREZ MONTERO M (1996) Geologiacutea del Cuaternario dela cuenca pleistocena de Tarija ndash 12ordm Congreso Geo-loacutegico de Bolivia Memorias 455-463


SUAacuteREZ-SORUCO R amp DIAZ-MARTINEZ E (1996) Leacutexicoestratigraacutefico de Bolivia ndash 227 pp Cochabamba

TAKAI F AROacuteZQUETA B P MIZUNO T YOSHIDA A ampKONDO H (1984) On fossil mammals from the TarijaDepartament Southern Bolivia ndash The Research Instituteof Evolutionary Biology Tokyo 4 1-63

TAKAI F MIZUNO T IWASAKI K TANAKA K amp YOSHI-DA A (1982) Tarija mammal-bearing Formation inBolivia ndash The Research Institute of Evolutionary Bio-logy Tokyo 3 1-72

TINAUT A amp RUANO F (2000) Biodiversidad Clasifi-cacioacuten y Filogenia ndash In SOLER M (Ed) Evolucioacuten labase de la Biologiacutea Editorial Proyecto Sur 6 297-306Madrid

TONNI E P SOIBELZON E CIONE A L CARLINI A ASCILLATO-YANEacute G J ZURITA A E amp PAREDES RIOS F(in press) Mammals from the Pleistocene of the TarijaValley (Bolivia) Correlation with the Pampean chrono-logical standard ndash Quaternary International

UBILLA M (2004) Mammalian biostratigraphy of Pleisto-cene fluvial deposits in northern Uruguay SouthAmerica ndash Proceedings of the Geologistsrsquorsquo Association115 347-357

UBILLA M PEREA D AGUILAR C G amp LORENZO N(2004) Late Pleistocene vertebrate from northernUruguay tools for biostratigraphic climatic and en-vironmental reconstruction ndash Quaternary International114 129-142

WERDELIN L (1991) Pleistocene vertebrates from TarijaBolivia in the collections of the Swedish Museum ofNatural History ndash In SUAREZ-SORUCO R (Ed) Foacutesilesy Facies de Bolivia Vertebrados ndash Revista teacutecnica deYacimientos Petroliacuteferos Fiscales Bolivianos 1 (12)673-684

ZURITA A E (2007) Sistemaacutetica y evolucioacuten de los Hoplo-phorini (Xenarthra Glyptodontidae HoplophorinaeMioceno tardiacuteo-Holoceno temprano) Importancia bio-estratigraacutefica paleobiogeograacutefica y paleoambiental ndashTesis Doctoral (unpublished) Universidad Nacional deLa Plata 367 pp

ZURITA A E CARLINI A A SCILLATO-YANEacute G J ampTONNI E P (2004) Mamiacuteferos extintos del Cuaternariode la provincia del Chaco (Argentina) y su relacioacuten conaquellos del este de la regioacuten Pampeana y de Chile ndashRevista Geoloacutegica de Chile 31 (1) 65-89

ZURITA A E SCILLATO-YANEacute G J amp CARLINI A A(2005) Palaeozoogeographic biostratigraphic andsystematic aspects of the genus Sclerocalyptus AME-GHINO 1891 (Xenarthra Glyptodontidae) of Argentina ndashJournal of South American Earth Sciences 20 120-129

Manuscript received July 7th 2008Revised version accepted by the Stuttgart editor August21st 2008

Addresses of the authors

ALFREDO EDUARDO ZURITA AacuteNGEL RAMOacuteN MINO-BOILINICentro de Ecologiacutea Aplicada del Litoral (CECOAL-CONICET) y Universidad Nacional del Nordeste Ruta 5km 25 (3400) Corrientes Argentinae-mail azuritacecoalcomarangelmioboiliniyahoocomar

ESTEBAN SOIBELZON ALFREDO ARMANDO CARLINI Divi-sioacuten Paleontologiacutea de Vertebrados Museo de La PlataFacultad de Ciencias Naturales y Museo Paseo del Bosquesnordm 1900 La Plata Argentinae-mail esoibelzonfcnymunlpeduaracarlinifcnymunlpeduar

FREDDY PAREDES RIacuteOS Museo Nacional Paleontoloacutegico-Arqueoloacutegico UniversidadAutoacutenoma Juan Misael Sarachocalle General Trigo 402 casilla 51 Tarija Boliviae-mail freddypar68hotmailcom




logical schemes always in comparison to the schemefor the Pampean region of Argentina (eg CIONE ampTONNI 1995 1999 COLTORTI et al 2007 CIONE et al2007)Thus from a historical perspective the sedi-

mentary sequences that crop out in Tarija Valley havebeen successively assigned to different ages andstages ranging from the Lower to the Upper Pleisto-cene (DE CARLES 1888 AMEGHINO 1902 ROVERETO1914 BOULE amp THEacuteVENIN 1920 KRAGLIEVICH 1934OPPENHEIM 1943 PATTERSON amp PASCUAL 1972TAKAI et al 1982 1984 MACFADDEN et al 1983MACFADDEN amp SHOCKEY 1997 MACFADDEN 2000COLTORTI et al 2007 MARSHALL et al 1984 33)More recently and on the basis of a preliminaryrevision SOIBELZON et al (2007) and TONNI et al (inpress) have indicated that from a biostratigraphicviewpoint Tarija includes taxa characteristic of theEnsenadan Bonaerian and Lujanian interval (Early-Middle Pliocene ndash Early Holocene) included in thePampean chronological standard (see CIONE amp TONNI2005)From a geological and stratigraphical perspective

the Tarija Valley is part of a Quaternary sedimentationbasin filled with fluvio-lacustrine sediments thatdiscordantly overlie a Paleozoic basement (SUAacuteREZ-

MONTERO 1996) The most outstanding feature of thearea is the characteristic badlands landscape withirregular relief produced by differential sedimenterosion (OPPENHEIN 1943 SUAacuteREZ-MONTERO 1996)In this context COLTORTI et al (2007) included

the entire Pleistocene sequence within the TolomosaFormation subdivided into two major Units AncoacutenGrande Unit (AG) and San Jacinto Unit (SJ) Thetransition from AG Unit to SJ Unit is evident alongmost of the badlands The upper part (SJ) is yel-lowish-gray and fine-grained The lower part (AG)is reddish and coarser-grained (see more details inTONNI et al in press)The fossil mammal fauna exhumed from the Tarija

Valley comprises a wide taxonomical diversity (ca 55species HOFFSTETTER 1963 MACFADDEN 2000)including taxa characteristic for both forested or sub-forested relatively warm and humid environments(eg Tapirus BRUumlNICH Myocastor KERR NeochoerusHAY) as well as others adapted to open arid orsemiarid and cold areas where grasslands are domi-nant (Equus LINNAEUS Camelidae Glyptodontidae)(MACFADDEN amp SHOCKEY 1997) Some palynologicalstudies (TAKAI et al 1982) suggest that during most ofthe Pleistocene the Valley was dominated by an opensemiarid environment occupied by grasslands with

Fig 1 Location map showing the Tarija Valley (Bolivia)





















































4 Conclusions







Acknowledgements






References

































































































































































4 Conclusions







Acknowledgements






References
















































































































References














































































































the diversity of glyptodontidae (xenarthra, cingulata) in the tarija valley (bolivia): systematic,...

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