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* 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
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631
al. 1984; Lavenu, 1986; Sbrier et al., 1988). In contrast, new data
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
632
,
,
633
(*'
LI
69'
noulhsm S J O Surfoca
i. ,
A
/ of Subandaon
\,
65E
\/
N
dsformdtlon
$3'
'i.
61I
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,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
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
( 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
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
638
$GE
M ~ )&
A L T I P L A N 3
GLACIAL LAKES ASSOCIATED
FORMATIONS
LANDMAMMAL AGE
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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
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
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
640
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~~
I I
641
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
643
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
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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).
645
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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