TIIE MESOZQIC AND PALEOCENE VERTEBRATES

OF BOLIVIA
AND THEIR STB2ATIGRAPHIC CONTEXT : A REVIEJN
MIREILLE GAYET", LARRY G. MARSHALL** & THIERRY SEMPERE***

*URA 11 du CNRS, Centre des Sciences de la Terre, Lyon I,2 7 4 3 boulevard du 11 novernbre. 69622 Villeurbanne, Fraice
** Itistitute of IIunian Origins, 2453 Ridge Road, Berkeley, California 94709, USA
***Convenio YPFB-Orstom (UR IH), Sanh Cruz de la Sierra, Bolivia.
Present addiess: Centre de GCologie GCnrale et Minire,
Ecole des Mines, 35 rue Saint-HonorC,77305 Fontninebleau, France

INTRODUCTION

Dolivia lias a rich nnd diverse record of fossil verkbratcs for the
Mcso7.oic aiid early Ceno7,oic. The majority of h e 56 knowii fossil
localities for this time interval occur in the soutlierri Altiplano and in
thc soutlicrii piut d the Cordillera 01ieiitnl (Figs. 1-4).
I n Illis pnpcr we review thc iniddlc Triassic llirougli 1)nlcocciic
fossil vertebrote record o f Bolivia in its strntigrnphic and
palcociiviroririicntal context aiid attempt to correlate tliese rocks aid
faunas with those in, adjaccrit Argeiitina, Brazil, Chile arid Pcru. As
documented below, tlie highlights of the Bolivian vertcbrale rccord
include: the taxonomically ricliest aiid best studied faunas of late
Cretaceous age (from tlie marine anti continental El Molino
Formation) aiid the spectacularly rich and only well known land
maminal fauna of e d y , but not earliest. Paleocene age (from the
Santa Luch Forination a l Tiupninpa) ti all of South Atiierica. We
show that many or tlie late Cretaceous taxa (especially selachians)
are of clironostratigrapliic value within the Andean basin. The
problem of tlie K/?; bouiidary in the Andean basin of Bolivia is
critically evaluated and based on this siudy is believed to occur
within the upper part of tIie middle mcinber of the EI Molino
Formation.
The following abbreviations are used: km, kilometers; 111, nieters;
Mil, nicgnniitiuin o r tiiillioiis of years ngo, n point in h i c ; Myr,
inillioiis of ycrus, a durntioii of tiine.

Tlic "Mesozoic" s1r:itigr:ipIiy of Nolivia cornprises two distinct

tirne intervals (Sciiipcrc, 1990): tlie lirst s p i s tlie middle Triassic most of Jurassic, and the secoiid latest Jurassic-l'aleoceiie (Fig. 1).
The fluvio-eolian middle Triassic-Jurassic scdinietitary rocks wcre
dcpositcd in wh:it is now tlic castcrn liolf of tlie Andean doinairi, the
Subandcari belt and a Iargc p:ut of tlic noliviali 1owl;iiids (Ollcr &
Seinpere, 1990); their litliologies arid depositional enviroiitneiits
show similarities with coeval rocks in tlie Pnranri basin of southeni

11:

~.

Brazil, eastern Paraguay, northern Uruguay aiid northeastern

Argentinn. In contrast, the latest Jurcissic-I'nlcocenc clcpositioiial arca
was mainly restricted to tile Andean doniairi aiid bcloiigctl to thc
back-atc basin of the Pacific margin (Sctnpcre et al., 1988),
although soine late Cretaceous-Paleocene uiiits are also known in
parts of the iioitlicrn and cciifrril Suhnndcnii bclt (Figs. 1, 2).
The "Coiido cvcnt" thnt scpnriilcs tlicsc two intcrvnls ciiii be
iiiterpretcd as tlic capturc of tlic Bolivian Andean doinaiil, which
until Uien liad belonged to die crotoriic area, by tlie Pacific back-wc.
This event, acconipanied by iiiiinerous synscdimentary tectonic
manifestations (Sempere, ri press), correlates with the
Kirnnieridgian age Araucaii event in wesicentral Argentina aiid
adjacent Chile (Seinpere et al., 1988).

A. WfIDDLE TRIASSIC TO MIDDI,& JURASSIC

(Serere supersequetice)
The rich Phanerozoic rccord of Bolivia was traditionally thought
to lack Jurassic deposits. However, recent studies have shown that
certain stratigraphic units from the Subaiidean belt and from the
Andean domain should be corTelalcd and assigned a middle Triassic
and Jurassic age (Fig. 1) (Oller & Sempre, 1990; Senipere, 1990).
The Serere supersequcnce consists of these units, and includes the
Eiitrc IUos Ilnsnlt, Ipngiirv..d. Tnpecun, Cnskll6ii, lclion nnil Y nntirtri
foriiintions, as well ns lhe pnrtly cquivnlciit eu, Tiquina, Seynri nnd
Ravelo fonnations (Fig. 1). The Tacur Group of the central arid
southern Subandcau belt is distinct froin the Scicrc supersequence
bccnusc it includes tlic Intcst Crctaccous Ciijoncs Fortuntion but not
tlic Ipagunz Forination (Scinpcrc, 1990). In gcnctal, scdiincntary
continuity charactcl izcs tlic Sererc superscquciice (Oller & Senipere,
1990). As will be docuinented below, its deposits to date are sparsely
rossiliferous.
The Cnstcll6ii Fortiiatioti is probably of late Triassic-cnrly Jurassic
age (Scinpcrc & Ollcr, 1987; Ollcr 2% Sernpcrc, 1990), althougli it
lias been tentatively assigned io the early Crelaceous based on four
new species of ostracods (Pinto & Sanguiilctti, 1987). It is correlated

FUSILISS Y FACIES DE DO1,IVIA - VOL. I VEICTEURADOS

SUAREZ-SORUCO, JI
(Ed.)
.

. . ....

303

with the PirainboiQ Formation of Brazil and tlie lower part of

Tacuarembd Forination of Uruguay (Seinpere & Oller, 1987).
The lower boundary of the Serere superscquemce is a diachronous
unconformity except in the southern Subandean belt where,tlie oldest
and elsewhere unknown Ipaguaz Formation overlies the late
Permian-early Triassic Vitiacua Formation with only a slight, nonerosional, lithologic discontinuity (Sempere et al., 1990, 1992). In
other areas of the basin, younger units of the Serere supersequence
unconformably overlie Permian or older strata, reflecting onlapping
.deposition on an altered, eroded and/or deformed substratum. The
upper boundary of the Serere supersequence is the surface that marks
the "Condo event" (see above). This unconformity shows different
characteristics which relate to its location and position in the Iahst
Jurassic-Paleocene basin.
I n suinmary, tlie middle Triassic-Jurassic geologic evolution of
Bolivia involves: 1) an initial rifting process with deposition of red
beds in narrow troughs, locally with gypsum and halite (middle to
Iate Triassic); 2) termination of rifting and onlapping fluvio-eolian
scdinicntntion (late Triassic to earliest Jurnssic'?); and 3) extensive
erg development (early and middle Jurassic'?) (Oller & Sempre,
1990).

accoinpanied by synscdinientnry tcnsioniil tectonic nnd mngiiiolic

rnnnifestations (Sempere et al., 1988; Seinpere, is press) and is
termed the "Vilcapujio event" in Bolivia (Ctihvcz, 1987) or k3 in tile
central Aniles (Jaillard & Sempere, unpublished). Thus, the
Miraflores Formation stands as a specific "hinge" unit in the
Bolivian latest Jurassic-Paleocene stratigraphic sequence.
Mainly red mudstones with minor sandstones and evaporites
(gypsum, anhydrite, halite) were deposited between discontinuities
k 3 and k 4 in fluvial and lacustrine environments [Aroifilla
Formation and most of Chaunaca Formation (Sempere, in press); the
stratigraphically equivalent Torotoro Formation consists mainly of
red sandstones of fluvial origin]. Two greenish levels, which consist
of very fine sandy, green to black mark, black to grey laminated
limestones and yellow dolomites, occur in the Chaunaca Fonnation.
The foriner is more conspicuous, traditionally marks the base of the
unit, and is krmed the "basal limestone" of the Chaunaca Formation.
This 30 m-thick sub-unit commonly yields abundant bivalves
(Mytilidae, Brochidontes sp.) which have been interpreted to
indicate an intertidal cnvironnient (Briutisn ct al., 19GG). However, II
restricted mnrine origin for the "basal limcstonc" is now questioncd
(G.Camoin, personal coinunication). Identical Bracltidoort/es sp. are
recorded from a limestonc level in the Loinas Negras Formntion of
nortlicrn Chile (Mnrinovic & Litlisen, 1984). The "bnsitl linlcstonc"
II. I,h'l'lSS'l'
J UIt ASS IC 'IO l'A I,EO CICNI C
of the Chaunaca Formation yielded a palynological assemblage of
(I'ucu supersequence)
Santonian to earliest Campanian age (Prez, 1987), and may be
partly equivalent to the Snntonian age shallow-marine limestones
Classic works on the Bolivian "Cretaceous" Puca Group are by
Lohmann & Branisa (1962). Russo & Rodrigo (1965), Kriz & known from the Querque and Omoye formations of southern Peru
(Jaillard & Sempere, 1989). The Aroifilla and Chaunaca formations
Clierroni (1966). Lohmann (1970), Reyes (1972) and Cherroni
are equivalent, respectively, to the lower and middle Vilquechico
(1977).
Formation of southeast Peru (Jaillard et al., in press) (Fig. 1). The
The "Cretaceous" sequence stratigraphy of Bolivia and Peru is
upper greenish level of tlie Chaunaca Formation is equivalent to the
currently being re-evaluated and synthesized (Jaillard & Seinpere,
1989; Seinpere, in press). The Puca supersequence (Scmpere, 1990) uppennost green level of the middle Vilquechico (Jailllard et al., i n
press; Sempere, in press), which has yielded the selachian
is bound below by the kO discontinuity (;.e, the Kimmeridgian
Araucan-"Condo" event at about 144 Ma; Cowie & Bassett, 1989). Schizorhiza slronieri. Thus, its age is not older than niddle
and above by tlie k5 discontinuity which records the functional onset Campanian (Jaillard el al., in press).
The EI Molino, Santa Luca and Impora formations occur between
of western Bolivia as a continental external foreland basin of the
discontinuities
k4 and k5 i n Bolivia (Fig. 1). The EI Molino
paleo-Andes (Sempere et al., 1989). The age of W is close to the
Fonnation is equivalent to the Lecho, Yacoraite, Tunal and Olmedo
Paleocene-Eocene boundary (53 Ma; Cowie & Bassett, 1989) and
formations of northwestcrn Argentina (Fig. 1); to the Areniscas de
could b e as young as 50-51 Ma (Senipere, 1990; Marshall &
Azcar, Vivinn and Cnchiyacu fornialions of Subnndenn Peru
Senipere, 1991).
(Sempere et al., 1987: Jnillard & Sempcrc, 1989); to thc upper
I n this pnper we, divide the Pucn supersequence into three
Vilquechico Formation of the Peruvian Altiplano (Jaillard et al., i n
megasequences. The Puca A megasequence includes the partly
press) (Fig. 1); and to the Estratos de Quebrada Blanca de Poquis,
equivalent Condo, Kosmina, La Puerta S.S., Sucre and Tarapaya
niid possibly parts of thc Loinns Negras. Pnjonalcs and Toncl
forinations (Fig. l), nnd consisLs of nzoic non-marine dcposits; the
niiuinc Miraflores Fdrntntioii constitillcs the I'ucn B nicgnscquc~icc; formations i n nortlicrn Cliile (Gordcwcg 8c l<nnilre~.1985;
Salfity e/ al., 1985; MuRoz et al., 1989; Charrier & Reutter,
and the Puca C megasequence include's the Aroifilla, Chaunaca, EI
1990).
Molino, Santa Luca and Impora formations, and their stratigraphic
The basal sandstones of the El Molino Formation are partly
and teniporal equivalents (Fig. 1). Unlike the Puca A, the Puca B and
equivalent to the Lecho and Vivian formations. The thickness of
C have yielded many marine (Puca B and C) and/or continental
these sandstones in Bolivia is higlily variable and usually decreascs
(Puca C) fossils.
basinwards. Several lines of evidence suggest h a t these sandstones
The marine Miraflores Formation consists of fossiliferous
and other associated sediinent,uy rocks are of latest Campanian age
limestones and minor mudstones, up to 28 m in total thickness; it is
(Jaillard & Sempcre, 1989). The lowest part of the EI Molino
of Cenomanian and Turonian age (Jaillard & Sempere, in press).
Formation records a major marine transgression of apparent latest
Within h e Puca supersequence, the Miraflores Formation is the only
Campanian and/or early Maasfrichtian age; coeval transgressions are
unit of undisputed marine origin because it contains ammonites and
echinoids (Branisa, 1968; Reyes, 1972). It is separated from the known in many parts of South Anierica and the world (Haq et al.,
1987).
overlying Aroifilla Formation by an erosional discontinuity which is

394

FOSILES Y FACIES DE BOLIVIA - VOL. I- VERTEBRADOS

ANDEAN DOMAIN
AGE

GEOLOGIC TIME SCALE

(Haq e l a1.,1987)

(Ma)

- O W
- 60 - 9 8
20
66.5
- 70 - 54

SUBANDEAN BELT
NORTHWEST

SOUTHEAST

ARGENTINA

PERU

Danian

Cajones i~

Maaslriclilian

- 80 - 90 -

u)

- 100 -

Conlaclan
Turonian

Cenoinnrii:rii

O
Q

Lu
I-

Albian

O
10-

-1

Aptlan

-120-

-130-

131

-140-

-150-

Yantata

-160-

o
-170-

u)
u)

3
7

lchoa

-180-

lchoa

-190Liassic

-200 -210-

-220-

210

I
!

Keuper

u)
u)

-230-

-240-

5
n

I-

Figure 1. Clironostratigrapllic chart showing vertebrate-bearing formations (*) of middle Triassic-Paleocene age in Bolivia, and in
adjacent Argentina and Peru.

395

-I

MIREILLE GAYET. LARRY O. MARSIIALL &TIIIIiKRY SEMl'liKI!

Rased on recognition of large-scale transgressive-regressive

sequences, the EI M o h o Formation is divided into lower, middle
and upper membcrs which are of stratigraphic value (Jaillard el al.,
in press; Jaillard & Sempere, unpublished; $empere, in press).
Common sedimentary facies in the El Molino Formation include:
mudstones and mark (green to black or purplish, reddish); grey to
light brown limestones (micritic to bioclastic and/or oolitic);
stromatolitic boundstones referred to as Piicaliiltus by Steinmann
(1923) and Ahlfeld & Branisa (1960); fine to medium-grained
sandstones, usually calcareous and light-colored; some yellow
dolomites, often with algal laminations; very rare evaporites
(gypsum and halite casts); and a few white altered tuffs of porcelain
aspect. Mudstones, mark and thin limestones largely predominate in
the basin; sandstones, dolomites and evaporites are more common
along its borders; and tuff levels are thicker and more frequent in the
west, i.e. in the direction of the active volcanic sources located in
present-day Chile and Peru. In the basinal localities, lime turbidites
are common and where fossiliferous the paleontological materials
were dcpositcd a stibstaiitial distance from their life environinent.
Coilcordant palcocurrciit data iiidicate that the basin deepened
toward the northwest in Bolivia (Sempere er al., 1987; Seinpere, in
press), and probably connected with a marine basin in the Peruvian
Subandcnii region. Aiiiiiioiiitcs tire reported 2 in bclow the top of the
Cachiyacu Foriiiation i n the HiinIlaga aren (7"s) of iiorthern l'cru
(Vargas, 1988). In Bolivia, several fish groups, which reputedly
indicate a marine environment (see below), are known from the
lower and basal middle El Molino. Dinoflagellate cysts are present in
at least some levels of the formation in Bolivia and in stratigraphic
equivalents in northwestern Argentina (Industry, unpublished; J.
Oller, personal communication). Ammonite embryos have been
reported from tlie lower El Molino (Sempere ef al., 1987), but their
identification needs confirmation. Foraminifera (Miliolidae,
Discorbidae) are present in many levels a i d facies (G. Tronchetti, in
Camoin et al., 1991), but no detailed study has yet been made of
their taxonomy or paleoenvironrnental implications. Molluscs are
reported from various localities (e.g. at El Molino; Pilsbry, 1939),
although the systematics of these taxa is outdated and the relevance
of described species has yet to be established. Echinoids are
apparently lacking and the proportion of brackish water ostracods is
locally high (J. F. Babinot. in Cainoin et al., 1991).
A pctrogrnphic study o f the Yacoraite Forination of northwestern
Argentina led Marquillas (1985) to interpret its depositional
environinent as a restricted carbonate basin. An oligohalirie
lacustrine environment has also been proposed (Camoin ef al.,
1991). However, paleontological d a t a d o suggest that
communication of the basin with an open marine realm was frequent
during deposition of the Ei Molino Formation, but that the basin
itself was never openly marine, at least in Bolivia and Argentina
(Gayet et al., 1992, Seinpere, i n press). For reasons discussed
below, we believe that most of the El Molino Formation was
deposited principally in a restricted marine environment which
connected with a more typically open marine realm located
northwest of Bolivia (see also Sempere, in press),
The overlying Santa Luca and Impora formations were deposited
in fluvial to lacustrine environments. No marine fossils or facies are
known from these units which consist principally of red to purplish
siliciclastic sedimentary rocks. Gypsum and "gypsifred" anhydrite (J.

M. Roucliy, ,personal conimunicntion), are abundant i n the iippcr

part of Uie Santa Luca Formatioli near Potosi. Some green maris and
light-colored limestones occur in the Impora Formation of southcrii
Boliv,ia.
T h e Santa Luca and Impora formations are, respectively,
stratigraphic?lly equivalent to the'Paleocene age Mealla and Maz
Gordo formations of northwestern Argentina (Marocco ef al., 1987)
(Fig. 1). The Mealla Formation c a n b e correlated with the
Tiupampian (early Paleocene) Land Mammal Age based on its
equivalence with the Sai?ta Luca Formation (see below) and the
Maiz Gordo Formation with the Riochican (late Paleocene) Land
Manimal Age based on palynology (Volkheimer ef al., 1984) and
stratigraphic position (Pascua1 et al., 1981).

VERTEBRATE FAUNAS AND LOCALITIES IN BOLIVIA

A. TACURU GROUP
Alilfcld & Brnnisa (1960) rcportcd rcinniiis of lurgc bones,
possibly of dinosaurs, from the Tacur Group in the Serrana de
Mandeyapecua. southeastern olivia. IIIthis area the Tacur Group
includes rocks of middle Triassic and Jurassic age, naiiiely tlie
Tnpccun, Cnslcll6ii aiid Ichoii forlnntiolis (Fig. 1).

U. CASI'ELLON FORMAlION
Scales and bones of semionotiform fishes were found in the late
Triassic-early Jurassic age CastelMn Formation on the western flank
of the Charagua anticline in tlie Quebrada de Charagua, 7 km west of
the town of Chbagua (Wenz, in Goi & Hoffstetter, 1964). Nothing
definite can be said of these bones, although a study of the scales
with Scanning Electron Microscopy (Gayet & Meunier, 1986) dnd of
their transverse sections permits reference to the family
Semionotidae, probably Lepidofes sp. (Gayet, 1991).
C . MIRAFLORES FORMATION
The Miraflores Formation is a marine limestone unit which, based
on knowledge of ammonites, bivalves, gastropods and echinoids,
was regarded as middle to late Ccnoiiinnian in age (ranisa et al.,
19GG; Jaillard & Scmpcrc, 1089), :itthough detailed scqucncc
sthtigrapliy now suggcsts that the entire rock unit spans Cenoinaniaii
and Turonian time (Jaillard & Sempere, in press). Near the locality
of Macha. 90 kni north-northwest of I'otos, L. Branisa found minute
pharyngeal teeth which are tentatively assigned to cf.
Cyprinodontiforines or Pycnodontiforrnes (Gayet, 1991).
I).AROIFILLA

T h e continental Aroifilla Formation overlies the marine

Cenomanian-Turonian age Miraflores Formation and underlies the
Santonian-late Campanian age Cliaunaca Formation; it is therefore
considered Coniacian and possibly early Santonian in age (Fig:l). A
series of four footprints of a large quadrupedal dinosaur were
discovered by the Brigada 10 of YPFB at Caleras, 4 in below the
"basal limestone" of the Chaunaca Formation. The footprints were
imprinted in soft sediment and are otherwise indeterminate.

396

FORMA'I'ION

FOSILFS Y FACIES DE B o L I m - VOL. I - VERTEBRADOS

1;

PERU

,I

,4
15"

100
km

.LA

PAZ

17"

*ChuIlDa Khasa
.COCHABAMBA
La Cabana*Parotan
pa,c~s~$~ipd
Est. Blanco Rancho
ORURO.
Arapampap
d!ilaVi'a
Lago Uru Uru
Torotoro-.
Tiupampa

-Rio Moile

USANTA CRUZ
DE LA SIERRA

19'

/-

*Chaup-Khocha

,f

echaragua
I

I'

*Calerias
*Tambo
Colorado

Salar de Uyi ni

*Camargo

Serrania
de
Manieyapecua

fI

,i

*Chocaya

21"

Chaupiuno

CHILE

Rancho Hovada-.
UTARIJA
*Villa Pacheco
Serpa\

.
((PARAGUI
'\
\

\
\

'

69"

67"

65"

63"

Figure 2. Map of Bolivia stlowine:vertebrate localities of middle Triassic-Paleocene age.

397

,
\'

MIRIIILLEGAYEI'. U R R Y G. hIARSIlALLLTIUERRY SEMI'EKI!

IC. CllAUNACA IWI<MA'lION

The Santonian-middle Campanian age Chaunaca Formation has

yielded microfossils (estheries, ostracods, charo hytes) and bivalves
(Mytilidae, Bracltidon/es sp.) (Branisa et al., 966). At La Palca,
about 8 km northwest of Potos, a tooth of an indeterminate
pycnodontiforrn fish (Pycnodontidae) was discovered in a red, very
fine-grained sandstone level (Gayet, 1991). At Agua Clara, about km
95 on tlie Potos-Challapata-Oruro road, Branisa reportedly found a
hypocoracoid of the clupeiform fish Gasteroclrrpea branisai,
although this needs conluination (Branisa et al., 1964).

F. EL MOLINO FORMATION
All three members of the E l Molino Formation have yielded
diverse vertebrate faunas which were long interpreted to indicate a
late Cainpanian-Maastriclitiaii age for this entire rock unit (Cappetta
1975, 1990; Muizon c/ al., 1983; Muizon, et al., 1984a; Marshall et
al., 1985; Gnyct, i986n, 19880; Cnrrnsco et al., 1989). However.
Palcoccnc palynological asscmblagcs ore rcportcd from the Tunal
Formation of northwestcrn Argentina (Quattrochio e/ al., 1986)
which is a stratigraphic equivalent of the upper EI Molino
Forniation, and securc Cretaceous age taxa arc rccognizcd only in
thc lower rncnibcr and basal rniddlc iricrnbcr of the EI Molino (scc
below). Van Valen (1988) has argued tliat at least some, if not all, of
the faunas of the l Molino Formation are of early, but not earliest,
Paleocene age. His arguments relied in large part on the belief by
Muizon et a l . (1983) and others, that the Tiupampa vertebrates all
come from the El Molino Formation. However, recent stratigraphic
studies conclusively demonstrate that the mammal-bearing level at
Tiupampa belongs to the Santa Luca Formation (Sempere and
Marshal, in press). For reasons documented below, the EI Molino
Formation as herein defined encompasses latest Campanian-early
Paleocene time (Fig. 1).

1. Agua Clara-Potosf Region

At La Palca, about 8 km northwest of Potos, a partial crocodile
jaw identified as Doliclwcltanlpsa ntininta (Dolichochampsidae) by
Buffclaut (1987) was collcctcd froin the uppcr EI Molino. This gcnus
and spccies were named by Gasparini & Buffetaut (1980) on the
basis of specimens collected from the upper part of the late
Cretaceous-early Paleocene age Yacoraite Formation in Salta
Province, northwest Argcntina. Kcmains of Siluriformes (Ariidac,
Rhineastes sp.) wcrc rccovcrcd from thc uppcr EI Molino at the sanie
La h l c a locality (Gayet, 199 1).
From the lower El Molino, about 67 in above the base of the
formation dong the road to Cayara, about 15 km west-northwest of
Potos, disarticulated Clupeiformes (Gasreroclrrpea branisai) were
recovered froni a black shale. At Cayara in the snme lower member,
but from a different level, has come a rich selachian fauna,
including Sclerorhynchidae (Pucaprislis branisi, Ischyrhiza
harrenbergeri, Schizorhiza aff. slronieri), Dasyatidae (Dasyalis
ntolinoensis, Dasyaris schaefleri), and Rhoinbodontidae (Pucabah
hoflstel/eri) (Cappetta, 1991). The type locality of the clupeiform
fish Gasferaclupea branisai Signeux (in Branisa el al., 1964) is in
the lower member at Cerro Muyurina south of Hacienda Cayara.

398

Othcr poorly preserved and indctcrrninnte tclcosts wcrc rccovcrcd in

association with Gasteroclrrpea in the sanie grey-greeii shales
(Wenz, 1969). From the upper EI Molino, at Uic same locality, are

several calcareous and liard sandstone levels with siluriforin

(Ariidae) fisJies, turtles and crocodiles.
From Agua Clara, about GO kni norlhwcst of Potosi, nunicrous
marine and/or freshwater taxa were recovered from levels belonging
to the lower EI Molino. Just below the uppermost stromatolite level
at this locality (Ahlfeld & Branisa, 1 9 6 0 ) is a calcareous
conglomerate of limited extension with a rich assemblage of marine
and freshwater fishes including: selachians (Cappetta, 1991)
(Dasyatidae, Dasyalis nov. sp. 1 ; Sclerorhynchidae, Iscltyrliiza
hartenbergeri, Schizorhiza aff. slronieri; Rhotnbodon tidae,
P u c a b a h nov. sp.); "holosteans" (Gayet, 1982a, 1987a, 1991)
(Pycnodontiformes. Pycnodontidae indet.; Semionotiformes.
Semionotidae, nov. gen.; Ginglyniodi, Lepisosteidae, Lepisosteus
sp.); teleosteans (Brnnisa el al., 1964; Gayet, 1982b, 1982c, 1991;
Gayet & Meunier. 1983); Clupciforincs, Clupcidac, Gas~eroclripen
brnnisai; Ostcoglossiforincs, Ostcoglossidnc, iticcrlnc sedis;
"Salmoniforincs", Enchodontidae, Enchodus sp., falllily incertae
sedis 1,?Apaleodrts sp., family incertae sedis 2, gen. and sp. indet.;
Cyprinifornies. family incertae sedis, Molinicltrltys inopinafus;
Ch aro c i for in c s . Erythrin id ne. c f . Il op lins, C li nrncidnc,
Tctrngonoptcrinae; Siluriformes, Ariidnc, Rliineastes sp.; cf.
Cyprinodontiformes; Tetraodontiformes, Eotrigonodontidae,
Stephanodus niinintus; and teleosts indet.
Just above the same uppermost stromatolite level are two levels of
green marls with Qasrerocltipea branisai and smooth-shelled
ostracods. The recovery of a complete fish specimen testifies to die
absence of postmortem movement. About 1 m above the saine
stromatolite level are two calcareous beds with tiny fish bones
mostly concentrated in a 2 min-tliick layer on the top of each bed, in
which remains of Clupeiformes (Clupeidae, Gasterocltipea
branisai). Cyprinifornies (family incerrae sedis, Moliniclitltys
inopinarus Gayet, 1982b), Siluriformes (Ariidae, R/iineas/es sp.) and
Tetraodontiformes (Eotrigonodontidae, S/eplran'odtrs niininiris;
Gayet, 1991) were recovered.
Further along Uie "quebrada" are several calcareous levels in Uie
lower EI Molino with Siluriforines, and green shales with
Clupcirornies (Gas/eroc/trpcn branisai) generally associalcd with
ostracods and soinctiincs fragincnts Qf iiidetcrcniiiiitcipliiiits.
Following the road to Challapata nre three additional localities.
The first is nem Wila Khasa in a curve of the road at about kin 100,
northwest of I'otosf, which corresponds to the basnl scquchce of the
lowcr EI Molino. Bctwccn the Inst two stroniatolitc horizons is a
restricted level with a few srnall complete cf. cyprinodontiforni
fishes (Gayet, 198Ga. 1991).
A little further, on tlie left side of the road GOO m south of the
village of Wila Khasn, is thc locality Hotcl Cordillcra (Gayet, 1982~1,
1982c) where, in a grey limestone bed of the basal middle EI Molino
were found a few reptiles (crocodiles, indeterminate vertebrae) and
fishes. Selachians are represented by Pucaprisfis branisi, Iscliyrhiza
Irartenbergeri, Schizorhizn aff. sfronieri, Dasyah branisai and D.
scltaefleri (Cappetta, 1991). The teleosteans and holosteans are
identical to those of Uie lower El Molino at Agua Clara (see above),
while some teeth of Characiformes (Serrasalmidae, Myleinae) are
also represented. Nearly ali bones found in this level belong to a

Cerro
Lama Chaqui
180

Torotor

Cruz Khasa

'

Ikm

'

Figure 3. Detailed map of tlie Torotoro area (for location, see Fig. 2) showing vertebrale localities in the CI Molino and'Santa Lucia
formations. The numbers correspond to those in Table 1. Based on Carta Nacional, Bolivia, Torotoro Quadrangle, Hoja 6439 IV,
serie 1731, 1/50,000, 1968 edition, Instituto Geogrhfico Militar, La Paz VIII-68; and San Vicente Quadrangle, Hoja 6439 I, serie
fi731, ,1/50,000, 1968 edition, Instituto Geogrhfico Militar, Ln Paz 111-68.
siluriforin (Ariidae, Rliineastes sp.) of very large size (more than 2 in
in total length). Vertebrae of Lepisosfeus sp. found in association
with the Siluriforines are also larger (more than 3 cm-long) than
those of Agua Clara (about 1cm-long). As in some levels at Agua
CI:lra, thc presence of sclaclii:ins, pycnodonti forms, tetradontiforins
and "saliiioniforins" (Enchodirs sp., ?Apateodirs sp.) testifies to some
conmunication to a sea (Gayet, 1991; Gayet et al, 1992).
Ten meters below the calcareous level just described, a restricted
conglomeratic unii was recently found with reinaius of Siluriformes
(Ariidae) and Teleostean indet. A little farher, at the crossing of the
Quebrada Saytu JoWiu with the road, are grey-green mudstones of
the upper EI Molino wilh remains of Clupeiformes (Casteroclitpea
branisai) and ostracods (Gayet, personal observation).
At Vilcapujio (=Wila Apacheta), at about hi 140 northwest of
Potosl on Uie Potosl-Challapata-Oruro road, is a bone bed probably
belonging to the basal middle EI Molino which yielded fishes

399

(Pycnodontiformes; Characiformes, Serrasalmidae, Myleinae;

Siluriforines, Ariidae, Rliineastes sp.) and reptiles (turtles and
crocodiles). In the vicinity have also been found several levels with
holostean Ginglyinodi (Lepisosteidne), teleostean Clupeiforrnes
(Clupeidae, Castcroclicpea Invnisai), Chnraciforines (Serrasahnidae,
Myleinae). Siluriforines (Ariidae, Rliineastes sp.), and/or reptiles
(crocodiles, turtles).
At Sevaruyo. south of Lake Poop6. about 120 kin west-norU!west
of Potos, are reported two levels with remains of Casteroclitpea sp.
(Branisa et al., 1964). The first level apparently belongs to Uie lower
EI Molino. Because of the large.(double) size of the hypocoracoids
found in the second level relative to .CasteroclupeaS holotype,
Branisa ef al. (1964) questioned the age of this level. However,
hypocoracoids of very large size have also been found at Agua Clara
where a transitional series exists that includes all sizes recorded at
otlier localities.

MIREILLE CIAYET, LARRY CI. M R S I U L L & 11 UliKKY SBMIEKB

2. Uyuni Region

At thc locnlitics of Jay-Jay, Calnzaya and Tomave, rcspcctively

north-northwest, east and northeast of Uyuni, were found remains of
Siluriformes (Ariidae, Rlrineasres sp.; Tomave only) and
Clupeiformcs (Gasreroclupea branisai) by the$rigada 10 of YPF
(specimens in CTP, Santa Cruz). At Tambo Colorado, squtheast of
Uyuni, was found a selachian (Pucapristis sp.) associated with
gastropods (Melania potosiensis; specimens in CTP, Santa Cruz).
With the exception of Calazaya which is upper El Molino, the
stratigraphic position of fossils at the other localities have not yet
been established.
3. Southern Region

In the Ro Angosto, 9 kin south-southwest of Chocaya, holostean

Ginglymodi (Lcpisosteidae, Lepisosteus sp.), Semionotiformes
(Semionotidae, nov,. gen.), teleostean Clupeiformes (Clupcidae,
Gasteroclirpea brunisni) and Siluri formes (Ariidae, Rltineasres sp.)
wcrc recovered froni,the lower EI Molino.
In Quebrada Tcxisca, 1 km west of Rancho Hoyada along the.Ro
San Juan del Oro, 29 kin north-northeast of Tojo, were recovered
rcninins of lioloslcnii Iyciiodontiforiiics (Pycnodontidac. indct.),
Ginglyinodi (Lcpisostcidae, Lepisosrens sp.), teleostean
Osteoglossiforrnes (Osteoglossidae, indet.), Siluriformes (Ariidae,
Rhineastes sp.) and Tetraodontiformes (Eotrigonodontidae,
Sfephunodirs niinkirs) (Gayet, 1991) from the lower EI Molino. The
saine Pycnodontidac,
Osteoglossidae, Ariidae and
Eotrigonodontidae, plus two new selachians (Dusyuris nov. sp. 2 et
3; Cappetta, 1991) and a crocodile tooth were recovered from the
basal middle El Molino at the same locality.
Fish remains, including Pucaprisfis brunisi, have also been found
by J. Blanco (YPFB, Brigada 10) near Serpa about 35 km south of
Tupiza, and dinosaur trackways were observed by Leonardi (1981)
near Camargo, 100 kin north of Rancho Hoyada.

4. Cochabamba Region
At Torotoro, aboui 95 kin south-southeast of Cochabamba, several
fossil lcvcls NC known (Fig. 3).From thc iniddlc part of thc lowcr EI
Molino sonic 3 kin soulti-soutlicnst of Torotoro, Cnppctla (1975)

dcscribcd numerous sclachians [Sclerorliyncliidae, fitcapristis

brnisi, Iscliyrhiza Irnrtenbergeri; Dasyatidae, Dasyaris branisai, D.
niolinoensis, D. schnefleri; Khoinbodontidac, Pucabah Imffstelleri
(Cappetta, 1987)J and Broin - ( 1991) recorded turtles
(Podocneinididac, ?Roxoclielyssp.).
In a higher level in the same inember, on the northeast side of
Torotoro, numerous dinosaur trackways were found that have been
callcd the pista dc danzas (Fig. 3).At lcast five types of dinosnurs
are represented, including sauropods, tlieropods and ornithopods
(Branisa, 1 9 6 8 ; Leonardi, 1981); o n e trackway named
Ligabueiclrniunr bolivianrtnr represents an ankylosaur or ceratopsian
(Leonardi, 1984).
Near the base of the lower EI Molino at Unia Jalanta, close to Uic
cave entrance, about 5 kin west-northwest of Torotoro, occur some
small tridactyle (Coelurosnuria?) footprints and isolated bones of
turtles (?Roxuclre[yssp.). At Cerro Llania Chaqui, about 4.3 km east

of Torotoro. occur scvcrnl scts of trackways of a sinall theropod

dinosaur (Coclurosaurin?). The siinic typc of trnckway occurs ncar
Uic top of tlic lowcr EI Molino on thc p t l to
~ Ulna Jetanta, about 2
km west of Torotoro.
From the transgressive base of the middle El Molino in the Ro
Cuchira Waykho, 3 kin south-southeast of Torotoro, a long rostrum
of a salmohiform (family indet. 2, gen. and sp. indet.; Gayet 1991)
is assigned to the marine Cretaceous suborder Ichtliyotringoidei
(semu Goody, 1969).
From an undetermined member at Cruz Khasa. on the west side of
the cemetary south of Torotoro, remains of Prtcapristis sp.,
crocodiles and turtles (?Ro.xoclielys sp.) have been recovered (R.
CCspedes, personal communication; specimens in Museo de Historia
Natural, Cochabamba).
At Tiupampa, located about 95 kin southeast of Cochabamba, two
fossil levels are known (Fig. 4). The lowest, near Hera Mokho, is
about 9 0 m above the base of t h e Cretaceous section from a
calcareous sandstone horizon of the middle EI Molino (see Marshall
et al., 1985. Fig. 4); vcrtcbrntcs nrc rcprcscntcd only by a sclochinn
(Dasyatidae, flasyaris sc11aejJeri) and an indctcrrninatc crocodile
(Marshall et al., 1985). There a r e also abundant gastropods
(Melanidne, Melania potosicnsis) which unTortunulcly nrc not useful
for ngc rcsotulion (Ahllcld & A r i i n i s n , t960). t4iglicr i n lhc scction,
along the Rio Pucarani, is an uppcr EI Molino bone bed with
Siluriforines (Ariidae, Rhineastes sp.), indeterminate turtles and
crocodiles.
At Vila Vila (=Villa Viscarra), about 9 0 krn southeast of
Cochabamba and 6 kin northwest from Tiupampa, a fossil level in
the lower El Molino about 100 ni above the base of the Cretaceous
section (see Marshall el al., 1985, Fig. 4) has yielded some of the
saine selachians recorded by Cappetta (1975) from t h t lower El
Molino at Torotoro (Prtcuprislis branisi, Isclryrliiza hartenbergeri,
Pircabaris hoffsfefreri);numerous actinopterygian fishes: holostean
Pycnodontiformes (Pycnodontidae, indet.), Semionotiformes
(Semionotidae. nov. gen.); teleostean Characifornies (Serrasalmidae,
Myleinae). Siluilformes (Ariidne, Rlrinensres sp.. Andinichthyidae
indet.), cf. Cyprinodontiformes; Brachiopterygii (=Cladistia),
Polyptcriforines (Polypteridae, Dnjerelln srrdmnericnnn; Gayet &
Meunier, 1991a, b. c); Dipnoi (Lepidosirenidne, Lepidosiren cf.
paradma; Schultzc, 199In); n turtlc (Iodocllciiiididrlc. ?I?o.toc/wI.ys
cf. v i l ~ ~ i l ~ t ~ fjroiii,
s i s ; 199I),und two iiidctcrniinotc ciocotlilcs
(Marshall et al., 1985).
Two new localities were discovered during the 1989 field scnson,
Estancia lnnco Rancho and Iiijcha Patii, niidwoy along the CliznAnznldo road, about 40 kin southeast of Cochabanibn. At Pajchn
Pata (lower EI Molino) were recovered reniaiils of selachians
(Pucubaris Iroffstetreri, Pucaprisfis brunisi), holostean
Pycnodontiformes (Pycnodontidac, Coelodrcs toncoensis;
Lep is o s tc id tic, Lep isosf e I I s s p. ; g II II o id li o 1os te nn sc a ICs ) ,
Clupeifornies (Gasreroclrrpea branisai), Siluriformes (Ariidae,
Rhineasfes sp.; Andinichthyidae, indet.), cf. Cyprinodontifoniies,
numerous bones belonging to as yet unidentified small teleosteans,
lungfish teeth (Lcpidosirenidae, Lepidosiren cf. L . paradoxo),
amphibian Anura, one vertebra which could be an atypical Urodela,
two other vertebrae which have the morphology of a Urodeln but are
procoelic (the vertebrae of Urodela are always ophisto- o r
amphicoelic), probably a snake, a tooth of a sninll tlieropod

400

Figure 4. Detailed map of tlie Tiupampa area (for location, see Fig. 2) showing localities (1-9)from the principal vertebrate level of Ille
Santa Luca Formation, a n d localities in lhe middle (A) and upper (B) El Molino Formation. Tiupampa is the local name used by
Quechua indiuns for tlie urea shown on lhe mup. Bused on Cnrln Nacionul, Ihliviu, Sun Vicenle Quudrungle (see cuption lo Fig. 3).
(Coelurosauria; Marshall,, 1989b), indeterminate turtles. various
crocodiles nnd indeterminate trackways. At Blanco Rancho (upper El
Molino) were found Clupeirormes (Gusleroclupeu brunisui).
Siluriformes (Ariidae, RltineasfPs sp.), cf. Cyprinodontilormes,
turtles and numerous ostracods.
Trackways of small bipedal dinosaurs (Coelurosauria?) are
recorded from Abpampa at Cerrito de Llamachaqui, Departrnent of
Polosi (Leonardil 1981).
Fragments of large bones, possibly of dinosaurs, and "dientes de
tipo cnico simple" were observed in a level with gastropods in tlie
El Molino Formation near La Cabaa, north of Parotani and west of
the Ro Rocha about 35 km southwest of Cochabamba (Ahlfeld &
Branisa, 1960).
Leonardi (1981: 935) reports a series of six poorly preserved
footprints of a bipedal dinosaur in the Santa Luca Fonnation along
the Cochabamba-La Paz road near Parotani in the Suticollo Syncline,

soutlieast of .Cochabamba. These are apparently the sanie foolprints

the lower EI Molino
described by Mnrshall & Molina (1990)
Formntion 4 km south-soulhenst of Satitivafiez rind 10 kin east of
Parotani: The best preserved footprints appear referable to an
ornithopod, probably of the family Hadrosauridae, while
others may represent a small theropod, possibly a
coelurosaur.
From Sayari at km 87 on the road from Cochabamba to Oruro, R.
CCspedes recovered remains of a selachian (Pucupristis brrurisi), a
siluriform (Ariidae, Rhineusles sp.) and an indeterminate turtle from
the lower EI Molino (specimens in the Museo de Historia Natural,
Cochabamba).
Chullpa Khasa, 6 km southwest of Morochata, has yielded a fish
mandible (Osteoglossiformes, Osteoglossidae, Phareodontinae) and
remains of reptiles (turtles, crocodiles) from an undetermined EI
Molino member.
401

Figure 5. Stratigrapliic sections of late Cretaceous-Paleocene rock units at Tiupampa, Pajclia Pata (+ Estancia
Ulanco Rancho), Torotoro (+ Caranota),' Potosl (road to La Falca and Cayara) and, in part, Agua Clara, showing
location of main fossiliferous levels. Undecompacted thicknesses. The base of the unconformable Cayara
Formation, is held horizontal. Fm, Formation; I, lower; m, middle; u, upper; Pz,Paleozoic: 1
,erosional surface; 2,
conglomerates; 3, conglomeratic sandstones; 4, sandstones; 5, fining and/or thinning-upward; 6, siltstones and
mudstones; 7, carbonates (mostly limestones); 8, coarsening and/or thickening-upward; 9, gypsum; 10, tuffite
bed; 1
1
,stromatolitic level; 12, dissolution breccia; 13, channels; 14, cross-bedding; 15, oiiids; 16, calcareous
cement; 17, rooting; 18, coalescent rhizolith-rich levels; 19, bioturbation; 20, red color predominant; 21, dinosaur
trackways; 22, vertebrate-bearing levels. All sections measured by T. Sempere, except parts of TorotoroCaranota and Potos (respectively by tlie Brigadas 10 and 9 of YPFU) and of Pajclia Pata and Agua Clara (by G .
Camoin and J. M. Rouchy); all were checked by the authors.
402

FOSILES Y FACIES DE BOU-

A vertebrate fauna descrilxd originally as of possible Palcocelie

age was reported forti Uie northern flank of tlie Huarachani syncline,
just north of tlie pueblo Huarachani along the Bolivia-Peru border
north of Lake Titicaca (Argollo el al., 1987). The fauna includes a
fish (Osteoglossoitiorplia, Osteoglossidae, Phareodontinae; Gayet,
1987b, 1991) and a crocodile vertebra (Mesosuchia, Dyrosauridae,
gen. arid sp. indet.). However, these fossils come from the middle
nieinber of the El Molino Formation (called Ococoya Formation in
this area; Fig. 1), and are therefore of late Cretaceous age (Martinez,
1980: 183; Gayet, 1987b; Mourier el al., 1986, 1988: 171; see
below).
6 . Northern Subaridean belt
About GO-70 in above the base of the late Cretaceous age EslaMn
Formation (=Flora Formation s.1. of Perry, 1963), from levels that
cnii bc correlated to the lower El Molino, nloiig the Rlo Flora, 315
kin north-northwest of La Paz, were collected rcmains of
Pucaprislis sp. (=anclioprisris), Gaslerocllrpea brunisai and
charophytes (including Peckichara compressa) (Perry, 1963; DBvila
& Ponce de Lc611, 1971). I n the saine area, Gasferoclupea branisai
was recovered froin the overlying Flora Forination S.S., an quivalent
of the upper El Molino (Vernet & Bokllo, 1975).

7. Central Subandeen belt

Sanjins-Saucedo (1982) and M p e z (1983) reported remains of
Gasteroclupea branisai from the Cajones Formation (partly or
totally equivalent to the EI Molino Formation; Fig. 1) in the Moile
Syncline along tlie Ro Moile about 95 kin west-northwest of Santa
Cruz. The level in which the fossils were foulid probably represents
the upper El Molino.

G. SANTA LUCIA FORMATION

The richest vertebrate fauna, in ternis of both number and quality
of specimens, is from Tiupampa about 9 5 k m southeast of
Cochabamba (Figs. 2, 4). Two fossil levels at Tiupampa belong to
the basal iniddlc and upper EI Molino (sec nbovc). A third fossil
level is located about 130 in above the base of the Cretaceous section
(see Marshall el al., 1985, Fig. 4). Sempere & Marshall (in press)
dcinonstratc that this innin rossil level, which represents the type
fauna of the Tiupainpian Land Mammal Age (Marshall. 19890)
belongs to the Santa Luca Forniation.(see Figure 5). The following
vertebrate taxa are recorded:
a) Fishes (Gayet. 1988b. 1990, 1991; Gayet & Meunier, 1991a.
199 1b, 1992; Schultze, 1991a): teleostean Clupeiforrnes (Clupeidae.
Gasteroclupeinae, Gasreroclupea branisai), Osteoglossiformes
(Osteoglossidae, Phareodontinae, Pliareodusichfltys favernei;
Osteoglossinae. incerfae sedis) (Hiodontidae sensu Muizon el al.,
1 9 8 3 is a confusion with Osteoglossinae; Gayet, 1991),
C h a r a c i f o r m e s , (Characidae, Tetragonopterinae, indet., cf.
Rhoadsiinae; Serrasalmidae, Serrasalminae, Myleinae; Erythrinidae,
Hoplias nov. sp.). Siluriformes (Ariidae, Rhineasles sp.;
AndinichUiyidae, Aridiniclllliys bolivianensis; families incerrae sedis

403

- VOL. I - VERIBBRMOS
1 and 2 , IfofJsfertericltrltyspucai and Incaiclirhys srrarezi),
Perciformes (Centropoinidnc), Polyptcriforines (Iolyptcridne,
Dajelella sudanicrictmn) and Dipiioi (Ccratodoiitidnc, Ccralodrts sp.
and cerntodont n. g., n. sp.; Lepidosirenidne, Lepidosiren cf.
paradoxa).
b) Amphibians (Rage, 1986, 1991a): Anura (Leptodactylidae);
Gymnophiona (family indet.).
c) Turtles (Broin, 1988, 1991): Podocnemididae, Roxockelys cf.
vilavilensis.
d) Lizards (Rage, 1991b): Lacertilia, ?Iguanidae and family indet.
e ) Snakes (Rage, 1991b): Aniliidae n. g.; Boidae, two species
indet.; ?Maqtsoiidae; Tropidopliiidae.
f) Crocodiles (Buffetaut, 1991; Buffetaut & Marshall, 1991):
Mesosuchia, Sebecidae, Sebecris querejazus; Dyrosauridae,
Sokorosucltus aff. iwiwilsoni. .
g) Mammals (Muizon el al., 1983; Muizon, el al., 1984a, b;
Marshall el al., 1983a, 1985. 1989; Muizon & Marshall, 1985,
198711, b, c. d, 1988, 1991. in press; Marshall, 1992: Marshall &
Muizon, 1988, 1992): Dcllotlicroidn (family indel,, J a ~ k h ~ d ~ / p l
ntinufus), Peradectia (Ieradcctidae, Peradectinae, Pcradcctes
a u s ~ r i n ~ n ~ ;Caroloainegliiiiiinne,
Roberllioffs~elleria
tiariotialgeograpliica), Microbiotheria (Microbiotheriidae, Kliasin
cordi llerensis), Did e l pli i ni or p Ii ia (Did e 1pli id oc, Did el pli i ri ne,
Piicadelpltys andinris, Incudelpliys anriqrrus, Mizquedelphys
pilpinensis; Eobrasiliinae, Tirilordia floresi), Sparassodonta
(H a t h 1i ac y n id a e, A I l qo k ir 11s a us I r a 1i s ) , Pa u c i tub e r c u 1at a
(Kollpaniidae. Kollpnnia fiupanipina), order and family indet.
(Andinodelpliys cochabanibensis), Leptictida (Palaeoryctidae?, cf.
Ciniofesles sp.), Pantodonta (Pantolambdidae, Alcidedqrbignya
inopinafa), Condylarthra (Hyopsodontidae, Mioclaeninae,
Tiuclaenus niinulus, Molinodus suarezi, Pucanodus gagnieri,
Andinodus boliviensis), and Notoungulata (cf. Henricosbomiidae or
Oldfieldthomasiidae, gen, et sp. indet.).
Criadero de Loro, a new locality about 1.2 km soutii-southeast of
the quarry at Tiupampa and from the sanie horizon (Fig. 4, locality
7). was discovered in 1989. The taxa seem to be the scarne as those
from the quarry and adjacent localities (Fig. 4, localities 1-6, 9) and
include, Siluriformes (Ariidae. Rhineasles sp.), Perciformes
(Centropornidae), Polypteriformes (Polypteridae, Dajetella
sudamericana), nu iiicrous 1a rgc lu iig fis li t ce t h (includi ti g
Ceratodoiitidiic and Lepidosircnidac), snnkcs (oidac sp. indct.),
turtles (?Roxochelys vilavilensis), crocodiles and a pantodont
(Alcidedorbignya inopinafa).
At Viln Vilb, om tin u p p x lcvcl lociilcd nbout LGO ni above the
base of tlie Cretaceous section (see Marshall el al., 1985, Fig. 4),
have come a turtle (Podocnemididae, ?Roxochelys vilavilensis;
Broin, 1971) and a crocodile (Sebecosuchia, Sebecidae, Sebecus
querejazus; Buffetnut & Marshall, 1991).
At Uie two newly discovered localities of Estancia Blanco Rancho
and Pajcha Pata, south of Cliza (Fig. 2), are bone beds of the lower
and/or upper members of the EI Molino described above, which are
overlain by red sandstones of the Santa Lucia Formation similar Lo
those at Tiupampa (Montao, 1968). A brief survey of Blanco
Rancho resulted in the recovery of numerous teleostean fishes
(Osteoglossiformes, Osteoglossidae; Siluriformes, Ariidae,
R liin ea st es sp., A n d i ni c li thy id ae , i nd e t er m i na te te leo st s),
amphibians, crocodiles (Dyrosauridae, complete skulls), turtles

MIREILLE CIAYET. LARRY O. MARSIMLL & IIUBKRY,SEMIEKI!

TABLE 1. VERTEBRATE FOSSIL LOCALITIES OF MESOZOIC AND PALEOCENE AGE 1N BOLIVIA ARRANGED BY
STRATIGRAPHIC OCCURRENCE.

UPPER MEMBER

TACURU GROUP
1. Serrana de Mandeyapecua

31. Calazaya
32. Cayara (Cerro Muyurina)
33.* La Paka (Doliciwciranipsamininia level)
34, Pajcha Pata
35. Estancia.Blanco Rancho
36. RoFlora
37. Ro Moile
38. Tiupampa (Ro Pucnrani)

CASTELLON FORMATION
2. Quebrada de Charagua
MIRAFLORES FORMAIION
3. Macha
AROIFILLA FORMATION
4. Calerlas (dinosaur footprints)

INDEIXIMINAIE MEMUER
CHAUNACA FORMATION
5. Aguaclara
6. La Paka

39. Arapampa (dinosaur footprints)

40. Camargo (diiiosiiur footprints)
41, Chullpn Khasa
42. Jay-Jay
43. Serpa
44. Tambo Colorado
45. Toinave
46. Torotoro (Cruz Khasa)

EL MOLINO FORMATION
LOWER MEMBER
7. Aguaclara
8. Cayara (Cerro Muyurina)
9. Chocaya (Ro Angosto)
10. Hotel Cordillera near Wila Khasa; Quebrada Saytu Jokhu with
Gasteroclupea
11. La Palca (black shale level with Gasteroclupea, and below
stromatolites)
12. Pajcha Pata
13. Kanclio Hoyada (Quebrada Texisca)
14. Sayari (km 87)
15. Scvaruyo
16. Torobro (Ceiro Llama Chaqui. dinosaur footprink)
17. Torotoro (Uma Jalantn cave, dinosaur Foolprints)
18. Torotoro (Uma Jalantn road, dinosaur footprints)
19. Torotoro (pista de danzas. dinosaur footprints)
20. Torotoro (selachian level)
21. Vila Vila (Piccaprisky icvel)
22. Wila Kliasa (km 100)
23. Ro Flora
I
24. Parotani (=Santivaez) (dinosaur footprints)

SANTA LUCIA FORMATION

47. Chaupi Khocha
48. Maragua
49. Pajcha Pata
50. Estancia Blanco Rancho
5 1. Tiupampa (quarry and associated localilies)
52. Tiupampa (Criadero de Loro)
53. Torotoro
54. Vila Vila

IMIOKA FORMATION
55. Cliaupiuno
56. Villa Iachcco

MIDDLE MEMBER

25. Hotel Cordillera (near Wila Khasa)

26. Huaracliani
27.
28.
29.
30.

Rancho Hoyada (Quebrada Texisca)

Tiupampa (Hera Mokho)
Torotoro (Ro Cuchira Waykho)
Vilcapujio m i l a Apacheta)

404

TAULE 2. SYSTEMATIC LIST OF VERTEBRATES FROM THE MESOZOIC AND I'ALEOCISNE O P

UOLIVlh WITH AUTHORSIIII'.

CLASS SELACHU
Order Rajiformes
Suborder Sclerorhynchoidei
Family Sclerorhynchidae
Pucaprisris branisi Schaeffer, 1963
Iscltyrltiza Iturtenbergeri Cappetta, 1975
Scltizorkiza
slronwri Cappetta, 1975
' Order Myliobatiformes
Family Dasyatidae
Dasyaris branisai Cappetta, 1975
Dagtis molinoensis Cappetb. 1975
Dasyatis scltucfleri Cnppeltn. I975
Dasyaris nov. sp. 1
Dasyatis nov. sp. 2
Dasyalis nov. sp. 3
Family Rhoinbodonlidnc
Pucabah ltoflslelteri Cnppclln. 1975
Pucabah nov. sp.

un.

CLASS OSTEICHTHYI
Subclass Actinopterygii
Superorder "Holostei"
Order F'ycnodontiormes
Family Pycnodontidae
Coelodus roncoensis Benedetto & Sanchez, 1972
Pycnodontidae indet.
Order Semionotiformes
Family Semionotidae
Lepidoles sp.
nov. gen.
Order Ginglymodi
Family Lepisosteidae
Lepisosteus sp.
Superorder Tclcostci
Order Clupciforincs
.
Family Clupeidae
Gasreroclupeo branisai Signeux in Branisa et al. 1964
Order Osteoglossiformes
Fnrnily Oskoglossidse
Subfamily Phareodontinae
Pkaerodusiclitlzys tavrnei Gayet, 1991
Subfsinily Osteoglossinae
nov. gen.
Order "Salmoniformes"
Family Enchodontidae
Emhodis sp.
Suborder ichthyotringoidei
Family incertae sedis 1
? Apdeodus sp.

Family incertae sedis 2

gen. and sp. indel.
Order Cypriniformes
Family incertae sedis
Moliniclitliys inopinatus Gayct, 1982c
Order Characiformes
Family Erythrinidae
Hoplias nov. sp.
cf. Hoplias
Family Senasalmidae
Subfamily Myleinae
gen. and sp. indel.
cf. Subfamily Serrasalminae
gen. and sp. indel.
Family Characidae
Subfamily Tetragonopterinae
gen. and sp. indel.
cf. Subfamily Rhoadsiinne
gen. and sp. indel.
Order Siluriformes
Family Ariidae
Rliineastes

Superfamily Andiniclithyoidea
Family indef.
nov. gen.
Family Andinichthyidae
Andiniclttliys bolivianensis Gayet, 1988b
Family incertae sedis 1
Incaichfhyssuarezi Gayet, 199Ob
Family incerlae sedis 2
Hoffsfettericlitkys pucai Gayet, 19Wb
at least five gen. and sp. nov.
Superorder Atherinomorpha
cf. Order Cyprinodontiformes
gen. and sp. indel.
Superorder Acnnthopterygii
Order Perciformes
Suborder Percoidei
Family Centropomidae
gen and sp. indel.
Order Tetraodontifonnes
Family Eotrigonodontidae
Slepl~anodrrsniinintus Gayet, 1991

Subclass Sarcopterygii
Order Dipnoi
Family Ceratodontidae
Ceratodirs sp.
gen. and sp. indel.
Family Lepidosirenidae
Lepidosiren cf. paradoxa

CLASS CLADISTM (= BRACHIOPTERYGII)

Order Polypteriformes'
Family Polypteridae
Dajetella sudaniericana Gayet & Meunier, 1991

406

FOSILES Y W a l s DE DOLMA

- VOL 1- VERIHBRAWS

CLASS AMPIIIUIA
Order A n u a
Family Lepldactylidae
gen. and sp. indel.
Order Gymnophiona
Family indel.
gen. and sp. indet.
Order Urodela
Family indel.
gen. and sp. indet.
CLASS REPTILIA
Order Ch,elonia
Family Podocnemididae

? Roxoclielys vilavilensis Broin, 1971

? ROXOC~ICIYS
CJ vilavilcrisis
Order Squaiiintn

Suborder Lncertilia
Family ?Iguanidae
gen, and sp. indel.
Family indet.
gen. and sp. indel.
Suborder Ophidia
Family Aniliidae
gen. and sp. indet.
Family Boidae
gen. and sp. 1indel.
gen. and sp. 2 indel.
Family ?Madtsoiidae
gen. and sp. indel.
Family Tropidopheidae
gen. and sp. indel.
Order Crocdilia
Suborder Mesosuchia
Family Sebecidae
Sebecus querejazus Buffetaut & Marshall, 1991
Family Dyrosauridae
Sokotosukus aff. ianwilsoni Halstead, 1975
Suborder Eusuchia
Family Dolichochampsidae
Dolickoclmnpsa niininla Gasparini & Buffclnut, 1980
Order Saurischin
Suborder Tlieropda
Infraorder Coelurosauria
gen. and sp. indet.
Suborder Sauropodomorpha
Infraorder Souropoda
gen, and sp. indel.
Order Omithischia
Suborder Ornithopoda
gen. and sp. indel.
Suborder Ankylosauna or Ceratopsia
Ligabueiclinium bolivianum Leonardi. 1984
Order indef.
gen. and sp. indel.

CLASS MAMMALIA
Infraclass Metatheria
Order Deltatheroida
Family inder.
Jaskliadelphys niinutus Marshall & Muizon, 1988
Order Peradectia
Family Peradectidae
Subfamily Peradectinae
Peradectes austrinuni (Sig, 1971)
Sublamily Caroloamegluniinae
RobertlioffstetferianafionalgeograpliicaMarshall et al., 1983
Order Microbiotheria
Family Microbiotheriidae
Khasia cordillerensis Marshall k Muizon, 1988
7 Order 'Didelphimorphia
Family Didelphidae
Subfamily Didelphinae
Pucadelpliys andinus Marshall & Muizon, 1988
Incadelpliys untiqrrrrs M ~ ~ l i a&l lMuizon. 1988
M i z y r t e d e l ~ ~ l i y s p i l ~ ~ iM
n e~~~i sl i i &
i l lMuizoii, 1988
Subfamily Eobrasiliinae
Tiulordiafloresi Marshall & Muizon, 1988
Order Sparassodonta
Family Hathliacynidae
Aflqokirrlsaustralis Marshall & Muimn, 1988
Order Polydolopoidea
Family Polydo Iopdae
Epidolops sp.
Order Paucituberculata
Family Kollpaniidae
Kollpania fiupanipinaMarshall & Muizon, 1988
Order indef.
Family indef.

Andinodelpliys cocliabunibensis Marshall & Muizon, 1988

Infraclass Eutheria
Order Lepticrida
Family Pnlnporyctidae?
cf. Ciniolesres sp.
Order Pantodonta
Family Pantolambdidae
Alcidedorbignyn inopinafa Muizon & Marshall. 1987a
Order Condylartlua
Family Hyopsodontidae
Subfamily Mioclaeninae
Tiuclaenus niinirfus Muimn & Marsliail, 1987b
Molinodus suarezi Muizon & Marshall, 1987c
Andinodus boliviensis Muizon & Marshall, 1987c
Pucanodus gagnieri Muizon &.Marshall, 1991
Order Notoungulata
Family cf. Henricosbomiidae or Oldfieldtliomasiidae
gen. and sp. indef.

408

~~

~
~

TABLE 3.SYSTEMATIC LIST OF VERTEBRATES FROM THE MESOZOIC A N D PALEOCENE OF BOLIVIA WITH INDICATION OF LOCALITIES AND
STRATIGWHIC OCCURRENCE BASED ON INFORMATION IN TABLE 1 AND TEXT. Abbreviations: Ca. Castelln Formation; Mi, Mirdores Formation;
Ar, Aroifilla Formation; Ch, Chaunaca Formation.

TAXA

EI Molino Formation
Ca
Fm
-

Mi

Ar

Ch
Fm

Fm Fm
-

h s Sclachii
Order Rajiforms
Suborder Sclemrhynchoidei
Family Sdcmrhynchidae
Puraprisrir branis
lschyrhiza harrenbergen
Schizorhiza 4.srromcri
Order Myliobatifonnes
Family Dasyatidae
D q a r i s bradsat'
D q a r i s molinoemis
D q a r i s Schaeffer
D q a t i s nov. sp. 1
Dasyaris nov. sp. 2
D q a t i s nov. sp. 3
Family Rhombodontidae
Purabarir hoffsreneri
Purabatis nov. sp.

lass Ostcichthyes
;uklass Actinoptcrygii
Maclass "Holostei"
Order Pycnodontiformes
Family Pycnodontidac
Coelodur roncoqsis
gen. and sp. inder.
Order Sanionotiforma
Family Smiqnotidac
Lepidores sp.
gen. and sp. nov.
Order Ginglymodi
Family Lepisostcidac
LepisosteUr sp.

nfmclass Teleostei
Order Clupciformes
Family Qupcidae
Gasreroclupeabranisa

lower member

middle mb

11.12 14.20.21

2.25
25

7,11,20,21
7.11

Santa Luca

upper member

Fm

25

20

25

11.20
11.20

25.28

7
27

n
11.20.21
7

3?

12
7.12.13.21

25,27,30

S?

----

1.9.21

25

7,9,13

25.21

7.8.9,10,11,12,.15

25

3235.36, 37

51

Impora
Fm

El Molino Formation
~

TAXA

Mi
Fm

lower member
.

Order Osteoglossifo~es
Family Osteoglossidae
Subfamily Phareodontinac
Phareodurichthystavermi
Subfamily Ostcoglossinae
gcn. and sp.nov.
Ordcr "Sahoniformes"
Family Enchcdontidae

I, 13

Rhineasres sp.
Superfamily Andinichthyoidca
Family indet.
gen. nov.
Family Andinichthyidae
Andinichthys boiivianensis
rtndm'chrhys5p.

Family incertae redis

Incaichrhyssuarezi
Hoffsfetrerichrhyspucai

mpxa

Fm
-

49.51.52

2521

51

26

51

25

SuborderIchthyoiringoidei
Family inccmc sedis 1
?Apareo& sp.
Family incenac sedis 2
gen. and sp. indct.
M e r Cypriniforms
Family incenae sedis
Mdlim'chrhysinopinafus
Order Characiformes
Family Erylhrinidac
Hoplias nov. sp.
cf. Hoplias
Family Semsahidac
Subfamily Myle/e
gen. and sp. inder.
d Subfamily Senasalminac
gen. and sp. indef.
Family Characidae
SubfamilyTetragonopterinac
gen. and sp. inder.
cf. Subfamily Rhoadsiinae
gen. and sp. indef.
Ordcr Siluriformes
Family Ariidae

Fm

upper member

Enchodussp.

middle mb

Santa Luca

25

29

25
51

25

'

25.30

51

51
51

56

51

25

51
7,9, 12, 13, 14.21

75,21.30

33,35,38

41-52

56

51

12

34.35

51
49,51,52

51
51

TAXA

Superorder Athcrinomorpha
d.Order Cyprinodontiforma
gen. and sp. inder.
SuperorderAcanthoptcrygii
Order Percifoms
SuborderR a i d c i
Family Centropomidae
gen. and sp. ider.
OrderTetraodonrifomes
Family Eotrigonodontidae
Srephamdus m ' m h u
'ubdas Sarwptcrygii
Order Dipnoi
Family Ccratodontidae
Cerotodur sp.
Family Lepidcsiddac
Lepidosiren cf. paraaka

Ar
Fm

El Molino Formation
lower member

middle mb

...

25

7,12,22

Santa Luca

upper member

Fm

35

50.51.52

I, 13

25.27

53.54

51.52
12,21

SI.52

ass Cladistia (= Brachioptcrygii)

Order Polyptcrifoms
Family Polyptcridae
Dajetella sudamericana

21

ass Amphibia

Orderh u r a
Family Leptodactylidae
gen. and sp. inder.
Order Gymnophiom
Family inder.
gen. and sp. inder.
Order Urodcla
.Family inder.
gen. and sp. inder.

ES Reptilia
>der Chelonia
Family Podocncmididae
? Roxochelys vilavilensir
? Roxochelys cfi vilavilensis
Order Squamata
SuborderLacertilia
Family ?Iguanidac
gen. and sp. inder.

Family inder.
gen. and sp. inder.
Suborder Ophidia
Family Aniliidae
gen. and sp. inder.

51
51
12

14

30

33.35.38

4849.50

51-54
20.21

51

51
51

TAXA

Ca

Mi

Fm

Fm

Ar
Fm

Family Boidac
gen. and sp. inder. 1
gen. and sp. inder. 2
Family ?MvIatroiidae
gen. and sp. inder.
Family Tmpidopheidac
gen. and sp. inder.
Order Cmcodilia
Suborder Mesmuchia
Family Scbecidae
Sebeeu querejazu

lass Mammalia
nfraclassMetatheria
Order Deltathemida
Family inder.
Jmkhadelphys minufus
Order Peradmia

Family Peradcctidae
Subfamily Pcradeainae
Peradectes aurrrinum
Subfamily Camloameghh/mae
Roberrhoffsreneria nafionalgeographica

Santa Luca

El ?olino Form:
lower member

middle mb

upper member

Fm

51
51
51

1221

?.5,27,28,30

33, 38

51
48-50, 51 .
51
54
51

26

Family Dyrosauridac
S o k o i o s h off.iamviLrod
Suborder Eusuchia
Family Dolichochampsidae
Doliehochampsa minima
Order Saurischia
SuborderThcmpoda
Infraorder Coeiumsauna
gen. and sp. inder.
SuborderSaumpcdomorpha
&order Saumpoda
gen. and sp. inder.
Order Omirhischia
Suborder Omithopoda
gen. and sp. inder.
Suborder Ankylosauria or Ceratopsia
Ligabueichnium bolivianum
Order Inder.
gen. and sp. inder.

Ch
Fm

51
33

19
12,16,17. 18
19

19.3
'

19

51

51
51

Impora
Fm

TAXA

lower member

EMolino For
middle mb

_ _

Order Microbiotheria
Family Microbiotheriidae
Khasia cordil!eremis
Order Didelphimorphia
Family Diddphidae
Subfamily Didclphinae
Pucadelphys andinus
Incadelphys m q m
Mizquedelphys pilpinensis
Subfamily Eobrasiliinac
Tiuloraafroren'
Order Spalassodonta
Family Hathliacynidae
-Allqokim ausrrab
Oder Polydolopoidea
Family Polydolopidae
Epidolops sp.
Order Paucituberntlata
Family Kollpaniidae
Kollpam'a riupampina
Oder indct
Family inder.
Andinodelphys cochabambenris
Infmclass Euthcria
Oder Lcptictida
Family Palacoryctidae?
cfi Cimolesressp.
Oder Pantadonta
Family Pdntolambdidac
Alcidedorbignya inopinata
Order Condylanhra
Family Hyopsodontidac
Subfamily Moclaeninae
Tiuclaenm minurus
Moli~durs~lnrezi
h d ~ d uboliviensis
r
Pucanadur gagnien

Order Notoungulata
Family cf. Hennmsborniidae
or Oldfieldthomasiidae
gen. and sp. indct

Santa Luca

ition
upper member

Fm
.

.
.

. .

51
50

51
51
51
51
51
50

51
51

51

51,52

51
51
51

51

51

Impor:
Fm

MIREILLE OAYET. LARRY O. MARSIIALI. &I'IUBRRY SBMI'LKI!

( ? R o x o c l ~ e l y sv i l a v i l e n s i s , complete shells), and mammals

(Didelphimorphia. Didelphidae, indet.; Polydolopoidea,
Polydolopidae. Epidolops sp.). Indeterminoted turtles and crocodiles
were observed at Pajcha Pata.
About 3 kin south of Torotoro, remains of lungfish
(Cerntodontidae, Ceratodus sp.; Lepidosirenidae, Lepidosiren cf.
paradoxa), turtles (?Roxochelys cf. vilavilensis), crocodiles and an
indeterminate mammal were collected from a sequence of fine to
medium grained red sands of the Santa Luca Formation (Marshall et
al., 1985; Broin 1991; Schultze, 1991a) (Fig. 3).
At Maragua (18 km west of Sucre) and Chaupi Khocha (11 km
west-northwest of San Lucas) were collected remains of silurifonns
fishes (Ariidae), turtles and crocodiles by M. Gayet and L. G .
Marshall in September 1989.

H. IMPORA FORMATION
Fossil vertebrates of possible late Paleocene age were recovered
from two localities of lhe Impora Formation in the Camargo
Syncline. southern Bolivia. One is on the north side of Villa I'acheco
near the Ro San Juan del Oro, about 14 km norlh of Tojo, which has
yielded remains of siluriform fishes. Just north of this locality
rcntiiins of crocodics yid turtles were collected. The other localily is
at Chaupiuno, about 60 km norUiwest of Tarija. where indeterminate '
fish remains were found.

IMPORT,GNT LATE CRETACEOUS-PALEOCENE

VERTEBRATE FAUNAS FROM ELSEWHERE IN SOUTH
AMERICA
A. ANDEAN BASIN

In northwestern Argentina, a sauropod (Titanosauridae,

Laplatasaurus sp.) and a theropod (family incerfae sedis,
Unqirillosaitrus ceibalii) are known from the Los Blanquitos
Formation (a temporal equivalent of the Chaunaca and/or Aroifilla
formations) in the upper part of the Pirgua Subgroup (Powell, 1979)
(Fig. 1). Frogs (Eokenepoidae, Eoxenepoides saltensis) are known
from the Las Curtiembres Formation, which transitionally underlies
the Los Blanquitos Formation (Parodi-Bustos, 1962).
From the Lecho Formation (a basal lower EI Molino equivalent;
Flg. 1) are reported sauropods (Titanosauridae, Salfasuirrus
loricatus), Coelurosauria (Noasauridae, Noasaurus leali) and
theropods (Carnosauria indet. and Avisauridae. Avisaurus
archibaldi) (Bonaparte et al., 1977; Bonaparte & Powell, 1980;
Brett-Surman & Paul, 1985; Bonaparte, 1986). The Yacoraite
Formation (nnothcr EI Molino equivalent; Fig.1) lias yielded a
crocodile (Dolichocliai~ipsidae,Doliclrochunipsa niininru: Gasparini
& Duffetaut, 1980) which was collected in association with a
pycnodont (Coelodits toncoensis; Benedetto & Sanchez, 1972), a
clupeid (Gasleroclirpea branisai) and turitellid gastropods (see
Gasparini & Buffetaut, 1980; Cione et al., 1985: 296). Other taxa
reported from various localities and levels of the Yacoraite
Formation include a selachian (Pucaprisfis brunisi) and a siluriform
(indet.) (Cione et al., 1985; Powell, 1979: 202); the tooth of an
indeterminate theropod (Powell, 1979: 203); and trackways of
carnosaurs (Sa~ifichnusnienfoor), ornithopods (Hadrosauridae?.

I~adrosauricliniisarisfralis, Taponiclinrrs donoftoi, 7'elosiclirt~is

saltensis), indeterminate reptiles, and birds (Yacoraifichnrrs avis)
(Alonso, 1980; Alonso & Marquillas, 1986).
The Mealla Formation (a Santa Lucia equivalent; Fig. 1) has
yielded remains of teleostean fishes (indet.), turtles (indet.) and two
no Io un gu 1a t e inn i n ni a 1s (Henri c o s bo r n i id ae , Sinip son o t u s
praecursor, S.nidjor; Pascual et al., 1978), whereas the Maz Gordo
Formation (an Impora Formation equivalent; Fig. 1) has yielded
specimens of teleostean fishes (Callichthyidae, Corydoras revelat us;
Poeciliidae, indet.), pleurodire turtles ("Podocneniis" argenfinensis;
Cattoi & Freiberg, 1958) and possibly a notoungulate mammal
(Henricosborniidae, Sinipsonofus sp.) (Pascual el al., 1981).
In northern Chile, the following vertebrates are reported from
Senonian age rock units: a selachian (Pucapristis brunisi) from the
Tonel Formation (=lower part of Purilactis Formation S.I.) (pide
Cione e / QI., 1985) and dinosaurs (sauropods indet.) from the
Estratos de Quebrada Blanca de Poquis (an EI Molino equivalent)
and the base of the Pajonnles Formation (Salinas ef al., 199la. b).
III bru, vertebrates from the Vilqucchico Formation at its type
locality near Vilquechico, include Pucaprisfis branisi and
Casteroclupeu branisai (DBvila & Ponce de Le6n, 1971; Cione CI
al., 1985: 297). A more detailed faunal list is provided by Jaillard et
01. (it1 press). From the hiisc o f llicir niidtllc Vilqucchico Forinii~ion
(a Chaunaca Formation equivalent) they report three fish
(P/ychofrygon sp., rhinobatid?, actinopterygian). Their upper
Vilquechico Formation is divided into three sequences which are
equivalent, 'respectively, to the lower, middle and upper EI Molino
members in Bolivia. From the lower sequence they report an
actinopterygian , fish and name two dinosaur trackways
(Coelurosauria, Ornithomimidae, Ornithoniiniipus jaillardi;
Ornithopda, Hadrosauridae. Nadrosaitriclinus f iticucaensis); from
the base of the middle sequence are two fish (Pucapristis brunisi and
an actinopterygian); and from the upper sequence are two fish
(Dusyafissp. and an actinopterygian) and numerous charophytes.
At Laguna Umayo, from what was called the Vilquechico
Formation and is now called the Umayo Formation (Laubaclier &
Marocco, 1990, Jaillard et al., in press) are reported charophytes,
actinopterygian fishes (Characiformes, Erythrinidae, Hoplias sp.,
Serrasalmidae, Myleinae, indet., Characidae. Tetragonopterinae,
indet.; Siluriformes iricertae sedis; Perciforiiics, family indet.),
lu n g fis li (Cer a todo n t id tie, Cera to dir s s p., Lep id os ir en id ne,
Lepidosiren cf. paradoxa), frogs (Leptodactylidae indef.), snakes
(Aniliidne), turtles (Podocnemididae, ?Roxochelys cf. vilavilemis),
crocodiles (indct.), egg shell fragments reportedly of dinosaurs,
marsupials (Pcrndcctidnc, Perudecles arrstrirrrinr; Didelphidac?;
Pedioinyidae or Microbiotheriidae), and two placentals
(Co nd y 1art lira, Dido lo don t id ae, undes c r ibed ; No to ling u Ia ta,
Perutlieriidae, Perutlterirrni alfiplanense) (Grambast cf al., 1967;
Sigf. 1968, 1971, 1972; Bonaparte & Powell, 1980; Riige. 1981;
Marshall ef al., 1983b, 1985; Kerourio & Sig& 1984; Marshall &
Muizon, 1988; Schultze, 1991a; Gayet, personal observation).

114

B. SOUTHERN ARGENTINA AND SOUTHERN BRAZIL

There are four important vertebrate faunas outside the Andean
basin that warrant special consideration: they are' from the Los
Alamitos Formation in Patagonia (late Cretaceous). the Adaitiantina

r-.
niid Mnrllin formations i n the Inruiil bnsin (Inle Crctaccous), tile
nnco Negro Infcrior in Pntngonin (middle Inlcocenc?), nnd ti~e

fissure fillings in the limestones of S o Jose de Itabornl near Rio de

Janeiro (middle Paleocene?).

1. Late Cretaceous
From the Los Alamitos Formation at Estancia Los Alainitos in Rlo
Negro Province, southern Argentina, numerous vertebrate fossils
were found in Ulis mainly brackish-lacustrine rock unit which was
tentatively assigned a late Campanian-early Maastrichtian age
(Bonaparte el al., 1987). The taxa include: selachians (Batoidei,
indet.), holosteans (Semionotidae, Lepidoles sp.; Lepisosteidae, cf.
Alractoslerrs sp.), teleosteans (Siluriformes, cf. Diplomystidae, cf.
Ariidae; Percifohnes, Percoidei, inder.), Neopterygii indef.
(Lepidofes or Sparidae), Dipnoi (Ptychoceratodotitidae, Cerafodirs
iheringi), amphibians (Pipidae, cf. Xenopus sp.; Leptodactylidae,
indef.),turtles (Meiolaniidae, cf. Niolamia; Chelidae, indef.),snakes
(oidnc, Mndtsoniinnc. Alaniiropliis argenfiniis, Patagoniophis
parviis, RionegropIris ntadtsoioides), dinosnurs (Titnnosauridnc,
Aelosarirus rionegrinris?: Hadrosauridac, Krilosauriis auslralis),
and innminals including Symmetrodontn (Bondesiidne. Bondesiiis
fcrox; Inm. in de^., Casaniiyiielia riomgrina: ?Spalncotlicriidac,
Brandonin interdiedia; B arbereniidnc, Barberenia araiijoae,
Qrtirogaflteriirna niajor), Dryolestoidea (Dryolestidae,
Groeberllieriunr slipanicici, G. novaci, Leonardus ciispidaliis;
Mesungulatidae, Mesiingulaluni Iioussayi; Reigitheriidae,
Reigilheriuni bunodonla), Triconodonta (Triconodontidae,
Arislrotriconodon nickennai), Multituberculata (Ferugliotheriidae,
Ferugliofheriuni wiridltauseni) and two Gondwanatheria
(Gondwanntlieriidae, Gondwanafheririnipalagoniciinr, Vucefichia
gracilis) (Albino. 1987; Baez, 1987; Bonaparte & Soria, 1983,
1985; Bonaparte, 198Ga, b, c, 1987, 1990; Bonaparte & Pascual,
1987; Bonaparte & Rougier, 1987; Broin, 1987; Cione, 1987;
Mones, 1987; Powell, 1987). In northern Patagonia, some levels of
the Los Alamitos Fonnation are probably associated with a marine
transgression from the west (Andreis, 1987). The Los Alarnitos
Formation is overlain by the transgressive marine Roca Formation of
Mnastriclitian-Daniail age (Bonnparte, 1990) which probably
corrclatcs with thc cocvnl nnd trnnsgrcssive lower EI Molino
Forniution of Bolivin. c c n u s c of their chnractcristics and
stratigraphic positions, we thus propose temporal correlation of the
Los Alarnitos Formation with the Chaunaca Formation. Hence, the
Los Alamitos Formation is likely to be of Cnmpanian age.
The vertebrate faunas from the Adainantina and Marilia
formntions (Upper Baum Group) in. the northern part of tlie Paran6
basin, southcentral Brazil, include fishes (Lepisosteiformes,
Lcpisostcidnc, Lcpisosreus coniinatoi; Osteoglossiformes,
Osteoglossidac; Characifornies; Silurifornlcs, cf. Doradidae;
Perciformes. ?Percichdiyidae), lungfish (Dipnoi, Neoceratodontidae,
Neocet afodris sp.), frogs (Lcptodactylitlac, Ba~irirbafracliiispricei).
turtles (Podocnemididae, Roxoclielys irarrisi, cf. R . elegam, aff.
Podocnentis brasiliensis), lizards (Iguanidae?, Prisligiiana
brasiliensis), snakes (inder.), crocodiles (Baurusuchidae,
Barrrrisuclius pachecoi; Peirosauridae, Peirosaurus forminni;
Goniopholidae, ?Goniopholis paulislanrts; Trematosuchidae,
Itasrichus jesuinoi; family indel.. Brasileosaurus pacltecoi,
Splragesaiirus huenei), dinosnurs (Carnosnuria, Abelisnuridae.
415

r I I d l y irdef.; Coclurosnurio, lnniily nov. I , nov. 2; Ssiiropd;\,

Titnnosauridnc. ?Aritarcfosnirriis brasiliensis, Iituriosnrtritscf.
australis; Ornithischia). niid mainrnals (Ilaccntalia. indct.) (Gayct 6t
rito, 1989; Bertini el al., in press). Both formations arc of
continental, mostly fluvial. origin and arc thought to bc Scnonian in
age.
2. Middle Paleocene?

A local fauna is known from tlie Banco Negro Inferior in the base
of the Ro Chico Formation about 1km southwest of Punta Peligro
(40 km norh-northeast of Comodoro Rivadavia along the Atlantic
coast) and another at Las Flores, 60 m above the Banco Negro
Inferior in the base of the Gran Barranca south of Lago ColhuHuap. Both local faunas are in the S a n Jorge basin, Chubut
Province, southern Argentina and were discovered in January 1979
during a National Geographic Society sponsored research program
under the direction of one of tlie authors (Marshall e l al.. 1981). The
fossils from Punta Icligro includc turtles (Chelidnc, at least four
taxa; roin, 1988), crocodiles (Scbecosuchia?; Crocodylidne.
Necrosirchus ionensis Simpson. 1937; Alligntoridac, Exainian sp..
Allognatosrichris? sp.), a possiblc edentnte or niultitubcrculate
Pnscuul.
i
(Sudnmcricidnc, Sudrinicrica cinrcghirioi Scilleto Ynnd
1985), condylarths (Arctocyonidae and/or Mioclncnidae) and several
indeterminate bone fragments (Marshall el al., 1981, n16; Pascual &
Ortiz-Jaurcguizar, 1991). Those from Las Flores include the
polydolopoid marsupial Epidolops sp. (Iascual C Bond, 1981). and
other taxa which show affinities with Itaboraian faunas in Brazil
(Poscunl & Ortiz-Jaureguizar, 1991). Many additional fossili of
marsupials and ungulates from Las Flores were collected by L. G.
Marshall and colleagues in 1979, and numerous unpublished
specimens from both Punta Pcligro niid Las Flores are now available
(Van Valen. 1988). The Banco Negro Infcrior is predominant& a
black bentonitic mudstone (Andreis el al., 1975). The Salamanca/Ro
Chico contact records a noteworthy marine regression which,
according to associated geochronologic age constraints (Marshall el
al., 198l), probably corresponds to the major regression identified
by Haq el al. (1987) in tlie early Thanetian (about 58.0-58.5 Ma).
The Itaborai fauna comes from fissure fillings in t h early
Pnlcoccnc (?) ngc SEO Jose de ltnbornf Forinntion locntcd about 25
kin eilst of Nitcr6i. stntc of Rio de Jnnciro, Brnzil. Taxonomic lists of
this exceptionally diverse fauna are provided by Paula Couto (1970).
Palma & Brito (1976) and Marshall el al. (1983). The vertebrates
wcre recovcred from numcrous karst cavitics in a lower liniestonc
unit and predate an upper limestone unit (Brito et al., 1972). For
most specimens, their association among themselves and/or with
specific cavities was never recorded. Although sonic nianiiiials
appear referable to what has been interprcted as middle Paleocene
time (i.e. the Itaboraian Land Maniinal nge; sensu Marsliall, 1985).
others appear more progressive and may represent faunas of late
Paleocene and possibly cai ly Eoccne iigc (i.e. the Kiocliican and
Casamayoran Land Mammal Ages, respectively; see Soria. 1987;
Van Valen, 1988; A. Cione, personal comunic?tion). In the
following discussion, we regard this fauna as middle Paleocene, but
caution that it may include taxa that accumulated between middle
Paleocene nnd early Eoccne time.
Although a hydrothermal origin has been clnssically favorcd for
the SEO Jose de Itabora lncustrinc limcstones (Francisco gZ Cunlin.

MIREILLE OAYCT. LARRY O. MARSIIALL LIIUUKRYSCMIBKI!

1978). water level i n the tectonic iicnr-shore lake where tlie

limestones were deposited might have been controlled by the level of
the nearby Atlantic Ocean through adjustment of marine and
continental phreatic levels. In such a case, the lacustrine episode
evidenced by the lower Itabora limestones would have been coeval
with a marine highstand. Since middle (and later?, see above)
Paleocene mammals are known from the infilling of the karstic
cavities Uiat f o r m a after lake dessication, it would be possible to
correlate this highstand period with the one in the late Danian, and
the subsequent dessication with the early Thanetian regression which
is also recorded near the Salamanca/Ro Chico contact in southern
Argentina (see above). A subsequent highstand period would have
resulted in the deposition of the upper limestones of So Jose de
Itaboraf.

DISCUSSION

A. UIOSTRATIGRAPIIY
The Mesozoic and Ioleoccne age vertebrate fossil localities in
Bolivia (Fig. 2) are arranged in T a b l e 1 according Lo their
stratigraphic context. A systematic list of all the known fossil
vertebrates is given in Tables 2, and in Table 3 with indication of
locality and stratigraphic occurrence.
In this section we review the known occurrence of the taxa listed
in Tables 2 and 3 in order to identify their cluonostratigraphic ranges
and hence potential usefulness for calibratin8 the Bolivian rock
sequence. Of particular interest is the apparent position of the
Cretaceous/Paleocene (K/T) boundary.within the 81Molino
Formation (see below).

Chile (Schultzc. 1981). niid existed uiitil the iiiiddlc Eocene i n

marine deposits around the world (Blot, 1987). Cione (1977)
tentatively referred specimens discussed by Wenz (1969) from
Bolivia to Coelodus loncoensis which was erected by. Benedetto &
SQnchez (1972) upon material from the Yacoraite Formation of
northwest Argentina. Gayet (1991) cautions that the spccics identity
of some Bolivian pycnodontids has still to be verified, while olhers
from the lower and basal middle EI Molino Formation can be
referred with confidence to Coelodus foncoensis which was
apparently endemic LO the Andean basin.
3. Semionotiformes

In Bolivia, Lepidofes sp. was collected securely from the late

Triassic-early Jurassic. Elsewhere in South America, Lepidofes is
reported from the Tithonian (=Portlandian) of Neuqun Province,
Argentina (Aramayo, 1981). the Aptian-Albian.of Brazil (Agassiz,
1841; Roxo & Lfgrcn, 1936; Woodward, 1895, 1908) and a
possible record froin the lntc Cniiipiiiiiiiii-eiirly Munslrichtiun ngc Los
Alainitos Forination in Argentina (Cione, 1987). Elsewhere i n the
world. Lepidotes is recorded from the Rlietian of Germany to
Cretaceous/Paleocene of India (Gayet et al., 1984).
Pillow-shape scales which rcprcscnt u new gctius of thc family
Semionotidae are known only in the lower and basal iniddle
members of the EI Molino Formation in Bolivia (Gayet & Meunier,
personal observation) and in the Yacoraite Formation in Argentina
I
(A. L. Cione, pers. com.).

4. Cinglymodi

In Bolivia, Lepisosfeus sp. is reported from the lower and basal

middle El Molino. The earliest Lepisosteidae are reported from the
early Cretaceous of Niger (Ariunbourg & Joleaud, 1943) and Congo
(Casier, 1943). Younger fossils, including Lepisosteus, are known
In Bolivia, selachians were recovered from the lower and basal
from the late Cretaceous of India (Jain & Sahni, 1983; Gayet el al.,
middle members of the El Molino Formation. Rhombodontidae are
1984) and from the Cenozoic of Euiope, India and North America
present only in the lower member, whereas Sclerorhynchidae and
(Wiley, 1976). In South America, Lepisosteidae are known in
Dasyatidae are present in both. Sclerorhynchidae range from Albian
Argentina from the late Campanian-early .Maastrichtian age Los
to Maastriclitian (Cappetta, 1990). Schizorhiza is known only in
Alamitos Formation (cf. Atracfosferrssp.; Cione. 1987) and from the
Maaskichtian age rocks from numerous localities around tlie world
EI Abra Formation of uncertain age (A.L. Cione, pers. com.)
(Texas, Morocco, Middle-East, Niger. Nigeria and Zaire), while
Ischyrliiza ranges from the Turonian to Maastrichtian in North (Lepisosferts; M. G.. pcrs. obs.); Lepisosteris is known froin the late
America and Niger (Cappetta, 1987, 1990, 1991). Pucaprisfis Cretaceous Adainantina and Marilia formations of the Upper Bauru
(Sclerorhynchidae), Pitcubatis (Rhombodontidae) and the three Group in Brazil (Santos, 1984; Gayet & Brito, 1989; Brito & Gayet,
named species of Dusyuris (D. molinoensis, D.scltaefferi, D. in press), and the late Cretaceous of Colombin (Gayet, 1991).
Lepisosfcits cxists to Kcccnt in southenstern USA and in Cciitrul
brunisui) (Dasyatidae) are currently endemic to the Andean basin
where they are proving exceptionally useful in intra-basin correlation America,
(Cappetta, 1990). Pircuprisfis branisi is also known from a level
below another which yielded dinosaurs in the lower part of the 5. Clupeiformes
Yacoraite Formation in northwest Argentina (Powell, 1979).
Gusferoclupeu brunisui is apparently endemic to the Andean basin
Dasyalis i s known in the Cenomanian of Texas, and from
where in Bolivia it is reported from the Chaunaca (possibly),
Maastrichtian to Recent world-wide.
Cajones, all three members of the El Molino, and the Santa Luca
formations; in Argentina it is known only from the Yacoraite
2. Pycnodontiformes
Formation (Cione el al., 1985) and has never been found in
overlying Paleocene formations. The oldest known Clupeidae are
Pycnodontiformes (Pycnodontidae) were collected in Bolivia from
from the Neocoinian of northern. Kyushu, Japan (Diplonlysrus
the Chaunaca Fondation and the lower and basal middle members of
the El Molino. Colodus was reported from the lower Cretaceous of prinzofiniis, D . kokirraensis} (Uyeno, 1979).
Brazil (Santos, 1963; Wenz, 1989). Colombia (Porta, 1970) and

1. Selachians

416

In Bolivia, the subfamilies Osteoglossinae and Phareodontinae of

the family Osteoglossidae are recorded from the EI Molino and
Santa Lucia formations (see Table 3). The earliest record for
Ostcoglossiformes in South America is from Uie Aptian age Areado
Fonnation in Brazil (Santos, 1985) where the taxon is referred to tlie
family Arapaemidae (Laeliichtliyinae) which lias African affinities.
Phnreodontinae were reported from the Paleocene and Eocene of
Africa (Cappetta, 1972) and Europe (Woodward, 1901; Taverne,
1978), Paleocene of southeast Asia (Sanders, 1934), late Paleocene
of Turkmenia (Danil'chenko, 1968), Paleocene of Australia
(Taverne, 1975; Gayet, 1991). and middle Eocene of North America.
Thus, Lhe Phareodontinae record from the EI Molino is the earliest
known to date.
Squamules (parts of scales) of Osteoglossidae (Phareodontinae
rind Osteoglossinae) were reported in the Maastrichtian of Niger,
CrelaceouslPaleocene of India and Paleocene of Pnkistan (Gayet &
Mcunicr, 1983). Ostcoglossinae wcrc rcportcd from tlic I'aleoccnc of
Sumatra (Sanders, 1934), and range lo Rcccnt in South America,
Asia, Africa and Australia.
7. "Sulmoniformes"

In Bolivia, Enchodontidne (Enclrudus sp.) and Ichthyotringoidei

(family indet. and 7Apafeudus sp.) were collected Gom the lower
and basal middle EI Molino. Encltudus is known only from marine
Maastrichtian ake rocks in Congo (Dartevelle & Casier, 1949).
Morocco (Arambourg, 1952). Brazil ( R e h u s a s & Santos, 1956),
Europe (Goody, 1968), North America (Goody, 19691, Lebanon
(Goody, 1969) and Israel (Chalifa, 1989) to note only the most
important in both preservation and quantity.
Apafeudus is a member of the suborder Ichthyotringoidei which
ranges from early Cretaceous (?Albian) to Maastrichtian (Goody,
1969).
8. Cypriniformes
The cypriniform (Mulinichfltys inopinafus: Gayet, 1982b) from
Ihc lower and basal middlc EI Molino rcprcscnts tlic only record,
fossil or extnnt, of this ordcr in South Amcrica. Elscwlicrc in lhe
world, this group is' first known from the Eocene of North America
(Grande ef al., 1982). Gayet (1986b. c) described Raniallichfhys
from the marine Cenomanian in Israel which may be considcred an
ancestral, if not true, cypriniform.

uinnyo i n Perii (Gnyct. 199 I), Mioccnc Loyoln Formntion of

Ecuador (Roberts, 1975) arid Mioccnc of Coloiiibin (Lundbcrg et al.,
1986); Characidae (Tetragonopterinae) from the Umayo Formation
at Laguna Umayo in Peru (Gayet, 1991). Miocene of Ecuador
(Roberts, 1975) and early Mioceile of Taubat in Brazil (Schaelfcr,
1947); Characidae (indef.) from the Paleocene of Morocco (Cappeta
ef al., 1978); Serrasalmidae (Serrasalminae) and Characidae
(Rhoadsiinae) had not yet been found as fossils. The oldest known
record of characiforms outside of South America is Salniinups
ibericus from the marine late Cenomanian of Portugal (Gayet,
1985a). Characiforms range.10 Recent in Africa and in South
America. One recent genus colonized Central America.

10. Siluriformes

In Bolivia. this order is represented by members of the families

Ariidae and Andinichthyidae from both the EI Molino and Santa
Luca formations, and at lcast by two families inccrfae scdis from
thc Snntn Luch Forinntion nt Tiupempn (Gnyc~,1991).
The other known siluriforliis elsewhere in Soutli America are
represented by cf. Diplomystidae and cf. Ariidae in tlie Carnpanian
age L o s Alamitos Formation in Argentina (Cionc, 1987);
Siluriforiiics indc/. from Ilic Mnns~riclitinnrtgc Coli Toro (Cionc C
Laffite, 1980) and Yacoraite formations (Cione et al., 1985) in
Argentina; cf. Doradidae from the late Cretaceous age Adamantina
and Maria formations in the Upper Bauru Group, Brazil (Gayet &
Brito. 1989; Brito & Gayet, in press); and Ariidae from the
Maastrichtian? age Umayo Formation at Laguna Umayo in Peru
(Gayet, 1991).
The only records for Cretaceous age Siluriformes outside of South
America are a single tooth referred to Arius sp. from the latest
Cretaceous of Central India (Jain & Salini, 1983) and doubtful
otolithes from the late Cretaceous of North America (Frizzell, 1965;
Fitch, 1975).
Siluriformes are known in the Paleocene of Argentina, [Arius?
and Baclintania in Patagonia (Dolgopol d e Saez. 1941), and
Prupygidirim (Bocchino, 1964) and Coryduras (which in Recent
limes is restricted to freshwater in South America) from the Maz
Gordo Formation (ardack, 1961)J. Africa (in Congo; Dartevelle &
Casicr, 1949; Cnsicr, 1960), Europc (clgium; Leriche, 1902).
Indoncsia (Sandcrs. 1934); nnd in tlic Eoccnc of North Anicricn
(Lundberg, 1975; Grande. 1987; Grande & Lundberg, 1988), Europe
(Woodward, 1901; Casier, 1946), Africa (Peyer, 1928) and India
(Salini & Misra, 1975). Ariidae range to Recent.
11. Cyprinodontiformes

9. Characiformes
In Bolivia, this ordcr is rcprcscnlcd by the families Erytluinidac,

Serrasalmidae (including Serrasalminac and Myleinae) and

Characidae (including Tetragonopterinae and Rhoadsiinae). All these
groups have been found in the Santa Luca Formation, while
members of the Erythrinidae, Myleinae and Tetragonopterinae were
also found in tlie lower and basal middle EI Molino Formation.
The oldest previous known Erytlirinidae (Nuplias) are reported
from the Miocene of Ecuador (Roberts, 1975); Serrasalmidae
(Myleinae) from the Maastrichtian? Umnyo Formation at Laguna

In Bolivia, pharyngeal teeth from the Miraflores Forniation are

tcntatively referrcd to indelcrininate cyprinodontiforms. In the lower
member of the EI Molino Forination are known several very
primitive cf. Cyprinodontiformes (Gayet, 199 1). The earliest
previous record of this group,was from the Oligocene of France
(Gaudant. 1988). and it ranges to Recent world-wide.
12. Perciformes

In Bolivia, perciforms of lhe family Cencropomidae are known

417

,
MIREIUE OAYIX. LARRY O.MARSIMLL EL IIIIBKRY SEMI'IJKC

from the Santa Ldca Formation (Gayet, 1991). Indeterminate

specimens of this order nre reported in South America from the
Campanian age Los Alamitos Formation of Argentina (Cione, 1987).
the Maastrichtian age Marlia Formation of Brazil (Gayet & Brito.
1989). and the Umayo Formation at Laguna Umayo in Peru (Gayet,
199 1). The earliest' Centropomidae were previously recorded from
the middle Eocene of Italy (Sorbin, 1970), and they range to Recent
world-wide.
13. Tetraodontiformes

In Bolivia, Slepltanodus (Eotrigonodontidae) is recorded from the

lower and basal middle El Molino Formation. Eotrigonodontidae
was reported from marine beds in the ?Albian-Aptian of Morocco
(Tabaste, 1964). Cenomanian of Egypt (Weiler, 1935), late
Cretaceous of Nigeria (White, 1934), Congo (Dartevelle & Casier,
1949), Niger (Tabaste, 1963; Cappetta, 1972). Morocco
(Arambourg, 1952), Israel (Raab, 1963), Upper Cretaceous of lndia
(Jain & Salmi, 1983) nnd Tertiary levels of Europe, (Leriche, 1906;
Casier, 1946), Angola (Dartevelle & Casier, 1959) and Egypt (Peyer,
1928). Eotrigonodontidae range from ?Albian-Aptian to middle
Eocene, whereas Sfeplianodus w a s reported only from the
Cretaceous (sanie rdccrcnces).

16. hmphiblans

In Bolivia, indeterminate -eptodactylidae are known only from

the Santa Luca Formation at Tiupampa. Elsewhere in South
Americn indeterminate Leptodactylidae are recorded from the
Campanian nge Los Alamitos Formation in Argentina (Baez, 1987),
and, questionably, from the Umayo Formation at Laguna Umayo in
Peru (Sig&, 1968). A probable Leptodactylidae, Bairrubafraclius
pricei has been recorded in the Maastrichtian age Marlia Formation
in Brazil (BQez, 1985; BQez & Per, 1989; Bertini et al., in press).
The oldest known Leptodactylidae are from the M a n a Formation
and the family extends to the Recent (BQez, 1987).
The oldest known Gymnophiona are represented by two vertebrae
from the Santa Luca Formation at Tiupampa (Rage, 1986, 1991b).
Previously this group was known from the middle Paleocene (at
Itabora) to the Recent.
Vertebrae of urodels are known from the lower El Molino
Formation at Pnjchn Pnta. This is the earliest record of this group in
the world (Kage, pers. com.).
17. Turtles

Dajefella sudamericana (Gayet & Meunier, 1991a, b, 1992) from

the lower El Molino and Santa Luca formations is the only record
for Polypteriformes outside of Africa, where members of this order
are known from the Senonian of In Becetem in Niger (Gayet et al.,
1988), Miocene of Tunisia and Kenya (Greenwood, 1951,,1973), and
Pleistocene of Ethiopia (Arambourg, 1948). They range to Recent in
Africa where they occur in freshwater.

Roxochelys was based on spccinicns collected [rom thc

Campaninn nge Adamnntinn Formation or the Upper Bauru Group in
Brazil (Broin, 1991; Bertini et al., in press). ?Roxoc/ielysvilavilensis
Broin, (1971), from the Santa Luca Formation is probably referable
to a genus distinct from Roxochelys (Broin, 1991). The species
vilavilensis is definitely known only from lhe Santa Luca Formation
in Bolivia and possibly frpm the Maastrichtian? Umayo Formation at
Laguna Umayo in adjacent Peru (see above), while very fragmentary
material from the lower EI Molino of Bolivia is referred to
?Roxoc/ielyscf. vilavilensis. This species was apparently endemic to
the Andean basin.

15. Lungfish

18. Lizards

In Bolivia, Ceratodontidae (Cerafodus sp. and ceratodont n. sp.)

and Lepidosirenidae (Lepidosiren cf. paradoxa) tooth plates occur
together in the early Paleocene of Tiupampa. In South America, the
first ceratodonts occur in the Upper Cretnceous of Patagonia
(Ameghino, 1906; Pascual & Bondesio, 1976; Cione, 1987) and
were also described from the Lower Cretaceous of northern Brazil
(Cunha & Ferreira, 1980). Ceratodonts were distributed world-wide
in tlie Mesozoic, still occurring during the Cretaceous in Africa,
Madagascar, Australia, North and South America, whereas the
Neoceratodontidae which occur in the early Cretaceous of Australia
persist there until today (Schultze, 1991a).
Lepidosirenidae a r e reported from the Maastrichtinn E l
Molino Formation a t Vila Vila and the Paleocene Santa Lucfa
Formation at Torotoro. Fossils are recorded from the
M a a s t r i c h t i a n ? a t L a g u n a U m a y o i n P e r u (Sig, 1968).
Eocene o f A r g e n t i n a (Fernandez et a l . , 1973; Fernandez,
1976) and Miocene of Brazil (Santos, 1987) and Colombia
(Bondesio & Pascual, 1977). Lepidosirenids occur today in
the Amazonian and Paraguayan regions of South America.

An unidentified member of what appears to be the family

Iguanidae is recorded from tlie Santa Luca Formation at Tiupampa
(Rage, 1991b). A possible iguanid (Prisfigrrattabrusiliensis; Estes &
Price, 1973) is known from the Manstrichtian agc Mnrflia Formation
of the Upper Bauru Group in Brazil, while h e earliest uncontested
members of this family are from the middle Paleocene of Brazil
(Estes, 1983) and North America (Sullivan, 1982).

14. Polypteriformes

19. Snnkes
The four faitlilies of snakes listed below are known only from the
Santa Luca Formation at Tiupampa in Bolivia.
In South Americn, Aniliidhe are first reported from the Umayo
Formation at Laguna Umayo in Peru (Rage, 1981) and middle
Paleocene at Itabora in Brazil (Rage, 1987).
The earliest Boidae are from the Maastrichtian of North America
(Estes & BBez, 1985), Europe (Rage, 1987) and India (Jain & Sahni,
1983), while in South America, previous to this work, they were first
reported in the middle Paleocene in Brazil (Rage, 1987) and early
Eocene of Argentina (Albino, 1986).

418

Madtsoiidne nrc first kiiown in the Suiitoiiian/Cninpnnian of

Mndngnscnr (Rage, 1984; Roiinparte. 1986) niid Campaninn age Los
Alainitos Formation in Argentina (Albino, 1986, 1987). In fact, Uie
oldest Madtsoiidae is probably from Niger (early Senonian) (Rage,
1984).
Previous to this work, Tropidoplieidae were first reported from the
middle Paleocene hi Brazil (Rage, 1987), middle Eocene in North
America (Hohnan, 1979), and Eocene-early Oligocene in Europe
(Rage, 1984).
20. Crocodiles

Three families are known in Bolivia: Dyrosauridae (basal middle

EI Molino and Santa Lucia formations), Dolichochampsidae (upper

EI Molino Formation) aiid Sebecidae (Santa Luca Formation).
Dyrosauridae are known from late Maastrichtian to late Eocene
rocks around the world (Buffetaut, 1982). In South America,
dyrosaurids are known fTom what appears to be the Mnastrichtian in
razil ,(Cope, 1886; Buffetnut, 1978), and a possible dyrosaurid is
kiiowii from tic Manstrichtinti in Argentina (Gnspnrini, per. coin.).
Sokotosuchus ianwilsoni (Halstend, 1975) is from the late
Maaskichtian Dukarnaje Forination in Nigeria.
ip~a
is known
Dolichochampsidae ( D o l i ~ l i ~ ~ l t a nniininta)
elsewhere only from tlie upper p u t of the Yacornitc Forination in
northwest Argentina and the upper El Molino Formation in Bolivia
(see above). The family was apparently endemic to the Andean
basin.
Sebecidae were previously known from the middle Paleocenemiddle Miocene (Buffetaut, 1982; Gasparini, 1984); they were
endemic to South America. Sebecus querejazus from the Santa
Luca.Formation is the earliest known member of the family
(Buffetaut & Marshall, 1991).
21. Dinosaurs

FOSOSILES Y FACIES bE noum VOL. I - VERTEBMWS

Dinosaur trackways are known from four localities in the lower EI

Molino near Torotoro and at Santivaez near Parohni, and from two
localities (Arapamba, Camargo) in an indeterminate EI Molino
member. The only dinosaur fossils in Bolivia are a tooth of a
Coelurosnurin (Marshall, 1989b) from the lowcr El Molino at Pnjchn
I'ntn, and undcscribcd boiics from two localitice of tlic Cajones
Formation near Santa Cruz (R.Suhez, pers. com.).
Although a detailed study of the trackways has yet to b e
undertaken, it appears that at least four suborders (Theropoda,
Sauropoda, Ornithopoda and Ankylosauria or Ceratopsia) are
represented within the lower EI Molino at Torotoro. This
demonstrates that dinosaurs were taxonomically diverse in lower EI
Molino time.
For cluonostratigraphic purposes we follow the general mnsensus
that dinosaurs became extinct at the end of Cretaceous time and that
there is no compelling evidence (contra Van Valen, 1988) to
substantiate Uiat they may have continued into the early Paleocene.
Recent work by Lofgren el al. (1990), for example, negates a
Paleocene age for dihosaurs from the upper Hell Creek Formation in
Montana. Our position is concordabt with the biostratigraphic data in
the Andean basin summarized above and below.

22. M,unimuls

Mammals are known only from the Santa L u c h Formation in

Bolivia. Peradectes airstriniim was based on a specimen collected
from tlie Umayo Formation at Laguna Uinayo in Peru (Sig&, 1971,
1972; Crochet, 1980) and a second specimen from Uie Santa Luca
Formation at Tiupampa in Bolivia was later referred to this species
(Marshall & Muizon, 1988). The earliest known Perudecfes is from
the base of the early Paleocene in North America (Van Valen, 1988:
29) and is common thereafter in early Tertiary age rocks in North
America and Europe (Crochet, 1980).
Robert/ioflsferteriais most closely related to Procuroloanreglrinia
from the middle Paleocene of Brazil (Marshall et al., 1983a),
although other more primitive members of the subfamily
Caroloaineghiniinae are known from the Maastrichtian in Norih
America (Marshall et a l , , 1989). The genera and species o f
Mi'crobiotlieriidae, Didelphidae, Hathliacynidae and Kollpaniidae are
known only Erom the Snnta Luca Formation at Tiupampa, and they
all represent the earliest known members of these faniilies in South
Ainericri (MarslinIl & Muiimii, 1988; MiuslinIl et d.,
1989).
Cf. Cintolestes sp. is the only known member of the order
Leptictida in South America. Cintolestes is recorded froin the
Mnnskichtinn and cnrly Paleocene in North America (Marshall &
Muizon, 1988).
The pantodont Alcidedorbignya is the only known member of this
order in South America. Pantodonts are f i s t known in the middle
Paleocene of North America where Panfolambda batlrniodon
represents a potential descendant of Alcidedorbignya (Marshall &
Muizon, 1988). Among the New World pantodonts, Alcidedorbignya
is the earliest and most primitive member yet known.
1
The mioclaenine Hyopsodontidae are the only representatiyes of
this family in South America. In North America, members of this
group are 6rst recorded in the early Paleocene (Marshall & Muizon,
1988; Van Valen, 1988).
The earliest known notoungulates are represented by PeruUieriimt
altiplanense (Perutheriidae) from the Umayo Formation at Laguna
Umayo in Peru (Marshall et al., 1983b), Sintpsonolus
(Henricosborniidae) from the Mealla Formation in northwest
Argentina (Pascua1 ef al., 1979), and cf. Henricosborniidae or
Oldlicldtlioinnsiidnc from lhe Siintri Luchi Forinntion nt Tiupniiipn in
Bolivia (Miusha11 & Muizori, 1988).
I

B, THE CRETACEOUS/PALEOCENE (KIT) BOUNDARY IN

THE ANDEAN BASIN
Based 011 the above discussion of chronostratigraphic ranges, the
taxa can be grouped into five categories (see Table 3):
1. Those which presently are known only from the EI Molino
Formation and its stratigraphic equivalents in tlie Andean basin.
These are currently regarde$ as "endemics" which are potentially
useful for intra-basin correlation' although their usefulness i n
calibrating these rocks witti the geologic time scale has yet to be
firmly established. Included are Pitcapristis (Sclerorhynchidae),
Prtcabafis (Rhombodontidae), the three species of Dasyafis
(Dasyatidae), Coelodrrs toncoensis (Pycnodon tidae). and
DoliclroclIanipsa (Dolichochampsidae).

419

2. Taxa from the EI Molino and Santa Luch formations (and their
stratigraphic cquivalcnls) which are "endemics" in the Andean basin.
Included are Gasteroclicpea bronisai (Clupcidae). Andinichtliys
bolivianensis , (Andinichthyoidea,
Andinichthyidae),
Hoffsterterichtliyspucai, Incaichthys suarezi (families incertae sedis)
and ?Roxochelys vilavilensis (Podocnemididae).

migrate bctwccn fresh water and marine; nnd 4) marinc fishes.

3 . Taxa known only from the lower and basal middle EI Molino
Fonnation which are currently regarded as late Cretaceous "guide
fossils" based on their records elsewhere in the world. Included are
Schizorhiza and Ischyrhiza (Sclerorhyncliidae), apparently Lepidot es
(Semionotidae) which has only o n e dubious record in the
Cretaceous/Paleocene of India, Enclzodiis (Enchodontidae),
?Apaleodris (family indet.), Ichthyotringoidei (gen. and sp. indet.),
Sleplranodiis (Eotiigonodontidae) and dinosaurs.

1. Primary freshwater fishes

The definition of these four categories is based on living fislies

and is not always evident with fossil forms (see below). For this
reason, the identification of paleoenvironment based only on one or a
few taxa is generally of dubious value.

a. Semionotidae

4. Taxa which occur in the EI Molino and/or Sant'a Luca

formations and, which based on records from elsewhere in the world.
lransgresscd the K/T boundary arid arc of dubious or no value in
defining this boundary. These include Lepisosteus (Lepisostcidae),
Os teog I os sid ae , Cyprin i form es , Ch araci f or mes, S i lur i for ni es,
Cypriiiodoiitiforrnes. Perciformes, Polypteriformes, Dipnoi.
Leptodactylidne. Podocncniididac. Laccrtilia, Boidae, Madtsoiidae,
Dyrosauridne. and Lcptictida.
5. Taxa recorded only in the Santa Luca Formation which are the
earliest or only records for these groups in South America. They do
not occur in association with diagnostic Cretaceous taxa and
therefore are regakded as Paleocene. Included are Ceratodontidae n.
gen. and sp., Gymnophiona, Aniliidae, Tropidopheidae, Sebecidae,
Peradectidae, Caroloameghiniidae, Microbiotheriidae, Didelphidae,
Hathliacynidae, Kollpaniidae, Pantodonta, Hyopsodontidae and
Notoungulata.

As demonstrated by group 3 (above), apparent late Pretaceous

taxa occur only in tlie lower and basal middle members of the EI
Molino Formation. It therefore appears that the K/r boundary occurs
above the basal middle member, while the upper El Molino member
may be regarded as early Paleocene based on palynological study of
a stratigraphic equivalent in norlhwestern Argentina (Quattrocchio el
al., 1986) and od the absence o f diagnostic Cretaceous taxa. We
therefore suggest &hatthe K/T boundary lies somewhere in the upper
part of the middle member of the EI Molino Formation. In addition,
tlie Sanla Lucla Formation, which also lacks diagnostic Cretaceous
taxa and conformably overlies the early Paleocene upper member of
the EI Molino Formation, may be regarded as early, but no earliest,
Paleocene in age.
C. PALEOENVIHONMENT

The fossil Semionotidae are generally considered as primary

freshwater fishes. A new genus is reported in Bolivia in the lower
and basal middle members of the EI Molino Formation. Except for
Uie questionable Cretaceous/Paleocene record from India (Gayet et
al., 19841, Semionotidae are not known to survive beyond the
Cretaceous/Paleocene boundary. Their absence at Tiupampa is
probably related to the Paleocene age of the Santa Luca Forination.
The new genus from Aguti CI:m und Hokl Cordillern, known only
by scales and referred to the Semionotidae, seems endemic to
Bolivia and northern Argentina where it is recorded in both marine
and fresh wakr environments.
b. Osleoglossidue

Living Osteoglossidae are restricted to freshwater environments in

tropical South America, Africa and Asia. Some fossil remains were
reported from fresh water (Green River basin, Wyoming, Grande
1984; Sumatra, Sanders, 1934; Australia, Taveme, 1979), some from
brackish water (middle Cretaceous of Zaire; Taverne, 1976) and
some from marine environments (Paleocene of Zaire; Dartevelle &
Casier, 1959; Turkmenistan, Danil'chenko, 1968; Niger, Cappetta,
1972, Gayet & Meunier, 1983; Eocene of Italy, Taverne, 1979;
England, Woodward, 1901; Morocco, Arambourg, 1952; and
Denmark, Bonde, 1966). In well known marine Cretaceous levels
(Lebanon, Israel, Morocco, Italy, Portugal and Yugoslavia) where
the ichthyofauna is abundant, no fossils of this order are represented.
Consequently, Osteoglossiformes are considered as primary
freshwater fishes following Nelson (1969) and Gayet (1987b). In
Bolivia, Ostcoglossinac wcrc reportcd in rill mcmbcrs of the EI
Molino and in the Santa Lucia foniiatons, wlicrcns' Phareodontinae
were reported in the Santa Luca Formation only (Tiupampa). In the
El Molino Formation, they occur in association with marine fishes.
c. Cyprinodontiformes

Nothing can be said about the paleoenvironinent of the cf.

Cyprinidontiforines in Bolivia, because this group is too poorly
know II.

d. Polypteridae

Pishes
Fishes have been divided into four main categories (Myers, 1938)
based on their tolerance to fresh or salt water: 1) primary freshwater
fishes found only in fresh water; 2) secondary freshwater fishes,
found normally in fresh water, but capable of entering and surviving
in marine water; 3) peripheral freshwater fishes, diadromous, which

420

Living Polypteridae are Ahican freshwater fishes as are all fossils

of this group known from Senoniai to Pleistocene. In Bolivia h e y
were reported at Tiupampa (Santa Luca Formation) and occur in
association with marine fishes at Vila Vila (lower EI Molino
Formation).

e.

Lungfishes

Ceratodonts a r e generally associated with fresh water

environments by extrapolation based on occurrence of extant forms.
NeverUieless, ceratodonts occur in undoubtedly marine deposits in
Zaire (Casier, 1961), Niger dnd Algeria (Martin, 1984), Mali
(Tabaste, 1963). Egypt (Schad, 1984) and Kansas (Schultze, 1981).
According to Uiese different authors, these faunas may be interpreted
as d e a h assemblages of marine, brackish and hesh water forms or
that they were tolerant of a wide range of salinity.
Lepidosirenids are restricted to fkesh water environments.
2. Secondary freshwater rilies

fresh water species. This family includes lhb gcnus Ccrr~roponrrrs

from the coastal waters of the New World, a species wliicli enters
Brazilian rivers (Berra, 1981). Centropomidae are present at
Tiupampa and Criadero de Loro in Bolivia.
d. Cypriniformes
Cypriniformes are living freshwater fishes and fossil remains
attest, as far as is known, to a similar environment. Only
Ranrallicfitliys from Ein Jabrud (Israel) which may be considered as
an ancestral Cypriniformes (but is not a true Cypriniformes) is
marine. Cypriniformes are reported in the lower and basal middle
members of the EI Molino Formation in association with marine
fishes.

a. Lepisosteidae
e. Characiformes
Most of the living Lepisosteidae are found in fresh water, but one
species may tolerate brackish or even marine conditions and is found
along Uie coast of the Gulf of Mexico (Sutlkus, 1963). Bccouse of
their Uiick scales covered with ganoine which are more resistan1 llinn
any oflier parts of th fish, the fossil Lepisosteidae are often reported
on scaks only, which may be transported far from their biotope
without damage. In Bolivia. Lepisosteidne are always found in
association with marine fislies including selachians, pycnodonts and
eotrigonodonts in the lower and basal middle members of the El
Molino Formation, but they are always disassociated, indicating
postmortem movement. In the only locality where no true marine
fishes were found (.e.Tiupampa), Lepisosteidae are absent but Ulis
absence may e the consequence of age (Paleocene) and not of
environment. episosteidae had a nearly world-wide distribution
during Cretaceous and early Tertiary times. Subsequently, they
disappeared from all the continents except North America where
they are now restricted to the southeastern part of the USA and
Central America. I

b. Clupeidae
Gasteroclicpea branisui is a Clupeiformes endemic to the Andean
basin. Clupeiform,es are generally marine fishes with the exception
of some South American li.ving genera (RnnmoRnsrer, frisrignsrcr;
G6ry. 1969; Cionc p l'ercirii, 1985). III Uic middle Eocene of tlic
Green River Formation (Wyoming, USA), Kniglrtia and
Diploniystus which are known to be marine, are reported in strictly
fresh water environments (Grande, 19820. b). Gaslerocfupea
branisui is reported at Tiupampa (Santa Lucia Formation) where no
marine fishes are found, and in some levels of the EL Molino
Formation (at Agua Clara) in association w i h marine fishes. In some
green marl levels a t Agua Clara, Gasferoclupea is found in
association only with brackish-water ostracods. Consequently,
Gasteroclupea may be considered, with some confidence, as a
secondary Jeshwater clupeid.
e. Centropomidae

'

The family Centropomidae, as defined by Greenwood (1976). is

composed of marine and fresh water genera. Lares, to which the
Bolivian genus may be compared, is known to include marine and

Living Characiformes are freshwater fishes (only a few

cxccptions enter brnckisli waters) while sonic fossils werc
apparcnlly adapted to brackish water (Cappetta el al., 1972;
Gaudant, 1980). The oldest true Characiformes, Sulnrinops
( G a y e t , 1 9 8 5 ) , w a s m a r i n e ( G a y e t , 1981). I n B o l i v i a .
Characifornies are reported i n the lower and basal middle
m e m b e r s o f t h e EI M o l i n o F o r m o t i o n ( S e r r a s a l i n i d a e ,
My l e in a e ; C li ar a c i d ae, T e t r a go nop ter i n a e ) , i n t li e S a n t a
Luca Formation at Tiupampa (Serrasalmidse, Myleiiiae and
S e r r as a I m in a e ; C li a r a c i d a e , T e t r a g o no p t e r i II a e a n d c f .
R h o a d s i i n a e ) a n d in t h e E o c e n e a g e C a y a r a Formation
(Serrasalmidae, Myleinae). In the El Molino Formation they
occur in the same levels as marine fishes.

f. Siluriformes
Most S i l u r i f o r m e s o c c u r i n fresh water. However; the
Plotosidae are an advanced marine family which today occur
around Auslralia and the Ariidae live along the coast or i n t e r
e s t u a r i e s and c o a s t a l rivers. T e r t i a r y S i l u r i f o r m e s were
reported in fresh water areas and/or in brackish water (see
.Gayet, 1991). In Bolivia, Andiniclithyoidea (3 families) are
known i n the Siinln Lucln Forincilion n t Tiupiiinpn aiid Rlnnco
Kiinclw (tlic ichtliyol'iiunii ol' the liiter lociility is not yet well
enough known to provide pnleoenvironmental interpretation);
they are not reported at Agua Clara o r Hotel Cordillera where
m a r i n e f i s h e s a r e p r e s e n t a n d may b e c o n s i d e r e d as
freshwater forms. Ariidae are reported everywhere in both
formations with or without marine fishes. They are present at
Tiupampa, but are few in number. At Hotel Cordillera they
a r e v e r y numerous a n d o c c u r in a s s o c i a t i o n with a few
marine taxa.
3. Marine fislies

a. Selachians
Selachians are, considered as marine fishes. Nevertheless, one
family of American batoids (Potamobygonidae) lives exclusively in
fresh waters. One pristid fish and a shark can also enter rivers or

421

MIKI?ILLI?tiAYE1. LARRY

U. MAWIIALL&'I'IIIBKKY SBMI't!KI!

Inkcs, but it is iniportnnt to note thnt these elasinobranch fishes are

marine species which temporarily enter fresh water. According to
Cappetta (1991), il is highly probable that the rocks containing
selachians in Bolivia were deposited in marine environments.

b. I'ycnodontiformes
Pycnodonts, a purely fossil group, are considered to have been
marine fishes (Cione 8c Pereira, 1985). They were flat and high
rishcs that apparently swam slowly along the coast and were l i n k d
to the substratum for their alimentation. Some could probably enter
waters of lesser salinity. Wenz (1989) noted that the Brazilian lower
Cretaceous pycnodontid Ieninnja may have lived in fresh water.
Traquair (19 1O) thought that pycnodonts found at Bernissart
(Relgium) with thc rcptilc Igrrunodon were living in fresh water
without any connection with the sea. Nevertheless, these examples
are sporadic and pycnodonts are typically found in more or less open
marine environments in rocks of Jurassic and Cretaceous age (Hake1
and Ht1jula in Lebanon; Ein Jabrud i n Israel; Jebcl Tselfnt in
Morocco; etc.). In Bolivia. pycnodonts were reported i n the
Chaunaca Formation (only one tooth of no paleoenvironmental
significance) and in,the lower and basal middle EI Molino.

More difficult is lhe interpretation of tlic ichtliyofaunn from the EI

Molino Formation. In nearly all tlie localities. niarine and fresh water
fishes occur in association and, as noted above, bones, teeth and
scales are never found articulated. It is probable h a t the calcareous
fossiliferous levels of the lower member of tlie EI Molino Formation
at Agua Clara were deposited in or near tlie sea (lagoon or coast).
Other levels at the same locality (e.g. some green mark) were
probably deposited farther from the sea (perhaps without any
connexion with it). Gusferoclupeu,which has been found articulated
in these levels. has also been rcportcd at Tiupampa and is possibly a
freshwater genus. Nevertheless, this does not dismiss the possiblity
of a brackish water habit for this genus.
Camoin el al. (1991) concluded that tlie EI Molino rocks were
deposited in a conl~ncntalenvironment. while Gayet ef al. (1992)
and Sempere (in press) noted that marine fish taxa occur in these
sedimentary rocks and discuss their depositional environment.,

c. "Snlinoniformes"

"Salinoniforines" are a paraphyletic group present in all kinds of

environ in e nt s (B krr a, 19 8 1). Nevertheless, Encho don tid ae
(Enchodus) and Ichthyotringoidei (including ?Apareodiis) are
known only in marine rocks (Goody, 1969). In Bolivia,
"S almoniformes" were reported in the lower and basal middle
members of the EI Molino Formation.
d. Tetraodontiformes

One living family of tlie o r d e r Tetraodontiformes, the

Tetraodontidae with about 9 0 species, has a few fresh water
representatives in South America, Africa and India (Berra, 1981).
However, Eotrigonodontidae is a fossil family so far known only in
marine deposits. Sfephuriodrrs has only been fouiid in Iate
C r e t x e o u s marine scdiincnts. I n Bolivia. the cotetragonopterid
Sfephanodrts as bdcn rcportcd in the first two rneinbcrs of the EI
Molino Forination.

In conclusion, ye note that at Tiupampa and Criadero de Loro

(Santa Lucia Formation, Paleocene), no' true marine fishes
(selachians, pycnodonts, Salmoniformes and Eotrigonodontidae)
have been reported. All of the primary freshwater fishes reported in
Bolivia, with the exception of Semionotidae (Lepidofes) which do
not cross the Cretaceous/Paleocene boundary, are present at these
localities. Although some Osteoglossidae may occur in marine
waters (i.e. Brychaerrrs), the subfamily Phareodontinae is known thus
far only in fresh water levels. Tiupampa is Uie only Bolivian locality
where nearly complete skulls (Osteoglossiformes, Siluriformes and
Characiformes) were found. In all other localities bones are always
separated, attesting to postmortem movement. Thus, based solely on
the ichthyofauna. we can confidently conclude that the Tiupampa
and Criadero de Loro fossiliferous sediments were deposited in a
fresh water environment.
b .

422

IIiglier ver1el)rules

Crocodiles of the family Dyrosauridae generally occur in siiallow

niarine near-shore deposits, but can exist i n fresh water as well
(Bufretaut, 1982). The ainpiiibians. lizards. snakes. turtles. otlicr
families of crocodiles and mammals all indicale terrestrial and/or
fresh water paleoenvironmcnts.

D. FALEOBIOGEOGRAPHY
Numerous vertebrate grou'ps in the late Cretaceous and Paleocene
of South America are also recorded in North America, and these
faunal "links" were used by paleontologists to postulate that a transCaribbean land bridge existed during part or all of this time interval
(see Gayet el al., in press; Marshall & Sempere, in press and
references therein). These concIusions have recently been
corroborated by geodynamic data (see Pindell ef d.,1988; Stephan
el ul., 1990). It now appears that the Caribbean plate began its eastnortheastward migration in Campanian time. The now submerged
Greater Antilles and the Aves Ridge formed a subaerial volcanic arc
established 'on the northe:istern edge of the Caribbean plate during
tlic late Crctaccous and early I'aleocciic. It is likely h a t this arc was
the land bridgc that providcd the pathway for dispersal of continental
(and fresh water) vertebrates. However, the land bridge provided by
the Aves Ridge was apparently strongly dependent on tlie inaginatic
and tcctoniC activity of this arc. and on sen level changes. Maginatictectonic quiescence and/or marine high-stand periods would have
promoted relative subsidence of part of the isthmus below sea level.
In view of the probable fragility of the Aves Ridge connection,
faunal interchange is likely to have occurred i n several "pulses"
beginning in the Campnninn until the land bridge finally disappeared
in late Paleocene time as is suggested by paleontological data. This
"pulse-IiypoUiesis" would have permitted opportunities for endemic
evolution of first-wave immigrants and for subsequent dispersal
during a later "pulse". For this reason, the direction of dispersal of
most groups is not certain (Marshall & Senipere, in press).
Some vertebrate groups which participated in this interchange(s)
include dinosaurs (Avisauiidae, Ceratopsidae, Hadrosauridae,
Titanosauridae), snakes (Anilioidea, Boidea). possibly lizards
(Teiidae) and sliore-dwelling turtles (Bohremydidae) (Gayet el al.,

i n prcss; MarslisIl & Senipere, i n prcss und references thcrcili).

Aliiong fishes arc the Lepisosteidne (Lepisosteus), Osteoglossidae
(I'hareodontinae) and possibly Cypriniforines and Ariidae (Gayet el
al.,in press).
Multiple taxonomic similarities among mammals (10 examples)
"link" late Cretaceous and/or Paleocene faunas in North America
with early and middle Paleocene (Tiupampian and Itaboraian,
respectively) faunas in South America. Among marsupials,
Peradecles is known in Tiupampian faunas of Bolivia and Peru, and
is present in early Paleocene faunas of North America;
caroloameghiniids (Glnsbirts) are present in tlie late Cretaceous of
North America and Tiupampian (Robertlioffstetleria) of South
Anierica; Didelphidne are present in the early Eocene of North
Anicric:i and Tiupampian of South America; and Pediornyidae (Iate
Cretaceous of North America) are a sister-group of
Microbiotlieriidae (first recorded in the Tiupampian of South
Anierica). Among placentals, a Leptictida (cf. Ciniolesres) is known
I
from Tiupampa while Ciniofestes occurs in the late Cretaceous and
carly Palcoceiic of North Anicrico; I'niitodontn (Alcidedor Digt~ya)at
Tiupampa a r e first known i n the middle Paleocene in North
America; mioclaenine hyopsodontid Condylwtlua at Tiupampa are
fist recorded in early Paleocene faunas of North America; Xenarlhra
(Dasypodidae) first appear in Itnboraian faunas in South Amcrica
and in the late Paleoccne in North America; aid Dinocerata which
first appear in the late Paleocene of North America are regarded as a
sister-group of Xenun d a t a which occur in the Itaboraian of South
America (Marshall & uizon, 1988; Marshall & Seinpere, in press;
Gayet ef al., in press and references herein).
Although latest Cretaceous and Paleocene interchanges are well
docuiiiented, it is of interest to note that several terrestrial groups of
vertebrates which were often rather widely distributed either in the
Laurasirui realm or on Gondwanian continents in the late Cretaceous
and Paleocene, never crossed the Caribbean nor Tethyan areas.
Acipenserid, polyodonlid and gonorhynchoid fishes, as well as
palaeobatrachid frogs, baenid turtles, and anguimorph lizards
inhabited North America during the Cretaceous; hiodontid and
esocid fishes were present on that continent during die Paleocene.
None of these northern groups rdached South America. Conversely,
polypterid and characifonns, as well as pipid frogs and araripeniyid
turllcs of Gondwunirin origin. known i n the Crctnccous of South
Anici cil. did not rcricli Noilli Anicricli. Tlic iilx~vclisted vcrlcbrnlcu
rank aniong tlie most signiliciiiit ones that did not tuke part' in the late
Cretaceous and Paleocene interchange(s). Other groups could be
added to this list but, for the time bcing, they cannot be considered to
be very conclusive bccause their distribution is still inadcquately
known.
The failure of various groups to cross the Caribbean area would
apparently favour the hypothesis of a discontinuous terrestrial route
or even an island hopping hypothesis. But the ascertained dispersals
of large-size animals as Titanosauridae and of freshwater fishes
(Lepisosteidae, Phareodontinae, and perhaps Ariidae) are
inconsistent with such possibilities as they require a continuous land
(fluviatile for the fishes) route. These failures could result from
climatic, faunal interactions (niches occupied by ecologically
vicarious autochthonous forms) or shortness of duration of the
unbroken nature of the land route.

423

CONCIJUSIONS
This paper is the first attempt to provide a detailed overview of h e
Mesozoic and Paleocene vertebrate faunas of Bolivia within Uicir
stratigraphic context. Much of the inforination presented here is new,
and includcs data obtuiiicd as a result of IIII iiitciise palcontological
program started in 1981 (M. G., L. G. M and others) and of geologic
studies initiated in 1984 (T. S. and others). Most of the fossils are
still being described and it will be many years before monographic
publications can be completed. In addition, magnetoslratigrapliic and
isotopic (Ar/Ar, K-Ar) studies of the El Molino and Santa Luca
formations and isotopic oxygen and strontium analyses on fishes Ne
in progress, and cornpletion of this gcochronologic program will
hopefully perinil secure calibrntioii of these rock units witti tlie
geologic time scale. Synthetic overviews for the biostratigraphy of
Argentinian, Chilean and Peruvian rocks in the Andean basin are
now urgently necdcd to permit refined correlation with those in
Bolivia.
Dcspile tlic fact tlint his is by iicccssity only ii "progrcss rclwri",
many advances in knowledge have been made 011 the Bolivian
Mesozoic-Paleocene scqucnce during the last few years, and in this
study we have been able to clarify some debated issues. The most
iinprtant of thcsc is thc apparcnt, although still ill-dcfincd, location
boundary and its position relative to Uic inanmal-bearing
of the
level at Tiupampa. Within the El Molino Formation, undoubted late
Cretaceous taxa occur only in Uie lower and basal middle members.
It therefore appears that the K/T boundary lies somewhere in the
upper part of the niiddlg member, while the upper member may be
regarded as earliest Paleocene based on the absence of diagnostic
Cretaceous taxa and on Uie palynological studies of a stratigraphic
equivalent in northwestern Argentina. The EI Molino Ihus appears to
range from latest Cainpanian to earliest Paleocene, while the
disconformably overlying Santa Luca Formation appears 'to be
early, but not earliest. Paleocene. Thus, confra Van Valen (f988),
the EI Molino dinosaurs were all late Cretaceous (apparently-early
Maastrichtian), while the land mammals from the Santa Luca
h
Fonnatioii at Tiupanpa were Paleocene.
ACKNOWLEDGMENTS
~ s l ) u~
r tilis
~ ~sttltiy
s
W C I C su1)l")ltcd I)Y rillitis fro111 IIIC cclltlc
National de la Keclicrclie Scientifique (UKA 12, Institut de
PalCoiitologie, Paris, 1981); ari "Action Spcifique" (1983 to 1989)
from the MusCuni National d'Histoire Naturcllc, Paris; National
Geographic Society (2467-82. 2908-84, 3381-86); URlH of Orstoni;
the Gordon Barbour Fund. Department of Geological and
Geophysical Sciences, Princeton University; and the National
Science Foundation (EAR-8804423). Logistic help was given by
Orstom, YPFB and VARIG. Fossil collecting was carried out under
the auspices of the Asociacin Boliviana de Paleontologa and
stratigraphic work under the YPFB-Orstom research prcgram on
Bolivian geology. Thanks to L. Barrios, J. Blanco, F. de Broiii,
E. Buffetaut. G. Camoin, H. Cappetta, R.' Cspedes, A. L. Cione.
J. Dejax, D. Dingerkus. E. Jaillard, M. Surez, R. SuBrez,
F. J. Meunier, J. Oller, J. C. Rage, B. Sig, C. de Muizon and S.
Wenz for help in aspects of Ulis study. Figures 1 to 4 were drawn by
E. Liebmm.

AGASSIZ, L. 1841. On the fossil fishes found by Mr. Gardner in

the province C e a r l , i n t h e north of Brazil. Edinburgh New
Philosophical Journal, 30 (59): 82-84. Edinburgh.
AHLFELD F. & L. BRANISA. 1960. Geologa de Bolivia. Instituto
Boliviano del Petrleo, p. 1-245, La Paz.
ALBINO, A. M. 1986. Nuevos Boidae Madtsoiinae en el Cretlcico
tardo d e Patagonia (Formaci611 Los Alamitos, Ro Negro,
Argentina). IV Congreso Argentino de Paleontologia y
BioestrafigraJfa,Actas 2: 15-21, Menclom.
ALBINO, A. M. 1987. The Late Cretaceous fauna of Los Alamitos,
Patagonia, Argentina. Part V - The Ophidians. Revista del Museo
Argentino de Ciencias Nalurales "Bernardino Rivadavia",
Paleontologa, 3 (3): 141-145, Buenos Aires.
ALONSO, R. N. 1980. Icnitas d e dinosaurios (Ornithopoda,
Hadrosauridae) en el Cretcico Superior del Norte de Argentina.
Acta Geolgica Lilloam, 15: 55-63, Tucumh.
ALONSO, R. N. & R. A. MARQUILLAS. 1986. Nucvn localidad
con huellas de dinosaurios y primer Iiallazgo de huellas de aves en la
Formaci6n Yacoraite (Maastrichtiano) del Norte Argentino. IV
Congreso Argentino de Paleontologfa y Bioestratigraffa, Actas 1:
3 3 4 0 , Mcndoza.
AMEGHINO, F. 1906. Lcs formations sCdimentaires du CrdtacC
Supbrieur et du Tertiaire d e Patagonie, avec un paralMe entre leurs
faunes mammalogiques et .celles de l'ancien continent. Anales del
Museo Nacional de1Historia Natural, 15, Serie 3, 8: 1-568, Buenos
Aires.
ANDREIS, R.'R. 1987. T h e l a t e C r e t a c e o u s f a u n a of Los
Alainitos, Patngonia, Argentina. P a r t . I - Stratigraphy and
Paleoenvironment. Revista del Museo Argentino de Ciencias
Naturales "Bernardino Rivadavia", Paleontologa, 3 ( 3 ) : 103110, Buenos Aires.
ANDRESIS, K. R., M. M. MAZZONI & L. A. SPALLETTI. 1975.
Estudio estratigrfico y paleoambiental de los sedimentos terciarios
entre Pico Salamanca,y Baha Bustamente, Chubut, Argentina.
Revista de la Asociacin Geolgica Argentina, 30: 85-103, Buenos
Aires.
ARAMAYO, S. A. 1981. Hallazgo de LRpidotes maxinius, Wagner
(Pisces) cri el Titoniano dc la Provincia de NeuquCn, Argentina. I
l
Congreso Latino-Americano de Paleontologfa, Anais: 322-329,
Porto Alcgre.
ARAMBOURG, C. 1948. Mission scientifique de l'Omo,, Gdologie
et Pa160ntologiei Firmin-Didot & Cie (ed.),562 p., Mesli.
ARAMBOURG, C. 1952. Les verttSbr6s fossiles des gisements de
phosphates (Maroc-AlgCrie-Tunisie).Notes et Mhoires du Service
gologique du Maroc, 92: 1-372, Paris.
ARAMBOURG, C. & L. JOLEAUD. 1943. Vertkbrds fossiles du
Bassin du Niger. Bulletin de la Direction des Mines de l'Afrique
Occidedale Franaise. 7: 1-74, Dakar.
ARGOLLO, J.,E. BUFFETAUT, H. CAPPETTA, M. FORNARI,
G. HERAIL, G. LAUBACHER, B. SIGB & G. VISCARRA. 1987.
Dgcouverte de, vert6brks prksumds p a l k o c h e s dans les Andes
septentrionales de Bolivie (Ro Suches, Synclinorium d e Putina).
Geobios, 20 (1): 123-127, Lyon.
BAEZ, A. M. 1,985. Anuro leptodactdo en el Cretcico Superior
(Grupo Bauru) de Brasil. Anreghiniana, 22: 75-79, Buenos Aires.

BAEZ, A. M. 1987. The lntc Crctnccous fauna of Los Alamitos, ,'

Patagonia, Argentina, Part III - Anurans. Revista del Museo
Argentino de Ciencias Naturales "Bernardino Rivadavia",
Paleontologa, 3 (3): 121-130, Buenos Aires.
BAEZ A. M. & S. PERI. 1989. Baurubatrachus price;, nov. gen. et
sp., un Anuro del Cretlcico superior de Minas Gerais, Brasil. Anais
da Academia Brasileira de Cincias, 6 1 (4): 447-458, Rio de
Janeiro.
BARDACR, D. 1961. New Tertiary teleosts from Argentina.
Anierican Museum Novitales, 2041: 1-27, New York.
BENEDETTO, J. L. & T. M. SANCHEZ. 1972. Coelodus
toncoensis nov. sp. (Pisces, Holostei, Pycnodontiformes) de la
Formacidn Yacoraite (Cretlcico Superior) de la provincia de Salta.
Ameghiniuna, 9 (1): 59-71, Buenos Aires.
BERRA, T. M. 1981. An Atlas of Distribution of the Freshwater
Fish Families in the World. University ofNebraska Press, 197 p.
BERTINI, R., L. G. MARSHALL, M.GAYET & P. BRITO. In
press. Vertebrate fauna from the Adamantina and Marlia Formations
(Upper Bauru Group), lntc Crctaccous, Brazil. Nerres Jalirbrrclr Jiir
Geologie irrui Palon/ologie Abhandlungen, Stuttgart.
BLOT, J. 1987. L'ordre des Pycnodontifonnes. Studi e riclterclre sui
giacimedi terziari di Bolca, 5: 1-211, Verona,
BOCCHINO, K. A. 1964. Sobrc tin I'ygidiidac (Pisces. Siluriformes)
del Eoccno dc Klo Negro. Ameglriniana, 1 I (7): 185-189, Buenos
Aires.
BONAPARTE, J. F. 1986a. A new and unusual late Cretaceous
mammal from Patagonia. Journal of Vertebrate Paleontology, 6:
264-270, Nonnan. '
BONAPARTE, J. F. 1986b. Sobre Mesungitlatirni hoirssayi y
nuevos mamferos crctdcicos de Patagonia, Argentina. I V Congreso
Argentino de Paleontologia y Bioestratigraifa, Actas 2: 48-61,
Mcndoza.
BONAPARTE, J. F. 1986c. History of the terrestrial Cretaceous
vertebrates of Gondwana. n/ Congreso Argentino de Paleontologfay
Bioestratigrafia,Achs 2: 63-95, Mendoza.
BONAPARTE. J. F. 1987. The late Cretaceous fauna of Los
Alamitos, Patagonia, Argentina, Part VU1 - The Mammals. Revista
del Museo Argentino de Ciencias Naturales "Bernardino
Rivadavia", Paleontologia, 3 (3): 163-169, Buenos Aires.
BONAPARTE, J. F. 1990. New late Crctaccous mammals from Uie
Los Alainilos Forinntion, northcrn Patngonin. National Gcograpliic
Research, 6 (11: 63-93.
BONAPARTE, J. F., A. M. BAEZ, A. L. CIONE, F. DE BROIN, J.
E. POWELL & A. ALBINO. 1987. The late Crctnccous fauna of Los
Alamitos, Patagonia, Argentina. Part IX - Resumen. Revista del
Museo Argenlino de Ciencias Naturales "Bernardino Rivadavia",
Paleontologa, 3 (3): 171-177, Buenos Aires.
BONAPARTE, J. F. & R. PASCUAL. 1987. Los niamiferos
(Eotheria, Allotheria y T h e r i a ) d e la Forniaci6n Los
Alamitos, Campaniano de Patagonia, Argentina. I V Congreso
Latinoamericano de Paleontolgfa, Actas 1: 361-378, Santa
Cruz.
BONAPARTE, J. F. & J. E. POWELL. 1980. A continental
assemblage of tetrapods from the upper Cretaceous beds of El Brete,
northwestern Argentina (Sauropoda-Coelurosauria-CarnosauriaAves). Mmoires de la Socit Geologique de France, nouvelle
s6rie. 59 (139): 19-28, Paris.

424

FOSOSILES Y FACIES DE DOUMA VOL.I VERIEUKI\IX)S

BONAIARTE. J. F. & G. I<OUGIER. 1987. The late Crctnccous

f n u n n of Los Alarnitos, Patagonia. Argentina, Part VI1 - The
Hadrosaurs. Revista del Museo Argentino de Ciencias Naturales
BernardinoRivadavia, Paleontologa, 3 (3) : 155-161. Buenos
Aires.
BONAPARTE, J. F., J. A. SALFITY, G. BOSSI & J. E. POWELL.
1977. Hallazgo de dinosaurios y aves cretlcicas en la Formaci6n
Lecho de El Brete (Salta), pr6ximo al lmite con Tucumin. Acla
GeoMgica Lilloana, 14 5-17, Tucumhi.
BONAPARTE, J. F. & M. SORIA. 1983. El primer mamfero del
Cretficico Argentino. C r c u l o Informativo, Asociacidn
Paleontolgica de Argettfina, 11: 5 , Buenos Aires.
BONAPARTE, J. F. & M. SORIA. 1985. Nota sobre el primer
mainfero del Cretlcico Argentino, Campaniano-Maastrichtiano
(Condylarthra). Anteghiniuna, 21: 177-183, Buenos Aires.
BONDE, N. 1966. The fishes of the Moy-Clay Formation (lower
Eocene). A short review. Meddelerser f r a Dansk Geologisk
Fororctiiq, 16: 198-2021
DONDESIO, I.& li.PASCUAL. 1977. Restos de Lcpidosircnidue
(Ostciclithyes, Uipnoi) del Grupo Hondn (Mioceno tardo) de
Colombia. Sus denotnciones paleoambientales. Asociacin
Geolgica Argentina, 32 (1): 34-43, Buenos Aires.
RRANISA, L. 1968. Hnllnzgo del ammonite Neolobites en Ia Calizn
Miriiflores y de huellns de dinosaurios en la Formaci6n EI Molino y
su significado para la determinacin de la edad del Grupo Puca.
Boletn del instituto Boliviano del Petrleo, 8: 16-29, La Paz.
BRANISA, L., L. GRAMBAST & R. HOFFSTETTER. 1969.
Quelques prcisions nouvelles, dapr8s les charophytes sur Ige du
Groupe Puca (Crtac-PalogEne, Bolivie). Coniptes Rendus
Soniniaires des Sances de la Socit Gologique de France, 8: 321322, Paris.
BRANISA, L., R . HOFFSTETTER & J . SIGNEUX. 1964.
Additions 14 faune ichthyologique du Crtac Suprieur de Bolivie.
Bulletin du Musum National dlfistoire Naturelle, section 2, 36 (2):
219-297, Paris.
RRANISA, L., R. HOFFSTETTER, S. FRENEIX, J.ROMAN & J.
SORNAY. 19GG. N o u v c l l c c o n t r i b u t i o n h Idtudc d e I n
p a l o n t o l o g i e e t d e I a g e d u G r o u p e P u c a (CrtacC d e
Bolivie). Bulletin du Musum National dHistoire Naiurelle,
section 2, 38 (3): 301-310, Paris.
BRETT-SUKMAN, M. K. & G . S. PAUL. 1985. A new family of
bird-like dinosaurs linking Laurasia and Gondwana. Journal of
VertebratePaleonfology,5: 133-138, Norman.
BRITO, I. M., H. E. C. M. FRANKE & D. de A. CAMPOS. 1972.
Geologla e Petrografa da Bacia de So.JosC de Itnboraf, Estado do
Rio de Jnneiio. Arrois da Aca&nria Bra.dcCa deCincim, 4 4 225-234.
BRlTO. I. & M. GAYET. in press. Dados preliminares sobre
a ictiolauna d o Grupo Bauru, Cretaceo Superior d o Estado de
S l o Paulo. Anais da Acadniia Brasileira de Cincias, Rio de
Janeiro.
BROIN, F. d e . 1971. Une esp8ce nouvelle d e tortue pleurodire
(?Roxocltelys vilavilensis n. sp.) dans le Crtac Suprieur de
Bolivie. Bulletin de la Socit Gologique de France, 13 (3-4): 445452, Paris.
BROIN, F. de. 1987. The late Cretaceous fauna of Los Alamitos.
Yatagonia, Argentina, Part IV - Chelonia. Revista del Museo
Argenfino de Ciencias Naturales Bernardino Rivadavia,
Paleontologa, 3 (3): 131-137, Buenos Aires.

BROIN, F. de. 1988. Les tortucs et ICGondwana. Exnnicii des

rapports elitre le Frnctioiineniei~tdu Gondwnnn e t In dispersion

gographique des tortues pleurodires a partir du CrBtac6. Studia

Palaeochelotiiologica, 2: 103-142, Salamanca.
BROIN. F. de. 1991. Fossil turtles from Bolivia. In: Fdsiles y
Facies de Bolivia. (R. Suirez-Soruco. ed.). Revista Tdcnica de
YPFB, 12 (3-4) 509-527, Santa Cruz.
BUFFETAUT, E. 1980. Histoire bioghgraphique des Sebecosuchia
(Crocodilia, Mesosuchia): un essai dinterpretation. Annales de
Palontologie des Vertbrs, 66 (1): 1-18, Paris.
BUFFETAUT, E. 1982. Radiation volutive, palCobologie et
biogCographie des Crocodiliens mbsosuchiens. Mmoire de ta
Scxit Gologique de France, 60 (142): 1-88, Paris.
BUFFETAUT, E. 1987. Occurrence of the crocodilian
Dolichocltanipsa niininla (Eusuchia, Dolichochmpsidae) in the El
Molino Formation of Bolivia. Bulleiin de la Socit Belge de
Gologie, 96 (2): 195-199, Bruxelles.
BUFFETAUT, E. 199 1. Fossil crocodilians froin Tiupnmpii,
Bolivin: u prcliininnry repoil. lu: F6sile.s y Ftrcics dc flolivin (K.
Sudrcz-Soruco, ed.). Revista ldcnica de YPFD 12 (3-4): 541-544,
Santa Cruz.
BUFFETAUT. E. & L. G. MARSHALL. 1991. A new crocodilian,
Sebecrrs querejazris nov. sp.( Mcsosuchiu, Scbccidnc) rrom tlic Santa
Luca Formation (early Ialcocenc) at Vila Vila, southcentral Bolivia.
In: Fsiles y Facies de Bolivia (R. Surez-Soruco, ed.). Revista
Tcnica de YPFB, 12 ( 3 4 5 4 5 - 5 5 7 , Santa Cruz.
CAMOIN, G., J. M. ROUCHY, J. F. BABINOT, J. F.
DECONNINCK & G. TRONCHETTI. 1991.: Dynamique
sdimentaire et contrle climatique des systmes continentaux
maastrichtiens de la Cordillre Orientale (Bolivie sud-occidentale).
Coniptes Rendus If ebdpniadaires des Sances de IAcaddmie des
Sciences de Paris, Srie II. 3 1 2 1335-1341, Paris.
CAPPETTA, H. 1972. Les poissons crtac6s et tertiaires du Bassin
de Iullemmeden (RCpublique du Niger). Palaeovertebrata, 5 (5):
179-243, Montpellier.
CAPPEITA. H. 1975. Sur quelques Sdlaciens nouveaux du Cr6tacC
Suphicur de nolivie ( A m 6 r i c p du Sud). Geoln~s,8 (1): 5-24, Lyon.
CAPPETTA, H. 1987. Mesozoic and Cenozoic Elasniobraichii. In:
Handbook of Paleoichthyology (H.-P. Schultze, ed.), Chondrichthyes
II, 3B. 193 p.. Stutlgnrt/Ncw York.
CAPPETTA, H. 1990. Tcrlinry dinosaurs in South America? A reply
to some of Van Valens assertions. Hisforical Biology, 3: 265-268,
London.
CAPPEITA. fI. 1991. Late Crelaccous selachiai faunas from Bolivia:
new data and sunimary. In: F6siles y Facies de Bolivia (K. SufitezSOIUCO,
ed.). Revista 7cnica de YPFB, 12 (3-4):435-440, Santa Cruz.
CAPPETTA, H.. J. J. JAEGER, M. SABATIER, B. SIG& J.
SUDRE & M. VIANEY-LIAUD. 1978. Dcouverte dans le
PalCoc?ne du Maroc des plus anciens niaminif5res euthdriens
dAfrique. Geobios, 2 (2): 257-262, Lyon.
CAPPETTA, H., D. E. RUSSELL & J. BRAILLON. 1972. Sur la
dcouverte de Characidae dans 1Eocne infrieur franais (Pisces,
Cypriniformes). Bulletin du Musuni National dHistoire Naturelle,
38me srie, 51, Sciences de la Terre, 89: 37-51, Paris.
CARRASCO, C.. R. BALLON, H. PEREZ & K. OKAMOTO. 1989.
Anllisis estratigrifico del sinclinal Cretlcico d e Vila Vila, zona
Tiupampa - Prov. Mizque-Dpto. Cochabamba. Boletfn del Servicio
Geolgico de Bolivia, Serie A, 4 (1): 2-14, La Paz.

425

CASIER, E. 1943. Contribution h I'Ctudc dcs poissons fossiles de

Belgique. II. Rcstes du gcnre Lepidosteris du Laiidiiien continental
de la Hesbaye. Bulletin du Musuni National d'Histoire Naturelle de
Belgique, 19 (1): 1-12, Bruxelles.
CASIER, E. 1946. Faune ichthyologique du London Clay. Trustee of
the British Museuni (Natural History): 1-496, London.
CASIER, E, 1960. Note sur la collection des poissons palCoches et
ocnes de l'Enclave de Cabinda (Congo). Annales du Muse Royal
du Congo Belge, A3, I(2): 1-48, Bruxelles.
CATTOI, N. & M.A. FREIBERG. 1958. Una nueva especie de
"Podocnentis" del Cretlceo argentino. Pl~ysis,W.58-67, Buenos Aires.
CHALIFA, Y. 1989. New species of Encliodus (Pisces:
Enchodontoidei) from the lower Cenomanian of Ein-Yabrud, Israel.
Journal of Vertebrate Paleontology, 63 (3): 356-364.
CHARRIER, R. & K. J. REUTTER. 1990. The Purilaclis Group of
northern Chile: link between arc and back-arc during late Cretaceous
and Paleogene. Internulional Symposium on Andean Geodynaniics:
249-252, Grenoble.
CHAVEZ. H. C . 1987. Interprctaci6n gcodinlmica de l a s
forniacioiies sedimentarias y estructuras tcct6nicas de la zona de
Challapata. Unpublished Thesis, Universidad Mayor de San Andrs,
Facultad de Ciencias Geolgicas, 163 p., La Paz.
CHEKKONI. C. 1977. EI sistcnia Crcthcico ci1 Ia parle boliviana de
la cuenca andina. ReviAta Tcnica de YPFB, 6: 5-46, La Paz.
CIONE, A. L. 1977. Algunas consideraciones sobre
Pycnodontifornies (Pisces, Holostei) procedentes de la Formaci6n
Yacoraite, Cretcico tardo de la provincia de Salta, Argentina.
Anieghiniana, 14: 315-316, Buenos Aires.
CIONE, A. L. 1987. The Late Cretaceous fauna of Los Alamitos,
Patagonia, Argentina, Part II - The fishes. Revista del Museo
Argentino de Ciencias Naturales "Bernardino Rivadavia",
Paleontologa, 3 (3): 111-120, Buenos Aires.
CIONE, A. L. & G. LAFITTE. 1980. EI primer Siluriforme (Pisces,
Ostariophysi) del Cretcico de Patagonia. Consideraciones sobre el
rea de diferenciacin de los Siluriformes. Aspectos biogeogrficos.
II Congreso Argentino de P aleontologla y Bioestratigrafa y I
Congreso Lofinoaniericano de Paleontologa (1978), Actas 2: 3546,
Buciios Aires.
CIONE, A. & S. M. PEKEIKA. 1985. Los peces de la Formaci611
Y acornitc (Crct6cico tardo-Tcrcinrio, Noroeste Argctitina) coiiio
indicadores dc saliiiidad. Nevista de la Asociacidtt Geolbgica
Argentina, 4 0 (1-2): 83-88, Buenos Aires.
CIONE, A., S. M. PEREIRA, R. ALONSO & J. ARIAS. 1985. Los
Bagres (Osteichthyes, Siluriformes) de la Fop"61i Yacoraite
(Cretlcico tardo) del Noroeste Argentino. Coiisidqracioiics
biogeogrlficas y bioestratigrlficas. Aniegliiniana, 21 (2): 294-304,
Buenos Aires.
COPE, E. D.1885. A contribution to the vertebrate paleontology of
Brazil. Proceedings of the American Pliilosophical Sociely, 11: 1516, Philadelphia.
COWIE, J. W. & M. G. BASSETT. 1989. Global Stratigraphic
Chart. Supplement to Episodes, 12 (2).
CROCHET, J. Y. 1980. Les marsupiaux du Tertiaire d'Europe.
Edition Fondation Singer-Polignac, 1: 1-279, Paris.
CUNHA, F. L. S. & C. S. FERREIRA. 1980. Um Dipnoi n a
formao Itapecuru (Cenomaniano), Maranho, Brasil. II Congreso
Argentina de Paleorrtologa y Bioestratigrafa y I Congreso

Latinoanicricano de Palco~ito[ogfn(1978). Actas 2: 1-9, Buenos

Aires.
DANIL'CHENCO. P. G . 1968. Ryby verklinego paleotsena
Turkmenii. In: Oclterki PO jlogenii isistentdike iskapaeniykli ryb i
bezsclieliustnykk ( D .V. Obruchev, ed.). Nauka Press; 113-156,
Moscow.
,
DARTEVELLE, E. & E. CASIER. 1949. Les poissons fossiles du
Bas-Congo e t des rCgions voisines ( d e u x i h e partie). Annales du
Muse dir Congo Belge. A . Minralogie, Gologie, Palontologie,
srie 3,2 (2): 201-256, Tervuren.
DARTEVELLE. E. & E. CASIER. 1959. Les poissons fossiles du
Bas-Congo et des rgions voisines (troisime partie). Annales du
Muse du Congo Belge. A . Minralgie, Gologie, Palontologie,
sCrie 3, 2 (3): 257-368. Tervuren.
,DAVILA, J. & V. PONCE DE LEON. 1971. La seccidn del Ro
Iiiambari en la faja subandina del Per y la presencia de sedimentitas
de,la Formacin CancaRiri (Zapla) del Silrico. Revista Tcriica de
YPFB, 1 (1): 67-85. La Paz.
D'ERASMO, G. 1934. Sopra iilcuiii iivanzi di vcrtcbrati fossili dcllii
Patagonia raccolti dal doll. E. Fcruglio. L'ittiofautia cretacea di
Comeno nel Carso triestino. Atti dellilccadentia di Scienzefisiclte e
naturale di Napoli, 2, 8: 1-136, Napoli.
DOLGOPOL DE SAEL, M. 1941. Noticiris sobrc pcccs f6silcs
argentinos. Siluroidcos tcrciarios del Chubut. Notas del Museo de La
Plata, 6: 451457, La Plata.
ESTES, R. 1983. Sauria terrestria. Amphisbaenia. Ifandbuch der
Palolterpetologie, Part 10A, xxii I- 249 p. G. Fischer Verlag,
Stuttgart/New York.
ESTES, R. & A. M. BAEZ. 1985. Herpetofaunas of North and South
America during the late Cretaceous and Cenozoic: Evidence for
Interchange? In: The Great Anierican Biotic Interchange (F. G.
Stelili & S. D. Webb, eds.), Plenrini Press: 139-196, New York.
ESTES, R. & L. PRICE. 1973. Iguanid lizard froin the upper
Cretaceous of Brazil. Science, 180: 748-751, Washington.
FERNANDEZ, J. 1976. Hallazgo de peces pulmoiiados fsiles
e n l a P u n a Jujeia. Anales d e la Sociedad Cientfica d e
Argentina, 201, S e r i e II,C i e n c i a s a p l i c a d a s , 41: 13-18,
Buciios Aires.
FERNANDEZ, J., P. BONDES10 & R. PASCUAL. 1973. Restos
de Lepidosircn / x i r ( h . w (Ostciclithycs. Dipiioi) dc In Foriiioci6ri
Luiiibrcra (Eogcno?, Eoccno?) dc Jujuy. hicgIii/iit~ti~~,
10: 152-172,
Buenos Aires.
FITCH, J. E. 1975. Vorhizia Frizzell, a living fossil. Abstract.
Program for Amcricaii Socicty of Ichtliyologists niid Hcrpctologists.
Annual Mcding: 96, Aiiii Arbor.
FRANCISCO, B. H. K. & F. L. de S. CUHNA. 1978. Geologia e
estratigrafia de Bacia de So Jos, Municipio de Itabora, RJ. Anais
da Acadniia Brasileira de Cincias, 50: 38 1 4 16.
FRIZZELL, D. L. 1965. Otoliths of a new fish (Vorhizia vulpcs, n.
gen., n. sp., Siluroidei?) from the upper Cretaceous of South Dakota.
Copeia, 2: 178-181, Ann Arbor.
GARDEWEG. M.& C. F. RAMREZ. 1985. Hoja Ro Zapaleri, II,
Regi611 de Antofagasta. Carta Geol6gica de Chile, escala 1: 250,000.
Servicio Nacional de Geologa y Minera, 66: 1-89. Santiago.
GASPARINI, Z. B. de. 1984. New Tertiary Sebecosuchia
(Crocodylia: Mesosuchia) from Argentina. Journal of Vertebrare
Paleontology, 4 (1): 85-95, Norinali.

426

.
I

'

F O S W Y FACIES DE BOUVIA VOL i - VERTEBRADOS

GASPARINI, Z. B. de & E. BUFFETAUT. 1980. Dolichochanrpsa

niininla, n. g., n. sp., a representative of a new family of eusuchian
crocodiles from the lote Cretaceous o f northern Argentina. Neues
Jalirbuclt jir Geologie und Palonfologie. Monatsheft, 5: 257-271,
Stuttgnrt.
GAUDANT, J. 1980. Eiirocharax roitrairii nov. gen.. nov. sp.
(poisson, t616ost6en, Ostariophysi), nouveau Cliaracidae fossile des
"Calcaires h Bythinies" du Var. Geobios. 13 (5): 683-703, Lyon.
GAUDANT, J. 1988. Les Cyprinodontiformes (Poissons
rClCost6ens) oligoc&nesde Rozon, Le Puy-en-Velay (Haute-Loire):
anatomie et signification pal6oobcologique. Geobios, 21(6): 773-782,
Lyon.
GAYET, M. 1981. Considerations relatives h la palkoecologie du
gisement cdnomanien de Laveiras (Portugal). Bulletin du Must?~"n
Nafional d'lfisfoire Nafurelle, 4 h n e serie (3), section C, 4 (1-2): 2141, Paris.
GAYET, M. 198211. Nouvelle extension gkographique et
stratigrapliique du genre Lepidores. Coniptes Rendus Webdoniadaires
des Skances de 1Xcaddmie des Sciences, serie 2, 294: 1387-1390,

GAYET, M. 1988b. Le plus ancien crne de Siluriforine:

Andiniclitliys bolivianetlsis nov. gen., nov. sp. (Andinichthyidae iiov.
fain.) du Maastrichtien de Tiupampa (Bolivie), Conipes Reridirs
Nebdoniadaires des Sances de llcadniie des Sciences, srie 2,
307: 833-836. Paris.
GAYET. M. 1990. Nouveaux Siluriformcs du Maastrichtien de
Tiupampa (Bolivie). Coniptes Rendits Ilebdoniadaires des Sances
de I'Acadt?niiedes Sciences, serie 2, 3 10: 867-872, Paris.
GAYET, M. 1991. "Holostean" and Telostean from Bolivia. In:
Fbsiles y Facies de Bolivia (R. Suiez-Soruco, ed.). Revista Tcnica
de YPFB, 12 (34): 453-494, Santa Cruz.
GAYET, M. & P. BRITO. 1989. Ichthyofaune nouvelle du Cr6tac6
Sup6rieur du groupe Bauru (Etats de S0 Paulo et Minas Gerais,
Bred). Geobios, 22 (6): 841-847, Lyon.
GAYET, M. & F. J. MEUNIER. 1983. Ecailles actuelles et fossiles
d'Ost6oglossiformes (Poissons, TCIostkens). Conililes Rendus
Hebdomadaires des Sances de llcadkniie des Sciences, serie 2:
297: 867-870, Paris.
GAYET, M.& F. J. MEUNIER. 198G. Apport d e l'elude de
l'ornementation microscopique de la ganone dans la d6tennination
Paris.
de l'appartenance g6n6rique et/ou spkcifique des kcailles isolkes.
GAYET, M. 1982b. Cypriniforme cr6tack en Amkrique du Sud.
Conipfes Rendus Ilebdoniadaires des Sbances de l'hadeniie des Coniptes Rendus Ilebdoniadaires des Sances de I'Acaddniie des
Srictrcrs, sfric 2, 303: (13): 1259-1261, Piiris,
Sciatrccs,sfric 2,295: 405407, Piuis.
GAYET, M. 1982c. Dbcouvertc dais le CrBtaa? de Bolivie des plus GAYET. M. & F. J. MEUNIER. 1991a. First discovery o f
anciens Characiformes connus. ConipfesRendus Hebdomadaires des
Polypteriformes (Pisces, Cladistia, Polypterifonnes) out of Africa.
Geobios, 24 (4): 463-466. Lyon.
Sances de llcadniie des Sciences, serie 2,294: 1037-1040, Paris.
GAYET, M. 1985. Contribution il ktude ailatoinique et systCinatique GAYET, M. & F. J. MEUNIER. 1991b. Polypteridae (Pisces,
Cladistia, Polypteriforrnes) from the Cretaceous and Paleocene of
de l'ichthyofauiie cCnoiiiaiiienne du Portugal. III - CoinplCment B
I'tude des Ostbriophysaires. ConiunicaGes dos ServiGos Geolgicos Bolivia In: Fsiles y Facies de Bolivia (R. Surez-Soruco, ed.),
Revista Zcnica de YF'FB, 12 (3-4):495-498, Santa Cruz.
deForfiigal,71 (1): 91-117,Lisboa.
GAYET, M. & F. J. MEUNIER. 1992. Polypteriforin& du
GAYET, M.1986a. Ictiofnuna cretacea y lerciaria de Bolivia. Adas
Maestrichtien et du I'alfocenc de Bolivie. Geobios, Memoire Special
del printer Siniposio de la Invesfigacibn Francesa en Bolivia: 67-74,
14, Lyon
La Paz.
F. J. MEUNIER & V. LEVRAT-CALVIAC. 1988.
GAYET, M. 198Gb. Ranlallichtliys Gayet du CBnonianien infrieur GAYET, M.,
Mise e n evidence des plus anciens Polypteridae connus dans le
inarin d e Ramallah (Judge), une introduction aux relation
gisement d'In Becetem (Niger). Contptes Rendrrs lfebdoniadaires des
phylogn6tiques des Ostariophysi. Mnioires d u Mitskimi National
d7listoire Naturelle, nouvelle skrie, C, Sciences de la Terre, 51: 1- Sances de I'Acadniie des Sciences, sdrie 2 307, 205-210, Paris.
GAYET, M., J. C. RAGE & R. S . RANA. 1984. Nouvelles
81, Paris.
ichthyofaune e t herptofaune de Gitti Khadan, le plus ancien
GAYET, M. 1986c. About Ostariophysan fishes: a reply to S. V.
Fink, I'. H.Greenwood & W . L. Fink's criticism. Llrrllcfin du giscinent connu du Dccciin (Crbtiicd/l'iilgoccii~) i\ inicrovcrt6brfs.
lmpliculionr; piilfogfogi iipliiqiies. Mhnoit c clc Itr SocIciLI Gcologicpc
M i r d i t n i Ntr~iorrtrld'lli.ifoit c Ntitwcllc, 4flllC Rerie, 8 , seclion C (3):
de France, nouvelle srie 147: 55-65, Paris.
393-409, Paris.
GAYET, M., J. C. RAGE, T. SEMPERE & P. Y. GAGNIER. in
GAYET, M. 1987a. Affinitks palCobiogCographiques des
press. Modalitfs des 6chniiges de vertCbrCs continentaux entre
Lcpisosteidae du Cr6tnc6 Brdsilien e t bolivien. Anais do X
Congresso brasileiro de Paleonfologia, 19-25 de Julho 1987, 1: 55- 1'AinCrique du N o r d et l ' A i n 6 r i q u e d u S u d a u CrCtac
Supkrieur 'et au P a l o c h e . Bitllelin de la Socitk Gologique
65, Rio de Janeiro.
GAYET, M. 1987b. Consideraciones preliminares sobre la de France, Paris.
paleobiogeografia d e los Osteoglossomorpha. I V Congreso GAYET, M., T. SEMPERE & H.CAPPETTA. 1992. A propos de
l'environnement marin restreint du bassin centro-andin au
Lafinoatnericano de Paletontologla, Bolivia, Actas 1: 379-398,
Maastrichtien. Conipfes Rendus Nebdoniadaires des Sceances de
Sarita Cruz.
I'Acadniie des Sciences, 2,3 14: 223-228, Paris.
GAYET, M. 1987c. Lower vertebrates from the Early-Middle
GERY. J. 1969. The freshwater fishes of South America. In:
Eocene Kuldana Formation of Kohat (Pakistan): Holostei and
Teleostei. Confribufionsof the Museunt of Paleontology. Universily Biogeography and Ecology in Sottfh Anierica (E. J. Fittkau, J. Illies,
H.Klinge. G . H. Schwabe & H. Sciolo, eds.). W.Junk Publication,
ofMichigan,27 (7): 15 1- 168, Ann Arbor.
the Hague.
GAYET, M. 1 9 8 8 ~ Los
.
peces del Cretcico d e Bolivia y los
GOI, J. C. & K. HOFFSTETTER. 1964. Lexique Sfraligrapltiqae
problemas paleogeogrLficos suscitados. Actas del Segundo
Simposio de la Investigacibn Francesa en Bolivia: 67-74, La Infernafional, V. Ainrique Latine, rasc. 9a, Uruguay (CNRS ed.),
Paris, 202 p.
Paz.
427

GOODY, I'. C. 1968. Tlic skull of Ericltodus fatcjasi froin tlic

Maastricht of Southern Holland. I. Proceedings of Koninklijke
Nederland Akademie van Wetenscltappen, Amsterdant,
Paleontology, srie B, 71 (3): 209-231.
GOODY, P. C. 1969. The relationships of certain upper Cretaceous
teleosts with special reference to the myctopliyoids. Bulletin of the
British Musertni ( N a t u r a l H i s t o r y ) , G e o l o g y , 7: 1 - 2 5 5 ,
London.
GRAMBAST, L., M. MARTINEZ, M. MATTAUER & L.
THALER. 1967. PetWterium altiplanense, nov. gen., nov. sp.,
premier inammifkre msozoque d'Amrique du Sud. Contptes
Rendus Hebdomadaires des Sances de l'Acadmie des Sciences,
264: 707-710, Paris.
GRANDE, L. 1982a. A revision of the fossil genus Diplonlyslus,
with comments on the interrelationships of clupeomorph fishes.
Aniericm MuseuniNovitates, 2728: 1-34, New York.
GRANDE, L. 1982b. A revision of the fossil genus Kniglifia, with a
description of a new genus from the Green River Formation
(Tclcostci: Clupcidnc). American Museiirn Novitates, 273 1: 1-22,
New York.
GRANDE, L. 1984. Paleontology of the Green River Formation,
with a review of the fish fauna. Tlie Geological Survey of Wyoming.
63: 1-333, Lnramic.
GRANDE, L. 1987. Redescription of Ilypsidoris farsonensis
(Teleostei: Siluriformes), with a reassessment of its phylogenetic
relationships. Journal of Vertebrate Paleontology, 7 (1): 24-54,
Norman.
GKANDE, L., J. T. EASTMAN & T. CAVENDER. 1982. Aniyzon
gosiutensis, a new catostomid fish from the Green River Formation.
Copeia: 523-532, New York.
GRANDE, L . & J. H. LUNDBERG. 1988. Revision and
redescription of the genus As/eplius (Siluriforrncs: Ictaluridae) with a
discussion of its phylogenetic relationships. Joirrnal of Verlebrate
Paleontology, 8 (2): 139-17 1, Norman.
GREENWOOD, P. H. 1951. Fish remains from Miocene deposits of
Rusinga Island hiid Kavirondo province, Kenya. Annals arid
Magazine ofNarirra1 Ilistory, series 12,4: 1192-1201, London.
GREENWOOD, P. N. 1973. Fish Fossils from the Late Miocene of
Tunisia. Notes du Service Gologique, 37: 70-72, Tunis.
GREENWOOD, P. H. 1976. A review of the family Centropomidae
(Pisces, I'erciformcs). Bulletin of the British Muserini (Natural
llistory), Zoology, 29 (1): 1-81, London.
HAQ, B. U., J. HAKDENBOL & P. R. VAIL. 1987. Chronology of
fluctuating sea levels since the Triassic. Science, 235: 1156-1 166.
Washington.
HALSTEAD, L. R. 1975. Sokotosuchus ianrvilsoni n. g., II. sp.. a
new tcleosaur crocodile from the upper Creaceous of Nigeria.
Journal of the Nigerian Mineralogical, Geological and
Meteorological Society, 11: 101-103, Ibadan.
HOLMANN, J. A. 1979. Paleontology and geology of the Badwater
Creek area, central Wyoming. Part 17. The late Eocene snakes.
Annals oftlie Carnegie Museiini. 48 (6): 103-110, Pittsburgh.
JAIN, S. L. & A. SAHNI. 1983. Sonic uppcr Cretaceous vertebrates
from Central India and their paleogeographical implications. In:
Cretaceous of India (H. K. Maheswari, ed.). Indian Association of
Palynosrrarigral~hers:66-83, Lucknow.

JAILLARD, E. & T. SEMI'EIIE. 1080. Crctirccous scqucncc

stratigraphy of I'cru and Bolivia. ConrriDtrciones de los Siniposios
sobre Cretdcico de Anierica Latina, Parte A: Evenlos y Registro
Sedimentario: 1-27, Buenos Aires.
JAILLARD. E. & T. SEMPERE. 1991. L a s s e c u e n c i a s
sedimentarins d e I n Formaci611 Miraflores y s u significado
cronoestratigr8fico. Revista Tecnica de Y P F B , 1 2 (2): 257264, Santa Cruz.
JAILLARD, E., H. CAPPETTA, P. ELLENBERGER, M. FEIST, N.
GRAMBAST-FESSARD, J. P. LEFRANC & B. SIGE. in press. Tlie
late Cretaceous Vilquechico Formation of southern Peru:
Sedimentology, Paleontology, Biostratigraphy, Correlations.
Newsletters on Srratigraphy. Berlin/S tuttgart.
KEROURIO, P. & B. SIGE. 1984. L'apport des coquilles d'oeufs de
dinosaures de Laguna Uinayo 1 I'ge de la Formation Vilquechico
(Prou) et 1 l a comprhension d e Perrrfherirrnt altiplanense.
Neivslefterson Strafigraphy, 13: 133-142, BerliiljStuttgart.
KRIZ. S. & C. CHERKONI. 1966. Diagramas correlativos de Ins
formacioncs crcthcicus del Sudocstc de Bolivia. GBOBOL, l l u j a
ItIJornialiva2, La kaz.
LAUBACHER, G. & R. MAROCCO. 1990. La cuenca crethcica
del Altiplano Peruano Litocstratigrnfa e intcgraci6n sccucncinl.
Boleliri de Ia Sociedad Gculdgicci de Peri, 81: 33-46
LEONAKDI, G. 1981. As localidades coni rastros fosscis de
tetrapodes na Amrica Latina. I I Congreso Latinoanwricano de
Paleontologia, Anais 2: 929-940, Porto Alegre.
LEONARDI, G. 1984. Le impronte fossili di dinosauri. In: Sirlle
Ornie dei Dinosairri, Erizzo Publishers: 162-186.
LERICHE G . 1902. Rvision de la faune ichthyologique des terrains
crtacs du Nord de la France. Annales de la Socit Gologique du
Nord, 31: 1-000, Lille.
LERICHE, G. 1906. Contribution B I'tudc des pissons fossiles du
Nord de la France et des rgions voisines. Mmoire de la Socii
Gologique du Nord, 5 (1): 1-423, Lille.
LOFGREN, D. L., C. L. HOTTON & A. C. RUNKEL. 1990.
Reworking of Crctnccous dinosaurs into Palcocciic chnnncl dcposits,
upper Hell Creek Formation, Montnila. Geology, 18: 874-877.
L O H M A " , H. H. 1970. Outline of tectonic history of Bolivian
Andes. Antericari Associarion of Petrolerim Geologists Bulletin, 54:
735-757, Tulsa.
LOHMANN, H. H. & L. BRANISA. 1962. Estrntigrafn y
peleontologa del Grupo Puca en el siiicliiial de Mirallores. Potosi.
Pc/rleo Boliviano, 4: 9- 16, Lii I'az.
LOPEZ, K. 1983. Informc cstr:1tigrdllcode los Kos Alto Moile, Alto
Eterazama, Ichoa, Alto Beni y Tequejc. Inforine Intcrno, YI'FB: 195, Santa Cruz.
LUNDBEKF J. G. 1975. Tlie fossil catfishes of North Anierica.
Papers on Paleonrology 11, C I . W. Hibbnrd Meniorinl, 2: 1-51,
Ann Arbor.
LUNDBEKG, J. G., A. MACHNO-ALLISON & R. F. KAY. 1986.
Miocene characid fishes from Colombia: evolutionary stasis and
extirpation. Science, 2 3 4 208-209, Washington.
MAROCCO, R. T. SEMPERE, M. CHIBIAN, & J. OLLER. 1987.
Mise en Bvidence d'une d6formation palCoc&neen Bolivie du Sud. Sn
place dans I'volution godynaniique des Andes centrales. Conples
Rendus Ilebdoniadairrs des Sances de I'Acadniie des Sciences,
sci& 2. 304: 1139-1142. Paris.

428

- VOL. I - VERTEB~ADOS

FOSILES Y FACIES DE BOW

MARINOVIC, N. & A. LAHSEN. 1984. Hoja Calama, Regi611 de

Antofagasta. Carta Geolgica de Chil&,58: 1-140, Santiago.
MARQUILLAS, R. A. 1985. Estratigrafa, sedimentologia y
paleoambientes de la Formaci6n Yacoraite (CretAcico superior) en el
tramo austral de la cuenca, Norte Argentino. Unpublished Thesis,
Universidad Nacional de Saltc, Salta, Argentina.
MARSHALL, L. G. 1985. Geochronology and land-mammal
biochronology of the transamerican faunal interchange: 49-85. In:
The (;rea American Biotic Interchange (J?. G. Stehli & S. D. Webb,
eds.). Plenum Press, New York.
MARSHALL, L. G.1989a. The K-T boundary in South America: on
which side is Tiupampa? National Geogral~hicResearch, 5 (3): 268210, Washington.
MARSHALL, L. G. 1989b. El primer diente de dinosaurio en
Bolivia. Revista Tcnica de YPFB, 10 (3-4): 129-130, Cochabamba.
MARSHALL, L. G . 1992. Skull Structure of the marsupial
Pucadelphys from the Santa Luca Formation (early Paleocene) of
Bolivia. Bulletin du Musum National d'Histoire Naturelle, Paris, in
press.
MARSHALL, L. G. & C. MOLINA. 1990. Huellas tridlctiles de
dinosaurios en la Formaci6n EI Molino Inferior, cerca a Santivaiiez,
Dpto. de Cochabamba. Revista Tbcnica de YPFB, 11 (2-3): 323-328,
Cochabamba.
MARSHALL, L. G. & C. de MUIZON. 1988. The dawn of the age
of mammals in South America. National Geographic Research, 4
(1): 23-55, Washington.
MARSHALL, L. G. & C. de MUIZON. 1992. Atlas photographique
(MEB) des Metatheria et d e quelques Eutheria du P a l o c h e
infbrieur de la formation Santa Luca B Tiupampa (Bolivie). Bulletin
du Musum National d'Histoire Naturelle, Paris. 4 srie, section C,
14 (1-2): 63-91.
MARSHALL, L. G. & T. SEMPERE. 1991. The Eocene to
Pleistocene vertebrates of Bolivia and their stratigraphic context: a
review. In: F6siles y Facies de Bolivia (R. Sulrez-Soruco, ed.).
Revista Tcnica de YPFB 12 (3-4): 631-652, Santa Cruz.
MARSHALL, L. G. & T. SEMPERE. in press. Evolution of the
Neotropical Cenozoic land mammal fauna in its geochronologic,
stratigraphic and tectonic context. In: Biological Relationsltips
Between Africa dnd South America (P. Goldblatt, ed.), Yale
University Press, New Haven.
MARSHALL, L. G., R. F. BUTLER, R. E. DRAKE & G . H.
CURTIS. 1981. Calibration of the beginning of the age of mammals
in Patagonia. Science, 212: 43-45, Washington.
MARSHALL, L. G., R. HOFFSTE'ITER & R. PASCUAL. 1983.
Geochronology of the mammal-bearing Tertiary of South America.
Palaeovertebrata, Mmoire Extraordinaire, 1983: 1-93, Montpellier.
MARSHALL, L. G., C. de MUIZON & B. SIGE. 1983a. Late
Cretaceous mammals (Marsupialia) from Bolivia. Geobios, 16 (6):
739-745, Lyon.
MARSHALL, L. G., C d e MUIZON & B. SIGE. 1983b.
Perutlierium altiplanense, un notongul du CrdtactS SupQieur du
PBrou. Palaeovertebrata, 13: 145-155, Montpellier.
MARSHALL, L. G., C. de MUIZON, M. GAYET, A. LAVENU &
B. SIGE. 1985. The "Rosetta Stone" for mammalian evolution in
South America. National Geographic Research, 1 ( 2 ) : 274-288,
Washington.
MARSHALL, L. G., J. A. CASE & M. O. WOODBURNE. 1989.
Phylogenetic relationships of the families of marsupials. Current

Mammalogy, 2: 433-502.
MARTIN, M. l 9 8 4 . Rvision des Argonidontidds e t des
NBocratodontids (Dipnoi, Ceratodontiformes) du CrtactS africain.
Neues Jahrbuchfiir Geologie und Palontologie Abhandlungen, 169
(2):225-277, Stuttgart.
MARTINEZ, C. 1980. Structure et &volution d e la chane
hercynienne et de la chane andine dans le nord de la Cordillhe des
Andes de Bolivie. Travaux et Documents de I'ORSTOM, 119: 1-352,
Paris.
MAURY, C. J. 1930. O Cretlceo da Parahyba do Norte, Brasil.
MOnogr@a do Servio Gedgicoe Mineraldgico, 8 1-305.Rio de JoneirO.
MONES, A. 1987. GondwanaUieria, un nuevo orden de mamiferos
sudamericanos (Mammalia: Edentata: ?Xenmthra). Comunicaciones
Paleontolhgicas del Museo de Historia Natural de Montevideo, 18:
237-240.
M O N T m O , M. 1968. Estudio geol6gico de Anzaldo-Irata-Vilque
y proyecto de investigaci611petrogentStica en los miembros basales
d e la secuencia cretlcica. Unpublished Thesis, Universidad
Mayor de San A n d r b , Facultad de Ciencias Geoldgkas, 83
p., La Paz.
MOURIER, T., E. JAILLARD, G. LAUBACHER, C. NOBLET,
A. PARDO, B. SIGE 62 P. TAQUET. 1986. Dcouvertes de restes
dinosauriens et mammaliens d'ge crtStac6 suprieur ?
laibase des
couches rouges du synclinal de Bagua (Andes nord-pruviennes):
aspects stratigraphiques, sdimentologiques et palogographiques
concernant la rBgression fini-crtace. Bidletin de la Sociktll
Geologique de France, srie 8 , 2 (1): 171-175, Paris.
MOURIER, T., P. BENGTSON, M. BONHOMME, E. BUGE,
H. CAPPETTA, J.-Y. CROCHET, M. F. FEIST, K. F. HIRSCH,
E. JAILLARD, G . LAUBACHER, J.-P. LEFRANC,
M. MOULLADE, C. NOBLET, D. PONS, J. REY, B. SIGE,
Y. TAMBAREAU & P. TAQUET. 1988. The upper Cretaceouslower Tertiary marine to continental transition in the Bagua basin,
northern Peru. Paleontology, Biostratigraphy, Radiometry,
Correlations. Newsletters on Stratigraphy, 19 ( 3 ) : 143-177,
BerlinlStuttgart.
MUIZON, C. de & L. G. MARSHALL. 1985. Sur la piste des
premiers mammiRres d'Amdrique du Sud. La Recherche, 16 (167):
812-815, Paris.
MUIZON C. de & L. G . MARSHALL. 1987a. L e plus ancien
Pantodonte (Mammalia), du Crbtac6 Supdrieur de Bolivie. Comptes
Rendus Hebdomadaires des Sances de I'Academie des Sciences,
srie 2,304 (5): 205-208, Paris.
MUIZON C. de & L. G. MARSHALL. 1987b. Le plus ancien
condylarthre (Mammalia) sud-amricain (CrtStac Supbrieur,
Bolivie). Comptes Rendus Hebdomadaires des Sances de
l'Acadmie des Sciences, sdrie 2,304 (13): 771-774, Paris.
MUIZON C. de & L. G. MARSHALL. 1987c. Deux nouveaux
condylarthres (Mammalia) du Maastrichtien de Tiupampa (Bolivie).
Comptes Rendus Hebdomadaires des Sbances de I'Acadkmie des
Sciences, serie 2,304 (15): 947-950, Paris.
MUIZON C. de & L. G.MARSHALL. 1987d. El alba de la edad de
los mamiferos en Sudamerica. N Congreso Latino-americano de
Paleontologfa,Actas 2: 757-791, Santa Cruz.
MUIZON C. de & L. G. MARSHALL. 1988. Tiupampa (Bolivia),
un punto clave en la historia de los mamiferos sudamericanos. Actas
del Segundo Simposio de la Investigacbn Francesa en Bolivia: 5360. La Paz.

429

MIREILLI!GAYET. LARRY O. MARSIIALL &IIIIBRRY SEMPERI!

MUIZON, C. d e & L. G. MARSHALL. 1991. Nouveaux

Condylarthres du Paldockne inf6rieur d e Tiupampa (Bolivie).
Dulletin du "!um
National d'Histoire Naturelle, Paris, 4' serie,
scctiotl C,13: 201-227.
MUIZON, C. de & L. G. MARSHALL. in press. Alcidedorbignya
inopinata (Mammalia, Pantodonta) from the early Paleocene of
Bolivia: phylogenetic and paleobiogeographic implications. Journal
of Paleontology.
MUIZON, C. de, M.GAYET, A. LAVENU, L. G. MARSHALL,
B. SIGE & C. VILLARROEL. 1983. Late Cretaceous vertebrates,
including mammals, from Tiupampa, southcentral Bolivia. Geobios,
16 (6): 747-753,Lyon.
MUIZON C. de, M. GAYET, A. LAVENU, L. G. MARSHALL &
C. VILLARROEL. 1984. Observation to the note by Muizon Ch. de,
Gayet M., Lavenu A., Marshall L. G., Sig6 B. & Villarroel, C.,
entitled "Late Cretaceous vertebrates including inammals from
Tiupampa, southcentral Bolivia". Geobios, 17 (3): 251-252,Lyon.
MUIZON, C. de, L. G. MARSHALL & B. SIGE. 1984. Tlie
mammal fauna from the El Molino Formation (Late CretaceousMnastrichtian) at Tiupampa. southcentral Bolivia. Uirllefin dir
MusCuni Nntional d'llistoire Naturelle, sfric 6, scctioii C, 4: 3 15327, Paris.
MUOZ, N.. R. CHARRIER, & S. PICHOWLAK. 1989. CretQcico
Superior volclnico-sediinentario (Formacidn Quebrada Mnln) en la
regi6n d e Antofagasta, Chile. y su significado geotCctonico.
Contribuciones de los Siniposios sobre Cretdcico de Andrica Lutina.
Parte A Eventos y Registro Sedimentario: 133-148, Buenos Aires.
MYERS, G. S. 1938. Fresh-water fishes and West Indian
zoogeography. Annual Report of Smithsonian Institution, (1937):
339-364,Washington.
NELSON, G. J. 1969. Infraorbital bones and their bearing on the
phylogeny and geography of osteoglossomorph fishes. Anierican
Mirseuni Novitates, 2394: 1-37,New York.
OLLER, J. & T. SEMPERE. 1990. A fluvio-eolian sequence of
probable middle Triassic-Jurassic age in both Andean and
Subandean Bolivia. International Syniposiuni on Andean
Geodynaniics:237-240,Grenoble.
PARODI-BUSTOS, R. 1962. Los anuros cretlcicos de Puente
Morales (Salta) y SUS vinculaciones con Shelartia pascuali
Casarniquela (Chubut) y Eoxenepoides reuningi Haugliton (Africa
del Sur). Revista de la Facacrtltad de Ciencias Naturales, 1 (3): 8 1-85,
Salta.
PASCUAL, R. & M. BOND. 1981. Epidolopinae subfam. nov. de
los Polydolopidae (Marsupialia. Polydolopoidea). I I Congreso
Latino-americano de Paleontologfa, Actas 2: 479-488, Porto
Alcgrc.

PASCUAL, H., M.BOND & M.(3. VUCETICH. 1981. EI subgrupo

Santa Bdrbara (grupo Salta) y sus vertebrados. Cronologia,
paleoambientes y paleobiogeografa. VI11 Congreso Geolgico
Argentino, Actas 3: 743-758,San Luis.
PASCUAL, R. LP. BONDESIO. 1976. Notas sobre vertebrados de
l a frontera Cretficica-Terciaria. III: Ceratodontidae (Peces
Osteichtliyes, Dipnoi) de la Formaci6n Coli-Toro y de otras unidades
del C r e t l c i c o Tardio d e Patagonia y S u r d e Mendoza. Sus
implicancias paleobiogeogrificas. V t Congreso Geoldgico
Argentino, Bahia Blanca, Actas 1: 565-577,Buenos Aires.
PASCUAL. R. & E. ORTIZ-JAUREGUIZAR. 1991. El ciclo

faunfstico Cocliabambiano (Paleoceno temprmio): su incidencia en la

historia biogeogr6fica de los mmnferos sudamericanos. In: Fbsiles y
Facies de Doliida (R. Salrcz-Soruco, ed.). Rcivktn Tdcnica fe Y f F B . .
12 (3-4): 559-574, Sanla Cruz.
PASCUAL, R., M. G. VUCETICH & J. FERNANDEZ. 1979. Los
primeros mamiferos (Notoungulata. Henricosborniidae) de la
Formaci6n Mealla (Grupo Salta, Subgrupo Santa Bdrbara). Sus
implicancias filogenCticas, taxon61nicas y cronol6gica.s.
Amegliiniana, 15 (34): 366-390,Buenos Aires.
PAULA C O U " , C. 1970. Novo Notoungulado no Riochiquense de
Itahraf. IIieringia (Geologia) 3: 7-86, Porto Alegre.
PALMA, J. M.& I. M. BRITO. 1976. Paleontologfa e estratigrafa
da bacia do So Jose de Itabora, Estado de Rio de Janeiro. Anais de
Acadhia Brasileira de Cincias, 46 (34): 383-406,Rio de Janeiro.
PEREZ, M.A. 1987. Dntos paliiiol6gicos del Cretficico Superior de
la scccidn cstrutigrslica de Curatu, Potosf. Bolivia. I V Congreso
Lalinoanrericano de P aleontologia, Actas 2: 739-756, Santa
Cruz.
PERRY, L. 1963. FIorn Forinntion (upper Cretaceous) of northern
Bolivia. '171~Anicricori Associaliori of PeIroleirni Geologists idletin,
47 (10): 1855-1860. Tulsii.
PEYER, B. 1928. Ergebnisse der Forscliungsrcisen Prof. E.
Stromers in den Wsten egyptens. V. Tertiiire Wirbeltiere. 2. Die
wclsc des gyptischcn Altcrliks. Ahlrciridliingcn der Uaycrisckcn
Akadeniie der kVissenscltaflen, Mathenilisclt-nutnrwissensclraliche
Abfeilung, 32 (3), 85 p., Miinchen.
PILSBRY, M. A. 1939. Fresh-water Mollusca and Crustacea from
near EI Molino, Bolivia. Tlie Johns Hopkins University Sludies in
Geology, 13: 69-72, Baltimore.
PINDELL, J. L., S. C. CANDE, W. C. PITMAN, D. B. ROWLEY,
J. F. DEWEY, J. LABRECQUE & W.HAXBY. 1988. A platekinematic framework for models of Caribbean evolution.
Tectortopliysics, 155: 121-138.
PINTO, I.D. t Y. T. SANGUINETTI. 1987. Lower Cretaceous
ostracodes from Bolivia. X Congresso Brasileiro de Paleonlologla,
Actas 2 761-781, Rio de Janeiro.
PORTA, J. de. 1970. Presencia de Pycnodontiformes en el CretAcico
Inferior de Colombia. Geologia Colonibiana, 7: 99-103, Bogotl.
POWELL, J. E. 1979. Sobre una asociacidn de dinosaurios y otras
evidencias de vertebrados del Cretacico Superior de la regi611de la
Cnndclnrin, Prov. (le Salta, Argciitinn. Anieghiriiann, 16 (1-2): 191204. Buenos Aires.
POWELL, J. E. 1987. The late Cretaceous fauria of Los Alainitos,
Patagonia. Argentina. Part VI - Tlie Titanosaurids. Revista del
Museo Argentino de Ciencias Naturales "Bernardino Rivadavia",
f ~ l l ~ ~ ~ J ~3 (3):
l l ~ 147~ / ~153,
) ~ ~f U~c l~l o, SAires.
QUATTHOCCHIO, M. E., H. A. MARQULLAS & W.
VOLKHEIMER. 1986. Palinologa, paleoainbientes y edad de la
Formaci6n Tunal, cuenca del Grupo Salta (Cretdcico-Eoceno),
Repblica Argentina. IV Congreso Argentino de P afeonlolog,izy
Bioes[raligrafia,Actas 3: 95-107,Mendoza.
RAAB, M. 1963. Fossil fish and reptiles from Late Campanian
phosphatic depsits of the Negev region of Israel. Israel Journal of
Earth Sciences, 12(1): 26-40,Tel Aviv.
RAGE, J. C. 1981. Les continents pt5ri-atlantiques du Crdtac6
SupBrieur: migrations des faunes continentales et problkines
palhg6ographiques. Cretaceous Research, 2 (1): 65-84,London.

430

FOSILES Y FACIES DE BOLIVIA - VOL,I - VERTEBRADOS

RAGE, J. C. 1984. Serpentes. Nandbuch der Palolterpefologie, part

11, xii t 80 p. G. Fischer Verlag, Stuttgarmew York.
RAGE, J. C. 1986. Le plus ancien Amphibien apode (Gymnophiona)
fossile, Remarques sur l a rpartition et l'histoire
pal6obiogograhique des Gymnophiones. Conipfes Rendus
Iiebdoniadaires des Sances de l'kadntie des Sciences, s6rie 2,
302 1033-1036, Paris.
RAGE, J. C. 1987. Fossil History. In: Snakes, Ecology and
Evolutionary Biology (R. A., Seigel, J. T. Colins, & S. S. Novak,
eds.): 5 1-76, MacMillan, New Yormoronto/London.
RAGE J. C. 1991a. Gymnophionan Amphibia froin the early
Paleocene (Santa Luca Formation) of Tiupampa (Bolivia). The
oldest known Gymnophiona. In: Fsiles y Facies de Bolivia (R.
Surez-Soruco, ed.). Revista Tcnica de YPFB, 12 (3-4): 499-501,
Santa Cruz.
RAGE, J. C. 1991b. Squamate Reptiles from the early Paleocene of
the Tiupampa area (Santa Luca Formation), Bolivia. III: Fsiles y
Facies de Bolivia (R. Surez, ed.). Revista Tcnica de YPFB, 12
(3-4):503-508, Santa Cruz.
REYES, F. C. 1972. Correlaciones en el Cretcico de la Cuenca
Andina de Bolivia, Per y Chile. Revista Tcnica de YPFB, 1 (2-3):
101-144, La Paz.
REBOUAS, J. C. & R. d a SILVA SANTOS. 1956. Fatma
ictiolgica do fosfato de Pemambouco, Brasil. Divis0 de Geologia
e de Mineralogia, 162: 1-29, Rio de Janeiro.
ROBERTS, T. R. 1975. Characoid fish teeth from Miocene deposits
in the Cuenca Basin, Ecuador. Journal of Zoology, 175: 259-271,
London.
ROXO, M. G. & A. LOFGREN. 1936. Lepidofes piaithyensis, sp.
nov. Servicio Geolgico e Mineralgico, Notas Preliniinares e
Estudos, 1: 7-12, Rio de Janeiro.
RUSSO, A. & L. A. RODRIGO. 1965. Estratigrafa y
paleogeografa del Grupo Puca en Bolivia. Boletin del Insfifuto
Boliviano delPefrleo, 5: 5-51, La Paz.
SAHNI, A. & V. P. MISRA. 1975. Lower tertiary vertebrates from
wcstcrn India. Monograplis of the Paleontological Socicfy ($I~ldia,
3: 1-48.
SALFITY, J.,R. MARQUILLAS, M. GARDEWEG, C. RAMIKEZ
& J. DAVIDSON. 1985. Correlaciones en el Cretcico Superior del
norte de Argentina y Chile. N Congreso Geolgico Chileno, Actas
1: 654-667,Antofagasta.
SALINAS, P., L. G. MARSHALL & P. SEPULVEDA. 1991a.
Vertebrados continentales del Paleozoico y Mesozoico de Chile. VI
Congreso Geolgico Chileno, Via del Mar, Actas 1: 310-313,
Santiago.
SALINAS, P., P. SEPULVEDA & L. G. MARSHALL. 1991b.
Hallazgo de restos Bseos de dinosaurios (saurpodos) en la
Formacin Pajonales (Cretcico Superior), Sierra de Almeyda, II
Regin d e Antofagasta, Chile: implicancia cronolgica. VI
Congreso Geolgico Chileno, Via del Mar, Actas 1: 534537, Santiago.
SANDERS, M. 1934. Die fossilen Fische der Alttertiren
S ss w ass er ablag er u n gen aus Mit te 1-S u m a tr a. Verk ande 1
Geologische Mijnbouwkundig Genootschap voor Nederland en
Kolonin, Geologische Serie, 11: 1-144,Amsterdam.
SANJINES-SAUCEDO, G. 1982. Estratigrafa del Carbnico,
Trilsico y Cr-:<rico boliviano en el borde oriental de las Sierras

subandinas centrales. V Congreso Lafinoaniericano de Geologa,

Actas'l: 301-318, Buenos Aires.
SANTOS da SILVA, R. 1963. Peixes do Cretcico do Rio Grande
do Norte. Anais da Acadniia Brasileira de Cincias, 33 (3-4): 32,
Rio de Janeiro.
SANTOS da SILVA, R. 1984. Lepisosfeus continufoi n: sp. da
Formab Bauru, Estado de SGo Paulo, Brazil. Anais da Acadniia
Brasileira de Cincias, 56 (2): 197-202, Rio de Janeiro.
SANTOS da SILVA, R. 1985. Laeliickfhysancestralis, novo gnero
e' espcie de Osteoglossiformes do Aptiano da Formao Areado,
Estado de Minas Gerais, Brasil. MME-DNPM, Geologa 27,
Paleonfologfae Esfratigrafla, 2: 161-167, Rio de Janeiro.
SANTOS da SILVA, R . 1987. Lepidosiren megalos n. sp. do
Tercilrio do Estado do Acre, Brasil (Paleoichtiologia: Dipnoi,
Lepidosirenidae: Terciario). Anais da Acadniia Brasileira de
Cincias, 59 (4): 375-384, Rio de Janeiro.
SCHAAL, S. 1984. Oberkretazische Osteichthyes (Knochenfische)
I
aus dein Bereich von Bahariya und Kharga, gypten, und ihre
Aussagen zur Palkologie und Stratigraphie. Berliner
Geowissenschaftilclte Abhandlrtngen, Reihe A/Band 53: 1-79,
Berlin.
SCHAEFFER, B. 1947. Cretaceous and Tertiary actinopterygian
fishes from Brazil. Billlefin of the American Museuni of Natural
History, 89 (1): 1-40,New York.
SCHAEFFER, B. 1963. Cretaceous fishes from Bolivia, with
comments on pristid evolution. Anierican Mciserrni Novilates, 2159:
1-20, New York.
1
SCHULTZE, H. P. 1981..A pycnodont dentition (Paramicrodon
volcanensis n. sp., Pisces, Actinopterygii) from the lower Cretaceous
of El Volcan region, southeast of Santiago, Chile. Revista Geolgica
de Chile, 12: 87-93, Santiago.
SCHULTZE, H.P. 1991a. Lungfish from the i Molino (Late
Cretaceous) and Santa Luca (early Paleocene) Formations in
southcentral Bolivia. In: Fbsiles y Facies de Bolivia (R. SurezSoruco, ed.). Revista Tcnica de YPFB.12 (3-4): 441-448, Santa
Cruz.
SCHULTZE, H. P. 1991b. Pycnodont fish (Actinopterygii,
Osteichthyes) from the El Molino Formation, late Cretaceous of
Bolivia. In: Fbsiles y Facies de Bolivia (R. SuLrez-Soruco, ed.).
Revista Tcnica de YPFB, 12 (3-4): 449-451, Santa Cruz.
SCILLATO-YANE, G. J. & R. PASCUAL. 1985. Un peculiar
Xenarthra del Paleoceno medio de Patagonia (Argentina). Su
importancia en la sistemtica de los Paratheria. Anieghiniana, 21:
173-176, Buenos Aires.
SEMPERE, T. 1990. Cuadros estratigrficos fanerozoicos de
Bolivia: propuestas nuevas. Revista Tcnica de YPFB, 11 (2-3): 215227, Cochabamba.
SEMPERE, T. in press. Kimmeridgian? to Paleocene tectonic
Andes. (J. A,
evolution of Bolivia. In: Cretaceous Tecfonics of
Salfity, ed.). Earth Evolution Sciences, Monograph Series, Vieweg
Publication, Wiesbaden.
S E M I " , T. & J.OLLER. 1987. Cuadro estratigrfico del Mesozoico
boliviano. Dociintenfo Interno, YPFB-Orsfom,Santa Cruz.
SEMPERE, T. & L. G. MARSHALL. In press. Location of the
Tiupampa vertebrates in the late Cretaceous-Paleocene stratigraphy
of Bolivia: resolution of a paleontological debate. Mewslefferson
Sfraigraplty,Berlin/S tuttgart.

43 1

MIREILLE GAYET, LARRY G. MARSIIAU & l l l I B R R Y SIJMI'IXE

SEMPERE, T., J. OLLER, C. CHERRONI, O. ARANIBAR. L.

BARRIOS. L. BRANISA, M. CIRBIAN & M. PEREZ. 1987. Un
ejemplo de cuenca cnrbonatada en un contexto distensiv.0 de
ichoii~co:I)iilcogcotliiidiiiicii del ChctBcico tcriiiiiiiil en III lteptblicii
de Bolivia (Formaci6n El Molino y equivalentes). X Congreso de
Geologfa Aigeritina, IGCP 242, Abstract: 18-19 (and unpublislied
manuscript), Tucum A n .
SEMI'ERE, T., J. OLLER & L. BARRIOS. 1988. Evoluci611
tectosedimentaria de Bolivia durante el Crethcico. V Congreso
Geolbgico Chileno, Actas 3: H37-H65, Santiago.
SEMPERE, T., G. HERAIL, J. OLLER & P. BABY. 1989.
G e o l o g i c s t r u c t u r e a n d t e c t o n i c history of the Bolivian
orocline. 28th International Geological Congress, 3: 72-73,
Washington.
SEMPERE, T., J. OLLER, E. AGUILERA, J. LOBO, D.
MERINO, R. MAROCCO & R. GARCIA. 1990. Elementos para
una estrotigraffa secuencia1 del Paleozoico Superior de Bolivia. IX
Congreso Geolbgico de Bolivia: 41-42, Cochabamba.
SEMPERE, T., E. AGUILERA., J. DOUBINGER, P. JANVIER, J.
LORO, J. OLLER & S. WENZ. 1992. La Formation de Vitiacun
(Permian Su$rieur-Trias 7InfCrieur. Bolivie du Sud): stratigraphic,
palynologie et pal6ontologie. Neires Jalirbrtcli fiir Geologie und
P aldontologie, Abhandlungen, 185: in press.
SIG& n. 1968. Dents de itiicri)iiiiii:ictiil&rcs ct friigincnts de
coquillcs d'oeufs de dinosilures dans III fiiune de VertfbrCs du
Cr6tad SupCrieur de Laguna Umayo (Andes pCruvicnncs). Coniptes
Rendus Hebdomadaires des Sances de I'Acadntie des Sciences,
serie D, 267: 1495-1498, Paris.
SIGB, B. 1971. Les Didelphoidea de Laguna Umayo (Formation
Vilquechico, Crtac6 Suprieur, PBrou), et le peuplement marsupial
&AmCrique du Sud. Coniptes Rendids Iiebdoniadnires des Sances
de l'Acadmie des Sciences, serie 2,273: 2479-2481, Paris.
SIG?,
B. 1972. La faunule de mammiRres du Cr61ac6 SupCrieur de
Laguna Umayo (Andes p6ruviennes). Bulletin du Musum Naional
d'lfistoire Naturelle de Paris, 99, Sciences de la Terre, 19: 375-409,
Paris.
SORBINI, L. 1970. Un nuovo genero fossil dell'ittiofauna di M.
Bolca: Eolates nov. gen. Meniorie del Museo Civico di Storia
Naturale di Verona, 18: 11-29, Verona.
SORIA, M. F. 1987. Estudios sobre los Astrapotlieria (Mammalia)
del Paleoceno y Eoceno. Parte I: Descripci6n de Eoastrapostylops
riolorertse Soria y Powell, 1982. Anlcghiniana, 24: 21-34, Buc~ios
Aires.
STEINMANN, G. 1906. Die Enstellung der Kupfererzlagcrstiittevon
Corocoro und Verwanter Vorkommnisse in Bolivia. Ferslirift 70,
Gebruder Il. Rosenbuslt: 335-368, Stuilgart.
STEINMANN, G. 1923. Umfang, Bezicliung und Besonderlicitcn
der andinen geosynklinale. Geologische Rundschau, 14 (1): 69-82.
STEPHAN, J. F., B. DE LEPINAY MERCIER, E. CALAIS, M.
TARDY, C. BECK, J. CH. CARFANTAN, J. L OLIVET, J.-M.
VILA, Pli. BOUYSSE, A. MAUFFRET, J. BOURGOIS, J. M.
W R Y , J.TOURNON, R. BLANCHET & J. DERCOURT. 1990.
Pnleogeodynamic maps o f tlie Caribbean: 14 steps from Lias to
Present. Bulletin de la SocitkGologiqiie de France (8), 6 (6): 915919, Paris.
SULLIVAN, R. M. 1982. Fossil lizards from Swain Quarry "Fort
Union Formation", Middle Paleocene (Torrejonian). Carbon
County, Wyoming. Journal of Paleontology. 56: 996-1010, Tulsa.

SU'ITKUS. R. D. 1963. Order Lepisostei. In: Fishes ojtlie western

North Atlantic: 61 -68. Memoir of Sears Foundation foi Marine
Rescorch 1, par1 3. New HIVCI~.
'I'AIIAS'IE, N. 1903. Etude de restes d e poissons d u Crdtiicd
saliarien. Mnioire IFAN 68, M6lange.s iclithyologiques: 438-485.
Dakar.
TAQUET, 1'. 1987. Ln 1'iilcontologi:i dei Vcrtcbriili. In: Dolivia
Lungo I Sentieri Incantati, (Erizzo, ed.), Explorazione c Richerclte,
XI: 3748, Roma/Venezia.
TAVERNE, L. 1975. Etablissement d'un genre nouveau Pliareoides
pour Phareodics queenslandictrs Hills, E. S., 1934 (Pisces,
Osteoglossiformes) du Tertiaire d'Australie. Bulletin de la Socit
Belge de Gologie, de Palontologie et d'Hydrologie, 82 (4), 1963:
497-499. Bruxelles.
TAVERNE, L. 1976. Les Tlostens fossiles du Crktac6 Moyen de
Kipala (Kwango, Zaire). Annales du Muse Royal d'Afrique
Ceiah-ale,Sciences Gi!ologiqries,79: i-ix + 50 p.
TAVERNE, L. 1978. Ost4ologie, pliylogCnEse et syst6matique des
tklCost6cns fossiles et actuels du super-ordre des
Ostfoglossoniorplics. Dcuxi&nic partie. Acaddniie Noycile de
Bclgigrte, Mfinoirc de la Clinse des Sciences. sfric 2,42 (6): 1-213,
Bruxelles.
TAVERNE, L. 1979. Ost6ologie, phylogni?se et systmatique des
t616ostfcns I'ossilcs et iictucls du super-ordre dcx
Os t Co gIossoi n or pli c s. Tro isi &nic p ii r t i e. A caddni i e Hoy a 1e de
Belgique. MCinoire de In Classe des Sciences, sfric 2, 43 (3): 1-168,
Bruxelles.
TRAQUA!R, H. 1910. Les poissons wealdiens de Rernissart.
Imprimerie Polleunis & Centrick. Bruxelles: 1-65.
UYENO, T. 1979. Early Cretaceous fxeshwater fishes from
northern Kyushu, J a p i . I. Description of two new species of lhe
clupeid genus Dildoniystus. Bulletin of the Kitakyrishrr Mitserini of
Natural Ilistory, 1:ll-24.
VAN VALEN, L. M. 1988. Paleocene dinosaurs or Cretaceous
ungulates in South Anlerich. Evolutionary Monographs, 10: 1-79,
Chicago.
VARGAS, J. M. 1988. Potencial petrolfero de la cuenca Huallaga,
Oriente peruano. I I I Siniposio Bolivariano sobre Gxploraci6n
P etrolera en las Cuencas Subandinas: 196-225, Caracas.
VERNET, R. & R. BOTELLO. 1975. Area de Madidi. Sintesis
gcol6gica. Unpublislied report, Total-olivie.
VOLwEIMEK, W., M.E. QUATTROCCHIO & J. A. SALFITY.
1984. Datos palinol6gicos de la Formaci611 Maz Gordo, Terciario
inferior de la Cuenca de Salta. IX Congreso Geolbgico Argentino,
Actas 4: 523-538, San Carlos de Bariloclie.
WEILER, W. 1935. Ergcbnissc der Forschungsreisen Prof.
E.Stromers hlden Wilstcn gyptens. I. Wirbeltierreste der BaharijeStufe (interstes Cenoman). 16. Neue Untersucliungen an den
Fischresten. Abhandlungen der Bayerischen Akademie der
Wissertsclraften,Matlreniatiscli-nafunvissenscliaftlicheAbteilung, II.
f. 1: 1-57. Mnchen.
WENZ, S. 1969. Note sur quelques Poissons ActiiioptCrygiens du
Cr6tacB SupCrieur de Bolivie. Bulletin de la Socidtk Gologique de
France, skrie 7, 11: 434-438, Paris.
WENZ, S. 1989. lenranja palnia n.g., n.sp., Gyrodontidae nouveau
(Pisces, Actinopterygii) du CrBtac6 Inf6rieur de la Chapada do
Araripe (N-E du Brisil), Coniptes Rendus lfebdoniadnires des
Sdances de l'haddinie des Sciences, sfrie II, 308: 975-980. Piiris.

432

FOSILES Y FACES DE BOI,IVU\ - VOL. I VERTEBRADOS

WHITE, E. I. 1934. Fossil fishes of Sokoto Province. Geological

Siirvcy cfNigcria, 14: 1-78, Lagos.
WILEY, E. O. 1976. The phylogcny and biogeography of fossil and
Recent gars (Aclinopterygii: Lepisosteidae). University of Kansas,
Mitseiini of Natitral History, Miscellaneous Publications, 64: 1-111,
Lawrence.
WOODWARD, A. S. 1895. Catalogue of the fossil fishes in the
British Museum (Natural History). British Museum (Natirral
lfistory)Part 3: 1-544,London.

WOODWARD, A. S. 1901. Catalogue of the fossil fishes in the

nritish Museuin (Natural History). British Mitscrrr?i (Nnftrral
lfistory), Part 4: 1-213, London.
WOODWARD, A. S. 1908. Cretaceous fishes from r a z i l .
Quarterly Journal of the Geological Society, 6 4 : 35'1-362,
London.

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