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THE EOCENE TO PLEISTOCENE VERTEBRATES

OF BOLIVIA 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, Santa Cruz de la Sierra, Bolivia. Present address: Centre de Gologie Gnrale et Minibre,
Ecole des Mines, 35 rue Saint Honor& 77305 Fontainebleau, France

INTRODUCTION

'I
I

The record of Cenozoic fossil vertebrates in Bolivia is extremely


good. Compared with other countries in South America, Bolivia is
second only to Argentina in the number of known localities and in
the wealth of taxa.
Of the different vertebrate groups, the mammals are by far the
most abundant and best known. In fact, the record of mammal
evolution in South America is so complete that these fossils are used
by geologists and paleontologists to subdivide geologic time. The
occurrence of unique associations of taxa that are inferred to have
existed during a restricted interval of time has resulted in the
recognition of discrete chronostratigraphic units called Land
Mammal Ages. These ages were established on the basis of
knowledge of stage of evolution of the taxa, on their time of first
and/or last appearahce in the fossil record, and on changing faunal
associations through time. The sequence of South American Land
Mammal Ages (SALMA) has been based largely on knowledge of
the exceptionally rich and complete record in Argentina (Marshall,
1985). Detailed systematic studies of taxa along with geochronologic
studies and magnetostratigraphy of the fossiliferous rock sequence
and/or radioisotopic (Ar/Ar, K-Ar) dating of associated volcanic
rock units (i.e. basalts, tuffs, ignimbrites) permit geologists and
paleontologists to correlate Bolivian mammal faunas with the
Argentine sequence atid to make distinctions between ecological and
temporal relationships of local faunas and faunal assemblages. The
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.
We provide a review of the Eocene to Pleistocene land mammal
record of Bolivia within its stratigraphic and tectonic context. The
use and chronology of South American Land Mammal Ages follows
Marshall (1985) as supplemented by Marshall, Drake e al. (1986),
and Marshall, Cifelli ef al. (1986). Recently, Tonni et al. (1987)
recognized a new ,Pleistocene Land.Mamma1 Age between the
Ensenadan and Lujanian which they called Centinelan. However, the
validity and utility of this new age has yet to be f m l y established in
Argentina, and unti this is done it is futile to attempt to recognize it
in Bolivia. In addition, Marshall & Salinas (1990) have shown that

the type fauna of the Friasiali Land Mammal Age (conventionally


middle Miocene) in southern Chile is temporally equivalent to the
Santacrucian Land Mammal Age. They thus use Colloncuran for the
land mammal age between Santacrucian and Chasicoan. For all
practical purposes, Friasian of previous workers is equivalent to
Colloncuran as used in this study.
This paper represents an expansion and updating of the Bolivian
land mammal record as provided by Robert Hoffstetter (in Marshall
el al. 1983, 1984). As documented below, the highlights of this
record include: the taxonomically richest and best studied faunas of
late Oligocene-early Miocene (Deseadan) and early Pleistocene
(Ensenadan) age in all o[ South America; and the exceptionally rich
record of late Miocene (Huayquerian) and early to middle Pliocene
(Montehemosan) age faunas from the northern Altiplano.
The Altiplano, including the Puna in adjacent Argentina and Chile,
is a high-altitude plateau which covers southwest Bolivia, southeast
Peru, northeast Chile and northwest Argentina. It is about 200 km
wide and about 1500 km long, and some 300,000 km2 at 3,6004,000 m (mean 3650 m; Isacks, 1988) of altitude. In Bolivia, the
Altiplano covers about 170,000 km2 and is situated between the
Cordillera Occidental and Cordillera Oriental.
The following abbreviations are used: GEOBOL, Servicio
Geolgico de Bolivia; ka, thousands of years ago, a point in time; kg,
kilograms; km, kilometers; m, meters; Ma, megaannum or millions
of years ago, a point in time; M y , millions of years, a duration of
time; yrbp, years before present.

GEOLOGICAL EVOLUTION OF BOLIVIA: EOCENE TO


PLEISTOCENE
The reader must first be aware that the geologic evolution of the
central Andes is currently being re-evaluated, and that traditional
models are being challenged. The latter envision that the central
Andes were subjected during most of Cenozoic time to a tectonic
regime characterized by long tensional periods (during which the
sedimentary basins formed), short synchronous compressional
pulses, and generally high-angle faulting (Mgard, 1978; Dalmayrac
el al., 1980; Martinez, 1980; Lavenu & Marocco, 1984; Mgard et

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LARRY G . MARSHALL & THIERRY SEMPERE

al. 1984; Lavenu, 1986; Sbrier et al., 1988). In contrast, new data

and new interpretations favor structural models dominated by


progressive crustal shortening, thrust propagation, and foreland basin
evolution (Jordan et al., 1983; Jordan & Alonso, 1987; Isacks, 1988;
Roeder, 1988; Sempere et al., 1988, 1989, 1990a, b; Baby et al.,
1990; Sheffels, 1990).
In Bolivia, the two models of Andean tectonics are sometimes at
odds with each other. For instance, the thick Cenozoic clastic strata
of the Altiplano were traditionally thought to have been deposited
during long epochs of extension with short interruptions by
compressive pulses. It is now believed that their deposition took
place mostly in intermontane foreland basins (i.e. in permanently
compressional settings; Baby et al., 1990; Sempere et al., 1990a, b,
1991). Predominance of compressive conditions in the central
Andean area since late Cretaceous time was first suggested i n a
general way by Marocco ef al. (1987).
The angular unconformities observed on the Altiplano and
elsewhere in the Bolivian Andes have traditionally been interpreted
as materializing the mentioned "compressional pulses". However,
this is not the only possible interpretation for these angular
unconformities (Sempere, 1991). Moreover, although details of some
important field observations relative to deformations in the Altiplano
have been well described, in particular in the works of Martinez
(1980), Lavenu (1986) and Lavenu et al. (1989), several independent
new chronologidal data make untenable the traditional timing of
deformation (Sempere et al., 1990b).
Therefore, the Calidity of the six "compressional tectonic pulses"
or "phases", as listed by Sbrier et al. (1988), may be questioned.
Intuitively, it seems unlikely that the enormous central Andes, which
today are second only to the Himalayas in size and height of the
mountain belt, crustal thickness, etc., developed by a small number
of short "compressional pulses".
In this paper we adopt the tectonic model and stratigraphy
currently being worked-out by the Orstom-YPFB research program
on the geology of Bolivia (Sempere et al., 1989, 1990a, b, c;
Sempere, 1990), which we summarize below. A map showing the
tectonostratigraphic domains presently recognized i n Bolivia
(Sempere ef al., 1988, 1990a). and the location of the main
sedimentary basins of late Oligocene-Pleistocene age are shown in
Figure 1.
The geological history of Bolivia since the Eocene consists of two
tectosediinentary periods, respectively spanning the Eocene-early
Oligocene and late Oligocene-Present intervals (Sempere et al.,
1989, 1990a; Sempere, 1990).
During the Eocene-early Oligocene time span, the paleo-Andes
were located west of the Bolivian territory, and their associated
continental external foreland b a s h , resembling the present-day
Chaco plain, occupied what is today tlie Altiplano and most of tlie
Cordillera Oriental. At 27 Ma, a major tectonic crisis began to
develop, in conjunction with important changes in plate convergence
velocity (Pardo-Casas & Molnar, 1987), and lasted until 19 Ma
(Sempere et al., 1990b, c). An important phase of magmatic activity
developed during the same interval. It must be stressed that the
Bolivian Andes started to exist as a mountain belt, albeit of limited
altitude, only at that time. Concurrently, the Andean external
foreland basin rapidly shifled eastwards, reaching the SubandeanLlanura domain, while several intermontane basins formed in the
Andean domain, including the large ones on the Altiplano.

After a time of relative quiescence (- 19-17 Ma), magmatism


slowly resumed a t 17 Ma (Grant et al., 1979; Redwood &
MacIntyre, 1989) and tectonism at 14 M a (Sempere, 1990). and
considerably increased at 11-10 M a with the onset of the second
major tectonic crisis (Sempere et al., 1989). This crisis lasted until
5 Ma (Sempere et al., 199Oa), but tectonic and magmatic activities
have lingered on to the present. There is apparently little relation
between the onset of this second tectonic crisis and plate
convergence history.
Each tectonic crisis was followed by the development of an
extensive planation surface, respectively the Chayanta surface in the
middle Miocene and the San Juan del Oro surface during the late
Miocene-early Pleistocene interval (Servant et al., 1989; Sempere et
al., 199Od).

STRATIGRAPHIC FRAMEWORK
The Cenozoic stratigraphy of Bolivia strongly relies on knowledge
of the basins where the strata were deposited. As already mentioned,
the Eocene-early Oligocene area of sedimentation was a large
foreland basin, and its paleo-landscape was a wide alluvial plain that
probably resembled the present-day Chaco-Beni lowlands. In
contrast, the late Oligocene-Pleistocene period was characterized by
numerous separate intermontane basins, many of small size, that
developed in deformed areas, and by the large Subandean-Llanura
external fxeland basin of the Chaco-Beni lowlands.
The lower boundary of the Eocene-early Oligocene period
coincides with the upper boundary of the Kimmeridgian-Paleocene
Puca Group (Sempere el al., 1988; Jaillard & Sempere, 1989;
Sempere, 1990). This locally angular unconformity (Marocco et al.,
1987) marks the initiation of the foreland basin which developed, in
what is today the Bolivian Altiplano and.Cordillera Oriental, during
Eocene-early Oligocene time (see above), and very likely resulted
from a deep modification of the geodynamics of the Andean margin
at that time. Although approximately located at the PaleoceneEocene boundary (Cirbin et al., 1986; i.e.- 53 Ma according to
Cowie & Bassett, 1989, and Odin, 1989; Fig. 2), this tectonic
change might be related to the plate rearrangement which occurred
in the Pacific area at anomaly 24N (Lonsdale, 1988), i.e. in the late
early Ypresian (erggren et al., 1985; Cavelier & Pomerol, 1986;
Haq ef af.,1987) at 5 1-50 Ma (according to updated information in
Cowie & Bassett, 1989, and Odin, 1989). Such an age for this major
change in Andean geodynamic conditions nearly coincides with the
50-49 Ma age for termination of emplacement of the coastal
batholith of Peru (Soler, 1987), which also reflects a niajor change in
modalities of subduction.
The boundary between these two tectosedimentary periods is 27
Ma (Sempere ef al., 1990b) which coincides with the early
Oligocene-late Oligocene boundary of Cowie & Bassett (1989). The
tectonic upheaval which started a't ttiat time resulted in development
of numerous small to mediuni-size basins in the area of the Altiplano
and, especially, Cordillera Oriental.
Within the Andean domain during both major tectonic crises,
thrust deformations developed more particularly in certain areas and
along specific structures, and intermontane foreland basins formed in
relation to them. Examples are the northern Altiplano basin, foreland
of the Huarina fold-thrust belt and Coniri thrust system; the Ro
Mulatos basin, foreland of Uie Sevaruyo-Chita fold-thrust belt; the

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. FOSILES Y FACIES DE BOLMA - VOL. I VERTEBRADOS

Rolvar-MondragBn basin, foreland of lhe Main Altiplanic thrust; the


Lpez basin, foreland of the Khenayani fault system; etc.... (Fig. 1).
Other small basins can be considered as piggyback basins,
completely or nearly isolated from others due to reliefs developed by
thrusting. Examples are the basins of Salla-Luribay, Lacayani,
Tarabuco, probably Muyu Huasi, Tipuani (Fornari et al., 1987),
Padilla, Tarija and San Isidro. The San Juan del Oro planation
surface and its related deposits originated by the action of a
longitudinal fluviatile system trapped within the Cordillera Oriental
(Sempere et al., 1990d), which can be considered as a giant
piggyback basin.
Other basins formed in relation to wrench-faults. Examples are the
Tupiza and Ro Khuchu basins, and the basin system of the
Cochabamba "valles".
Present knowledge of the Eocene-Pleistocene chronostratigraphy
of Bolivia, including some unpublished data, has been summarized
by Sempere & Oller (1987) and Sempere (1990), and is shown in
Figures 2 and 3. Partly due to the numerous basins or sub-basins,
multiplication of formational names during the 1960's has
unnecessarily complicated the stratigraphic nomenclature. Many
names of local value have therefore been eliminated from our
stratigraphic charts. Some of these are mentioned in the text when
used in the cited literature, and their preferred equivalent is given.
Of the two tectosedimentary periods defined above, the second
one, which spans the late Oligocene-Pleistocene interval is by far the
richest in fossil vertebrates, as documented below.
VERTEBRATE FAUNAS AND LOCALITIES
EOCENE
(Casamayoran-Mustersan)
A probable Casamayoran age fauna reported by Sig6 et al. (1984)
includes remains of actinopterygian fish (Characiformes,
Serrasalmidae, Myleinae; Siluriformes), indeterminate turtles (Broin,
1991), crocodile teeth, and a single upper premolar of a primitive
notoungulate mammal close to Camargomendesia (sensu Cifelli,
1983) from a locality they designated Vela Pachita, about 57 km
southeast of Challapata, in what they considered to be the Santa
Luca Formation. However, the correct name of this locality is Wila
Apacheta and the fossils are from the basal Cayara Formation which
rests unconformably on the El Moliio and/or Santa Luca formations
(Marocco et al., 1987); the age of this unconformity shouldsbe 53 or
51-50 Ma (see above).
Also from the Cayara Formation at La Cabaa on the
Cochabamba-Oruro road, about 25 km southwest of Cochabamba,
were recovered remains of fish and reptiles (crocodiles, turtles) from
just below a level with abundant plants; turtles were observed in the
upper levels of the Cayara Formation at Tiupampa, about 95 km
southeast of Cochabamba; and indeterminate fish and turtles wcre
recovered Iiom Ulis formation about 5 kin north-northeastof Chita.
A dubious record of a Casamayoran age locality was given in an
unpublished report by E. Ortega in 1968. This report was seen by
Hoffstetter in GEOBOL, La Paz, although attempts to relocate it by
one of us (L.G.M.) in 1989 were unsuccessful. According to
Hoffstetter (in Marshall et al., 1983, p. 41), Ortega recorded remains
of mammals, including a fragment of a jaw of a Notoungulata
(Interatheriidae, Nofpithecus sp.), from the "areniscas rojas del

Anticlinal de San Andr6s" (=San Andres Formation of Evemden el


al., 1966 which has a dubious associated K-Ar age of 38 Ma on
"glauconite") to the west of San Andres de Machaca on the Altiplano
just south of Lago Titicaca. The fossils reputedly came from a unit
equivalent to a part of the Mauri Formation (s.[.; YF'FB, unpublished
data) and is thus late Oligocene-Miocene in age (Lavenu et al.,
1989), though Eocene age sediments have been traditionally mapped
in this area based on the one age obtained on "glauconite" (Martinez,
1980). Glauconite is widely considered to be a marine mineral,
whereas only continental strata have been deposited in the Altiplano
since the Paleocene. Hence, the age determination by Evemden et al.
is dubious because the dated clay mineral is probably not authigenic.
Unfortunately, these fossils were never described, Ortega (1970)
made no mention of their existence in his review of Bolivian fossil
faunas, and they cannot be located in the GEOBOL collections.
LATE OLIGOCENE-EARLY MIOCENE
(Deseadan)
The most spectacular land mammal fauna of Deseadan age in all
of South America is from the Salla-Luribay basin, about 95 km
southeast of La Paz in the Cordillera Oriental (Hoffstetter, 1968b).
The principal fossil-bearing unit lias been called the Estratos de Salla
(Evemden et al., 1966, 1977; Villarroel & Marshall, 1982). Strates
o u Couches de Salla (Hoffstetter, 1976a) and Salla Beds
(MacFadden et al., 1985) which consist of 540 m of well-indurated
red-brown, pink-tan and'whitish claystones and siltstones deposited
primarily in a fluviatile environment. The Estratos de Salla represent
a unit partly equivalent to the Coniri Formation (Hoffstetter, 1968b;
Sempere et al., 1990b).
The geology and fossil localities are discussed by Hoffstetter
(1968b, 1976a), Villarroel & Marshall (1982) and MacFadden et al.
(1985). The latter provide a magnetostratigraphic and radioisotopic
study of the Salla Beds and conclude that they span a time interval
from about 28.5 - 24.0 Ma, making this the best dated rock unit of
Cenozoic age in Bolivia. MacFadden et al. (1985) also demonstrate
that the fission-track dates of 54.0 k 2.6 and 52.0 It 2.1 Ma rehrted
by Hayashida et al. (1984) for the Salla Beds are too old and
apparently represent dates obtained on detrital (reworked) zircon
grains. Naeser et al. (1987) published three fission-track (34.5 f 4.0,
24.2 f 3.6, 23.5 f 2.2 Ma) and four K-Ar(biotite- 28.0 f 0.9, 27.2
rt 0.9, 27.9 f 0.9, 25.1 f 0.7 Ma) dates from four ash levels in the
lower part of the Salla Beds and conclude that the principal fossil
horizons range from 27-24 Ma, while the complete section ranges
from about 28-22 Ma. These dates are consistent with those of
Hayashida & Danhara (1985) who report four fission-track dates
(27.2f1.6,25.0k1.5,26.1f 1.9,24.0flSMa)onzirconsf?oman
unspecified level(s) in the Salla Beds. The geochronology of the
Salla Beds is summarized by MacFadden (1990b) and McRae
(1990). Sempere ef al. (1990b) reinterpret the geochronologic data
presented by MacFadden et al. (1985) and suggest that their dated
section may be 3.0-4.5 Myr younger.
The diverse mammal fauna includes: Marsupialia (Borhyaenidae,
Notogale mitis, Paraborhyaena boliviana, Pharsophorus lacerans,
Sallacyon hoffsletteri: Caenolestidae, Palaeothentes boliviensis;
?Polydolopidae or Palaeothentinae. gen. et sp. indet.; Argyrolagidae,
Proargyrolagus bolivianus), Edentata (Dasypodidae, gen. et sp.
indet.; Peltephilidae, cf. Peltiphilus sp.; Glyptodontidae, cf.

633

LARRY G . MARSHALL &THIERRY SEMPERE

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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,Holocene; e, early; m, middle; 1, late.
Abbreviations for fhults and surfaces: CALP, Cabalgamiento Altiplsnico Principal (Main Altiplanic Thrust); CANP, Cabalgamiento
Andin Principal (Main Andean Thrust); CCR, Cabalgamiento de la Cordillera Real (Cordillera Real Thrust); CFP, Cabalgamiento
Frontal Principal (Main Frontal Thrust); FAT, Palla Aiquile-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. e t n. sp; Mylodontidae, 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), Primates (Cebidae, 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
small quarry along the railroad about 3 km southwest of Corocoro
Branisamys luribayensis), Litopterna (Macraucheniidae,
?Coniopternium printitivunz; Adianthidae, Tliadanius Itoffstetteri, from rocks which belong to the middle part of the Coniri Formation
Tricoelodus boliviensis), Notoungulata (Isotemnidae, gen. et sp.
(Cherroni & Cirbiln, 1970). This footprint is figured by Singewald
indet.; Notohippidae, ?Rltynchippus cf. eqiiinirs, ?R. cf. puntidis;
& Berry (1922, pl. 6, p. 52), but its true identity has yet to be
established. Because of its stratigraphic position, this edentate foot
Toxodontidae, Nesodontinae, gen. et sp. indet.; Hegetotheriidae,
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
Astrapotheria (Astrapotheriidae, gen. et sp. indet.), Pyrotheria fossil turtle from along the Rio Cesarsama at a locality they called
(Pyrotheriidae,Pyrotheriuni ronieri). order and family incertae sedis 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 Lpez (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
he calls (p. 24) the Quendeque Formation, of probable late Miocene
Marshall, 1978; Patterson & Wood, 1982; Petter & Hoffstetter,
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 primate known i n 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
(Colhuehuapian-Chasicoan)
come from a n unnamed rock u n i t include: Marsupialia
A relatively rich fauna of middle Miocene age is known from an
(Rorhyaenidae), Edentata (Dasypodidae, cf. Prozaedyus;
Glyplodontidac, G l y p f n f e l u s ;Orophodonlidae), Litopterna
unnamed unit at Quebrada Honda (3500 m),about 55 km southwest
(M acr a u ch en i id a c , c f. Coniopt ern iuni) , N oto u IIg u 1a t a of Tarija, just north of the Argentine border in southernmost Bolivia.
This unit is of fluvio-lacustrine origin and is only slightly deformed.
(Mesotheriidae, 7rachytherus spegazzinianus; Hegetotheriidae,
Prohegetoherirmi),and caviomorph Rodentia (Cephaloinyidae, gen. The fauna was assigned a Colloncuran (Friasian) age by Hoffstetter
(1977), and subsequent study by Frailey (1988) suggested that it was
et sp. indet.; Chinchillidae, Eoviscaccia boliviana) (Hoffstetter et
Santacrucian. However, recent geochronologic work by MacFadden
al., 1971; Hartenberger, 1975; Vucetich, 1988, 1989).
(1990a) and MacFadden et al. (1990) shows that most of the
A poorly known fauna from the base of the Petaca Formation at
Quebrada Honda strata were deposited between 13 and 12 Ma,
Quebrada Saguayo in the Subandean belt, about 60 km westnorthwest of Santa Cruz, is probably of Deseadan age. The known indicating that the fauna is late Colloncuran in age. A brief
description of the geology is given by MacFadden & Wolff (1981).
fauna includes a right maxilla of a mammal identified as
The fauna includes: Marsupialia (Borhyaenidae; Caenolestidae;
?Rhynchippus sp. (Notoungulata, Notohippidae, Rhynchippinae) by
Villarroel (in Sanjins & Jitn-nez, 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, Chaco basin (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 Sacaba 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, Uyuni basin (10-H); 26, Salinas de Gard Mendoza basin (?-H); 27, Carangas basin
(M-H); 28, Charaa 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-Chapadmalalan) South American Land
Mammal Ages (modified after Marshall, 1985) and principal vertebrate-bearingformations (*) discussed in text (modified after
Sempere, 1990). Geologic time scale 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
maximum thickness of 160 in. The reported fauna includes 10 taxa
sp.), Litopterna (Proterotheriidae, Diadiaphorus sp.;
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
sp.; Glyptodontidae, Propalaelioplophorics sp., Neothoracoplioriis?
Caviidae) (Hoffstetter, 1977; MacFadden & Wolff, 1981; Takai et
sp.), Notoungulata (Mesotheriidae, Plesiotypollzeriuni sp.;
al., 1984; Frailey, 1987, 1988; Villarroel & Marshall, 1988).
The 300 m-thick Quebrada Honda strata accumulated by infilling Hegetotheriidae, lfegetofheriuni sp.), Litopterna (Proterotheriidae,
Diadiapliorus sp.), and Rodentia (Octodontidae, gen. et sp. indet;
of tlie Pasajes syncline which formed just west of the CANP major
fault (Fig. 1). This syncline is bounded on its western side by a Chinchillidae. Prolagosfonwssp.).
skinger of en-chelon anticlines which involve Cambrian and early
Also from the Nazareno Formation at Suipacha, 3 km northnorthwest of Nazareno, an armadillo was collected during
Ordovician strata and document a regional right-lateral component
construction of a railway cut at km 69 on the line from Villazn to
for the CANP (Sempere, unpublished). The formation of the Pasajes
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 erosion of Ordovician shales and Cambrian Hoffstetter (1977) tentatively regarded this fossil as Huayquerian.
Ois0 (1991) regards Dasypodon alnvris as a small species of
quartzites, and probable local faulting; led to formation of a hollow
depression in the central part of the Pasajes syncline were fluvial and Slenotafus (i.e. S. afnvus), and given its provenance from the
Nizareno Formation is also almost certainly of Colloncuran age
lacustrine sediments. as described by MacFadden & Wolff (1981),
(sensu Marshall & Salinas, 1990).
gently accumulated. These deposits are thus partly coeval with the
Huxley (1860; see also Blakk, 1861) described the partial skeleton
development of the middle Miocene Chayanta planation surface
of a litoptern (Macratcchenia boliviensis) from what Singewald &
(Sempere et al,, 1990e). and with the deposition bf the Quehua
Berry (1922, p. 44) call the Ramos Series at Corocoro. a copper mine
Form ation.
on the Altiplano south of La Paz. This unit is equivalent to the
A second relatively rich fauna of Colloncuran age is reported by
Kollukollu Formation (Cherroni, 1974) which has a K-Ar date of
Ois0 (1991) from the middle and lower members of the Nazareno
16.6 f 0.4 Ma on a sanidine concentrate from a reworked tuff
Formation about 10 km south of Nazareno (on the west side of
(Swanson el al., 1987). Hoffstetter & Paskoff (1966, p. 483) cotifinn
Estacin Arenales; 21"40'S, GS"3S'W) and 30 kin southeast of
Tupiza in the Cordillera Oriental, department of Potos, 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


specimen is poorly prescrvcd and a secure generic identification
appears improbable (Ahlfeld & ranisa, 1960, p. 165). Thus, the true
gencric idcntity and age of this species remains unknown. Because
of its stratigraphic position, this macraucheniid might be of
Colhuehuapian or Santacrucian age.
There are two poorly known faunas of possible Colloncuran age in
Bolivia. The first is from the Choquecota Formation (=Caquiaviri
Formation, which near its base contains blocks of dacite lava K-Ar
dated at 14.2 f 0.4 Ma; Swanson et al., 1987) about 3 km north of
Choquecota, department of Oruro, on the Altiplano (see Hoffstetter
et al., 1972). This was the site of collection of a small notoungulate
(Mesotheriidae) which Villarroel (1974b) named Microfypofheriuni
choquecoteme. Remains of Dasypodidae (Prozaedyus),
Glyptodontidae, Hegetotheriidae and Chinchillidae are also known
(Hoffstetter e l al., 1972). The second locality is in southwestern
Bolivia between Cerdas and Atocha, where Microtypolherirmz cf. M.
choquecoteme was collected from tlie lower part of the Quehua
Formation S.S. (mapped as "Formacin Quehua Superior'' on Hoja
6331, Quechisla) (see Hoffstetter, 1977; Villarroel, 1978, p. 166;
Hoffstetter in Marshall et al., 1983, p. 41).
A small species of Notoungulata (Mesotheriidae,
Plesiofypofheriuni niinus Villarroel, 1978) was collected 3 kin
southeast of Cerdas from the upper part of the Quehua Formation S.S.
(mapped a s "Formacin Quehua Superior" on Hoja 6331,
Quechisla). This species appears to be more specialized than
Microfyyotheriuni choquecoteme suggesting that it may be
Chasicoan in age, although this has yet to be demonstrated
(Villarroel, 1978; Hoffstetter in Marshall et al., 1983, p. 41).
A locality from the uppermost part of the Petaca Formation
o n the Ro Yapacani, about 9 0 kin west-northwest of Santa
Cruz in the Subandean belt yielded a single right mandible
with five teeth which was identified by Pascual and Odreman
Rivas (in Sanjins 6c Jimnez, 1976) as cf. Vassallia minuta
(Edentata, Dasypodidae, Pampatheriinae). This identification
indicates a Chasicoan to Montehermosan age for this level of
the Petaca Formation.
A few vertebrate fossils have been recovered from the Yecua
Forination, which overlies the Petaca Formation, along the Ro Alto
Moile, about 95 km west-northwest of Santa Cruz in southcentral
Bolivia (Lpez, 1983, p. 18). The known fauna includes a limb bone
o f a litoptern tentatively identified as cf. Theosodon sp.
(Macraucheniidae), an indeterminate rodent tooth, the first fossil
remains of the fish order Gyninotiformes (Apteronotidae, gen. et sp.
indet.) (electric eels) (Gayet, 1986) and indeterminate Siluriformes
(Gayet, 1991). If the identification of Tlieosodon is correct, then the
age of this fauna is somewhere between Colhuehuapian and
Chasicoan, which is the known stmtigraphic range of this genus
elsewhere in South America.
The occ,urrence o f the Chasicoan to Montehermosan cf.
Vassallia niinula below a level with the Colhuehuapian to
Chasicoan cf. Theosodon sp. implies that the top of the
Petaca Formation and at least a part of the Yecua Formation
are Chasicoan in age (Fig. 2). This age assignment is supported by
the fact that the Yecua Formation represents the northern extent of
the Paranense epicontinental sea which is, at least in part, temporally
equivalent to Chasicoan (Sempere, 1990). The fish fossils reported
by Lbpcz (1975) from tlie base of the Tariqua Formation along the

Alto Rio M o i l e a r e a p p a r e n t l y a l s o of Cllasiconri or


Huayquerian age.
A fauna of probable Chasicoan age is reported by Russo (1959, p.
29) from "un afluente derecho, sin nombre, de la Quebrada de
Palmar" at Agua Salada near Yacuiba in southeast Bolivia just north
of the Argentine border. The fossils were found in sediments which
Russo called the "Chaco Inferior" and are now called the Tariqua
Formation, which overlies the Yecua Formation. As noted by Russo
(p. 29), "en una arenisca situada directamente sobre el banco tobaceo
gris, se hallaron huesos, dientes y un bozo de columna vertebral de
unos 40 centmetros de longitud, con apfisis dorsales y costillas an
sujetas a l, de un mamfero fsil de hbito terrestre. Estos fsiles no
han podido ser determinados en el campo, su clasificacin requerira
el trabajo de un especialista y a h a s sera difcil hacerlo debido a la
pobreza del material recolectado".
From t h e b a s e o f t h e G u a n d a c a y F o r m a t i o n a few
kilometers north of Bermejo along the Rio Bermejo, Ing. Antonio
Sadud collected a partial skull and skeleton of a notoungulate
(Hegetotlieriidae; determined by R. Pascual). This formation is
apparently Chasicoan to Huayquerian in age.

LATE MIOCENE
(Huayquerian)
There are numerous land mammal faunas of Huayquerian age in
Bolivia, particularly from the Mauri 6, Rosa Pata and Quehua
formations on the Altiplano. At localities where these formations
were deformed during Huayqueriaii time they are disconformably
overlain by rocks of Pliocene and/or Pleistocene age. These
deformations on the Altiplano correspond to the second major
tectonic crisis mentioned above (11-5 Ma), and apparently to what is
termed the Quechua deformation in Argentina (Hoffstetter, 1986, p.
222) which began about 10 +_ 2 Ma (Jordan & Alonso, 1987). '
A sample which yielded a K-Ar age of 10.5 Ma (Evernden
e f al., 1966, 1977) was apparently collected from a tuff about 1500
m above the base of the Mauri 6 Formation at Cerro Jakokota
(=Hakakota) (4,000 m), 12 kni southeast of Santiago de Machaca.
This locality may be the same as Cerro Jancocota where Berry
(1922) collected and described a fossil flora that includes leaves of
Pteris, Pliragmites, Alnus, Osteomeles, Polylepis, Calliandra,
Cassia, Cesalpinia and Melastoniites; fossil wood of
Prosopis (Mimosaceae) occurs below the tuff at this locality
(Hoffstetter, 1986). The Mauri 6 is disconformably overlain by the
subhorizontal Prez Ignimbrite which has a mean radioisotopic
age of 2.8 Ma (Evernden et al., 1966, 1977; Lavenu et al.,
1989; Marshall e r al., 1992) and probably belongs to the same
igiiimbritic flow as the Chijini and Ayo Ayo tuffs which are
also 2.8 Ma (Marshall el al., 1992).
The richest of the mammal-bearing Mauri 6 localities is at Acliiri
(especially at Cerro Pirapi Grande, about 6 km from Achiri) which
occurs in the upper part of this formation. Ofier noteworthy Mauri 6
localities include Canacho, Jankojakhe (=Jankoaqui) and Rosario.
The known fauna from these four localities includes: Marsupialia
(orhyaenidae, Borliyaenidirim altiplanicus), Edentata
sp.;
(Dasypodidae, Chorobates sp., cf. Kraglievchia
Glyptodontidae, Sclerocalyptinae; Megalonychidae; Mylodontidae),
and Notoungulata (Toxodontidae, Toxodontinae, gen. et sp. indet.,
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
reputed Huayquerian age in Bolivia, yet all need confirmation.
Rio Mauri, a tributary of the Rio Desaguadero. Villarroel (1977, p.
First, an edentulous skull of a Mesotheriidae (aff.
29) notes that the Desaguadero Formation of previous workers has
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 Potos, near the Argentine,border in
southernmost Bolivia (Anaya et al., 1989).
probably from the Mauri 6. This specimen was identified by C. W.
Second, is a fauna from the "Estratos Muyu Huasi" at Muyu
Andrews of the British Museum (Natural History), London as
Huasi, about 50 km northeast of Sucre in southcentral Bolivia. The
Nesodon sp., a genus known securely only from rocks of
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
caviomorph Rodentia (Abrocomidae, Profabroconta sp.; Caviidae,
Toxodontidae, gen. et sp. indet.
?Orflioniycferasp., ?Cardionlys sp.; Chinchillidae, Lagosromopis
Land mammals have been collected from the upper part of the
sp.; Dinoinyidae, gen. et sp. indet.; Eretliizontidae, gen. et sp. indet.;
Totora Group, 2 km north of Ulloma at Minita Chocopini near
Octodontidae, cf. Sciamys sp.) (Villarroel & Marshall, 1989).
Torini, 4 km west of Callapa, north of Chacarilla, and in the vicinity
Third, is the fauna from the Cobija Formation collected along the
of Curahuara de Carangas (Hoffstetter, 1986). Two tuffs from the
Rosa Pata Formation (upper part of the Totora Group) named toba Ro Acre between Cobija and Bolpebra, department of Pando in
northwest Bolivia. The vertebrates include fish (Chondrichthyes),
Callapa and toba Ulloma, have been K-Ar dated at about 9.0 and
10.4 Ma, respectively (Marshall et al., 1992). The Rosa Pata reptiles (Chelonia, Podocnenzis sp.; Crocodilia, Gavialosrcclius sp.),
Formation is either disconformably overlain by the latest Miocene birds (indet.), and mammals (Rodentia indet.) (Carrasco, 1986). The
Pomata Formation, Pliocene Umala Formation which has a mean jaw of an enormous crocodile was collected from this same
formation at Candelaria, to the west of Riberalta along the Ro
radioisotopic date of 5.4 Ma for its basal tuff (=toba 76, toba Umala
or toba Jankhomarka; Thouveny & Servant, 1989; Marshall et al., Madre de Dios (Leytn & Pacheco, 1989).
And fourth, a CTagment of a right dentary witli well preserved M*
1992), Prez Ignimbrite dated at 2.8 Ma (see above), or Pleistocene
and M' of cf. Psertdoliegefoflieriuni sp. (Notoungulata,
Ulloma Formation (see below; Marshall et al., 1983; Hoffstetter,
1986). The known mammals include: Edentata (Dasypodidae, Hegetotheriidae) was recently collected at the Estacin Abaroa,
Pampatheriinae; Glyptodontidae), Litopterna (Macraucheniidae?) about 25 km east-northeast of Charaa, department of La Paz, The
specimen is in GEOBOL collections in La Paz and has the catalogue
and Notoungulata (Mesotheriidae, P seudofypotlierirrni sp.)
number GB-089 (C. Villarroel, personal communication, 1990).
(Martinez & Rosales, 1972; Hoffstetter et al., 1972).
From the base of a 205 m section of what are informally called the
EARLY AND MIDDLE PLIOCENE
Micaia beds at Micaia in the Cordillera Oriental (17'30's. 67"24'W;
(Monteliennosan-Cliapadinalalaii)
38004000 in) were recovered a niediuni-sized niegatheriid ground
sloth (gen. et sp. indet.) and a small notoungulate (Mesotheriidae,
The most important mammal fauna of Montehermosan age i n
Microtypotlierirmi cf. M.choquecotense) (MacFadden et al., 1990).
Bolivia is from tlie Umala Formation at and near Ayo Ayo (3800 in),
A magnetostratigraphic study and an associated fission-track date of
Vizcacliani and Umala, 65-75 km south of La Paz. A tuff from the
6.9 k 1.1 Ma, indicate that these rocks and fauna span fToin 7.5 base of the Umala Formation (toba 7 6 ) has yielded mean
6.7 Ma (MacFadden ef al., 1990), making thein late Huayquerian.
Remains of Notoungulata (Mesotheriidae, Plesiofyi~oflieriuni) radioisotopic dates of 5.4 Ma (Evernden ef al., 1966, 1977; Lavenu
c/ al., 1989; Marshall et al., 1992). whilc thc Ayo Ayo tuff in thc top
havc bccn collcclcd from two localitics of thc Quehua Foi inat ion S.S.
of this foriiiatioii has yicldcd a iiiean age of 2.8 Ma (Lavenu CI al.,
to thc east-southcast of Quchua in the southern part of the Altiplano
1989; Marshall et al., 1992). The fauna includes: Marsupialin
(Villarroel, 1978, p. 166; Hoffstcttcr & Villarroel, unpublished as
cited in Marshall et al., 1983, p. 43). Plesiotypotheririni cf. P. niajrts (Didelphidae, Sparassocyniiiae, Sparassocynrrs lieterotopicrrs;
?Caeiiolestidae; Argyrolagidae, Microtragrrlrrs bolivianrrs),Edentata
or niinrrs and scutes of Dasypodidae were collected from
(Dasypodidae. Pampatheriinne, cf. Plaina sp., Euphractinae,
approximately 3 km southeast of Quehua; and cf. P. achirense was
Macroeirplzractiis sp.; Megatheriidae, cf. Megt~lieriiinr;
collected approximately 7 km southeast of Quehua (the first locality
Mylodontidae, Mylodontinae), Notouiigulata (Toxodontidne,
is from a level 40 ni stratigraphically higher than the second).
Xotodontinae, Posnanskytlieriiini desagrraderoi), Litopterna
Rodentia (Caviidae, Dolichotinae, Orflioniycferasp.) are also
(Macraucheniidae, cf. Proniacrauclienia sp.), and cavioniorph
recorded from Quehua (Hoffstetter, 1986, p. 222). The presence of
these taxa establishes that the Quehua Formation at these localities is Rodentia (Octodontidae, Ctenomyinae, Praecfenoniys rhonibidens,
P. vagus; Chinchillidae, Lagostominae; Caviidae, Caviinae;
temporally equivalent to the Mauri 6 and Rosa Pata, formations
' Hydrochoeridae. Cltapalntarlieriuni saavedrai) (Liendo-Lazarte,
fartlier to the north.

638

FOSILES Y FACIES DE BOLMA -VOL 1 VERTEJHUDOS

$GE
M ~ )&

A L T I P L A N 3
GLACIAL LAKES ASSOCIATED
FORMATIONS

LANDMAMMAL AGE

:I
I-

1.04

SOUTH AMERICAN

ENSENADAN

GLACIATIONS

Ulloma Fm.*
Kaluya

!Purapurani T. 1.6Ma

OCHABAMBP

Anzaldo

Anzaldo

Anzaldo
Sacaba I

*
*

Cabana
Mataro

:ORDILLERA
ORIENTAL
(Central 1

Betanzos

2.5
30

MONTEHERMOSAN

Umala Fm.*

Prez Ign.2.8Mr:

r--Tari)o*, Padcayo

i;luapua~*

*,

ConceEcn2------ -----

Charaa Fm.*

- ?-?-?-

CHACO

American Faunal Interchange

Figure 3. Chronostratigrapliic chart showing late Pliocene-Pleistocene (Uquian-Lujanian) South American Land Mammal Ages
(modified after Marshall, 1985) and principal vertebrate-bearingformations (*) discussed in text. Geologic time scale follows
Berggren et al. (1985).
1943; Hoffstetter, 1986; Hoffstetter et al., 1971, 1984; Hoffstetter &
Villarroel, 1974; Villarroel, 1975, 1977; Villarroel & Marshall,
1983).
Numerous mammals have been collected from the La Paz
Formation around La Paz at Achocalla, Alto Obrajes, Alto
Segencoma, Calacoto, Gualberto Villarroel, Kenko and
Tembladerani. This formation consists of some 700 m of fine to
coarse sediments of fluvial to fluvio-lacustrine origin (Lavenu et al.,
1989) which Dobrovolny (1962, pp. 20-27) subdivided into three
units (lower, middle, upper). The fossils are from below a tuff (toba
Chijini) in the upper part of the La Paz Formation which has yielded
a mean radioisotopic date of 2.8 Ma (Clapperton, 1979; Lavenu el
al, 1989; Marshall et al., 1992). This tuff appears to belong to the
same ignimbrite flow that deposited the PCrez Ignimbrite and Ayo
Ayo tuff (see above). Dates of 5.5 Ma have also been obtained on
a tuff (= toba Cota Cota) in the lower part of this formation at Cota
Cota, southeast of La Paz (Lavenu et al., 1985, 1989; Servant et al.,
1989). This tuff seems to belong to the same event that deposited the
Jankhomarka tuff (= toba 76), which apparently correlate with
collapse and eruption of the Soledad Caldera at 5.4 Ma (Redwood,
1987), but was probably emitted somewhere in the western
Cordillera as suggested by its thickness increasing toward the west.
A magnetostratigraphic study of the La Paz Formation shows that
deposition of the middle and uppe; units occurred during the Gauss
epoch (i.e.3.4 to 2.48 Ma) (Thouveny & Servant, 1989). The fauna
includes: Edenlata (Dasypodidae, Euphractini, Macroertphractus
aff. nioreni; Glyptodontidae, Sclerocalyptinae,cf. Sclerocalypfussp.,
cf. Plolrophorops; Megalonychidae, gen. nov.), Notoungulata
(Toxodontidae, Xotodontinae, Posnanskytkeriitm desagrraderoi, P .
nov. sp.) and Litopterna (Macraucheniidae, cf. Proniacrauchenia
sp.) (Ahlfeld & Branisa, 1960, p. 135; Clapperton, 1979; Radelli,
1964, p. 842; Hoffstetter, 1986; Villarroel, 1977; Villarroel & GrafMeier, 1979; Mones & Mehl, 1990). The vast majority of these
fossils are from Dobrovolny's (1962) middle unit (Mones & Mehl,
1990, p. 22).
A few vertebrates have been collected from north of Pomata Ayte,

639

about 200 km south of La Paz, from rocks designated as the


Formacin Mauri (Hoja 6038, Corque; Hoja 6039, Choquecota; see
Hoffstetter et al., 1972, Fig. on p. 740) which unconformably overlie
the Huayquerian age Rosa Pata Formation (see above) or more
specifically its terminal facies, the Conglomerado Pomata, which has
a K-Ar date of 6.4 Ma obtained on a rhyolite (Evernden et al,, 1966,
p. 78). The fauna occurs aboqe a tuff
1977; Marshall et al., l9!3,
which may represent the toba Jankhomarka (Martinez, in Marshall
et al., 1983, p. 78) and includes: Notoungulata (Toxodontidae,
Xotodontinae, Posnanskytheriuni desaguaderoi), Litopterna
(Macraucheniidae, cf. Pronaacrauchenia sp.), Edentata
.(Mylodontidae), caviomorph Rodentia, and a large terrestrial
carnivorous phororhacoid bird (Hoffstetter et al., 1972; Hoffstetter,
1986).
From the Remedios Formation at Cerro Canasa near Corque and at
Orinoca south of Oruro are reported remains of Edentata
(Mylodontidae), Notoungulata (Toxodontidae, Xotodontinae,
Posnanskytherium sp.), and Litopterna (Macraucheniidae, cf.
Pronzacrauchenia sp.) (Lavenu, 1984; Hoffstetter, 1986) from above
a tuff with dates of 5.2 - 4.6 Ma (Marshall et al., 1992). This
formation is locally separated by an angular unconformity from the
overlying mammal-bearing "lacustre Minchn" of late Pleistocene
age (see below).
A Montehermosan age for the mammal faunas from the Umala, La
Paz, Remedios and "Mauri" (at Pomata) formations is established on
faunal content and radioisotopic dates. These Pliocene faunas differ
markedly from those of the late Miocene (Mauri 6, Rosa Pata,
Quehua formations) on the Altiplano by the absence of
Mesotheriidae and in the predominance of Macrkheniidae (cf.
Proniacrauchenia sp.), other Toxodontidae (Xotodontinae,
Posnanskytlterium sp.), other Edentata (Megatheriidae;
Mylodontinae; Pampatheriinae, Kraglievichia sp., Plaina sp.), and
Hydrochoeridae (Chapalmatkeriuni sp.) (Hoffstetter, in Marshall et
al., 1983, p. 44; Hoffstetter, 1986). Furthermore, these faunas are
bracketed below by the 5.4 Ma toba Jankl>omarkafrom the base of
the Umala Formation and the 5.5 Ma Cota Cota tuff in the lower
part of the La Paz Formation, and above by the 2.8 Ma Chijini tuff in

LARRY G.MARSHALL &THIERRY SEMPERE

the upper part of the La Paz Formation and 2.8 Ma Ayo Ayo tuff in
the top of the Umala Formation.
There are three other reports of possible early to middle Pliocene
a g e m a m m a l f a u n a s i n B o l i v i a , a l l o f which need
confirmation.
First, Pachynodon validus Burmeister (1892, pp. 433-434)
(Notoungulata, Toxodontidae) is probably from Bolivia and may be
of Pliocene age. As noted by Simpson (1940, p. 705), 'I ... the only
available information as to provenience is that the specimen was sent
to Vaca Guzmn Blanco (the Bolivian Minister in Buenos Aires)
from Santa Cruz de la Sierra, Santa Cruz, Bolivia. The implication is
that the specimen was found at that place, but this is not definitely
established".
Second, Monta0 (1968) described a mammal fauna from Anzaldo
(3000 m), about 55 kni southeast of Cochabamba, which he assigned
to the Pleistocene, A later visit to this locality by Hoffstetter,
Montaiio and Ortega resulted in recovery of a specimen of cf.
Proniacrarrclienia sp. (Notoungulata, Macraucheniidae) from a 2 mthick siltstone in the lower level at Quebrada Tijascka, about 3 km
southwest of Anzaldo. This is a small macraucheniid and is of
Pliocene aspect (Hoffstetter, in Marshall et al., 1983, p. 44). A true
Pleistocene mammal fauna occurs at the same localitiy (see below).
And third, from,a rock unit informally called "lac de Sacaba" or
"Sacaba 1"(Lavenu, 1986, pp. 114-115) south of Sacaba in the area
of Laba-Laba Alta (Quebrada Fierro Churu) about 18 km eastsoutheast of Cochabamba, were recovered remains of mammals
identified by Villarroel (in Mancilla, 1979, p. 32) as Plohophorus sp.
(Edentata, Glyptodontidae).

LATE PLIOCENE
(Uquian)
There is only ode potential fauna of this age in Bolivia. It is from
near Anzaldo, about 55 km southeast of Cochabamba. In Quebrada
Tijascka, 3 km southwest of Anzaldo, Montaiio (1968) recovered a
complete caudal tube of Prodaedicurus cf. P . divincenzi (Edentata,
Glyptodontidae) identified by Hoffstetter (in Marshall et al., 1984, p.
36) from a gravel layer. This is the only record of Prodaedicurus for
Bolivia, a taxon originally described from beds designated as the
"Piso Castellanosense" in Uruguay, the age and faunal content of
which are discussed by Mones (1979, pp. 16-17). Mammals of
possible Monteheimosan (see above) and Pleistocene (see below)
age are also known from Anzaldo.
L A TI C T 15R3'1A KY <?It 11
' ,151 S'I'OCEN I Z

From a locality called Cruz Vilque to the northwest of the pueblo


Chilahuala on the west bank of the Ro Desaguadero, L. Branisa
collected mammals from a 20 in section composed of cream. gray
to reddish clays. Branisa (correspondance in Princeton University)
believes there are two distinct faunas: one in the lower part of the
section is of Tertiary aspect, while the other higher in the section
yielded specimens r e s e m b l i n g Megatlzerium and is of
Pleistocene aspect.

EARLY AND MIDDLE PLEISTOCENE


(Ensenadan)
A fauna regarded by Hoffstetter (1986, p. 226) as possibly early
Ensenadan was collected by Blanco (1980, p. 99) from the Charaa
Formation just east of Charaa (4000 m) on the Altiplano near the
Chilean border. This formation overlies the Pbrez Ignimbrite and is
thus younger than 2.8 Ma (Lavenu et al., 1989; Marshall et al., 1992)
and is older than the Brunhes Normal chron (i.e. >0.73 Ma)
(Servant-Vildary & Blanco, 1984). The fossils, as yet undescribed,
include: Edentata (Glyptodontidae, Doedicurinae, Plwliaplous sp.;
Mylodontidae, Glossotlzerium sp.), Litopterna (Macraucheniidae,
Macrauclienia cf. ullonzensis), and Artiodactyla (Cervidae, gen. et
sp. indet.) (Hoffstetter, in Marshall el al., 19M, p. 36; Hoffstetter,
1986; Blanco, 1980, Anexo 2).
Another Altiplano fauna of probable Ensenadan age is from Ayo
Ayo (3800 m), about 70 km south of La Paz. Specimens of Edendata
(Mylodontidae, Scelidodon cf. tarijensis), Litopterna
(Macraucheniidae, Macrauchenia cf. pafaclionica), and Artiodactyla
(Cervidae, gen. et sp. indet.) were recovered from a sandy horizon
above a tuff and palcosol at the top of the Urnala Formation, which
has yielded a Monteliermosan age fauna at the same locality
(Hoffstetter ef al., 1971; Hoffstetter, 1986; Lavenu, 1978).
Hoffstetter (1986, p. 224) notes that Lavenu (1978) regards the Ayo
Ayo tuff (which is 2.8 Ma; Lavenu et al., 1989; Marshall et al.,
1992) as probably equivalent to the pre-Calvario tuff at Patapatani
(which gave a K-Ar age of 2.8 Ma; Lavenu et al., 1989) north of La
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
(Marshall et al., 1992).
The most spectacular vertebrate fauna of late Ensenadan age in
South America is known from numerous localities and from various
stratigraphic levels in the basins of Tarija (1950 m; i.e. Armaos, San
Blas, San Pedro, Santa Ana, Pueblo Viejo, Tolomosa), Concepcin
(= Uriondo, 1800 m; about 20 km south-southeast of Tarija), and
Padcaya (2100 m; about 40 km south of Tarija) in southernmost
Bolivia (Ahlfeld & Branisa, 1960, Fig. 11, pp. 168-171). The
predominant lithologies within the Tarija basin are silty clays, sands,
conglomerates and volcanic ash which were apparently deposited in
a fluvial environment consisting of an accreting stream and
floodplain system (MacFadden & Wolff, 1981).
The first citation of fossil mammals from Tarija, and for that
matter the first written record of fossil mammals from Bolivia, was
by Diego d e Avalos (1602; fide d'orbigny, 1842, 111-4,
Palontologie, p. 145). Joseph de Jussieu, in a letter to his brother
Bernard ici 1761(J& Cuvicr, 1823, pp. 266-267), noted the prescnce
of fossil bones and teeth at Tarija, and in 1842 Laurillard (in
d'orbigny. 1842, 111-4, Palontologie, p. 145. PIS. 10-11) described
mastodonts. Specimens collected by Weddell in 1845 as a meinber
of the F. de Castelnau expedition (see Weddell, 1851, pp. 196-207)
were sent to the Musbuni Natiqnal d'Histoire Naturelle, Paris and
subsequently studied and described by P. Gervais (1855) and H.
Gervais & Ameghino (1880) (Hoffstetter, 1970). The collection
made by de Carles (1888) for the Museo Argentino de Ciencias
Naturiles, Buenos Aires, was studied by Ameghino (1902), and

640

~~

~~

FOSILES Y FACIES DE BOLIVIA - VOL. I VERTEBRADOS

I I

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

641

LARRY G. MARSHALL &THIERRY SEMPERE

later papers on this collection were provided by Kraglievich (1928,


1930b), Cabrera (1929, 193l), and Lpez Aranguren (1930). In 1902
Nordenskild sent a small collection to Uppsala, Sweden which
resulted in publications by Nordenskild (1902, 1903) and Sefve
(1912, 1915b). A large collection of specimens purchased in 1903 by
the Marquis de Crqui-Montfort and E. Snchal de la Grange from
the Echaz family was donated to the Musum National dHistoire
Naturelle, Paris and served as the basis for the now classic
monograph by Boule & Thvenin (1920) (see also Cottreau, 1921);
the stratigraphy was described by Mortillet (1922), a member of the
1903 expedition. Between 1894 and 1915 members of the Echaz
family of Tarija made an impressive private collection of fossil
mammals from Tarija and surrounding areas @chaz, 1905, 1921).
Some of these fossils were sold to various foreign institutions (see
above, below), although the major part of the collection, consisting
of some 2,630 specimens which remain virtually unstudied, was sent
to La Paz in the 1930s. In 1977 part of this collection was returned
to Tarija and deposited in the Museo de la Universidad de Tarija; it
is planned that the remainder of the collection will be returned to
Tarija in the ilex future (Takai ef al., 1982, 1984). In 1924 and 1927
expeditions from the Field Museum of Natural History, Chicago
under the direction of E. S.Riggs made a spectacular collection from
Tarija and Padcaya, and purchased some specimens from the Echaz
family (Riggs, 1928, 1930); this magnificent collection remains
virtually undescribed (Marshall, 1978). During 1978 and 1980
paleontologists from the Research Institute of Evolutionary Biology,
Tokyo made a small collection which was taken to Japan (Takai et
al., 1982, 1984). The most recent work by foreign researchers was
carried-out by a team from the University of Florida, Gainesville in
1980 and 1986 (MacFadden, 1981; MacFadden & Azzaroli, 1987;
MacFadden & Wolff, 1981; MacFadden ef al., 1983; Frailey et al.,
1980).
A history of the debated age of the Tarija fauna is provided by
Marshall et al. (1984). Ameghino (1902) and Boule & Thvenin
(1920) regarded the fauna as Pliocene; Rovereto (1914) assigned it
to a new land mammal age, the Tarijense; Kraglievich (1930a, 1934)
stressed the post-Ensenadan (s.s.) and pre-Lujanian (s.s.) aspect of
the fauna which means that it is comparable to the Bonaerense as
defined in Argentina (see Marshall ef al., 1984, Fig. 1). Palterson &
Pascua1 (1972, p. 249) regarded the fauna as Lujanian, while Webb
(1974, p. 176) considered it Ensenadan. This debate was resolved, at
least in part, by MacFadden et a l . (1983) who provided a
geochronologic study of the 250 m-thick Tarija Formation in the
Tarija basin using magnetostratigraphy and radioisotopic dating.
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
current standards. Iakai et al. (1982, p. 5) report dates of 0.25 - 0.20
Ma on some fossil bones using electron spin resonance technique,
but these dates are at odds with the more secure conclusions of
MacFadden et al. (1983). Nevertheless, Hoffstetter (1986) cautions
that some fossils from upper levels of the Tarija Formation may be
younger, perhaps early Lujanian in age. It is apparent that additional
detailed stratigraphic studies of the type provided by Oppenheim
(1943), additional magnetostratigraphic and radioisotopic studies as
provided by MacFadden et al. (1983), and more refined systematic
and biostratigraphic studies of the faunas from Tarija, Concepcin
and Padcaya are needed to establish the absolute and relative ages of
the numerous fossil levels.

This fauna is in dire need of a systematic revision. In addition to


the papers cited above, the mammals are discussed by Berta (1981,
1985,,1987, 1988), Liendo-Lazarte (1946) and Hoffstetter (1963a, b,
1968a, 1986). The following taxa appear to be securely identified:
Marsupialia (Didelphidae, Lutreolina crassicaudata), Edentata
(Dasypodidae, Chaetophractus tarijensis, C . villosus, Eupliractus
sexcinctus, Propraopus cf. sulcatus or grandis, Pampatherium cf.
Itumboldti or typum; Glyptodontidae, Chlamydotherium cf. sellowi,
Glyptodon ret iculat us, Hop loplwr us ecliazui, Neothoracopliorus cf.
elevatus, Panoclitus cf. tuberculatus; Megalonychidae, Nothropiis
tarijensis; Megatheriidae, Megatherium tarijense; Mylodontidae,
Glossotheriuni tarijense, Lestodon armat us, Scelidodon tarijensis),
Rodentia (Cricetidae, Andinomys cf. edea, Calomys cf. lancha,
Kunsia fronto, Necfoniyscf. squamipes, Oxymycterus cf. paranensis,
Pltyllotis cf. darwini; Octodontidae, Ctenonzys subassentiens, C .
brachyrhinus; Echimyidae, Euryzygpmatomys hoffstetteri;
Myocastoridae, Myocastor perditus; Erethizontidae, Coendou
magnus, C . sp.; Caviidae, Cavia sp., Galea cf. musteloides;
Hydrochoeridae. Hydrockoerus sp., Neoclioerus tarijensis),
Carnivora (Canidae, Canis dilus, Cltrysocyon brachyurus,
Pseudalopex gymnocercus, Theriodictis tarijensis, Protocyon
troglodytes; Ursidae, Arctodus tarijensis, A. wingei; Procyonidae,
Nasua sp.; Mustelidae, Conepatus suffocans; Felidae, Felis
concolor, F. yaguaroundi, Panthera onca, Sniilodon populator),
Litopterna (Macraucheniidae, Macrauchenia patachonica),
Notoungulata (Toxodontidae, Toxodon cf. platensis), Proboscidea
(Gomphotheriidae, Cuvieronius liyodon, Notiomastdon sp.,
Haplontastodon or Stegopiastodon sp.), Perissodactyla (Equidae,
Equus insulatus, Hippidion principale, H . bonaerense,
Onohippidiuni devillei; Tapiridae, Tapirus tarijensis), Artiodactyla
(Tayassuidae, Platygonus tarijensis; Camelidae, Lama cf. onteni,L.
glania, L . provicugna, Palaeolania sp.; Cervidae, Charitoceros
tarijensis, Hippocamelus sp.). Undescribed specimens of fish and
reptiles are reported by Hoffstetter (1963~1,pp. 201-202); frogs
(Bufonidae, Bufo cf. niarinus liorribilis; Leptodactylidae, Ceratoprys
sp.) from Tarija are described by Vergnaud-Grazzini (1968); buds
(Ciconiiformes, Vulturidae, Vultur gryplius = V . pratruus Lnnberg,
1905; see Emslie, 1988) and undescribed specimens of birds are
known from Tarija (locality of Turumayu, in Peabody Museum, Yale
University) and Padcaya (Villarroel, in Marshall et al., 1984, p. 33;
specimens also in Peabody Museum, Yale University).

642

LATE PLEISTOCENE-HOLOCENE
(Lujanian-Recent)
A faana of large body-size mammals of possible early Lujanian
age has long been known from the now classic locality of Ulloma
(3880 m), situated about 40 km south of Corocoro on the south side
of the Rio Desaguadero. The fossils are from the Ulloma Formation
which consists of well stratified fine-grained sands rich in calcareous
nodules; it is bounded below by an undated tuff, above by a
diatomite, and rests unconformably on the folded Totora Group (late
Miocene). The Ulloma Formation was apparently deposited during
retreat of the Sorata glaciation as part of Lake Ballivian, an ancient
southern extension of Lake Titicaca (Servant & Fontes, 1978;
Hoffstetter, 1986, p. 224; Hoffstetter, in Marshall et al., 1984, p. 35).
The fauna includes: Edentata (Glyptodontidae, Glyptodon sp.;
Megatheriidae, Megatherium cf. americanum, Megatherium

FOSDXS Y FACLES DE B O L M A - VOL. I VERTEBRADOS

(Pseudomegatherium) medinae = M. sundti; Mylodontidae,


Sceli#odon cltliense Lydekker 1886 = Scelidotheriunt?
bolvianum), Litopterna (Macraucheniidae, Macraucltenia
ullontensis), Proboscidea (Gomphotheriidae, Cuvieronius ltyodon =
Mastodon bolivianus), and Perissodactyla (Equidae, Onoltippidium
(Paraltipparion) bolivianuni = Scelidotlterium? compressunt)
(Sundt, 1900; Philippi, 1893a, b; Pompeki, 1902; Oliver, 1934;
Sefve, 1914, 1915a, b; Douglas, 1914; Kozlowski, 1923;
Casamiquela & Seplveda, 1974; Ahlfeld & Branisa, 1960; Servant
& Fontes, 1978; Hoffstetter, 1986; Marshall & Salinas, 1991). A
recently discovered fauna at a locality 4 km west of Carsani is also
probably from the Ulloma Formation, and the taxa include: Edentata
(Mylodontidae?, Megatheriidae), Notoungulata (Toxodontidae),
Perissodactyla (Equidae), and Rodentia (R. Cspedes, personal
communication, 1989).
The richest Lujanian fauna in Bolivia is from uapua, between
Carandait and Capiranda. Hermann (see Schiller, 1913, p. 180, n2)
first reported mammals from this locality, and Bonarelli (1921, p.
80) later recorded mastodonts and other mammals. Hoffstetter
(1968a) published the first study on the stratigraphy, sedimentology
and vertebrate paleontology based on work he did with L. Branisa
between 1962 and 1965. MacFadden & Wolff (1981) made a similar
study based on wdrk done in 1978.
The uapua Formation consists of 9 m of sediments which rest
unconforinably on non-fossiliferous gray and red clays of the
Tertiary age Chaco Series. Hoffstetter (1968a) divided the uapua
Formation into three members (from oldest to youngest, uapua 1,
uapua 2, uapua 3) based on color of sediments and faunal
content. MacFadden & Wolff (1981) provide a magnetostratigraphic
study of the uapua Formation and show that all three members are
located within the Brunhes Normal chron which begins at 0.73 Ma
and are thus Lujanian in age or younger (see below).
The lower uapua 1 member, which consists of 2-3 m of pink to
reddish consolidated cinerite, is probably the level from which
mammals were collected by early workers. The fauna (see
Hoffstetter, 1968a; MacFadden & Wolff, 1981; Berta, 1981, 1985,
1987, 1988) includes: Reptilia (turtles, Chelonia, Testudinidae, large
Chelonoidis sp., a small testudinid? and an undetermined chelid?;
Broin, 1991). Edentata (Dasypodidae, Panipatlterium sp.;
Glyptodontidae, Chlamydotherium sp., Glyptodon cf. reticulatus,
Panoclitus cf. tuberculatus, Sclerocalyptus cf. ornatus;
Megalonychidae, cf. Notlirotlterium sp.; Megatheriidae,
Megatherium cf. americanum; Mylodontdae, Mylodon darnhi),
Rodentia (Hydrochoeridae, Hydrocltoerus cf. hydrocltoeris;
Ctenomyidae; Capromyidae; Cricetidae; Octodontidae), Carnivora
(Canidae, Protocyon troglodytes, Pseudalopex sp.; Ursidae,
Arctodiis sp.; Felidae, Panthera onca, Smlodon popiflator),
Litopterna (Macraucheniidae, Macraucltenia cf. patacltonica),
Notoungulata (Toxodontidae, Toxodon ensenadensis), Proboscidea
(Gomphotheriidae, Stegomastodon sp.), Perissodactyla (Equidae,
Equus curvidens, Hippidion sp.), and Artiodactyla (Camelidae,
Palaeolama sp.; Tayassuidae; Cervidae). The uapua 1 fauna is
clearly of Lujanian age.
The middle uapua 2 member consists of 1.5 - 1.0 m of reworked
gray tuff. This fich fauna, discovered by L. Branisa in 1955,
includes: frogs (Bufonidae, Bufo cf. paracnemis; Leptodactylidae,
Leptodactylus cf. ocellatus, Ceratrophrys cf. ornata), reptiles
(Teiidae, Tupinambis teguixin; Amphisbaenidae, Leposternon sp.;

Boidae; Colubridae, Colubrinae; Viperidae, Crotalinae;


undetermined turtles), birds (Rheidae, Rhea cf. americana;
Podicipidae, Podiceps minor, P . cornutris; Plataleidae, Plareleu sp.;
Ajaja rosea; Anseridae, Dendrocygna sp., Anas sp.; Rallidae;
Jacanidae, Jacana sp.; Tinamidae, Cryptiirus lataupa, cf.
Rltynclrotus sp.; Columbidae; Falconidae; Cuculidae, Coccyzus sp.;
Caprimulgidae, Nyctibus sp.; Passeriformes); the mammals include
Chiroptera (Molossidae, Euntops perotis), Edentata (Dasypodidae,
Chaetopliractus cf. vellerosus or villosus, Zaedyus pichiy , Dasypus
cf. novenicinctus, Eupltractus sexcinctus, Tolypeutes matacus,
Propraopus cf. punclatus), Rodentia (Cricetidae, Caloniys sp.,
Graomys sp., Holochilus brasiliensis, Zygodontomys lasiurus;
Octodontidae, Ctenomys sp.; Myocastoridae, Myocastor coypus;
Chinchillidae, Lagostomus maximus; Caviidae, Galea cf.
musteloides; Capromyidae; Hydrochoeridae, Hydrochoerus
hydrocltaeris), Lagomorpha (Leporidae), Carnivora (Canidae,
Dusicyon griseus), Artiodactyla (Cervidae, Morenelapltus sp.), and
a human skull (Hoffstetter, 1968a; MacFadden & Wolff, 1981;
Vergnaud-Grazzini, 1968; Marshall et al., 1984). A carbon-14 date
of 7200 f 400 yrbp was obtained on the human skull, and another
date of 6600 f 370 yrbp was obtained on unspecified associated
mammal bones (MacFadden & Wolff, 1981; Marshall el al., 1984, p.
34). Some fossils of extinct megafauna ( i . e . Glyptodontidae,
Scelidodon, Toxodon, Palaeolama, and mastodonts) were recovered
from the uapua 2 member, but aspects of their preservation suggest
that these specimens were reworked from the uapua 1 member and
are thus not contemporaneous with the rest of the uapua 2 fauna
(Hoffstetter, 1968a, pp. 832-833; but see MacFadden & Wolff, ,1981
who did not recognize this fact). Thus, the carbon-14 dates and the
absence of securely associated megafauna suggest that the uapua 2
fauna is Holocene in age.
Hoffstetter (1968a, p. 834) reports the recovery of an ulna of
Toxodon sp. from the basal contact of uapua 3 with the underlying
uapua 2, but not within the uapua 3 tuff which is otherwise
unfossiliferous and which MacFadden & Wolff (1981) regard as
Quaternary alluvium. This specimen was also probably reworked
from uapua 1.

PLEISTOCENE sensu lato


(Ensenadan-Lujanian)
There are numerous other scattered reports of Pleistocene
mammals in Bolivia, but the known specimens have been little
studied and for many their age and stratigraphic context have yet to
be securely established.
On the Altiplano at La Paz and Tambillo (west of La Paz) have
been collected remains of mastodonts and glyptodonts (Hoffstetter,
1986). "Pleistocene fossils" have also been reported at San Andrs
and Achiri (Cantn Achiri, i.e. at Challncollo and Huana Jahuira),
although the exact stratigraphic position and identification of these
fossils need confirmation (Hoffstetter, 1986, p. 224). Furthermore,
the reputed Pleistocene fossils at Achiri may, in fact, represent a
confusion with those from theqlate Miocene Mauri 6 Formation
(Hoffstetter, 1986, p. 224). A glyptodont carapace (now in
GEOBOL, La Paz) was collected by Anaya (1988) from the
Comunidad Segueri, about 1.5 km east of Tiahuanacu; a possible
Megatherium sp. was collected in 1985 about 600 in to the
southwest of the ruins of Tiahuanacu (Anaya. personal

643

LARRY G.,MARSHALL & THIERRY SEMPERE

communication, 1989); and L. Branisa (correspondence now in


Peabody Museum, Yale University) records a fossil horse (Equidae)
from at or near Tiahuanacu.
On the western slope of the Cordillera Oriental, about 110 km
southeast of La Paz, have been recorded remains of Panthera onca at
Yaco (3600 m) and Macrauclzenia sp. at Puchuni (Hoffstetter, 1986,
p. 226). Kozlowski (1923) records discovery by miners of
"Pleistocene mammals" from Calacoto, Cuesta de Chacarilla and
Paria; while Lavenu (1984) records camelids (Camelidae) and L.
Branisa (unpublished notes) records remains of artiodactyls and a
horse from Culluri near Corque, southwest of Oruro. Mastodonts are
reported from northwest of Oruro, from Paria (16 km northeast of
Oruro), and from Pampa Aullagas on the southwest bank of Lake
Poop opposite Crqui-Montfort Island (Hoffstetter, 1986, p. 224).
Ahlfeld & Branisa (1960, p. 165) report remains of a mastodont tusk
and a lower jaw of Hippidion from "las minas de estao de
Carguaycollo, e n la Cordillera de los Frailes" (4100 m); the
identification of the horse needs confirmation (Hoffstetter, 1986, p.
226) and it is possibly referable to Onohippidiuai.
Near Anzaldo on the east slope of the Cordillera Oriental, about
55 kin southeast of Cochabamba, Monta0 (1968) recovered a
Pleistocene mammal fauna from sands, silts and gravels in an upper
level at Quebrada Tijascka, 3 km southwest of Anzaldo. This fauna
is from a level above the Uquian age gravel unit with Prodaedicurus
(see above), and includes: Edentata (Glyptodontidae, Glyptodon sp.;
Megatheriidae, Megatlteriuni sp.), Proboscidea (Gomphotheriidae,
Cuvieroniris sp.), and Artiodactyla (Camelidae, Lama sp.)
(Hoffstetter, in Marshall et al., 1984, p. 36; Hoffstetter, 1986; p.
223).
I
Ahlfeld & Branisa (1960, p. 167) record fragment5 of glyptodonts
from along the Ro Rocha near Sacaba about 7 km east-southeast of
Cochabamba, and R . Cspedes and R . Suarez (personal
communication, 1989) recovered remains of glyptodonts and
camelids in 1986 from along the Ro Loro Mayu, just south of the
Ro Rocha, about 10 km east-southeast of Cochabamba. Reports of
mastodonts and glyptodonts from near Sacaba and Cliza in the
Cochabamba-Sacaba basin (2600-2800 m), and of mastodonts in the
Padilla basin (2120 m) (which has a tuff dated at 3.36 k 0.3 Ma) are
given by Hoffstetter (1986, p. 226). A femur of an edentate is also
known from near Toco (R. Cspedes, personal communication,
1989).
Near Sucre in the small Sucre basin (2800 in) "esti ubicada una
meseta que constituye la lnea divisoria de Ias aguas entre los ros
Pilcomayo y Grande" where remains of Glypfodon sp. have been
found (Ahlfeld & Branisa, 1960, p. 167; Hoffstetter, 1986, p. 226).
This meseta is part of the San Juan.de1 Oro planation surface (see
above) and the fossils are from its associated deposits.
In the Betanzos basin (3390 in) between Potos and Sucre have
been found remnids of Edentata (Glyptodontidae, Sclerocalyptinae,
Panochrltus sp.), Proboscidea (Gomphotheriidae, Cuvieronius
Ityodon = Mastodon andinun!), Perissodactyla (Equidae, gen. et sp.
indet.), and Artiodactyla, (Cervidae; Camelidae) (Pick, 1944;
Hoffstetter, 1986, p. 226). About 4 km to the south of Betanzos have
been collected Mylodontidae?, Glyptodontidae, Gomphotheriidae
and Equidae which are now in the Universidad Tomis Fras, Potos
(Anaya, personal communication, 1989). At Khoa-Paya near
Betanzos, were collected remains of Equidae from between a tuff
dated at 1.9 Ma in the base of the section and another tuff dated at

less than 1.0 Ma near the top of the section (Servant et al., 1989). A
mastodont has recently been found at the pueblo of Salitre, situated
to the east of Villazon near the Argentine border in southernmost
Bolivia (reported by Ing. Camacho to F. Anaya, personal
communication, 1989) and remains of glyptodonts are known from
Mojo (R. CBspedes, personal communication, 1989). All these
fossils come from sediments related to the San Juan del Oro
planation surface.
In the Amazonian region at Fortin Madidi on the Ro Madidi and
at Cara Cara on the Ro Maniqui (in the region called YucumoMaracas near San Borja) in the Ro Beni basin have been found
remains of Toxodontidae in the former and Mixotoxodon sp. in the
lalter (Hoffstetter, 1968a; Hoffstetter, in Marshall el al., 1984, p. 36;
Anaya, personal communication, 1989). Further to the southeast,
d'Orbigny (1842, p. 205) recovered a collection of bones of large
mammals "en las barrancas cerca de la confluencia de los ros Piray
y Grande" which were lost during the same expedition "en un vuelco
de la canoa" (Ahlfeld & Branisa, 1960). In the Ro Piray basin at
Santa Rosa north-northwest of Santa Cruz and to the southwest of
Santa Cruz, Hoffstetter (1968a) recovered mastodont teeth in 1960.
Remains of mastodonts have also been recovered by L. Branisa
(unpublished) from Quebrada Chorrillos, about 12 km northeast of
Charagua, and Trujillo (1984) reports Cuvieronius andiuni from the
Valle de Carohuaycho, 28 km north-northeast of Camiri and 30 km
west of Charagua.

644

EVOLUTION OF THE BOLIVIAN LAND MAMMAL


FAUNA
There are four interrelated phenomena that were of profound
importance in influencing and/or determining the taxonomic
composition of late Cenozoic land mammal faunas in Bolivia:
Andean uplift and related climatic changes, glaciations, the Great
American Faunal Interchange, and megafaunal extinctions.

1. Andean uplift and relaed climatic changes. The main climatic


effect of the central Andean uplift in Bolivia was the creation of: (a)
a rain shadow that prevented the northeast-proceeding moisture-rich
winds to reach the coastal areas of southern Peru and northern Chile,
and (b) dry "high"-altitude climates on the Altiplano and adjacent
cordilleras.
As demonstrated by geomorphological studies in southernmost
Peru, uplift began 20 Ma, but considerably decreased in this area
by
8 Ma (Tosdal ef al., 1984). According to Benjamin et al.
(1987), uplift rates of the Bolivian Cordillera Real were about 0.1 0.2 mm/yr between 40 and 20 Ma, and increased significantly
between 15 and 10 Ma to reach values perhaps as high as 0.7 mm/yr
by 3 Ma. Hyperarid conditions, due to a permanent rain shadow,
became established in northern Chile (24" S) in middle Miocene
time (Alpers & Brimhall, 1988). Middle Miocene climatic
dessication in northern Chile and southern Peru was accentuated by
oceanic cooling related to the contemporaneous development of the
Antarctic ice cap (Alpers & Brimhall, 1988), and this must also have
liad some influence on Altiplano climates. Although it is difficult to
calculate precisely the paleoaltitudes of the central Andes at a given
time, it seems that the mountain belt had attained at least half its
present elevation prior to about 15 Ma (Alpers & Brimhall, 1988). It
is likely that in Bolivia, Andean uplift was accelerated by the

~-

POSILES Y FACIES DE BOLIVIA - VOL. I VERTEBRADOS

tectonic load produced by the first major tectonic crisis (see above)
starting at about 27 Ma. However, there may have been more than
one uplift mechanism subsequent to 45 Ma.
Thus, the significant elevation of the central Andes during at least
the past 20 Myr must have influenced the land mammal fauna in two
ways. First, the rising central Andes came to serve as a barrier tothe
moisture-rich northeastem winds, resulting in the formation of a rain
shadow effect along its western flank. Second, ecological changes
resulted from the uplift, and true desert, high montane and extensive
grassland habitats at lower elevations came into existence during this
time. These ecological changes provided new evolutionary
opportunities for taxa which could adapt to colder high-elevation
habitats, and caused other taxa to abandon these emerging
environments in favor of more hospitable warmer environments at
lower elevations. For example, Mesotheriidae disappeared from the
Altiplano at the end of Huayquerian time, and Hydrochoeridae and
Pampatheriinae disappeared from high-elevation faunas at the end of
Montehermosan time (Hoffstetter, 1986).
The net result of Andean uplift is that it produced markedly
diverse geographic regions in Bolivia and many of the differences
seen particularly in Pleistocene faunas can be attributed to elevation
and hence ecologies. As examples, some Pleistocene taxa apparently
thrived at elevations of 3800-4000 m (i.e. Megatherium, Scelidodon,
Macraucltenia, Cuvieronius, Onohipyidium, Lama, Pantliera,
Glypfodon) and occur at low elevations as well; other taxa (i.e.
Palaeolama, Equus) are unknown above 3300 m; while others are
either unknown or extremely rare above 2000 m (i.e. Toxodontinae,
Pampatheriinae, Dasypodinae, Hydrochoeridae, Erethizontidae,
Dasyproctidae, Agutidae, Dinomyidae, Echimyidae, Tapiridae,
Tayassuidae, most Cervidae, Haplomastodon, Chrysocyon,
Smilodon) (Hoffstetter, 1986).

2. Glaciations. Four major Pleistocene glaciations in Bolivia are


recognized by Servant & Fontes (1978) (from oldest to youngest: I
Calvario, II Kaluyo, III Sorata, IV Choqueyapu 1 and 2). A tuff
(Purapurani tuff = toba Sopari of Thouveny & Servant, 1989, p. 332)
interbedded within the glacial and interglacial deposits that overlie
the Calvario Drift, has yielded a K-Ar date of 1.6 Ma (Lavenu et al.,
1989). A fifth glaciation called the Patapatani (Hoffstetter, 1986) is
reported in Pliocene sediments from below the Chijini tuff (which
has a mean age of 2.8 Ma; Lavenu et al., 1989; Marshall ef al., 1992)
of the La Paz Formation.
During times of glacial advance, ice sheets extended to about 1000
m below their present day limits (Hoffstetter, 1986). During glacial
retreat, large lakes formed on the Altiplano (see Stoertz & Ericksen,
1974; Lavenu et al., 1984) and three of these have been named: Lake
Bnllivifii was n southern expansion of Lake Titicaca (which
presently covers 8,400 km) in the northern Altiplano and covered an
area of 12,600 km; Lake Minchn and Lake Tauca occupied the
same area in the central and southern Altiplano, including presentday Lake Poop, Salar de Coipasa and. Salar de Uyuni, and had
maximum extensions respectively at 30,000 yrbp and 13,00011,OOO yrbp (Servant & Fontes, 1978). Two lake levels called Lake
Cabana and Lake Mataro, which occurred prior to the Sorata
glaciation, have also been identified (Lavenu et al., 1984). Campbell
ef al. (1985) have suggested that glacial lakes which formed during
the last glacial advance may have been united to form one large lake
which Campbell (1989) named Lake Carrasco, that covered an
excess of 150,000 km2, or nearly all of the surface area of the

645

Bolivian Altiplano. During times of glacial retreat, as today, these


lakes contracted or disappeared. The overall result is that the
climates on the Altiplano were greatly altered during times of glacial
advance and retreat.
Climates to the east of the Cordillera Oriental were likewise
effected by glacial advances and retreats. The tropical and
subtropical lowlands of eastern Bolivia experienced cold-dry
climates during glacial advance, resulting in contraction of tropical
forest biomes and expansion of savanna biomes. During times of
glacial retreat, as today, these areas had warm-wet climates with
extensive tropical forest biomes and restricted savanna biomes
(Arroyo el al., 1988). These climatic changes affected the
distribution of animals living in these habitats, and the fauna which
occurs at any one locality during times of glacial advance may be
different from that which occurs in the same locality during times of
glacial retreat. Unfortunately, the chronology of glacial advances and
retreats is not well documented in Bolivia, and the identity of glacial
faunas versus interglacial faunas has yet to be firmly established.
Nevertheless, this feature must be kept in mind when attempting to
explain differences in taxonomic composition of mammal faunas, as
they may be due to differences in ecology, altitude, and/or time.
3. Tlie Great American Faunal Interchange. Prior to about 2.5 Ma,
North and South America were separated by a narrow seaway called
the Bolvar Trough which connected the Caribbean and Pacific
Oceans across what is today northwestem Colombia and southern
Panama. About 2.5 Ma this seaway began to disappear as a result of
tectonic uplift of southem central America and/or a glacioeustatic
drop in sea level (Webb & Barnosky, 1989; Keller et al., 1989). The
Panamanian Land Bridge came progressively into existence, uniting
North and South America. The Pacific-Caribbean marine gateway
finally closed by 1.8 Ma (Keller et al., 1989). The land connection
permitted the reciprocal intermingling of the long-isolated biotasof
these continents, an event of spectacular importance known as the
Great American Faunal Interchange (Marshall, 1988).
The land mammals which walked across the newly emergent land
bridge froth North to South America included members of the
Tayassuidae and Mustelidae which are first known from rocks of
Chapadmalalan age in Argentina (Marshall, 1985). However, it is in
rocks of Uquian through Lujanian age that we begin to see in South
America a major contingent of these North American immigrants
which include: Proboscidea (Gomphotheriidae), Lagomorpha
(Leporidae), Carnivora (some Procyonidae, more Mustelidae,
Canidae, Felidae, Ursidae), Perissodactyla (Equidae, Tapiridae),
Artiodactyla (more Tayassuidae, Camelidae, Cervidae) and
eventually man (Homo sapiens) about 12 ka (Marshall, 1985,1988;
Marshall e f al., 1984). The appearance of these taxa in South
America is thus an important immigrant datum event for recognizing
faunas of Uquian through Lujanian age.

4. Megafaunal Exfinciions.The end of Pleistocene time in Bolivia,


as elsewhere in South America, is marked by the extinction of most
large body size mammals. The vast majority of these now extinct
mammals had body weights in excess of 100 kg, and because of their
large size are commonly referred to as megafauna. The peak of these
extinctions occur between 12-10 ka (Marshall et al., 1984). In
Bolivian faunas these now extinct megafauna include all members of
the families Glyptodontidae, Megatheriidae, Mylodontidae,
Megalonychidae, Macraucheniidae, Gomphotheriidae and Equidae;
and some Dasypodidae (Pampafherium),Camelidae (Palaeolama),

LARRY G. MARSHALL &THIERRY SEMPERE

Cervidae {Morenelaplius) and Felidae {Sniilodon). As demonstrated


at uapua, these extinctions occurred prior to 7200 k 400 yrbp as
indicated by the oldest carbon-14 date on the uapua 2 fauna which
includes only extant taxa. The exact cause of these extinctions is
unknown, although they have been attributed to excessive hunting
(overkill) by early man, climatic change (a sharp climatic drying
occurred in Bolivia and adjacent regions starting at -10,000 - 9,000
yrbp in the Andes and 8000 yrbp in the lowlands; Servant et al.,
1989b; Absy et al., 1991; M. Servant, personal communication), and
a combination of the two (Hoffstetter, 1986).

ACKNOWLEDGMENTS
Aspects of this study were supported by grants from the National
Geographic Socicly (2467-82, 2908-84, 3381-86); the Gordon
Barbour Fund, Department of Geological and Geophysical Sciences,
Princeton University; and the National Science Foundation (EAR8804423). Stratigraphic work was funded by the Institut Franais de
Recherche Scientifique pour le Dveloppement en Coopration
(Orstom) and realized in collaboration with field geologists of
Yacimientos Petrolferos Fiscales Bolivianos (YPFB). For help on
the geology and/or identification of specimens we thank D. Anaya,
A. Berta, R. Cspedes, M. Gayet, A. Lavenu, R. Hoffstetter, R.
Marocco, J. Oller, J. Pacheco, R. Pascual, G. Sanjins, R. Suirez &
C. Villarroel.

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