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Home , Achocalla, Altiplano Basin, Anzaldo, Ayo Ayo, Branisella, Callapa

THE TO VERTEBRATES OF AND THEIR STRATIGRAPHIC CONTEXT A REVIEW

LARRY G. MARSHALL" & THIERRY SEMPERE** * Institute of Human Origins, 2453 Ridge Road, Berkeley, California 94709, U.S.A. ** Orstom, UR lH, Casilla 4875, , Bolivia. Present address: Centre de Géologie Générale et Minibre, Ecole des Mines, 35 rue Saint Honor& 77305 Fontainebleau, France

INTRODUCTION the type fauna of the Friasiali Land Age (conventionally middle ) in southern is temporally equivalent to the The record of fossil vertebrates in Bolivia is extremely Land Mammal Age. They thus use for the good. Compared with other countries in , Bolivia is land mammal age between Santacrucian and . For all second only to in the number of known localities and in practical purposes, of previous workers is equivalent to the wealth of taxa. Colloncuran as used in this study. Of the different vertebrate groups, the are by far the This paper represents an expansion and updating of the Bolivian most abundant and best known. In fact, the record of mammal land mammal record as provided by Robert Hoffstetter (in Marshall evolution in South America is so complete that these fossils are used el al. 1983, 1984). As documented below, the highlights of this by geologists and paleontologists to subdivide geologic time. The record include: the taxonomically richest and best studied faunas of occurrence of unique associations of taxa that are inferred to have late -early Miocene () and early Pleistocene existed during a restricted interval of time has resulted in the () age in all o[ South America; and the exceptionally rich recognition of discrete chronostratigraphic units called Land record of late Miocene () and early to middle Mammal Ages. These ages were established on the basis of (Montehemosan) age faunas from the northern . knowledge of of evolution of the taxa, on their time of first The Altiplano, including the Puna in adjacent Argentina and Chile, and/or last appearahce in the fossil record, and on changing faunal is a high-altitude plateau which covers southwest Bolivia, southeast associations through time. The sequence of South American Land , northeast Chile and northwest Argentina. It is about 200 km Mammal Ages (SALMA) has been based largely on knowledge of wide and about 1500 km long, and some 300,000 km2 at 3,600- the exceptionally rich and complete record in Argentina (Marshall, 4,000 m (mean 3650 m; Isacks, 1988) of altitude. In Bolivia, the 1985). Detailed systematic studies of taxa along with geochronologic Altiplano covers about 170,000 km2 and is situated between the studies and magnetostratigraphy of the fossiliferous rock sequence Cordillera Occidental and Cordillera Oriental. and/or radioisotopic (Ar/Ar, K-Ar) dating of associated volcanic The following abbreviations are used: GEOBOL, Servicio rock units (i.e. basalts, tuffs, ignimbrites) permit geologists and Geológico de Bolivia; ka, thousands of ago, a point in time; kg, paleontologists to correlate Bolivian mammal faunas with the kilograms; km, kilometers; m, meters; Ma, megaannum or millions Argentine sequence atid to make distinctions between ecological and of years ago, a point in time; My, millions of years, a duration of temporal relationships of local faunas and faunal assemblages. The time; yrbp, years before present. result of such studies is that land mammals have proven very useful for dating and correlating rock units within Bolivia in particular and with other South American countries in general. GEOLOGICAL EVOLUTION OF BOLIVIA: EOCENE TO I We provide a review of the Eocene to Pleistocene land mammal PLEISTOCENE 1 record of Bolivia within its stratigraphic and tectonic context. The

I use and chronology of South American Land Mammal Ages follows The reader must first be aware that the geologic evolution of the 'I Marshall (1985) as supplemented by Marshall, Drake e¿ al. (1986), central is currently being re-evaluated, and that traditional and Marshall, Cifelli ef al. (1986). Recently, Tonni et al. (1987) models are being challenged. The latter envision that the central

I recognized a new ,Pleistocene Land.Mamma1 Age between the Andes were subjected during most of Cenozoic time to a tectonic Ensenadan and which they called Centinelan. However, the regime characterized by long tensional periods (during which the validity and utility of this new age has yet to be fmlyestablished in sedimentary basins formed), short synchronous compressional Argentina, and unti this is done it is futile to attempt to recognize it pulses, and generally high-angle faulting (Mégard, 1978; Dalmayrac 1 in Bolivia. In addition, Marshall & Salinas (1990) have shown that el al., 1980; Martinez, 1980; Lavenu & Marocco, 1984; Mégard et

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l Y l LARRY G. MARSHALL & THIERRY SEMPERE al. 1984; Lavenu, 1986; Sébrier et al., 1988). In contrast, new data After a time of relative quiescence (- 19-17 Ma), magmatism and new interpretations favor structural models dominated by slowly resumed at - 17 Ma (Grant et al., 1979; Redwood & progressive crustal shortening, thrust propagation, and MacIntyre, 1989) and tectonism at - 14 Ma (Sempere, 1990). and evolution (Jordan et al., 1983; Jordan & Alonso, 1987; Isacks, 1988; considerably increased at 11-10 Mawith the onset of the second al., al., Roeder, 1988; Sempere et 1988, 1989, 1990a, b; Baby et major tectonic crisis (Sempere et al., 1989). This crisis lasted until - 1990; Sheffels, 1990). 5 Ma (Sempere et al., 199Oa), but tectonic and magmatic activities In Bolivia, the two models of Andean are sometimes at have lingered on to the present. There is apparently little relation odds with each other. For instance, the thick Cenozoic clastic strata between the onset of this second tectonic crisis and plate of the Altiplano were traditionally thought to have been deposited convergence history. during long epochs of extension with short interruptions by Each tectonic crisis was followed by the development of an compressive pulses. It is now believed that their deposition took extensive planation surface, respectively the Chayanta surface in the place mostly in intermontane foreland basins (i.e. in permanently middle Miocene and the San Juan del Oro surface during the late al., al., compressional settings; Baby et 1990; Sempere et 1990a, b, Miocene-early Pleistocene interval (Servant et al., 1989; Sempere et 1991). Predominance of compressive conditions in the central al., 199Od). Andean area since late time was first suggested in a general way by Marocco ef al. (1987). STRATIGRAPHIC FRAMEWORK The angular unconformities observed on the Altiplano and elsewhere in the Bolivian Andes have traditionally been interpreted The Cenozoic stratigraphy of Bolivia strongly relies on knowledge as materializing the mentioned "compressional pulses". However, of the basins where the strata were deposited. As already mentioned, this is not the only possible interpretation for these angular the Eocene-early Oligocene area of sedimentation was a large unconformities (Sempere, 1991). Moreover, although details of some foreland basin, and its paleo-landscape was a wide alluvial plain that important field observations relative to deformations in the Altiplano probably resembled the present-day Chaco-Beni lowlands. In have been well described, in particular in the works of Martinez contrast, the late Oligocene-Pleistocene period was characterized by (1980), Lavenu (1986) and Lavenu et al. (1989), several independent numerous separate intermontane basins, many of small size, that new chronologidal data make untenable the traditional timing of developed in deformed areas, and by the large Subandean-Llanura deformation (Sempere et al., 1990b). external fxeland basin of the Chaco-Beni lowlands. Therefore, the Calidity of the six "compressional tectonic pulses" The lower boundary of the Eocene-early Oligocene period or "phases", as listed by Sébrier et al. (1988), may be questioned. coincides with the upper boundary of the Kimmeridgian- Intuitively, it seems unlikely that the enormous central Andes, which Puca Group (Sempere el al., 1988; Jaillard & Sempere, 1989; today are second only to the Himalayas in size and height of the Sempere, 1990). This locally angular unconformity (Marocco et al., mountain belt, crustal thickness, etc., developed by a small number 1987) marks the initiation of the foreland basin which developed, in of short "compressional pulses". what is today the Bolivian Altiplano and.Cordillera Oriental, during In this paper we adopt the tectonic model and stratigraphy Eocene-early Oligocene time (see above), and very likely resulted currently being worked-out by the Orstom-YPFB research program from a deep modification of the geodynamics of the Andean margin on the geology of Bolivia (Sempere et al., 1989, 1990a, b, c; at that time. Although approximately located at the Paleocene- Sempere, 1990), which we summarize below. A map showing the Eocene boundary (Cirbián et al., 1986; i.e.- 53 Ma according to tectonostratigraphic domains presently recognized in Bolivia Cowie & Bassett, 1989, and Odin, 1989; Fig. 2), this tectonic (Sempere ef al., 1988, 1990a). and the location of the main change might be related to the plate rearrangement which occurred sedimentary basins of late Oligocene-Pleistocene age are shown in in the Pacific area at anomaly 24N (Lonsdale, 1988), i.e. in the late al., Figure 1. early Ypresian (ßerggren et 1985; Cavelier & Pomerol, 1986; af., The geological history of Bolivia since the Eocene consists of two Haqef 1987) at - 5 1-50 Ma (according to updated information in tectosediinentary periods, respectively spanning the Eocene-early Cowie & Bassett, 1989, and Odin, 1989). Such an age for this major Oligocene and late Oligocene-Present intervals (Sempere et al., change in Andean geodynamic conditions nearly coincides with the 1989, 1990a; Sempere, 1990). 50-49 Ma age for termination of emplacement of the coastal During the Eocene-early Oligocene time span, the paleo-Andes batholith of Peru (Soler, 1987), which also reflects a niajor change in were located west of the Bolivian territory, and their associated modalities of . continental external foreland bash, resembling the present-day The boundary between these two tectosedimentary periods is 27 Chaco plain, occupied what is today tlie Altiplano and most of tlie Ma (Sempere ef al., 1990b) which coincides with the early Cordillera Oriental. At - 27 Ma, a major tectonic crisis began to Oligocene-late Oligocene boundary of Cowie & Bassett (1989). The develop, in conjunction with important changes in plate convergence tectonic upheaval which started a't ttiat time resulted in development velocity (Pardo-Casas & Molnar, 1987), and lasted until - 19 Ma of numerous small to mediuni-size basins in the area of the Altiplano (Sempere et al., 1990b, c). An important phase of magmatic activity and, especially, Cordillera Oriental. developed during the same interval. It must be stressed that the Within the Andean domain during both major tectonic crises, Bolivian Andes started to exist as a mountain belt, albeit of limited thrust deformations developed more particularly in certain areas and altitude, only at that time. Concurrently, the Andean external along specific structures, and intermontane foreland basins formed in foreland basin rapidly shifled eastwards, reaching the Subandean- relation to them. Examples are the northern , foreland Llanura domain, while several intermontane basins formed in the of the fold-thrust belt and Coniri thrust system; the Río Andean domain, including the large ones on the Altiplano. Mulatos basin, foreland of Uie Sevaruyo-Chita fold-thrust belt; the

632 ~ ,

,.FOSILES Y FACIES DE BOLMA - VOL. I VERTEBRADOS

Rolívar-MondragBn basin, foreland of lhe Main Altiplanic thrust; the Anticlinal de San Andr6s" (=San Andres Formation of Evemden el Lípez basin, foreland of the Khenayani system; etc. ... (Fig. 1). al., 1966 which has a dubious associated K-Ar age of 38 Ma on Other small basins can be considered as piggyback basins, "glauconite") to the west of San Andres de Machaca on the Altiplano completely or nearly isolated from others due to reliefs developed by just south of Lago Titicaca. The fossils reputedly came from a unit thrusting. Examples are the basins of Salla-, Lacayani, equivalent to a part of the Mauri Formation (s.[.; YF'FB, unpublished Tarabuco, probably Muyu Huasi, (Fornari et al., 1987), data) and is thus late Oligocene-Miocene in age (Lavenu et al., Padilla, and San Isidro. The San Juan del Oro planation 1989), though Eocene age sediments have been traditionally mapped surface and its related deposits originated by the action of a in this area based on the one age obtained on "glauconite" (Martinez, longitudinal fluviatile system trapped within the Cordillera Oriental 1980). Glauconite is widely considered to be a marine mineral, (Sempere et al., 1990d), which can be considered as a giant whereas only continental strata have been deposited in the Altiplano piggyback basin. since the Paleocene. Hence, the age determination by Evemden et al. Other basins formed in relation to wrench-faults. Examples are the is dubious because the dated clay mineral is probably not authigenic. and Río Khuchu basins, and the basin system of the Unfortunately, these fossils were never described, Ortega (1970) "valles". made no mention of their existence in his review of Bolivian fossil Present knowledge of the Eocene-Pleistocene chronostratigraphy faunas, and they cannot be located in the GEOBOL collections. of Bolivia, including some unpublished data, has been summarized by Sempere & Oller (1987) and Sempere (1990), and is shown in LATE OLIGOCENE-EARLY MIOCENE Figures 2 and 3. Partly due to the numerous basins or sub-basins, (Deseadan) multiplication of formational names during the 1960's has unnecessarily complicated the stratigraphic nomenclature. Many The most spectacular land mammal fauna of Deseadan age in all names of local value have therefore been eliminated from our of South America is from the Salla-Luribay basin, about 95 km stratigraphic charts. Some of these are mentioned in the text when southeast of in the Cordillera Oriental (Hoffstetter, 1968b). used in the cited literature, and their preferred equivalent is given. The principal fossil-bearing unit lias been called the Estratos de Salla Of the two tectosedimentary periods defined above, the second (Evemden et al., 1966, 1977; Villarroel & Marshall, 1982). Strates one, which spans the late Oligocene-Pleistocene interval is by far the ou Couches de Salla (Hoffstetter, 1976a) and Salla Beds richest in fossil vertebrates, as documented below. (MacFadden et al., 1985) which consist of - 540 m of well-indurated red-brown, pink-tan and'whitish claystones and siltstones deposited VERTEBRATE FAUNAS AND LOCALITIES primarily in a fluviatile environment. The Estratos de Salla represent a unit partly equivalent to the Coniri Formation (Hoffstetter, 1968b; EOCENE Sempere et al., 1990b). (-) The geology and fossil localities are discussed by Hoffstetter (1968b, 1976a), Villarroel & Marshall (1982) and MacFadden et al. A probable Casamayoran age fauna reported by Sig6 et al. (1984) (1985). The latter provide a magnetostratigraphic and radioisotopic includes remains of actinopterygian fish (Characiformes, study of the Salla Beds and conclude that they span a time interval Serrasalmidae, Myleinae; Siluriformes), indeterminate turtles (Broin, from about 28.5 - 24.0 Ma, making this the best dated rock unit of 1991), teeth, and a single upper premolar of a primitive Cenozoic age in Bolivia. MacFadden et al. (1985) also demonstrate notoungulate mammal close to Camargomendesia (sensu Cifelli, that the fission-track dates of 54.0 k 2.6 and 52.0 It 2.1 Ma rehrted 1983) from a locality they designated Vela Pachita, about 57 km by Hayashida et al. (1984) for the Salla Beds are too old and southeast of , in what they considered to be the Santa apparently represent dates obtained on detrital (reworked) zircon Lucía Formation. However, the correct name of this locality is Wila grains. Naeser et al. (1987) published three fission-track (34.5 f 4.0, Apacheta and the fossils are from the basal Cayara Formation which 24.2 f 3.6, 23.5 f 2.2 Ma) and four K-Ar (biotite- 28.0 f 0.9, 27.2 rests unconformably on the El Moliio and/or Santa Lucía formations rt 0.9, 27.9 f 0.9, 25.1 f 0.7 Ma) dates from four ash levels in the (Marocco et al., 1987); the age of this unconformity shouldsbe 53 or lower part of the Salla Beds and conclude that the principal fossil 51-50 Ma (see above). horizons range from 27-24 Ma, while the complete section ranges Also from the Cayara Formation at La Cabaña on the from about 28-22 Ma. These dates are consistent with those of Cochabamba-Oruro road, about 25 km southwest of Cochabamba, Hayashida & Danhara (1985) who report four fission-track dates were recovered remains of fish and reptiles (, turtles) from (27.2f1.6,25.0k1.5,26.1f 1.9,24.0flSMa)onzirconsf?oman just below a level with abundant plants; turtles were observed in the unspecified level(s) in the Salla Beds. The of the upper levels of the Cayara Formation at Tiupampa, about 95 km Salla Beds is summarized by MacFadden (1990b) and McRae southeast of Cochabamba; and indeterminate fish and turtles wcre (1990). Sempere ef al. (1990b) reinterpret the geochronologic data recovered Iiom Ulis formation about 5 kin north-northeast of Chita. presented by MacFadden et al. (1985) and suggest that their dated A dubious record of a Casamayoran age locality was given in an section may be 3.0-4.5 Myr younger. unpublished report by E. Ortega in 1968. This report was seen by The diverse mammal fauna includes: Marsupialia (, Hoffstetter in GEOBOL, La Paz, although attempts to relocate it by Notogale mitis, Paraborhyaena boliviana, Pharsophorus lacerans, one of us (L.G.M.) in 1989 were unsuccessful. According to Sallacyon hoffsletteri: Caenolestidae, Palaeothentes boliviensis; Hoffstetter (in Marshall et al., 1983, p. 41), Ortega recorded remains ?Polydolopidae or Palaeothentinae. gen. et sp. indet.; Argyrolagidae, of mammals, including a fragment of a jaw of a Proargyrolagus bolivianus), Edentata (Dasypodidae, gen. et sp. (, Nofòpithecus sp.), from the "areniscas rojas del indet.; Peltephilidae, cf. Peltiphilus sp.; Glyptodontidae, cf.

633 LARRY G.MARSHALL &THIERRY SEMPERE

.- (*' noulhsm SJO Surfoca \, / of Subandaon L- \/ dsformdtlon I i. A R 69' , G 65E N T I I I $3' 'i. 61-I 23"

Figure 1. Sketch-map of principal late Oligocene-Pleistocene age sedimentary basins and tectonostratigrapllic boundary faults in Bolivia. Abbreviations for time: O, Oligocene; M, Miocene; P, Pliocene; PI, Pleistocene; H,; e, early; m, middle; 1, late. Abbreviations for fhults and surfaces: CALP, Cabalgamiento Altiplsnico Principal (Main Altiplanic Thrust); CANP, Cabalgamiento Andinò Principal (Main Andean Thrust); CCR, Cabalgamiento de la Cordillera Real (Cordillera Real Thrust); CFP, Cabalgamiento Frontal Principal (Main Frontal Thrust); FAT, Palla -Tupiza (Aiquile-Tupiza Fault); FC, Falla Cochabamba (Cochabamba

634 FOSILES Y FACIES DE BOLIVIA - VOL. IVERTEBRADOS

Glyptatelus sp.; family indet., Pseitdoglyptodon sallaensis; sp.; Broin, 1991). This age assignment is based on the presence of Megalonychidae. II. gen. et n. sp; , cf. Rltyncltippus, a genus known only from Deseadan faunas elsewhere Ocfodonfotheriiint sp.), Condylarthra (Didolodontidae. Salladolodlts in South America (Marshall, Cifelli el al., 1986). In addition, the deuterotheroides), (, Branisella boliviana), subfhily Rhynchippinae is unknown after Deseadan time, at least caviomorph Rodclilia (Octodontidae, Migraveranius beatus; in Argentina. Echimy idac, Sallaniys pascitali; Dasyproctidac, Incaniys bolivianus, III the Corocoro area, a cast of a single footprint of a large edentate cf. Neoreoniys sp., Cephaloniys bolivianus; Dinomyidae, was collected from what was called the Desaguadero Series, in a Branisamys luribayensis), (, small quarry along the railroad about 3 km southwest of Corocoro ?Coniopternium printitivunz; Adianthidae, Tliadanius Itoffstetteri, from rocks which belong to the middle part of the Coniri Formation Tricoelodus boliviensis), Notoungulata (, gen. et sp. (Cherroni & Cirbiln, 1970). This footprint is figured by Singewald indet.; , ?Rltynchippus cf. eqiiinirs, ?R. cf. puntidis; & Berry (1922, pl. 6, p. 52), but its true identity has yet to be , Nesodontinae, gen. et sp. indet.; , established. Because of its stratigraphic position, this edentate foot Prohegetotlieriunt sp.; Arcliaeohyracidae, Archaeohyrax sp.; print may be of late Deseadan age. Interatlieriidae, Argyroltyrax sp.; Mesotheriidae, Tracltytkerus sp.), Lastly, geologists of the Gulf Oil Corporation collected a partial (, gen. et sp. indet.), fossil turtle from along the Rio Cesarsama at a locality they called (Pyrotheriidae, Pyrotheriuni ronieri). order and family Icharco, about 30 km south of Todos Santos and 115 km northeast of (cf. Acanta sp.) (Cifelli & Soria, 1983a, b; Engelmann, 1987; Cochabamba. The fossil was sent to Dr. Donald Baird of Princeton Hartenberger, 1975; Hoffstetter, 1968b, 1969, 1976a; Hoffstetter & University who believed that it was of Tertiary age. This fossil Lavocat, 1970; Hoffstetter & Petter, 1983; Lavocat, 1976; probably came from what López (1983, p. 21) either calls the Bala MacFadden et al., 1985; MacFadden & Frailey, 1984; Patterson & Formation which is equivalent to the Petaca Formation or froin what Marshall, 1978; Patterson & Wood, 1982; Petter & Hoffstetter, he calls (p. 24) the Quendeque Formation, of probable late Miocene 1983; Soria & Hoffstetter, 1983; Villarroel & Marshall, 1982; Wolff, age (Fig. 2). Correspondance on the discovery and identification of 1984, 1985). Of noteworthy importance is Branisella boliviana, the this fossil is in the archives of the Museo Nacional de Historia oldest known in South America (Hoffstetter, 1969; Natural, Cochabamba (letter from T. A. Kibby to D. Baird, May 10, Rosenberger, 1981; Wolff, 1985). Turtles (Chelonia, 1961). Podocnemididae, ?Podocnemis sp.) are also recorded (Broin, 1991). A second Deseadan fauna is from Lacayani, located about 30 km EARLY AND MIDDLE MIOCENE southeast of La Paz (Hoffstetter ef al., 1971). The fossilsi which (-Chasicoan) come from an unnamed rock unit include: Marsupialia (Rorhyaenidae), Edentata (Dasypodidae, cf. Prozaedyus; A relatively rich fauna of middle Miocene age is known from an Glyplodontidac, Glypfnfelus;Orophodonlidae), Litopterna unnamed unit at Quebrada Honda (3500 m), about 55 km southwest (Macr a u ch en i id ac, c f. Coniopt ern iuni), Noto u II g u 1at a of Tarija, just north of the Argentine border in southernmost Bolivia. (Mesotheriidae, 7rachytherus spegazzinianus; Hegetotheriidae, This unit is of fluvio-lacustrine origin and is only slightly deformed. Prohegetoherirmi), and caviomorph Rodentia (Cephaloinyidae, gen. The fauna was assigned a Colloncuran (Friasian) age by Hoffstetter et sp. indet.; , Eoviscaccia boliviana) (Hoffstetter et (1977), and subsequent study by Frailey (1988) suggested that it was al., 1971; Hartenberger, 1975; Vucetich, 1988, 1989). Santacrucian. However, recent geochronologic work by MacFadden A poorly known fauna from the base of the Petaca Formation at (1990a) and MacFadden et al. (1990) shows that most of the Quebrada Saguayo in the Subandean belt, about 60 km west- Quebrada Honda strata were deposited between 13 and 12 Ma, northwest of Santa Cruz, is probably of Deseadan age. The known indicating that the fauna is late Colloncuran in age. A brief fauna includes a right maxilla of a mammal identified as description of the geology is given by MacFadden & Wolff (1981). ? sp. (Notoungulata, Notohippidae, Rhynchippinae) by The fauna includes: Marsupialia (Borhyaenidae; Caenolestidae; Villarroel (in Sanjinés & Jitn-énez, 1975; see also Hoffstetter, 1977, Argyrolagidae, Ifondalagus altiplanensis), Edentata (Dasypodidae; p. 1519), a small plate of an indeterminate armadillo (Dasypodidae) Glyptodontidae, Propalaeohoplopltorus andinus; Mylodontidae; and remains of a tortoise (Chelonia, Testudinidae, cf. Cltelonoidis Megalonychidae, Napalops angustipalafus), Notoungulata

~~ Fault); FCA, Falla Chita-Arica (Chita-Arica Fault); FCC, Frente de Cabalgamiento Coniri (Coniri Thrust Front); FE, Falla Eucaliptus (Eucaliptus Fault); FSI, Falla Sevaruyo-Incapuquio (Sevaruyo-Incapuquio Fault); SFK, Sistema de la Falla Khenayani (Khenayani Fault System); SJO, San Juan del Oro planation surface. Basins and ages: 1, Beni basin (M-H) and adjacent northern Subandean basin (IO-P); 2, (M-H) and adjacent southern Suhandean basin (10-P); 3, Chiquitano basin (?-H); 4, Tipuani basin (IM-P); 5, San Isidro basin (IP-ePI); 6, Muyu Huasi basin (1M); 7, Padllla basin (P-PI); 8, Tarija basin (PI); 9, Tarabuco basin (?-?); 10, Culpina basin (?-H); 11, Quebrada Honda basin (mM); 12, San Juan del Oro surface related deposits (IM-PI); 13, Cochabamba basin system, including Anzaldo and basins (IM-Pl); 14, La Paz basin (P-PI); 15, Ulla-Ulla basin (?-Pl); 16, Lacayani basin (10-eM); 17, Salla-Luribay basin (IO-eM); 18, Northern Altiplano basin (10-H); 19, noIlvar basin (IO-eM); 20, Mondragh I)asin (10-eM); 21, Rfo Mulatas basin (M); 22, Llpez basin (IO-H); 23, Rfo Khuchu basin (10-M); 24, Tupiza basin (10-M); 25, basin (10-H); 26, Salinas de Gard Mendoza basin (?-H); 27, Carangas basin (M-H); 28, Charaña basin (M-H). The Andean domain lies wesf of the CANP, whereas the Subandean belt (S.S.) extends between the CFP and the present-day edge of deformation to the east.

635 LARRY G.MARSHALL & 17IIERRY SEMPERE

(s/d 'unnamed

Figure 2. Clironostratigrapliic chart sliowing Eocene-middle Pliocene (Casamayoran-) South American Land Mammal Ages (modified after Marshall, 1985) and principal vertebrate-bearingformations (*) discussed in text (modified after Sempere, 1990). follows Berggren et al. (1985)

(Hegetotheriidae, Hegefofheriunisp.; In teratheriidae; Mesotheriidae; mainly of conglomerates, reddish-brown siltstones and mudstones, Toxodontidae), Astrapotheria (Astrapotlieriidae, ?Xenasrr~~otheririni and volcanic ash. These sediments are poorly indurated and reach a sp.), Litopterna (, sp.; maximum thickness of - 160 in. The reported fauna includes 10 taxa Macraucheniidae), and caviomorph Rodentia (Octodontidae; referable to S orders and 8 families: Marsupialia (Borhyaenidae, Echiniyidae; Capromyidae; Chinchillidae, Prolagosloniiis sp.; Acrocyon?), Edentata (Dasypodidae, Stemtatus sp., cf. Pelfej~hiliis Caviidae) (Hoffstetter, 1977; MacFadden & Wolff, 1981; Takai et sp.; Glyptodontidae, Propalaelioplophorics sp., Neothoracoplioriis? al., 1984; Frailey, 1987, 1988; Villarroel & Marshall, 1988). sp.), Notoungulata (Mesotheriidae, Plesiotypollzeriuni sp.; The 300 m-thick Quebrada Honda strata accumulated by infilling Hegetotheriidae, lfegetofheriuni sp.), Litopterna (Proterotheriidae, of tlie Pasajes syncline which formed just west of the CANP major Diadiapliorus sp.), and Rodentia (Octodontidae, gen. et sp. indet; fault (Fig. 1). This syncline is bounded on its western side by a Chinchillidae. Prolagosfonwssp.). skinger of en-échelon anticlines which involve and early Also from the Nazareno Formation at Suipacha, 3 km north- strata and document a regional right-lateral component northwest of Nazareno, an armadillo was collected during for the CANP (Sempere, unpublished). The formation of the Pasajes construction of a railway cut at km 69 on the line from Villazón to syncline is thus likely to have been coeval with the first major Atocha. This specimen was described by Castellanos (1925) and tectonic crisis mentioned above (27-19 Ma). After deformation had named Dasypodon afavus (Edentata, Dasypodidae, Euphractinae). ceased, differential of Ordovician shales and Cambrian Hoffstetter (1977) tentatively regarded this fossil as Huayquerian. quartzites, and probable local faulting; led to formation of a hollow Ois0 (1991) regards Dasypodon alnvris as a small of depression in the central part of the Pasajes syncline were fluvial and Slenotafus (i.e. S. afnvus), and given its provenance from the lacustrine sediments. as described by MacFadden & Wolff (1981), Nizareno Formation is also almost certainly of Colloncuran age gently accumulated. These deposits are thus partly coeval with the (sensu Marshall & Salinas, 1990). development of the middle Miocene Chayanta planation surface Huxley (1860; see also Blakk, 1861) described the partial skeleton (Sempere et al,, 1990e). and with the deposition bf the Quehua of a litoptern (Macratcchenia boliviensis) from what Singewald & Form ation. Berry (1922, p. 44) call the Ramos Series at Corocoro. a copper mine A second relatively rich fauna of Colloncuran age is reported by on the Altiplano south of La Paz. This unit is equivalent to the Ois0 (1991) from the middle and lower members of the Nazareno Kollukollu Formation (Cherroni, 1974) which has a K-Ar date of Formation about 10 km south of Nazareno (on the west side of 16.6 f 0.4 Ma on a sanidine concentrate from a reworked tuff Estación Arenales; 21"40'S, GS"3S'W) and 30 kin southeast of (Swanson el al., 1987). Hoffstetter & Paskoff (1966, p. 483) cotifinn Tupiza in the Cordillera Oriental, department of Potosí, in the Tupiza that this specimen is indeed a macraucheniid. but note that it is not basin of southern Bolivia. Here the Nazareno Formation consists referable to the Pleistocene genus Macrarcclienia and is smaller than

636 FOSILES Y FACES DE BOLIVIA - VOL. I VERTEBRADOS thc carly to middle Miocene gcnus Tlieosodon. Furthermore, the Alto Rio Moile are apparently also of Cllasiconri or specimen is poorly prescrvcd and a secure generic identification Huayquerian age. appears improbable (Ahlfeld & ßranisa, 1960, p. 165). Thus, the true A fauna of probable Chasicoan age is reported by Russo (1959, p. gencric idcntity and age of this species remains unknown. Because 29) from "un afluente derecho, sin nombre, de la Quebrada de of its stratigraphic position, this macraucheniid might be of Palmar" at Agua Salada near Yacuiba in southeast Bolivia just north Colhuehuapian or Santacrucian age. of the Argentine border. The fossils were found in sediments which There are two poorly known faunas of possible Colloncuran age in Russo called the "Chaco Inferior" and are now called the Tariquía Bolivia. The first is from the Choquecota Formation (= Formation, which overlies the Yecua Formation. As noted by Russo Formation, which near its base contains blocks of dacite lava K-Ar (p. 29), "en una arenisca situada directamente sobre el banco tobaceo dated at 14.2 f 0.4 Ma; Swanson et al., 1987) about 3 km north of gris, se hallaron huesos, dientes y un bozo de columna vertebral de Choquecota, department of Oruro, on the Altiplano (see Hoffstetter unos 40 centímetros de longitud, con apófisis dorsales y costillas aún et al., 1972). This was the site of collection of a small notoungulate sujetas a él, de un mamífero fósil de hábito terrestre. Estos fósiles no (Mesotheriidae) which Villarroel (1974b) named Microfypofheriuni han podido ser determinados en el campo, su clasificación requeriría choquecoteme. Remains of Dasypodidae (Prozaedyus), el trabajo de un especialista y ahasísería difícil hacerlo debido a la Glyptodontidae, Hegetotheriidae and Chinchillidae are also known pobreza del material recolectado". (Hoffstetter el al., 1972). The second locality is in southwestern From the base of the Guandacay Formation a few Bolivia between Cerdas and Atocha, where Microtypolherirmz cf. M. kilometers north of Bermejo along the Rio Bermejo, Ing. Antonio choquecoteme was collected from tlie lower part of the Quehua Sadud collected a partial skull and skeleton of a notoungulate Formation S.S. (mapped as "Formación Quehua Superior'' on Hoja (Hegetotlieriidae; determined by R. Pascual). This formation is 6331, Quechisla) (see Hoffstetter, 1977; Villarroel, 1978, p. 166; apparently Chasicoan to Huayquerian in age. Hoffstetter in Marshall et al., 1983, p. 41). A small species of Notoungulata (Mesotheriidae, LATE MIOCENE Plesiofypofheriuni niinus Villarroel, 1978) was collected 3 kin (Huayquerian) southeast of Cerdas from the upper part of the Quehua Formation S.S. (mapped as "Formación Quehua Superior" on Hoja 6331, There are numerous land mammal faunas of Huayquerian age in Quechisla). This species appears to be more specialized than Bolivia, particularly from the Mauri 6, Rosa Pata and Quehua Microfyyotheriuni choquecoteme suggesting that it may be formations on the Altiplano. At localities where these formations Chasicoan in age, although this has yet to be demonstrated were deformed during Huayqueriaii time they are disconformably (Villarroel, 1978; Hoffstetter in Marshall et al., 1983, p. 41). overlain by rocks of Pliocene and/or Pleistocene age. These A locality from the uppermost part of the Petaca Formation deformations on the Altiplano correspond to the second major on the Río Yapacani, about 90 kin west-northwest of Santa tectonic crisis mentioned above (11-5 Ma), and apparently to what is Cruz in the Subandean belt yielded a single right mandible termed the Quechua deformation in Argentina (Hoffstetter, 1986, p. ' with five teeth which was identified by Pascual and Odreman 222) which began about 10 +_ 2 Ma (Jordan & Alonso, 1987). Rivas (in Sanjinés 6c Jiménez, 1976) as cf. Vassallia minuta A sample which yielded a K-Ar age of 10.5 Ma (Evernden (Edentata, Dasypodidae, Pampatheriinae). This identification ef al., 1966, 1977) was apparently collected from a tuff about 1500 indicates a Chasicoan to age for this level of m above the base of the Mauri 6 Formation at Cerro Jakokota the Petaca Formation. (=Hakakota) (4,000 m), 12 kni southeast of . A few vertebrate fossils have been recovered from the Yecua This locality may be the same as Cerro Jancocota where Berry Forination, which overlies the Petaca Formation, along the Río Alto (1922) collected and described a fossil flora that includes of Moile, about 95 km west-northwest of Santa Cruz in southcentral Pteris, Pliragmites, Alnus, Osteomeles, Polylepis, Calliandra, Bolivia (López, 1983, p. 18). The known fauna includes a limb bone Cassia, Cesalpinia and Melastoniites; fossil wood of of a litoptern tentatively identified as cf. sp. Prosopis (Mimosaceae) occurs below the tuff at this locality (Macraucheniidae), an indeterminate , the first fossil (Hoffstetter, 1986). The Mauri 6 is disconformably overlain by the remains of the fish order Gyninotiformes (Apteronotidae, gen. et sp. subhorizontal Pérez Ignimbrite which has a mean radioisotopic indet.) (electric eels) (Gayet, 1986) and indeterminate Siluriformes age of - 2.8 Ma (Evernden et al., 1966, 1977; Lavenu et al., (Gayet, 1991). If the identification of Tlieosodon is correct, then the 1989; Marshall er al., 1992) and probably belongs to the same age of this fauna is somewhere between Colhuehuapian and igiiimbritic flow as the Chijini and tuffs which are Chasicoan, which is the known stmtigraphic range of this genus also - 2.8 Ma (Marshall el al., 1992). elsewhere in South America. The richest of the mammal-bearing Mauri 6 localities is at Acliiri The occ,urrence of the Chasicoan to Montehermosan cf. (especially at Cerro Pirapi Grande, about 6 km from ) which Vassallia niinula below a level with the Colhuehuapian to occurs in the upper part of this formation. Ofier noteworthy Mauri 6 Chasicoan cf. Theosodon sp. implies that the top of the localities include Canacho, Jankojakhe (=Jankoaqui) and Rosario. Petaca Formation and at least a part of the Yecua Formation The known fauna from these four localities includes: Marsupialia are Chasicoan in age (Fig. 2). This age assignment is supported by (ßorhyaenidae, Borliyaenidirim altiplanicus), Edentata the fact that the Yecua Formation represents the northern extent of (Dasypodidae, Chorobates sp., cf. Kraglievìchia sp.; the Paranense epicontinental sea which is, at least in part, temporally Glyptodontidae, Sclerocalyptinae; Megalonychidae; Mylodontidae), equivalent to Chasicoan (Sempere, 1990). The fish fossils reported and Notoungulata (Toxodontidae, Toxodontinae, gen. et sp. indet., by Lbpcz (1975) from tlie base of the Tariquía Formation along the and a new form with a frontal horn, see Hoffstetter, 1976b, pp. 124-

637 LARRY G.'hURSIIALL 8r'I'IUERRY SEMPERE

125 for photo: Mesotheriidae, Plesiotypotlieriunt aclrirense, P. In summary, the large herbivqres in Huayquerian faunas on the niajus) (Villarroel, 1974a; Villarroel & Marshall, 1983; Hoffstetter, Altiplano include Mesotheriidae (Plesiofypolheriunz),Toxodontidae 1986). (medium-sized Toxodontinae and a large forni with a frontal horn), Douglas (1914, p. 24) collected a fragment of the symphysis-of a several ground sloths, Glyptodontidae (Sclerocalyptinae) and large notoungulate (Toxodontidae) from "a few miles below the Mauri Dasypodidae (Pampatheriinae) (Hoffstetter, 1986, p. 236). bridge" from what he called the Desaguadero Formation along the There are four additional mammal-baring faunas of possible or Rio Mauri, a tributary of the Rio Desaguadero. Villarroel (1977, p. reputed Huayquerian age in Bolivia, yet all need confirmation. 29) notes that the Desaguadero Formation of previous workers has First, an edentulous skull of a Mesotheriidae (aff. no significance and it is not possible to securely know what Plesiotypofheririmachirense) was recently collected from the poorly stratigraphic units ih current use were assigned to it. Nevertheless, known Casira Formation to the southeast of the pueblo Casira on the Ahlfeld & Branisa (1960, p. 134) believe that the specimen collected southwest side of Cerro Khellu Khakha Loma, Modesto Omiste by Douglas came from the upper part of the Mauri Formation, and is Province, department of Potosí, near the Argentine,border in probably from the Mauri 6. This specimen was identified by C. W. southernmost Bolivia (Anaya et al., 1989). Andrews of the British Museum (Natural History), London as Second, is a fauna from the "Estratos Muyu Huasi" at Muyu sp., a genus known securely only from rocks of Huasi, about 50 km northeast of in southcentral Bolivia. The Santacrucian and Colloncuran age in Argentina (Marshall et al., taxa include: Edentata (Dasypodidae, cf. Doellofafussp., cf. 1983). Simpson (1940) reported that this specimen is no longer in Paleupltracfus sp.; Mylodontidae, gen. et sp. indet.), Litopterna the collections of the British Museum and as such its identity cannot (Proterotheriidae, gen. et sp. indet.), Notoungulata (Hegetotheriidae, be confirmed. For the present this taxon can only be regarded as Pseridohegetotlteriunt sp.; Interatheriidae, Profypoflieriuni sp.) and Toxodontidae, gen. et sp. indet. caviomorph Rodentia (Abrocomidae, Profabroconta sp.; Caviidae, Land mammals have been collected from the upper part of the ?Orflioniycferasp., ?Cardionlys sp.; Chinchillidae, Lagosromopis Totora Group, 2 km north of Ulloma at Minita Chocopini near sp.; Dinoinyidae, gen. et sp. indet.; Eretliizontidae, gen. et sp. indet.; Torini, 4 km west of , north of Chacarilla, and in the vicinity Octodontidae, cf. Sciamys sp.) (Villarroel & Marshall, 1989). of (Hoffstetter, 1986). Two tuffs from the Third, is the fauna from the Cobija Formation collected along the Rosa Pata Formation (upper part of the Totora Group) named toba Río Acre between Cobija and Bolpebra, department of Pando in Callapa and toba Ulloma, have been K-Ar dated at about 9.0 and northwest Bolivia. The vertebrates include fish (Chondrichthyes), 10.4 Ma, respectively (Marshall et al., 1992). The Rosa Pata reptiles (Chelonia, P odocnenzis sp.; , Gavialosrcclius sp.), Formation is either disconformably overlain by the latest Miocene (indet.), and mammals (Rodentia indet.) (Carrasco, 1986). The Pomata Formation, Pliocene Formation which has a mean jaw of an enormous crocodile was collected from this same radioisotopic date of 5.4 Ma for its basal tuff (=toba 76, toba Umala formation at Candelaria, to the west of Riberalta along the Río or toba Jankhomarka; Thouveny & Servant, 1989; Marshall et al., Madre de Dios (Leytón & Pacheco, 1989). 1992), Pérez Ignimbrite dated at 2.8 Ma (see above), or Pleistocene And fourth, a CTagment of a right dentary witli well preserved M* Ulloma Formation (see below; Marshall et al., 1983; Hoffstetter, and M' of cf. Psertdoliegefoflieriuni sp. (Notoungulata, 1986). The known mammals include: Edentata (Dasypodidae, Hegetotheriidae) was recently collected at the Estación Abaroa, Pampatheriinae; Glyptodontidae), Litopterna (Macraucheniidae?) about 25 km east-northeast of Charaíía, department of La Paz, The and Notoungulata (Mesotheriidae, P seudofypotlierirrni sp.) specimen is in GEOBOL collections in La Paz and has the catalogue (Martinez & Rosales, 1972; Hoffstetter et al., 1972). number GB-089 (C. Villarroel, personal communication, 1990). From the base of a 205 m section of what are informally called the Micaiía beds at Micaiía in the Cordillera Oriental (17'30's. 67"24'W; EARLY AND MIDDLE PLIOCENE 38004000 in) were recovered a niediuni-sized niegatheriid ground (Monteliennosan-Cliapadinalalaii) sloth (gen. et sp. indet.) and a small notoungulate (Mesotheriidae, Microtypotlierirmi cf. M.choquecotense) (MacFadden et al., 1990). The most important mammal fauna of Montehermosan age in A magnetostratigraphic study and an associated fission-track date of Bolivia is from tlie Umala Formation at and near Ayo Ayo (3800 in), 6.9 k 1.1 Ma, indicate that these rocks and fauna span fToin - 7.5 - Vizcacliani and Umala, 65-75 km south of La Paz. A tuff from the al., 6.7 Ma (MacFadden ef 1990), making thein late Huayquerian. base of the Umala Formation (toba 76) has yielded mean Remains of Notoungulata (Mesotheriidae, Plesiofyi~oflieriuni) radioisotopic dates of 5.4 Ma (Evernden ef al., 1966, 1977; Lavenu havc bccn collcclcd from two localitics of thc Quehua Foi inat ion S.S. c/ al., 1989; Marshall et al., 1992). whilc thc Ayo Ayo tuff in thc top to thc east-southcast of Quchua in the southern part of the Altiplano of this foriiiatioii has yicldcd a iiiean age of 2.8 Ma (Lavenu CI al., (Villarroel, 1978, p. 166; Hoffstcttcr & Villarroel, unpublished as 1989; Marshall et al., 1992). The fauna includes: Marsupialin cited in Marshall et al., 1983, p. 43). Plesiotypotheririni cf. P. niajrts (Didelphidae, Sparassocyniiiae, Sparassocynrrs lieterotopicrrs; or niinrrs and scutes of Dasypodidae were collected from ?Caeiiolestidae; Argyrolagidae, Microtragrrlrrs bolivianrrs),Edentata approximately 3 km southeast of Quehua; and cf. P. achirense was (Dasypodidae. Pampatheriinne, cf. Plaina sp., Euphractinae, collected approximately 7 km southeast of Quehua (the first locality Macroeirplzractiis sp.; Megatheriidae, cf. Megt~lieriiinr; is from a level 40 ni stratigraphically higher than the second). Mylodontidae, Mylodontinae), Notouiigulata (Toxodontidne, Rodentia (Caviidae, Dolichotinae, Orflioniycferasp.) are also Xotodontinae, Posnanskytlieriiini desagrraderoi), Litopterna recorded from Quehua (Hoffstetter, 1986, p. 222). The presence of (Macraucheniidae, cf. Proniacrauclienia sp.), and cavioniorph these taxa establishes that the Quehua Formation at these localities is Rodentia (Octodontidae, Ctenomyinae, Praecfenoniys rhonibidens, temporally equivalent to the Mauri 6 and Rosa Pata, formations P. vagus; Chinchillidae, Lagostominae; Caviidae, Caviinae; fartlier to the north. ' Hydrochoeridae. Cltapalntarlieriuni saavedrai) (Liendo-Lazarte,

638 FOSILES Y FACIES DE BOLMA -VOL 1 VERTEJHUDOS

SOUTH AMERICAN ALTIPLAN 3 :ORDILLERA $GE OCHABAMBP ORIENTAL CHACO & LANDMAMMAL AGE GLACIATIONS GLACIAL LAKES ASSOCIATED M~) FORMATIONS (Central 1 i;luapua~* Fm.* 1 Ulloma Anzaldo * I- Tari)o*, Padcayo *, 1.04 Cabana ------ConceEcn2- :I ENSENADAN I Kaluya

Mataro * Charaña Fm.* !Purapurani T. 1.6Ma - ?-?-?- Betanzos * Anzaldo *

2.5 r------jAmerican Faunal Interchange Pérez Ign.2.8Mr: * 30 MONTEHERMOSAN Anzaldo Umala Fm.* Sacaba I *

Figure 3. Chronostratigrapliic chart showing late Pliocene-Pleistocene (-Lujanian) South American Land Mammal Ages (modified after Marshall, 1985) and principal vertebrate-bearing formations (*) discussed in text. Geologic time scale follows Berggren et al. (1985).

1943; Hoffstetter, 1986; Hoffstetter et al., 1971, 1984; Hoffstetter & about 200 km south of La Paz, from rocks designated as the Villarroel, 1974; Villarroel, 1975, 1977; Villarroel & Marshall, Formación Mauri (Hoja 6038, ; Hoja 6039, Choquecota; see 1983). Hoffstetter et al., 1972, Fig. on p. 740) which unconformably overlie the Huayquerian age Rosa Pata Formation (see above) or more Numerous mammals have been collected from the La Paz specifically its terminal facies, the Conglomerado Pomata, which has Formation around La Paz at , Alto Obrajes, Alto a K-Ar date of 6.4 Ma obtained on a rhyolite (Evernden et al,, 1966, Següencoma, , Gualberto Villarroel, Kenko and 1977; Marshall et al., l9!3, p. 78). The fauna occurs aboqe a tuff Tembladerani. This formation consists of some 700 m of fine to which may represent the toba Jankhomarka (Martinez, in Marshall coarse sediments of fluvial to fluvio-lacustrine origin (Lavenu et al., et al., 1983, p. 78) and includes: Notoungulata (Toxodontidae, 1989) which Dobrovolny (1962, pp. 20-27) subdivided into three Xotodontinae, Posnanskytheriuni desaguaderoi), Litopterna units (lower, middle, upper). The fossils are from below a tuff (toba (Macraucheniidae, cf. Pronaacrauchenia sp.), Edentata Chijini) in the upper part of the La Paz Formation which has yielded .(Mylodontidae), caviomorph Rodentia, and a large terrestrial a mean radioisotopic date of - 2.8 Ma (Clapperton, 1979; Lavenu el carnivorous phororhacoid (Hoffstetter et al., 1972; Hoffstetter, al, 1989; Marshall et al., 1992). This tuff appears to belong to the 1986). same ignimbrite flow that deposited the PCrez Ignimbrite and Ayo From the Remedios Formation at Cerro Canasa near Corque and at Ayo tuff (see above). Dates of - 5.5 Ma have also been obtained on Orinoca south of Oruro are reported remains of Edentata a tuff (= toba Cota Cota) in the lower part of this formation at Cota (Mylodontidae), Notoungulata (Toxodontidae, Xotodontinae, al., Cota, southeast of La Paz (Lavenu et 1985, 1989; Servant et al., Posnanskytherium sp.), and Litopterna (Macraucheniidae, cf. 1989). This tuff seems to belong to the same event that deposited the Pronzacrauchenia sp.) (Lavenu, 1984; Hoffstetter, 1986) from above Jankhomarka tuff (= toba 76), which apparently correlate with a tuff with dates of 5.2 - 4.6 Ma (Marshall et al., 1992). This collapse and eruption of the Soledad Caldera at 5.4 Ma (Redwood, formation is locally separated by an angular unconformity from the 1987), but was probably emitted somewhere in the western overlying mammal-bearing "lacustre Minchín" of Cordillera as suggested by its thickness increasing toward the west. age (see below). A magnetostratigraphic study of the La Paz Formation shows that A Montehermosan age for the mammal faunas from the Umala, La deposition of the middle and uppe; units occurred during the Gauss Paz, Remedios and "Mauri" (at Pomata) formations is established on epoch (i.e. 3.4 to 2.48 Ma) (Thouveny & Servant, 1989). The fauna faunal content and radioisotopic dates. These Pliocene faunas differ includes: Edenlata (Dasypodidae, Euphractini, Macroertphractus markedly from those of the late Miocene (Mauri 6, Rosa Pata, aff. nioreni; Glyptodontidae, Sclerocalyptinae,cf. Sclerocalypfussp., Quehua formations) on the Altiplano by the absence of cf. Plolrophorops; Megalonychidae, gen. nov.), Notoungulata Mesotheriidae and in the predominance of Macrkheniidae (cf. (Toxodontidae, Xotodontinae, Posnanskytkeriitm desagrraderoi, P. Proniacrauchenia sp.), other Toxodontidae (Xotodontinae, nov. sp.) and Litopterna (Macraucheniidae, cf. Proniacrauchenia Posnanskytlterium sp.), other Edentata (Megatheriidae; sp.) (Ahlfeld & Branisa, 1960, p. 135; Clapperton, 1979; Radelli, Mylodontinae; Pampatheriinae, Kraglievichia sp., Plaina sp.), and 1964, p. 842; Hoffstetter, 1986; Villarroel, 1977; Villarroel & Graf- Hydrochoeridae (Chapalmatkeriuni sp.) (Hoffstetter, in Marshall et Meier, 1979; Mones & Mehl, 1990). The vast majority of these al., 1983, p. 44; Hoffstetter, 1986). Furthermore, these faunas are fossils are from Dobrovolny's (1962) middle unit (Mones & Mehl, bracketed below by the 5.4 Ma toba Jankl>omarkafrom the base of 1990, p. 22). the Umala Formation and the - 5.5 Ma Cota Cota tuff in the lower A few vertebrates have been collected from north of Pomata Ayte, part of the La Paz Formation, and above by the 2.8 Ma Chijini tuff in

639 LARRY G. MARSHALL &THIERRY SEMPERE the upper part of the La Paz Formation and 2.8 Ma Ayo Ayo tuff in EARLY AND MIDDLE PLEISTOCENE the top of the Umala Formation. (Ensenadan) There are three other reports of possible early to middle Pliocene age mammal faunas in Bolivia, all of which need A fauna regarded by Hoffstetter (1986, p. 226) as possibly early confirmation. Ensenadan was collected by Blanco (1980, p. 99) from the Charaña First, Pachynodon validus Burmeister (1 892, pp. 433-434) Formation just east of Charaña (4000 m) on the Altiplano near the (Notoungulata, Toxodontidae) is probably from Bolivia and may be Chilean border. This formation overlies the Pbrez Ignimbrite and is al., of Pliocene age. As noted by Simpson (1940, p. 705), 'I ... the only thus younger than 2.8 Ma (Lavenu et 1989; Marshall et al., 1992) available information as to provenience is that the specimen was sent and is older than the Brunhes Normal chron (i.e. >0.73 Ma) to Vaca Guzmán Blanco (the Bolivian Minister in Buenos Aires) (Servant-Vildary & Blanco, 1984). The fossils, as yet undescribed, from Santa Cruz de la Sierra, Santa Cruz, Bolivia. The implication is include: Edentata (Glyptodontidae, Doedicurinae, Plwliaplous sp.; that the specimen was found at that place, but this is not definitely Mylodontidae, Glossotlzerium sp.), Litopterna (Macraucheniidae, established". Macrauclienia cf. ullonzensis), and Artiodactyla (Cervidae, gen. et Second, Montañ0 (1968) described a mammal fauna from Anzaldo sp. indet.) (Hoffstetter, in Marshall el al., 19M, p. 36; Hoffstetter, (3000 m), about 55 kni southeast of Cochabamba, which he assigned 1986; Blanco, 1980, Anexo 2). to the Pleistocene, A later visit to this locality by Hoffstetter, Another Altiplano fauna of probable Ensenadan age is from Ayo Montaiio and Ortega resulted in recovery of a specimen of cf. Ayo (3800 m), about 70 km south of La Paz. Specimens of Edendata Proniacrarrclienia sp. (Notoungulata, Macraucheniidae) from a 2 m- (Mylodontidae, Scelidodon cf. tarijensis), Litopterna thick siltstone in the lower level at Quebrada Tijascka, about 3 km (Macraucheniidae, cf. pafaclionica), and Artiodactyla southwest of Anzaldo. This is a small macraucheniid and is of (Cervidae, gen. et sp. indet.) were recovered from a sandy horizon Pliocene aspect (Hoffstetter, in Marshall et al., 1983, p. 44). A true above a tuff and palcosol at the top of the Urnala Formation, which Pleistocene mammal fauna occurs at the same localitiy (see below). has yielded a Monteliermosan age fauna at the same locality And third, from,a rock unit informally called "lac de Sacaba" or (Hoffstetter ef al., 1971; Hoffstetter, 1986; Lavenu, 1978). "Sacaba 1" (Lavenu, 1986, pp. 114-115) south of Sacaba in the area Hoffstetter (1986, p. 224) notes that Lavenu (1978) regards the Ayo of Laba-Laba Alta (Quebrada Fierro Churu) about 18 km east- Ayo tuff (which is 2.8 Ma; Lavenu et al., 1989; Marshall et al., southeast of Cochabamba, were recovered remains of mammals 1992) as probably equivalent to the pre-Calvario tuff at Patapatani al., identified by Villarroel (in Mancilla, 1979, p. 32) as Plohophorus sp. (which gave a K-Ar age of 2.8 Ma; Lavenu et 1989) north of La (Edentata, Glyptodontidae). Paz. Lavenu et al. (1989) also believe that these tuffs are equivalent to the Chijini tuff which has a mean radioisotopic age of 2.8 Ma al., LATE PLIOCENE (Marshall et 1992). (Uquian) The most spectacular vertebrate fauna of late Ensenadan age in South America is known from numerous localities and from various There is only ode potential fauna of this age in Bolivia. It is from stratigraphic levels in the basins of Tarija (1950 m; i.e. Armaos, San near Anzaldo, about 55 km southeast of Cochabamba. In Quebrada Blas, San Pedro, Santa Ana, Pueblo Viejo, Tolomosa), Concepción Tijascka, 3 km southwest of Anzaldo, Montaiio (1968) recovered a (= Uriondo, 1800 m; about 20 km south-southeast of Tarija), and complete caudal tube of Prodaedicurus cf. P. divincenzi (Edentata, Padcaya (2100 m; about 40 km south of Tarija) in southernmost Glyptodontidae) identified by Hoffstetter (in Marshall et al., 1984, p. Bolivia (Ahlfeld & Branisa, 1960, Fig. 11, pp. 168-171). The 36) from a gravel layer. This is the only record of Prodaedicurus for predominant lithologies within the Tarija basin are silty clays, sands, Bolivia, a taxon originally described from beds designated as the conglomerates and volcanic ash which were apparently deposited in "Piso Castellanosense" in Uruguay, the age and faunal content of a fluvial environment consisting of an accreting stream and which are discussed by Mones (1979, pp. 16-17). Mammals of floodplain system (MacFadden & Wolff, 1981). possible Monteheimosan (see above) and Pleistocene (see below) The first citation of fossil mammals from Tarija, and for that age are also known from Anzaldo. matter the first written record of fossil mammals from Bolivia, was by Diego de Avalos (1602; fide d'orbigny, 1842, 111-4, Paléontologie, p. 145). Joseph de Jussieu, in a letter to his brother LATIC T 15R3'1A KY

640 ~~ ~~

FOSILES Y FACIES DE BOLIVIA - VOL. I VERTEBRADOS

II

I

Figure 4. Map of Bolivia showing vertebrate localities of Eocene-Pleistocene age.

64 1 LARRY G. MARSHALL &THIERRY SEMPERE later papers on this collection were provided by Kraglievich (1928, This fauna is in dire need of a systematic revision. In addition to 1930b), Cabrera (1929, 193l), and López Aranguren (1930). In 1902 the papers cited above, the mammals are discussed by Berta (1981, Nordenskiöld sent a small collection to Uppsala, Sweden which 1985,,1987, 1988), Liendo-Lazarte (1946) and Hoffstetter (1963a, b, resulted in publications by Nordenskiöld (1902, 1903) and Sefve 1968a, 1986). The following taxa appear to be securely identified: (1912, 1915b). A large collection of specimens purchased in 1903 by Marsupialia (Didelphidae, Lutreolina crassicaudata), Edentata the Marquis de Créqui-Montfort and E. Sénéchal de la Grange from (Dasypodidae, Chaetophractus tarijensis, C. villosus, Eupliractus the Echazú family was donated to the Muséum National d’Histoire sexcinctus, Propraopus cf. sulcatus or grandis, Pampatherium cf. Naturelle, Paris and served as the basis for the now classic Itumboldti or typum; Glyptodontidae, Chlamydotherium cf. sellowi, monograph by Boule & Thévenin (1920) (see also Cottreau, 1921); Glyptodon ret iculat us, Hop loplwr us ecliazui, Neothoracopliorus cf. the stratigraphy was described by Mortillet (1922), a member of the elevatus, Panoclitus cf. tuberculatus; Megalonychidae, Nothropiis 1903 expedition. Between 1894 and 1915 members of the Echazú tarijensis; Megatheriidae, Megatherium tarijense; Mylodontidae, family of Tarija made an impressive private collection of fossil Glossotheriuni tarijense, Lestodon armat us, Scelidodon tarijensis), mammals from Tarija and surrounding areas @chazÚ, 1905, 1921). Rodentia (Cricetidae, Andinomys cf. edea, Calomys cf. lancha, Some of these fossils were sold to various foreign institutions (see Kunsia fronto, Necfoniyscf. squamipes, Oxymycterus cf. paranensis, above, below), although the major part of the collection, consisting Pltyllotis cf. darwini; Octodontidae, Ctenonzys subassentiens, C. of some 2,630 specimens which remain virtually unstudied, was sent brachyrhinus; Echimyidae, Euryzygpmatomys hoffstetteri; to La Paz in the 1930s. In 1977 part of this collection was returned Myocastoridae, Myocastor perditus; Erethizontidae, Coendou to Tarija and deposited in the Museo de la Universidad de Tarija; it magnus, C. sp.; Caviidae, Cavia sp., Galea cf. musteloides; is planned that the remainder of the collection will be returned to Hydrochoeridae. Hydrockoerus sp., Neoclioerus tarijensis), Tarija in the ilex future (Takai ef al., 1982, 1984). In 1924 and 1927 Carnivora (Canidae, Canis dilus, Cltrysocyon brachyurus, expeditions from the Field Museum of Natural History, Chicago Pseudalopex gymnocercus, Theriodictis tarijensis, Protocyon under the direction of E. S. Riggs made a spectacular collection from troglodytes; Ursidae, Arctodus tarijensis, A. wingei; Procyonidae, Tarija and Padcaya, and purchased some specimens from the Echazú Nasua sp.; Mustelidae, Conepatus suffocans; Felidae, Felis family (Riggs, 1928, 1930); this magnificent collection remains concolor, F. yaguaroundi, Panthera onca, Sniilodon populator), virtually undescribed (Marshall, 1978). During 1978 and 1980 Litopterna (Macraucheniidae, Macrauchenia patachonica), paleontologists from the Research Institute of Evolutionary Biology, Notoungulata (Toxodontidae, cf. platensis), Proboscidea Tokyo made a small collection which was taken to Japan (Takai et (Gomphotheriidae, liyodon, NotiomastÓdon sp., al., 1982, 1984). The most recent work by foreign researchers was Haplontastodon or Stegopiastodon sp.), Perissodactyla (Equidae, carried-out by a team from the University of Florida, Gainesville in Equus insulatus, Hippidion principale, H. bonaerense, 1980 and 1986 (MacFadden, 1981; MacFadden & Azzaroli, 1987; Onohippidiuni devillei; Tapiridae, Tapirus tarijensis), Artiodactyla MacFadden & Wolff, 1981; MacFadden ef al., 1983; Frailey et al., (Tayassuidae, Platygonus tarijensis; Camelidae, Lama cf. onteni, L. 1980). glania, L. provicugna, Palaeolania sp.; Cervidae, Charitoceros A history of the debated age of the Tarija fauna is provided by tarijensis, Hippocamelus sp.). Undescribed specimens of fish and Marshall et al. (1984). Ameghino (1902) and Boule & Thévenin reptiles are reported by Hoffstetter (1963~1,pp. 201-202); frogs (1920) regarded the fauna as Pliocene; Rovereto (1914) assigned it (Bufonidae, Bufo cf. niarinus liorribilis; Leptodactylidae, Ceratoprys to a new land mammal age, the Tarijense; Kraglievich (1930a, 1934) sp.) from Tarija are described by Vergnaud-Grazzini (1968); buds stressed the post-Ensenadan (s.s.) and pre-Lujanian (s.s.) aspect of (Ciconiiformes, Vulturidae, Vultur gryplius = V.pratruus Lönnberg, the fauna which means that it is comparable to the Bonaerense as 1905; see Emslie, 1988) and undescribed specimens of birds are defined in Argentina (see Marshall ef al., 1984, Fig. 1). Palterson & known from Tarija (locality of Turumayu, in Peabody Museum, Yale Pascua1 (1972, p. 249) regarded the fauna as Lujanian, while Webb University) and Padcaya (Villarroel, in Marshall et al., 1984, p. 33; (1974, p. 176) considered it Ensenadan. This debate was resolved, at specimens also in Peabody Museum, Yale University). least in part, by MacFadden et al. (1983) who provided a geochronologic study of the - 250 m-thick Tarija Formation in the LATE PLEISTOCENE-HOLOCENE Tarija basin using magnetostratigraphy and radioisotopic dating. (Lujanian-Recent) They show that this formation spans a time interval between 1.0 and 0.7 Ma, or perhaps younger, making this fauna late Ensenadan by A faana of large body-size mammals of possible early Lujanian current standards. ‘I‘akai et al. (1982, p. 5) report dates of 0.25 - 0.20 age has long been known from the now classic locality of Ulloma Ma on some fossil bones using electron spin resonance technique, (3880 m), situated about 40 km south of Corocoro on the south side but these dates are at odds with the more secure conclusions of of the Rio Desaguadero. The fossils are from the Ulloma Formation MacFadden et al. (1983). Nevertheless, Hoffstetter (1986) cautions which consists of well stratified fine-grained sands rich in calcareous that some fossils from upper levels of the Tarija Formation may be nodules; it is bounded below by an undated tuff, above by a younger, perhaps early Lujanian in age. It is apparent that additional diatomite, and rests unconformably on the folded Totora Group (late detailed stratigraphic studies of the type provided by Oppenheim Miocene). The Ulloma Formation was apparently deposited during (1943), additional magnetostratigraphic and radioisotopic studies as retreat of the glaciation as part of Lake Ballivian, an ancient provided by MacFadden et al. (1983), and more refined systematic southern extension of (Servant & Fontes, 1978; and biostratigraphic studies of the faunas from Tarija, Concepción Hoffstetter, 1986, p. 224; Hoffstetter, in Marshall et al., 1984, p. 35). and Padcaya are needed to establish the absolute and relative ages of The fauna includes: Edentata (Glyptodontidae, Glyptodon sp.; the numerous fossil levels. Megatheriidae, Megatherium cf. americanum, Megatherium

642 FOSDXS Y FACLES DE BOLMA - VOL. I VERTEBRADOS

(Pseudomegatherium) medinae = M. sundti; Mylodontidae, Boidae; , Colubrinae; Viperidae, Crotalinae; Sceli#odon cltìliense Lydekker 1886 = Scelidotheriunt? undetermined turtles), birds (Rheidae, Rhea cf. americana; bolìvianum), Litopterna (Macraucheniidae, Macraucltenia Podicipidae, Podiceps minor, P. cornutris; Plataleidae, Plareleu sp.; ullontensis), Proboscidea (Gomphotheriidae, Cuvieronius ltyodon = Ajaja rosea; Anseridae, Dendrocygna sp., Anas sp.; Rallidae; Mastodon bolivianus), and Perissodactyla (Equidae, Onoltippidium Jacanidae, Jacana sp.; Tinamidae, Cryptiirus lataupa, cf. (Paraltipparion) bolivianuni = Scelidotlterium? compressunt) Rltynclrotus sp.; Columbidae; Falconidae; Cuculidae, Coccyzus sp.; (Sundt, 1900; Philippi, 1893a, b; Pompeki, 1902; Oliver, 1934; Caprimulgidae, Nyctibus sp.; Passeriformes); the mammals include Sefve, 1914, 1915a, b; Douglas, 1914; Kozlowski, 1923; Chiroptera (Molossidae, Euntops perotis), Edentata (Dasypodidae, Casamiquela & Sepúlveda, 1974; Ahlfeld & Branisa, 1960; Servant Chaetopliractus cf. vellerosus or villosus, Zaedyus pichiy , Dasypus & Fontes, 1978; Hoffstetter, 1986; Marshall & Salinas, 1991). A cf. novenicinctus, Eupltractus sexcinctus, Tolypeutes matacus, recently discovered fauna at a locality 4 km west of Carsani is also Propraopus cf. punclatus), Rodentia (Cricetidae, Caloniys sp., probably from the Ulloma Formation, and the taxa include: Edentata Graomys sp., Holochilus brasiliensis, Zygodontomys lasiurus; (Mylodontidae?, Megatheriidae), Notoungulata (Toxodontidae), Octodontidae, Ctenomys sp.; Myocastoridae, Myocastor coypus; Perissodactyla (Equidae), and Rodentia (R. Céspedes, personal Chinchillidae, maximus; Caviidae, Galea cf. communication, 1989). musteloides; Capromyidae; Hydrochoeridae, Hydrochoerus The richest Lujanian fauna in Bolivia is from Ñuapua, between hydrocltaeris), (Leporidae), Carnivora (Canidae, Carandaití and Capiranda. Hermann (see Schiller, 1913, p. 180, n2) Dusicyon griseus), Artiodactyla (Cervidae, Morenelapltus sp.), and first reported mammals from this locality, and Bonarelli (1921, p. a human skull (Hoffstetter, 1968a; MacFadden & Wolff, 1981; 80) later recorded mastodonts and other mammals. Hoffstetter Vergnaud-Grazzini, 1968; Marshall et al., 1984). A carbon-14 date 400 (1968a) published the first study on the stratigraphy, sedimentology of 7200 f yrbp was obtained on the human skull, and another and vertebrate paleontology based on work he did with L. Branisa date of 6600 f 370 yrbp was obtained on unspecified associated between 1962 and 1965. MacFadden & Wolff (1981) made a similar mammal bones (MacFadden & Wolff, 1981; Marshall el al., 1984, p. study based on wdrk done in 1978. 34). Some fossils of extinct megafauna (i.e. Glyptodontidae, The Ñuapua Formation consists of 9 m of sediments which rest Scelidodon, Toxodon, Palaeolama, and mastodonts) were recovered unconforinably on non-fossiliferous gray and red clays of the from the Ñuapua 2 member, but aspects of their preservation suggest Tertiary age Chaco Series. Hoffstetter (1968a) divided the Ñuapua that these specimens were reworked from the Ñuapua 1 member and Formation into three members (from oldest to youngest, Ñuapua 1, are thus not contemporaneous with the rest of the Ñuapua 2 fauna Ñuapua 2, Ñuapua 3) based on color of sediments and faunal (Hoffstetter, 1968a, pp. 832-833; but see MacFadden & Wolff, ,1981 content. MacFadden & Wolff (1981) provide a magnetostratigraphic who did not recognize this fact). Thus, the carbon-14 dates and the study of the Ñuapua Formation and show that all three members are absence of securely associated megafauna suggest that the Ñuapua 2 located within the Brunhes Normal chron which begins at 0.73 Ma fauna is Holocene in age. and are thus Lujanian in age or younger (see below). Hoffstetter (1968a, p. 834) reports the recovery of an ulna of The lower Ñuapua 1 member, which consists of 2-3 m of pink to Toxodon sp. from the basal contact of Ñuapua 3 with the underlying reddish consolidated cinerite, is probably the level from which Ñuapua 2, but not within the Ñuapua 3 tuff which is otherwise mammals were collected by early workers. The fauna (see unfossiliferous and which MacFadden & Wolff (1981) regard as Hoffstetter, 1968a; MacFadden & Wolff, 1981; Berta, 1981, 1985, alluvium. This specimen was also probably reworked 1987, 1988) includes: Reptilia (turtles, Chelonia, Testudinidae, large from Ñuapua 1. sp., a small testudinid? and an undetermined chelid?; Broin, 1991). Edentata (Dasypodidae, Panipatlterium sp.; PLEISTOCENE sensu lato Glyptodontidae, Chlamydotherium sp., Glyptodon cf. reticulatus, (Ensenadan-Lujanian) Panoclitus cf. tuberculatus, Sclerocalyptus cf. ornatus; Megalonychidae, cf. Notlirotlterium sp.; Megatheriidae, There are numerous other scattered reports of Pleistocene Megatherium cf. americanum; Mylodontìdae, Mylodon darnhi), mammals in Bolivia, but the known specimens have been little Rodentia (Hydrochoeridae, Hydrocltoerus cf. hydrocltoeris; studied and for many their age and stratigraphic context have yet to Ctenomyidae; Capromyidae; Cricetidae; Octodontidae), Carnivora be securely established. (Canidae, Protocyon troglodytes, Pseudalopex sp.; Ursidae, On the Altiplano at La Paz and Tambillo (west of La Paz) have Arctodiis sp.; Felidae, Panthera onca, Smìlodon popiflator), been collected remains of mastodonts and (Hoffstetter, Litopterna (Macraucheniidae, Macraucltenia cf. patacltonica), 1986). "Pleistocene fossils" have also been reported at San Andrés Notoungulata (Toxodontidae, Toxodon ensenadensis), Proboscidea and Achiri (Cantón Achiri, i.e. at Challncollo and Huana Jahuira), (Gomphotheriidae, Stegomastodon sp.), Perissodactyla (Equidae, although the exact stratigraphic position and identification of these Equus curvidens, Hippidion sp.), and Artiodactyla (Camelidae, fossils need confirmation (Hoffstetter, 1986, p. 224). Furthermore, Palaeolama sp.; Tayassuidae; Cervidae). The Ñuapua 1 fauna is the reputed Pleistocene fossils at Achiri may, in fact, represent a clearly of Lujanian age. confusion with those from theqlate Miocene Mauri 6 Formation The middle Ñuapua 2 member consists of 1.5 - 1.0 m of reworked (Hoffstetter, 1986, p. 224). A carapace (now in gray tuff. This fich fauna, discovered by L. Branisa in 1955, GEOBOL, La Paz) was collected by Anaya (1988) from the includes: frogs (Bufonidae, Bufo cf. paracnemis; Leptodactylidae, Comunidad Segueri, about 1.5 km east of Tiahuanacu; a possible Leptodactylus cf. ocellatus, Ceratrophrys cf. ornata), reptiles Megatherium sp. was collected in 1985 about 600 in to the (Teiidae, Tupinambis teguixin; Amphisbaenidae, Leposternon sp.; southwest of the ruins of Tiahuanacu (Anaya. personal

643 LARRY G.,MARSHALL & THIERRY SEMPERE

communication, 1989); and L. Branisa (correspondence now in less than 1.0 Ma near the top of the section (Servant et al., 1989). A Peabody Museum, Yale University) records a fossil (Equidae) mastodont has recently been found at the pueblo of Salitre, situated from at or near Tiahuanacu. to the east of Villazon near the Argentine border in southernmost On the western slope of the Cordillera Oriental, about 110 km Bolivia (reported by Ing. Camacho to F. Anaya, personal southeast of La Paz, have been recorded remains of Panthera onca at communication, 1989) and remains of glyptodonts are known from (3600 m) and Macrauclzenia sp. at Puchuni (Hoffstetter, 1986, Mojo (R. CBspedes, personal communication, 1989). All these p. 226). Kozlowski (1923) records discovery by miners of fossils come from sediments related to the San Juan del Oro "Pleistocene mammals" from Calacoto, Cuesta de Chacarilla and planation surface. Paria; while Lavenu (1984) records camelids (Camelidae) and L. In the Amazonian region at Fortin Madidi on the Río Madidi and Branisa (unpublished notes) records remains of artiodactyls and a at Cara Cara on the Río Maniqui (in the region called Yucumo- horse from Culluri near Corque, southwest of Oruro. Mastodonts are Maracas near San Borja) in the Río Beni basin have been found reported from northwest of Oruro, from Paria (16 km northeast of remains of Toxodontidae in the former and sp. in the Oruro), and from Pampa Aullagas on the southwest bank of Lake lalter (Hoffstetter, 1968a; Hoffstetter, in Marshall el al., 1984, p. 36; PoopÓ opposite Créqui-Montfort Island (Hoffstetter, 1986, p. 224). Anaya, personal communication, 1989). Further to the southeast, Ahlfeld & Branisa (1960, p. 165) report remains of a mastodont tusk d'Orbigny (1842, p. 205) recovered a collection of bones of large and a lower jaw of Hippidion from "las minas de estaño de mammals "en las barrancas cerca de la confluencia de los ríos Piray Carguaycollo, en la Cordillera de los Frailes" (4100 m); the y Grande" which were lost during the same expedition "en un vuelco identification of the horse needs confirmation (Hoffstetter, 1986, p. de la canoa" (Ahlfeld & Branisa, 1960). In the Río Piray basin at 226) and it is possibly referable to Onohippidiuai. Santa Rosa north-northwest of Santa Cruz and to the southwest of Near Anzaldo on the east slope of the Cordillera Oriental, about Santa Cruz, Hoffstetter (1968a) recovered mastodont teeth in 1960. 55 kin southeast of Cochabamba, Montañ0 (1968) recovered a Remains of mastodonts have also been recovered by L. Branisa Pleistocene mammal fauna from sands, silts and gravels in an upper (unpublished) from Quebrada Chorrillos, about 12 km northeast of level at Quebrada Tijascka, 3 km southwest of Anzaldo. This fauna Charagua, and Trujillo (1984) reports Cuvieronius andiuni from the is from a level above the Uquian age gravel unit with Prodaedicurus Valle de Carohuaycho, 28 km north-northeast of Camiri and 30 km (see above), and includes: Edentata (Glyptodontidae, Glyptodon sp.; west of Charagua. Megatheriidae, Megatlteriuni sp.), Proboscidea (Gomphotheriidae, Cuvieroniris sp.), and Artiodactyla (Camelidae, Lama sp.) EVOLUTION OF THE BOLIVIAN LAND MAMMAL (Hoffstetter, in Marshall et al., 1984, p. 36; Hoffstetter, 1986; p. FAUNA 223). I Ahlfeld & Branisa (1960, p. 167) record fragment5 of glyptodonts There are four interrelated phenomena that were of profound from along the Río Rocha near Sacaba about 7 km east-southeast of importance in influencing and/or determining the taxonomic Cochabamba, and R. Céspedes and R. Suarez (personal composition of late Cenozoic land mammal faunas in Bolivia: communication, 1989) recovered remains of glyptodonts and Andean uplift and related climatic changes, glaciations, the Great camelids in 1986 from along the Río Loro Mayu, just south of the American Faunal Interchange, and megafaunal . Río Rocha, about 10 km east-southeast of Cochabamba. Reports of mastodonts and glyptodonts from near Sacaba and in the 1. Andean uplift and relaed climatic changes. The main climatic Cochabamba-Sacaba basin (2600-2800 m), and of mastodonts in the effect of the central Andean uplift in Bolivia was the creation of: (a) Padilla basin (2120 m) (which has a tuff dated at 3.36 k 0.3 Ma) are a rain shadow that prevented the northeast-proceeding moisture-rich given by Hoffstetter (1986, p. 226). A femur of an edentate is also winds to reach the coastal areas of southern Peru and northern Chile, known from near Toco (R. Céspedes, personal communication, and (b) dry "high"-altitude climates on the Altiplano and adjacent 1989). cordilleras. Near Sucre in the small Sucre basin (2800 in) "esti ubicada una As demonstrated by geomorphological studies in southernmost meseta que constituye la línea divisoria de Ias aguas entre los ríos Peru, uplift began - 20 Ma, but considerably decreased in this area Pilcomayo y Grande" where remains of Glypfodon sp. have been by - 8 Ma (Tosdal ef al., 1984). According to Benjamin et al. found (Ahlfeld & Branisa, 1960, p. 167; Hoffstetter, 1986, p. 226). (1987), uplift rates of the Bolivian Cordillera Real were about 0.1 - This meseta is part of the San Juan.de1 Oro planation surface (see 0.2 mm/yr between 40 and 20 Ma, and increased significantly above) and the fossils are from its associated deposits. between 15 and 10 Ma to reach values perhaps as high as 0.7 mm/yr In the Betanzos basin (3390 in) between Potosí and Sucre have by 3 Ma. Hyperarid conditions, due to a permanent rain shadow, been found remnids of Edentata (Glyptodontidae, Sclerocalyptinae, became established in northern Chile (24" S) in middle Miocene Panochrltus sp.), Proboscidea (Gomphotheriidae, Cuvieronius time (Alpers & Brimhall, 1988). Middle Miocene climatic Ityodon = Mastodon andinun!), Perissodactyla (Equidae, gen. et sp. dessication in northern Chile and southern Peru was accentuated by indet.), and Artiodactyla, (Cervidae; Camelidae) (Pick, 1944; oceanic cooling related to the contemporaneous development of the Hoffstetter, 1986, p. 226). About 4 km to the south of Betanzos have Antarctic ice cap (Alpers & Brimhall, 1988), and this must also have been collected Mylodontidae?, Glyptodontidae, Gomphotheriidae liad some influence on Altiplano climates. Although it is difficult to and Equidae which are now in the Universidad Tomis Frías, Potosí calculate precisely the paleoaltitudes of the central Andes at a given (Anaya, personal communication, 1989). At Khoña-Paya near time, it seems that the mountain belt had attained at least half its Betanzos, were collected remains of Equidae from between a tuff present elevation prior to about 15 Ma (Alpers & Brimhall, 1988). It dated at 1.9 Ma in the base of the section and another tuff dated at is likely that in Bolivia, Andean uplift was accelerated by the

644 ~-- -

POSILES Y FACIES DE BOLIVIA - VOL. I VERTEBRADOS tectonic load produced by the first major tectonic crisis (see above) Bolivian Altiplano. During times of glacial retreat, as today, these starting at about 27 Ma. However, there may have been more than lakes contracted or disappeared. The overall result is that the one uplift mechanism subsequent to 45 Ma. climates on the Altiplano were greatly altered during times of glacial Thus, the significant elevation of the central Andes during at least advance and retreat. the past 20 Myr must have influenced the land mammal fauna in two Climates to the east of the Cordillera Oriental were likewise ways. First, the rising central Andes came to serve as a barrier to‘the effected by glacial advances and retreats. The tropical and moisture-rich northeastem winds, resulting in the formation of a rain subtropical lowlands of eastern Bolivia experienced cold-dry shadow effect along its western flank. Second, ecological changes climates during glacial advance, resulting in contraction of tropical resulted from the uplift, and true desert, high montane and extensive forest biomes and expansion of savanna biomes. During times of grassland habitats at lower elevations came into existence during this glacial retreat, as today, these areas had warm-wet climates with time. These ecological changes provided new evolutionary extensive tropical forest biomes and restricted savanna biomes al., opportunities for taxa which could adapt to colder high-elevation (Arroyo el 1988). These climatic changes affected the habitats, and caused other taxa to abandon these emerging distribution of living in these habitats, and the fauna which environments in favor of more hospitable warmer environments at occurs at any one locality during times of glacial advance may be lower elevations. For example, Mesotheriidae disappeared from the different from that which occurs in the same locality during times of Altiplano at the end of Huayquerian time, and Hydrochoeridae and glacial retreat. Unfortunately, the chronology of glacial advances and Pampatheriinae disappeared from high-elevation faunas at the end of retreats is not well documented in Bolivia, and the identity of glacial Montehermosan time (Hoffstetter, 1986). faunas versus interglacial faunas has yet to be firmly established. The net result of Andean uplift is that it produced markedly Nevertheless, this feature must be kept in mind when attempting to diverse geographic regions in Bolivia and many of the differences explain differences in taxonomic composition of mammal faunas, as seen particularly in Pleistocene faunas can be attributed to elevation they may be due to differences in ecology, altitude, and/or time. and hence ecologies. As examples, some Pleistocene taxa apparently thrived at elevations of 3800-4000 m (i.e. Megatherium, Scelidodon, 3. Tlie Great American Faunal Interchange. Prior to about 2.5 Ma, Macraucltenia, Cuvieronius, Onohipyidium, Lama, Pantliera, North and South America were separated by a narrow seaway called Glypfodon) and occur at low elevations as well; other taxa (i.e. the Bolívar Trough which connected the Caribbean and Pacific Palaeolama, Equus) are unknown above 3300 m; while others are Oceans across what is today northwestem and southern either unknown or extremely rare above 2000 m (i.e. Toxodontinae, Panama. About 2.5 Ma this seaway began to disappear as a result of Pampatheriinae, Dasypodinae, Hydrochoeridae, Erethizontidae, tectonic uplift of southem central America and/or a glacioeustatic , Agutidae, Dinomyidae, Echimyidae, Tapiridae, drop in sea level (Webb & Barnosky, 1989; Keller et al., 1989). The Tayassuidae, most Cervidae, Haplomastodon, Chrysocyon, Panamanian Land Bridge came progressively into existence, uniting ) (Hoffstetter, 1986). North and South America. The Pacific-Caribbean marine gateway finally closed by 1.8 Ma (Keller et al., 1989). The land connection 2. Glaciations. Four major Pleistocene glaciations in Bolivia are permitted the reciprocal intermingling of the long-isolated biotas’of recognized by Servant & Fontes (1978) (from oldest to youngest: I these continents, an event of spectacular importance known as the Calvario, II Kaluyo, III Sorata, IV Choqueyapu 1 and 2). A tuff Great American Faunal Interchange (Marshall, 1988). (Purapurani tuff = toba Sopari of Thouveny & Servant, 1989, p. 332) The land mammals which walked across the newly emergent land interbedded within the glacial and interglacial deposits that overlie bridge froth North to South America included members of the the Calvario Drift, has yielded a K-Ar date of 1.6 Ma (Lavenu et al., Tayassuidae and Mustelidae which are first known from rocks of 1989). A fifth glaciation called the Patapatani (Hoffstetter, 1986) is Chapadmalalan age in Argentina (Marshall, 1985). However, it is in reported in Pliocene sediments from below the Chijini tuff (which rocks of Uquian through Lujanian age that we begin to see in South has a mean age of 2.8 Ma; Lavenu et al., 1989; Marshall ef al., 1992) America a major contingent of these North American immigrants of the La Paz Formation. which include: Proboscidea (Gomphotheriidae), Lagomorpha During times of glacial advance, ice sheets extended to about 1000 (Leporidae), Carnivora (some Procyonidae, more Mustelidae, m below their present day limits (Hoffstetter, 1986). During glacial Canidae, Felidae, Ursidae), Perissodactyla (Equidae, Tapiridae), retreat, large lakes formed on the Altiplano (see Stoertz & Ericksen, Artiodactyla (more Tayassuidae, Camelidae, Cervidae) and 1974; Lavenu et al., 1984) and three of these have been named: Lake eventually man (Homo sapiens) about 12 ka (Marshall, 1985,1988; Bnllivifíii was n southern expansion of Lake Titicaca (which Marshall ef al., 1984). The appearance of these taxa in South presently covers 8,400 km’) in the northern Altiplano and covered an America is thus an important immigrant datum event for recognizing faunas of Uquian through Lujanian age. area of 12,600 km’; Lake Minchín and occupied the same area in the central and southern Altiplano, including present- 4. Megafaunal Exfinciions. The end of Pleistocene time in Bolivia, day Lake PoopÓ, Salar de Coipasa and. , and had as elsewhere in South America, is marked by the of most maximum extensions respectively at - 30,000 yrbp and 13,000- large body size mammals. The vast majority of these now extinct 11,OOO yrbp (Servant & Fontes, 1978). Two lake levels called Lake mammals had body weights in excess of 100 kg, and because of their Cabana and Lake Mataro, which occurred prior to the Sorata large size are commonly referred to as megafauna. The peak of these glaciation, have also been identified (Lavenu et al., 1984). Campbell extinctions occur between 12-10 ka (Marshall et al., 1984). In ef al. (1985) have suggested that glacial lakes which formed during Bolivian faunas these now extinct megafauna include all members of the last glacial advance may have been united to form one large lake the families Glyptodontidae, Megatheriidae, Mylodontidae, which Campbell (1989) named Lake Carrasco, that covered an Megalonychidae, Macraucheniidae, Gomphotheriidae and Equidae; excess of 150,000 km2, or nearly all of the surface area of the and some Dasypodidae (Pampafherium),Camelidae (Palaeolama),

645 LARRY G. MARSHALL &THIERRY SEMPERE

Cervidae {Morenelaplius) and Felidae {Sniilodon). As demonstrated AVALO, D. de. 1602. Fósiles óseos de Tarija (Bolivia). Miscellanea at Ñuapua, these extinctions occurred prior to 7200 k 400 yrbp as Austral, Lima 1602, Colloquia XXXIII: 147. indicated by the oldest carbon-14 date on the Ñuapua 2 fauna which BABY, P., T. SEMPERE, J. OLLER, L. BARRIOS, G. HERAIL & includes only extant taxa. The exact cause of these extinctions is R. MAROCCO. 1990. Un bassin en compression d'âge oligo- unknown, although they have been attributed to excessive hunting miocène dans le sud de I'Altiplano bolivien. Comptes-Rendus 1 (overkill) by early man, climatic change (a sharp climatic drying 1' Académie des Sciences, Paris, série II, 311: 341-347. occurred in Bolivia and adjacent regions starting at -10,000 - 9,000 BENJAMIN, M. T., N. M. JOHNSON & C. W. NAESER. 1987. yrbp in the Andes and - 8000 yrbp in the lowlands; Servant et al., Recent rapid uplift in the Bolivian Andes: evidence from fission- 1989b; Absy et al., 1991; M. Servant, personal communication), and track dating. Geology, 15: 680-683. a combination of the two (Hoffstetter, 1986). BERGGREN, W. A., D. V. KENT, J. J. FLYNN & J. A. VAN COUVERING. 1985. Cenozoic geochronology. Geological Society ACKNOWLEDGMENTS of America Bulletin, 96: 1407-1418. BERRY, E. W.1922. Late Tertiary plants from Jancocota, Aspects of this study were supported by grants from the National Bolivia. The Johns Hopkins University Studies in Geology, Geographic Socicly (2467-82, 2908-84, 3381-86); the Gordon 4: 205-220. Barbour Fund, Department of Geological and Geophysical Sciences, BERTA, A. 1981. Evolution of large canids in South knerica Anais II Princeton University; and the National Science Foundation (EAR- CongresoLatinoamericano de Paleontología, Porto Alegre, 2: 835-845. 8804423). Stratigraphic work was funded by the Institut Français de BERTA, A. 1985:The status of Sniilodori in North and South Recherche Scientifique pour le Développement en Coopération America. Natural History Museum of Los Angeles County, (Orstom) and realized in collaboration with field geologists of Contributions in Science, 370: 1-15. Yacimientos Petrolíferos Fiscales Bolivianos (YPFB). For help on BERTA, A. 1987. Origin, diversification, and zoogeography of the the geology and/or identification of specimens we thank D. Anaya, South American Canidae. Fieldiana: Zoology, n.s. 39: 455471. A. Berta, R. Céspedes, M. Gayet, A. Lavenu, R. Hoffstetter, R. BERTA A. 1988. Quaternary evolution and biogeography of Marocco, J. Oller, J. Pacheco, R. Pascual, G. Sanjinés, R. Suirez & the large South American Canidae (Mammalia: Carnivora). C. Villarroel. University of California Publications in Geological Sciences, 132: 1-149. REFERENCES BLAKE, C. C. 1861. On the discovery of Mucrauchenia in Bolivia. 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La Paz, 83 p. I RADELLI, L. 1964. Ensayo de reconstrucción de la MORTILLET, A. de. 1922. Le gisement fossilif5re de Tarija cronoestratigrafía y de la paleogeografía del Altiplano y de las (Bolivie). Compte-Rendu, Association Française pour l'Avancement cordilleras orientales de Bolivia después del Paleozoico superior. des Sciences, Paris, 45: 911-919. Rivista Italiana di Paleontologia, Milano, 70 (4): 833-868. NAESER, C.W., E. H. McKEE, N. M. JOHNSON & B. J. MAC REDWOOD, S. D. 1987. The Soledad Caldera, Bolivia: a Miocene FADDEN. 1987. Confirmation of a late Oligocene-early Miocene caldera with associated epithermal Au-Ag-Cu-Pb-Zn mineralization. age of the Deseadan Salla Beds of Bolivia. Journal of Geology, 95: Geological Society of America Bulletin, 99: 395404. 825-828. REDWOOD, S. D. & R. M. MacINTYRE. 1989, K/Ar dating of NORDENSKIOLD, E. 1902. ober die Säugetierfossilien in Miocene magmatism and related epithermal mineralization of the Tarijathal, Südamerika. Bulletin of the Geological Institution of the northeastern Altiplano of Bolivia. Economic Geology, 84: 618-630. University of Uppsala, 5( 10): 261-266 (Spanish translation: Sobre REVILLIOD, P. 1922a. Note préliminaire sur un Mastodon de los mamíferos fosiles del Valle de Tarija. Anales de la Sociedad Bolivie. Actes de la Société Helvétique des Sciences Naturelles, Científica Argentina, Buenos Aires, 55: 255-262). Aarau, 103e Session annuelle, Berne, 1: 226-227. NORDENSKIOLD, E. 1903. ober die Säugetierfossiliell des REVILLIOD, P. 1922b. Sur un crâne de mastodonte de Bolivie. Tarijatales, Südamerika. I Mastodon andiuni Cw. Kungliga Svenska Eclogae Geologiae Helvetiae, Lausanne, 17 (3): 357-358. Vetenskaps Akademiens Handlingar, Stockholm, 37 (4): 1-30. REVILLIOD, P. 1931. Sur un Mastodon de Bolivie. Mémoires de la NORDENSKIOLD, E. 1905. Exploration scientifique au Pérou et en Société Paléontologique Suisse, Bâle, 51: 1-21.

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~ 1927. Field Museum of Natural History, Geology Memoir, area. International Symposium on Andean Geodynamics, Grenoble, l(1): 7-34. p. 167-170. ROEDER, D. 1988. Andean-age structure of eastern Cordillera SEMPERE, T., G. HERAIL, J. OLLER & G. BONHOMME. 1990b. (Province of La Paz, Bolivia). Tectonics, 7: 23-29. Late Oligocene-early Miocene major tectonic crisis and related ROSENBERGER, A. L. 1981. A mandible of Brunisella boliviana basins in Bolivia. Geology, 18: 946-949. (Platyrrhini, Primates) from the Oligocene of South America. SEMPERE, T., G. HERAIL, J. OLLER & P. BABY. 1990~. International Journal of Primatology, 2 (1): 1-7. Nacimiento de los Andes bolivianos en el Oligoceno superior. IX ROVERETO, C. 1914. Los estratos araucanos y sus fósiles. Anales Congreso Geológico Boliviano, p.1, Cochabamba. del Museo Nacional de Historia Natural de Buenos Aires, 25: 1-250. SEMPERE, T., E. VARGAS & G. HERAIL. 1990d. Las superficies RUSSO, A. 1959. 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