Geobios 37 (2004) 771–794

http://france.elsevier.com/direct/GEOBIO/

Original article

Age and stratigraphic reassessment of the fossil-bearing Laguna Umayo

red mudstone unit, SE Peru, from regional stratigraphy, fossil record,
and paleomagnetism
Âge et stratigraphie des pélites rouges fossilifères de Laguna Umayo,
SE Pérou, reconsidérés d’après la stratigraphie régionale,
le registre fossile et le paléomagnétisme
Bernard Sigé a,*, Thierry Sempere b, Robert F. Butler c, Larry G. Marshall d, Jean-Yves Crochet e
a
Paléoenvironnements et Paléobiosphère (UMR 5125 du CNRS), Université Claude Bernard Lyon-1,
43, boulevard du 11 novembre, 69622 Villeurbanne cedex, France
b
Laboratoire Mécanismes de Transfert en Géologie (LMTG), institut des Sciences de la Terre,
14, avenue Edouard-Belin, 31400 Toulouse France
c
Department of Geosciences, The University of Arizona, Tucson, AZ, USA
d
Department of Paleontology, Mesa Southwest Museum, Mesa, AZ, USA
e
Institut des Sciences de l’Évolution (UMR 5554 du CNRS), Université Montpellier-2, Place Eugène Bataillon, Montpellier, France
Received 4 October 2002; accepted 16 June 2003
Available online 06 November 2004

Abstract

The thick red mudstone unit that crops out at Laguna Umayo (Puno department, southern Peru), here referred as LURMU, has yielded in
different levels a fossil assemblage with plants and vertebrates (including mammals). On the basis of charophytes, the unit was initially
assigned to the Vilquechico Formation (Maastrichtian-Danian), of regional extension, and the dinosaurian structure of egg fragments was
interpreted as consistent with that age. Revision of the regional stratigraphy leads to reassignment of this unit to the Lower Muñani Formation
(Early Tertiary). Mammals from the LU-3 and Chulpas levels present affinities with forms from the Upper Paleocene of South America
(Patagonia, Brazil). A bunodont marsupial, Chulpasia, is evidence for chronologic proximity to a transantarctic interchange with Australia at
the end of the Paleocene. Furthermore, magnetostratigraphy of the LURMU reveals a single reverse polarity zone of 300 m thickness. Because
of the new stratigraphic and paleomammalogic data, this long reverse polarity zone is likely correlative to Chron 26r (early Late Paleocene) or
Chron 24r (latest Paleocene–earliest Eocene), or, less likely, to Chron 29r (latest Cretaceous–earliest Paleocene). The arguments previously
invoked in favor of a Cretaceous age (charophytes, dinosaurian eggs) are critically evaluated, and correlation to Chron 24r is favored.
© 2004 Elsevier SAS. All rights reserved.

Résumé

L’épaisse unité de pélites rouges de Laguna Umayo (département de Puno, Sud Pérou), désignée ici par l’acronyme LURMU, qui a livré en
différents niveaux des assemblages fossiles à végétaux et vertébrés (dont mammifères), a été d’abord attribuée à la Formation régionale
Vilquechico (Maastrichtien-Danien) sur la base de charophytes. La structure dinosaurienne de fragments d’œufs fut aussi produite à l’appui de
cette acception d’âge. La révision de la stratigraphie régionale conduit à réattribuer ces terrains à la Formation Muñani inférieure (Tertiaire
inférieur). Les mammifères décrits des niveaux LU-3 et Chulpas ont des affinités avec des formes connues du Paléocène supérieur d’Amérique
du Sud (Patagonie, Brésil). Un marsupial bunodonte, Chulpasia, témoigne de la proximité chronologique d’un échange transantarctique,
fini-Paléocène, avec l’Australie. D’autre part, l’étude du paléomagnétisme de la LURMU montre une polarité inverse continue. Vu les

* Corresponding author. Paléoenvironnements et Paléobiosphère (UMR 5125 du CNRS), Université Claude Bernard Lyon-1, 43, boulevard du 11 Novem-
bre, 69622 Villeurbanne cedex 02, France.
E-mail address: bernard.sige@univ-lyon1.fr (B. Sigé).

0016-6995/$ - see front matter © 2004 Elsevier SAS. All rights reserved.
doi:10.1016/j.geobios.2003.06.006
772 B. Sigé et al. / Geobios 37 (2004) 771–794

nouvelles données stratigraphiques et paléomammalogiques, cette longue section magnétostratigraphique inverse correspond vraisemblable-
ment aux chrons 26r (partie inférieure du Paléocène supérieur) ou 24r (Paléocène terminal—Eocène basal), ou, moins favorablement, au
Chron 29r (Crétacé terminal—Paléocène basal). Les arguments précédemment allégués pour l’âge crétacé (charophytes et œufs dinosauriens)
sont envisagés de façon critique.
© 2004 Elsevier SAS. All rights reserved.

Resumen

La espesa unidad de pelitas rojas de Laguna Umayo (departamento de Puno, Sur del Perú), aquí referida como LURMU, ha proporcionado
en distintos niveles asociaciones fósiles con vegetales y vertebrados (incluyendo mamíferos). En base a carófitas, fue inicialmente atribuida a
la Formación Vilquechico (Maastrichtiano-Daniano), de extensión regional, y se ha invocado la estructura dinosauriana de fragmentos de
huevos para sustentar esta edad. La revisión de la estratigrafía regional conduce a reatribuir esta unidad estratigráfica a la Formación Muñani
inferior (Terciario inferior). Los mamíferos descritos en los niveles LU-3 y Chulpas presentan afinidades con formas conocidas del Paleoceno
superior de Sudamérica (Patagonia, Brasil). Un marsupial bunodonto, Chulpasia, atestigua la proximidad cronológica de un intercambio
transantárctico con Australia a fines del Paleoceno. Por otra parte, la magnetoestratigrafía de la LURMU indica una polaridad inversa continua.
Dado los nuevos datos estratigráficos y paleomamalógicos, esta larga sección magnetoestratigráfica inversa corresponde más probablemente
a los crones 26r (parte inferior del Paleoceno superior) o 24r (Paleoceno terminal - Eoceno basal), o, menos favorablemente, al Cron 29r
(Cretácico terminal—Paleoceno basal). Los argumentos anteriormente invocados en favor de una edad cretácica (carófitas, huevos dinosau-
rianos) se evaluan de manera crítica.
© 2004 Elsevier SAS. All rights reserved.

Keywords: Laguna Umayo; Peru; Fossils; Stratigraphy; Paleomagnetism; ?Early Tertiary

Mots clés : Laguna Umayo ; Pérou ; Fossiles ; Stratigraphie ; Paléomagnétisme ; ?Tertiaire ancien

Palabras claves: Laguna Umayo; Perú; Fossiles; Estratigrafía; Magnetoestratigrafía; ?Terciario inferior

1. Introduction vided apparently secure grounds for a Late Cretaceous age of

The brownish red fossiliferous strata that crop out on the
northwestern shore of Laguna Umayo, southeastern Peru,
were first reported by Grambast et al. (1967). On the basis of
their charophyte content, these authors assigned the strata to
the Vilquechico Formation (Newell, 1949), which was, at
that time, thought to be of Late Cretaceous age. Because it
has yielded vertebrate remains, including mammals, Laguna
Umayo is among the few localities in South America that
document continental biotic aspects of the critical period
encompassing the Cretaceous/Tertiary (K/T) boundary.
On the basis of the charophytes, the Laguna Umayo local
fauna (LU-3) was first regarded as Late Cretaceous, most
probably Maastrichtian (Grambast et al., 1967; Sigé, 1968,
1972; Clemens et al., 1979). In the late 1970s, for reasons of
suspected fossil age discrepancy, the Late Cretaceous age
was disputed, and some authors preferred to regard the La-
guna Umayo fossil assemblage as Early Tertiary, most prob-
ably Paleocene. For such authors, for example Van Valen
(1978, 1988) or Kielan-Jaworowska et al., (1979), the single
argument for a Tertiary age always remained the allegedly
advanced evolutionary stage of a single mammal species,
Perutherium altiplanense.
A later synthesis of the latest Cretaceous stratigraphy in
southeastern Peru (Jaillard et al., 1993) involved refinement
in sedimentologic and paleontologic data concerning the
Vilquechico Formation in its classical outcrops 75 km NNE
of Laguna Umayo. Jaillard et al. (1993) elevated this unit to
the rank of Group and tentatively correlated it with other
units in the Andean basin. This multidisciplinary study pro-
the majority of the Upper Vilquechico Formation (UVF). The
Laguna Umayo red mudstones of the nearby Puno area
(Fig. 1) were tentatively correlated with the “third lower-
scale sequence” of the UVF on the basis of a shared charo-
phyte assemblage.
Another type of fossil from level LU-3 that was used to
support a Late Cretaceous age is eggshell material of alleged
dinosaurian structure. This material was recently re-
evaluated by Vianey-Liaud et al. (1997), whose conclusions
are discussed below.
In contrast with the charophytes and alleged dinosaurian
eggshells, other fossils suggest an Early Tertiary age for the
Laguna Umayo assemblage. These taxa include several euth-
erian and metatherian mammals with rather derived tooth
patterns, among other vertebrates of less precise biochrono-
logic significance. They come either from the LU-3 level or
the Chulpas level (sensu Sigé, 1972; Crochet and Sigé, 1993,
respectively), the later being ~200 m stratigraphically higher
than the former within the Laguna Umayo red mudstone
section (Fig. 2).
New stratigraphic information is presented below and
considered with regard to biostratigraphic data. Formal re-
strictions and opposite interpretations have been stated. Nev-
ertheless, based on the present state of knowledge of the
Laguna Umayo and more general fossil record, none of the
reported data could be eluded, as they were in most previous
statements. As we comment below, if the Laguna Umayo red
mudstone section appears to be more probably Early Tertiary
in age, this unavoidably calls into question the charophyte
and eggshell data.
 B. Sigé et al. / Geobios 37 (2004) 771–794
Fig. 1. Geographic location of Laguna Umayo locality at different scales.
The small scale sketch shows the following structural units: 1, accreted
terranes; 2, coastal and western domain; 3, axial zone; 4, eastern domain.
The large scale map locates the section A–B of the fossil-bearing red
mudstone unit. Samples for paleomagnetic analysis were collected along the
new road cut in this section.
Localisation des affleurements de Laguna Umayo à différentes échelles.
L’esquisse à petite échelle montre les unités structurales suivantes : 1, unités
d’accrétion ; 2, domaine côtier et occidental ; 3, domaine axial ; 4, domaine
oriental. La carte à grande échelle situe la coupe A–B de l’unité des pélites
rouges fossilifères. L’échantillonnage paléomagnétique a été réalisé le long
de la coupe exposée par la tranchée de la nouvelle route.
Given the current paleontological debate, constraining
chronostratigraphic data from the Laguna Umayo fossil
record are urgently needed. Until recently, the stratigraphic
context appeared unfavorable in different ways: the Laguna
Umayo red mudstones outcrops were thought to be isolated
and of limited extent; the underlying and overlying beds lack
fossils; no reliable radioisotopic data could be obtained from
overlying volcaniclastic beds; and until now no magneto-
stratigraphic study had been undertaken.
We use here a recently updated regional stratigraphy
(Sempere et al., 2000a), and report the first magnetostrati-
graphic data from the Laguna Umayo red mudstones. These
new data shed light on the fossil calibration problem and
constrain the possible correlations to rather precise time
intervals, mostly Early Tertiary in age.
This paper is part of an ongoing chronostratigraphic pro-
gram to calibrate Late Cretaceous and Paleogene units in the
southern part of the Andean basin based on compiled and
new knowledge of regional stratigraphy, magnetostratigra-
773
phy, radioisotopic dating and biostratigraphy (see Marshall et
al., 1997; Sempere et al., 1997). We use the chronostrati-
graphic chart of Hardenbol et al. (1998). As agreed by the
International Union of Geological Science, the age of the
Paleocene–Eocene boundary is 54.9 Ma, and refers to Norris
and Röhl (1999) and Röhl et al. (2000).
2. Geological setting and descriptive lithology
The Laguna Umayo red mudstone unit (LURMU as acro-
nym) crops out in the southeastern Peruvian Andes, ~20 km
NW of Puno, west of Lake Titicaca (Fig. 1). This formation
contributes locally to the steep slope, which dominates the
restricted and almost isolated NE portion of Lake Umayo
(15.75° S; 289.85° E). The stratigraphic sequence forms a
monoclinal succession with an average dip of 30° to the SSE
(Fig. 2). This succession is unconformably overlain by thick,
subhorizontal, dark grey shoshonite flows, which form char-
acteristic plateaus (“mesas”) in the local landscape. At lower
elevations, the older strata are covered by bottom loam of the
Lake Titicaca system, which produces the typical subdued
Altiplano topography. These younger deposits, including
modern slope scree and breccias, restrict the LURMU out-
crop area.
The lowest strata of the monoclinal succession consist of
several hundred meters of hard sandstone beds with no sig-
nificant mudstone intercalations. These beds are exposed
along the road from Atuncolla to the Sillustani Peninsula
(Sigé, 1972: 378, Fig. 2), which protrudes westward into the
northern part of Lake Umayo. On this peninsula is located the
eponymous archeological site with preincaic funeral towers;
the same spelling having been misleadingly used for the
fossil site in the original report of Grambast et al. (1967).
The basal sandstones grade upward into soft red mud-
stones that become dominant and reach a thickness of
~500 m. Some hard sandstone levels of meter thickness
intercalate in the lower part of the red mudstone unit. Also
intercalated are thin, 10–50 cm-thick, red cross-bedded mi-
croconglomerate levels which consist of rounded carbonate
grains of millimeter size range within a variably calcified
clay matrix. Some bedding surfaces display desiccation
cracks. According to R. Simon-Coinçon’s thin section and
X-ray diffraction study, this dominantly sandy and muddy
sediment contains abundant calcite and iron oxide concre-
tions as irregular nodules, various included clasts, bioturba-
tion and numerous reworked charophyte gyrogonites, reveal-
ing both reworked materials and in situ pedogenic evolution.
These low energy sediments were deposited in channelized
areas having long drought episodes.
Such thin lenticular channelized levels repeatedly occur in
the lower half of the red mudstone unit. Separated from these,
the highest exposed channel level occurs ~100 m below the
upper limit of the red mudstone unit. These recurrent micro-
conglomeratic facies are the fossiliferous levels within the
red mudstone unit. With variable abundance, but in no place
774 B. Sigé et al. / Geobios 37 (2004) 771–794

Fig. 2. View of the Laguna Umayo red mudstone unit section (LURMU), drawn from a photograph taken from the eastern side of the Sillustani peninsula: 1,
underlying thick hard sandstones; 2, transitional zone with alternating sandstones and mudstones; 3, red mudstone unit containing various detrital intercalations
among which the fossil-bearing levels occur, the richest of them are shown by arrows (see Fig. 7 for detail); the upper two-thirds of the slope is scree-covered;
4, coarse-grained, sandstone-dominated unit, including conglomerates with volcanic clasts; 5, Umayo high-K basalts (shoshonites; latest Miocene–earliest
Pliocene); 6, remote landscape; 7, bottom lake loam deposit; A–B locates the section shown in Fig. 1; P, parking place of the Sillustani precolombian site; slight
or doted lines correspond to tracks, terrace remains or corral walls.
Vue d’ensemble de l’affleurement des pélites rouges de Laguna Umayo (LURMU), reprise d’une photographie prise de la rive orientale de la presqu’île
Sillustani. 1, grès indurés sous-jacents ; 2, alternances de grès et pélites ; 3, pélites rouges contenant des intercalations variées, parmi lesquelles les niveaux
fossilifères, les plus riches signalés par des flèches (Fig. 7 pour le détail) ; les deux tiers supérieurs de la pente recouverts d’éboulis ; 4, unité détritique grossière
à dominance gréseuse, incluant des conglomérats volcano-clastiques ; 5, basaltes Umayo (shoshonites ; Miocène terminal - Pliocène) ; 6, paysage
d’arrière-plan ; 7, dépôts de vases lacustres ; A–B situe la coupe ; P, zone de parking du site précolombien de Sillustani ; les lignes légères ou discontinues
correspondent à des sentiers, restes de terrasses ou murets d’enclos à bétail.

very rich, these levels contain fragments of small teeth and
bones, disassociated eggshell fragments, and charophyte ma-
terial. The uppermost ~100 m of the red mudstone unit
occurs in an eastward gully of the scree-covered slope and
has no clean outcrops. A detailed description of the lithology
of the lower portion of the red mudstone unit was given by
Sigé (1972: 377, Fig. 1).
The red mudstone unit is sharply overlain by a conglom-
erate with dark brown to black-coloured clasts of volcanic
origin, of decimeter size range. This hard level produces a
marked topographic step, which forms the northern side of
the Sillustani Peninsula. It is followed by a succession of
dark grey to dark violet volcaniclastic conglomerates and
sandstones, which likewise contrast with the underlying red
mudstone unit. Some indurated bedding surfaces display
bioturbation. The general dip remains the same (nearly 30°)
then goes slightly increasing. This markedly sandstone-
dominated succession is well exposed on the narrow isthmus
of the Sillustani Peninsula. The dark grey volcaniclastic unit
continues southward, forming the eastern side of Lake
Umayo. Similar rocks are seen on the western side of the
Lake Umayo near Vilque, as documented by Portugal (1974).
Different volcaniclastic levels and several large volcanic
clasts from this unit were sampled at the Sillustani isthmus.
Study of thin sections shows that these rocks consist prima-
rily of weathered andesitic material. Attempts at different
institutions to obtain radioisotopic dates from these altered
samples so far have been unsuccessful.
The subhorizontal shoshonites (Umayo Basalt of Portu-
gal, 1974; Umayo Formation of Klinck et al., 1986; Palacios
et al., 1993) that unconformably overlie older rocks in the
Puno region, including the Laguna Umayo-Sillustani sec-
tion, have yielded K-Ar ages of 5.7 ± 0.3 Ma (Bellon and
Lefèvre, 1976, 1977) and 5.97 ± 0.20, 5.94 ± 0.32, 5.92 ±
0.33, 5.91 ± 0.16, 5.82 ± 0.20, and 5.05 ± 0.17 Ma (Kaneoka
and Guevara, 1984). These latest Miocene–earliest Pliocene
rocks significantly post-date the red mudstone unit.
3. Stratigraphic units: previous attributions
and revised identifications
The first question that arises from the issue discussed
above concerns the stratigraphic unit in which the fossils
were found: does the red mudstone unit at Laguna Umayo
actually belong to the Vilquechico Group? If not, which
regional stratigraphic unit does it represent? Is such a revised
identification compatible with the fossil data? In order to
answer these questions, an updated knowledge of the re-
gional stratigraphy is necessary to assess the stratigraphic
position of the Laguna Umayo strata.
3.1. The revised Late Cretaceous–Paleogene stratigraphy
north and northwest of Lake Titicaca
For decades, the Mesozoic-Paleogene stratigraphy of the
Peruvian Altiplano has suffered from some confusion due to
serious discrepancies between authors. This stratigraphy was
recently criticized, restudied, redefined, and successfully
tested by Sempere et al., (2000a). The Late Cretaceous–
 B. Sigé et al. / Geobios 37 (2004) 771–794
Fig. 3. Late Cretaceous–Paleogene stratigraphy northwest of Lake Titicaca
(Azángaro-Putina-Moho region; see Fig. 5 for location). See text for des-
criptions of lithologies. Lateral thickness variations of all units may be
considerable. The term “La Cabaña Member” is taken from the Bolivian
stratigraphic nomenclature.
Stratigraphie du Crétacé supérieur et Paléogène au NE du Lac Titicaca
(région d’Azángaro-Putina-Moho, localisation voir Fig. 5). Voir le texte
pour la description des lithologies. Les variations latérales d’épaisseur des
différentes unités peuvent être considérables. Le terme « Membre La Ca-
bana » est repris de la nomenclature stratigraphique en Bolivie.
Paleogene portion is illustrated in Fig. 3. Regional correla-
tion of the corresponding units appears in Fig. 4.
Late Cretaceous–Paleogene strata in the Peruvian Late
Titicaca region are divided into:
• The Ayabacas Formation (Newell, 1949; spelling
emended by Sempere et al., 2000a), which consists of an
olistostrome, and, more precisely, of a limestone mega-
breccia (sensu Spence and Tucker, 1997) (De Jong, 1974;
Sempere et al., 2000a, 2000b). The unit mainly consists of
reworked interstratified red mudstones and grey to black
limestones of Middle Cretaceous age, as well as lithified
deposits from older Mesozoic units and, less commonly,
blocks of Paleozoic basement. This characteristic and
markedly reworked unit is several hundreds of meters
thick in the Puno area. In southern Peru, the Ayabacas
Formation is known to exist over a >50,000 km2 area. It
was deposited in an extensional tectonic setting, as docu-
mented by outcrops of normal faults controlling sliding of
limestone beds (Sempere et al., 2000b). The reworked
limestones yielded (Albian?-) Cenomanian fossils (New-
ell, 1949; Palacios et al., 1993; Jaillard, 1995). The avail-
able chronologic constraints indicate that deposition of
the Ayabacas Formation occurred during a period of time
within the Cenomanian–Campanian interval.
The Ayabacas Formation is overlain by the Late Creta-
ceous Vilquechico Group (north, northwest, and south of
775
Titicaca Lake), the Puno Group (as west and southwest of
Titicaca Lake), or the Late Oligocene–Miocene Tacaza
Group (farther southwest of Lake Titicaca) (Fig. 5).
• The Vilquechico Group (Jaillard et al., 1993) overlies the
Ayabacas Formation, and thus postdates the extensional
event responsible for the deposition of the limestone
megabreccia (Sempere et al., 2000b). The contact is a
marked disconformity that probably represents a temporal
hiatus. The Vilquechico Group includes three formations,
respectively referred to as the Lower, Middle, and Upper
Vilquechico formations (Jaillard et al., 1993).
The Lower Vilquechico Formation consists of dominantly
red mudstones. It is identical to the “variegated shale” de-
scribed by Newell (1949) below his “Vilquechico Forma-
tion”. This characteristic aspect results from interstratifica-
tion of the red mudstones with subordinate greenish levels. In
the NE part of the Putina fold-thrust belt (Sempere et al.,
2000a), the Lower Vilquechico Formation has at its base a
5–30 m-thick sandstone member, which consists of poorly
bedded, red argillaceous sandstones distributed in beds sev-
eral meters thick. This irregular member grades upward into
the Lower Vilquechico typical mudstones. The Lower
Vilquechico Formation is equivalent to the Chaunaca Forma-
tion of Bolivia and was thus deposited prior to 73 Ma (Sem-
pere et al., 1997, 2000a).
The Middle Vilquechico Formation overlies the Lower
Vilquechico with a simple sedimentary discontinuity. Its
conspicuous basal part consists of a ~15 m-thick, well-
bedded, slightly thickening-upward, sandstone member. It is
overlain by several tens of meters of grey to greenish mud-
stones, in which a characteristic member consisting of black
shales with thin limestone intercalations of restricted-marine
origin occurs. The succession continues with alternations of
greenish and red mudstones. In the upper part of the unit a
several meters-thick intercalation of limestones and marls
that yielded marine fossils (Jaillard et al., 1993) generally
occurs. The Middle Vilquechico Formation is equivalent to
the Lower plus basal Middle El Molino Formation of Bolivia
and was thus deposited during the ~73–66.5 Ma interval
(Sempere et al., 1997, 2000a; contra Jaillard et al., 1993).
The Upper Vilquechico Formation overlies the Middle
Vilquechico with a slightly erosional sedimentary disconti-
nuity. Its conspicuous basal part consists of a ~15–20 m-thick
sandstone unit made up of cross-bedded channels of fluvial
origin. It is overlain by mostly green mudstones intercalated
with numerous and thin, very fine-grained, often quartzitic,
sandstone beds. In the uppermost 20–60 m of the unit, the
mudstones turn bluish purple, which provides a conspicuous
guide-level in the area. The Upper Vilquechico Formation is
equivalent to the Middle (basal part excepted) plus Upper El
Molino Formation of Bolivia and was thus deposited during
the ~66.5–60 Ma interval (Sempere et al. 1997, 2000a; contra
Jaillard et al., 1993).
• The Muñani Formation (Newell, 1949) is the northeastern
equivalent of the Puno Group (sensu Newell, 1949) and
stratigraphically coincides with most of its lower portion
776 B. Sigé et al. / Geobios 37 (2004) 771–794

Fig. 4. Correlation chart of Late Cretaceous–Early Paleogene stratigraphic units between the Cusco area and Andean Bolivia. The stratigraphies of Cusco and
Sicuani areas are modified from Carlotto (1998). Note: shown position of the LURMU is approximate.
Corrélations des unités stratigraphiques du Crétacé supérieur et Paléogène inférieur entre la région de Cusco et la Bolivie andine. Les stratigraphies des régions
de Cusco et de Sicuani sont reprises et modifiées de Carlotto (1998). La position donnée à la LURMU est approximative.

(Sempere et al., 2000a). All preliminary paleocurrents
measured in the Muñani Formation and Lower Puno
Group indicate that all the clastic material was then de-
rived from the south, which, at a regional scale, is consis-
tent with paleocurrents measured in laterally equivalent
units to the northwest and southeast.
The Muñani Formation consists of a thick coarsening- and
thickening-upward succession of “red beds”, the top of
which is unknown in this area. The red color of its mudstones
and siltstones contrasts with the green (to purple) color of the
underlying Upper Vilquechico Formation. Its ≤10–50 m-
thick basal portion is dominated by cross-bedded sandstone
channels, generally interbedded with red siltstones and mud-
stones. These sandstone facies, the erosional surface present
at the base of this unit, and its stratigraphic relationships are
strongly reminiscent of the Cayara Formation of Bolivia, the
base of which is an erosional surface magnetostratigraphi-
cally dated at 58.2 Ma (Sempere et al., 1997). This
sandstone-dominated unit locally overlies a ≤30 m-thick unit
of dominantly red color, that correlates with the Santa Lucía
Formation of Bolivia because of its stratigraphic position. It
is here included in the Lower Muñani Formation for practical
reasons; its facies vary from red mud-flat deposits to fluvial
deposits including red siltstones and channels filled with
sandstones and microconglomerates.
Above its sandstone-dominated basal part, the ~50–
≥700 m-thick Lower Muñani Formation mostly consists of
thick red mudstones, which were deposited in distal alluvial
plain to mud flat and/or lacustrine environments; subordinate
siltstones and channel sandstones also occur. The Lower
Muñani Formation is strongly reminiscent of the Lower
Potoco Formation of Bolivia.
In the Putina-Azángaro area, the Lower Muñani includes a
10–40 m-thick green to dark grey intercalation that locally
includes limestone beds and is apparently of lacustrine ori-
gin. Because of its colors, this sub-unit markedly contrasts
with the thick red mudstones within which it is observed. A
similar sub-unit, the La Cabaña Member, is known in Bolivia
in a similar stratigraphic context (i.e. a thin greenish interca-
lation within the red, thick, mudstone-dominated, Lower
 B. Sigé et al. / Geobios 37 (2004) 771–794 777

Fig. 5. Distribution of Late Cretaceous–Early Paleogene facies and stratigraphic relationships in the Lake Titicaca region.
Distribution des faciès du Crétacé supérieur et du Paléogène inférieur et relations stratigraphiques dans la région du Lac Titicaca.

Potoco Formation) and was magnetostratigraphically dated of rather short duration (ibid.). Because a world-wide, short-
between 56 and 55 Ma (Sempere et al., 1997). This charac- lived, noteworthy climatic perturbation, caused by a large-
teristic sub-unit was deposited during a dry climatic anomaly scale methane release, is dated at 55.5 Ma (Bains et al.,
778 B. Sigé et al. / Geobios 37 (2004) 771–794
1999), it is tempting to identify in Bolivia and southern Peru
this global event as the La Cabaña Member (Sempere et al.,
2000a). This chronologic consistency may be seen as a con-
firmation of the validity of the upper part of Sempere et al.’s
(1997) magnetostratigraphy.
The Upper Muñani Formation is dominated by sandstones
that were deposited in fluvial channels; they generally show
thickening- and coarsening-upward, as well as rare and spo-
radic conglomeratic levels. To the north and in some locali-
ties south of Lake Titicaca, the Lower Muñani rapidly grades
into the Upper Muñani, whereas these units are separated by
a discontinuity in many localities south of Lake Titicaca.
When unconformable, the base of the Upper Munãni sand-
stones coincides with the base of the Tiahuanacu Formation
of the northern Bolivian Altiplano, and with the base of the
San Jerónimo Group of the Cusco area (Fig. 4), but remains
undated. The Upper Muñani Formation and lateral equiva-
lents represent a noteworthy abrupt invasion of the basin by
clastic material derived from a southern direction.
Charophytes suggest an Early Tertiary (Eocene?) age for
the Muñani Formation (Audebaud et al., 1976; Mourier et al.,
1988), which is consistent with the chronology used in this
paper.
3.2. The revised Late Cretaceous–Paleogene stratigraphy
south of Lake Titicaca
The Late Cretaceous–Paleogene stratigraphy defined
north of Lake Titicaca remains valid to the south, where the
Puno Group was originally described by Newell (1949). As
made fairly clear by this author, this group consists of a very
thick (≥5000 m) coarsening- and thickening-upward succes-
sion of “red beds”. Although accurate, this definition, how-
ever, was rigorously followed by only some of the later
authors (Klinck et al., 1986; Palacios et al., 1993; Sempere et
al., 2000a), whereas most of the others viewed the Puno
Group as exclusively composed of conglomerates, i.e. re-
stricted its definition to part of the Upper Puno Group of
Newell (1949).
The Puno Group mostly consists of fluvial and alluvial
plain facies that were deposited in a broad foreland-type
depositional system. Its lower portion includes a significant
amount of red mudstones, and can even be dominated by
such facies (as is observed for example on the Cabanillas-
Mañazo road at UTM 354-8262), which makes it lithologi-
cally similar to the Muñani Formation. South of 15°30′ S, the
Lower Puno Group presents more proximal, coarser facies
than the Muñani, reflecting the overall S>N polarity of the
basin. It is evident that the Muñani Formation is the northern,
more distal, equivalent of the Lower Puno Group, and that it
must therefore be included in this group.
This basin polarity and architecture are also reflected by
the stratigraphic relationships at the base of the Puno Group
(Fig. 5):
• To the northeast, the Puno Group uniformly overlies the
apparently complete Vilquechico Group, and locally in-
cludes at its base a thin equivalent of the Santa Lucía
Formation (see above). The known outcrops of the Peru-
vian equivalent of the La Cabaña Member are found only
in this domain.
• To the southwest, the Puno Group directly overlies the
Ayabacas Formation, as at Callanca and Sillustani (see
discussion below).
• Between these two domains occurs an elongated area
where sections are found in which the Puno Group uncon-
formably overlies a partially eroded Vilquechico Group
(as Chutane and Uyuccasa).
• Finally, to the extreme southwest, a domain is found
where the volcanic and volcaniclastic rocks of the Tacaza
Group directly overlie the Ayabacas Formation or older
strata. The Tacaza Group is of Late Oligocene–Miocene
age, overlies the Puno Group near Puno, and post-dates
significant deformation of Mesozoic strata to the south-
west. Deformation of these strata was probably coeval
with deposition of at least the conglomeratic parts of the
Upper Puno Group, as the lithologies observed in these
conglomerates match those of the deformed strata. It is
therefore likely that this domain was part of an active
deformation belt that bounded the Upper Puno foreland
basin to the southwest.
3.3. Discussion: stratigraphic units present at Laguna
Umayo
Although the contact is not exposed in the Laguna Umayo
area, geologic mapping and satellite imagery make it clear
that the red mudstone unit overlies the Ayabacas Formation
in this area (Fig. 6). The small limestone outcrop east of
Atuncolla, along the road that connects Laguna Umayo and
Sillustani to the Juliaca-Puno highway, and the larger one
farther east at Hacienda Ilpa belong to the Ayabacas Forma-
tion.
Due to their profound facies differences, the fossil-
bearing red mudstone unit at Laguna Umayo can no longer
be assigned to the Vilquechico Formation. The only regional
unit that matches the characteristics of the >500 m-thick red
mudstone unit at Laguna Umayo is the Lower Muñani For-
mation.
The red mudstone unit had been given the name Umayo
Formation by Laubacher and Marocco (1990), which was
followed by some subsequent authors (Muizon, 1991; Jail-
lard et al., 1993). However, the same name had been previ-
ously applied to the overlying subhorizontal Umayo Basalt
of Portugal (1974) by Klinck et al. (1986), who was followed
by Palacios et al. (1993). This confusion in the stratigraphic
nomenclature is resolved here by the identification of the
LURMU with the Lower Muñani Formation.
On the basis of regional correlations, this unit appears to
be of late Late Paleocene (Thanetian)–Early Eocene (early?
Ypresian) age (Fig. 4). The LU-3 and Chulpas fossil-bearing
levels are located in the upper part of the local Lower Muñani
Formation. These levels consist of microconglomerates of
 B. Sigé et al. / Geobios 37 (2004) 771–794
Fig. 6. Geologic sketch map of the Laguna Umayo area. F1= ‘LU-3’ site,
lower fossiliferous level; F2 = ‘Chulpas’ site, upper fossiliferous level.
Esquisse géologique de la région de Laguna Umayo. F1 = site ‘LU-3’,
niveau fossilifère inférieur ; F2 = site ‘Chulpas’, niveau fossilifère supérieur.
rounded millimeter-size limestone grains with clastic con-
tents. Near Cusco, similar limestone-grains, vertebrate-
bearing microconglomerates occur in the Quilque Forma-
tion, a unit that is correlated with the Lower Muñani on
lithostratigraphic grounds (Fig. 4); the Cusco fossils, how-
ever, remain unstudied. The limestone composition and grain
size of these microconglomerates reflect coeval erosion, in a
somewhat distant area, of a widespread limestone unit. Given
the regional geological constraints, we suggest that this erod-
ing unit was the Ayabacas Formation.
Geologic mapping in the Laguna Umayo area (Fig. 6)
shows that the sandstones observed at the base of the mea-
sured section (Fig. 2) are stratigraphically located within the
thick Lower Muñani Formation: they represent fluvial sedi-
mentation within the alluvial plain on which the Lower
Muñani was deposited.
The volcaniclastic sandstone-dominated unit that discon-
formably overlies the red mudstone unit at Laguna Umayo is
part of the Upper Muñani Formation. This is consistent with
the regional stratigraphic relationships and facies of this unit
(see above), and with the observation that, in the Puno-Santa
Lucía area of southern Peru, the abundance of volcanic mate-
rial in the Puno Group increases southwestwards (Portugal,
1974).
779
Because of its proximal position in the basin, the La
Cabaña Member guide-level (~55.5 Ma) is unfortunately not
present in the Lower Muñani Formation at Laguna Umayo.
4. Paleontology
4.1. General data
Oogones of charophyte algae in the Laguna Umayo red
mudstones were previously known to some Peruvian oil
geologists. During field work in 1965, these charophyte-
bearing beds were shown to the French geologist Maurice
Mattauer (University of Montpellier) who found in one of
them (then designated LU-3) a partial mammal jaw. This
mammal was described as Perutherium altiplanense (in
Grambast et al., 1967). In the same paper Louis Grambast
described the LU-3 charophyte species (in current nomencla-
ture Feistiella gildemeisteri, F. gildemeisteri forma minor
and Amblyochara peruviana). He considered this assem-
blage to be chronologically equivalent to that described by
Peck and Reker (1947) from the neighbouring Vilquechico
Formation (in current nomenclature Platychara perlata and
Feistiella ovalis). Until the reassessment provided by Jail-
lard et al. (1993), the classic Late Cretaceous age for the
Vilquechico Formation (Newell, 1949) was based primarily
on knowledge of this flora.
During field work in 1967, several fossiliferous microcon-
glomerate levels were prospected in the lower half of the red
mudstone unit. The most productive and the principally exca-
vated level (LU-3) yielded the assemblage subsequently re-
ported by Sigé (1968, 1971, 1972). According to previous
and new data, level LU-3 was assumed to represent a Late
Cretaceous biotic record of plants (charophytes) and ani-
mals. At that time, it was the first locality in South America to
have yielded bone and dental remains of alleged pre-Tertiary
mammals.
LU-3 fossils include fish, frogs, reptiles, possibly birds,
mammals, eggshells and charophytes. Among the non-
mammal taxa, some were subsequently described and illus-
trated, or have been identified by specialists (Table 1). The
turtle Roxochelis vilavilensis was reported by Bonaparte and
Powell (1980) on the basis of an undescribed plastron (F. de
Broin, pers. com. to B.S.) unprecisely located in the section.
Although including some terrestrial taxa, most LU-3 species
are completely or partially dependent on aquatic environ-
ments and none indicates a marine environment nor suggests
any marine influence in the sedimentation.
The LU-3 level has yielded eggshell material associated
with bone and dental remains of vertebrates. Two eggshell
types have been formerly identified (Dughi and Sirugue in
Sigé, 1968), then restudied (Kerourio and Sigé, 1984;
Vianey-Liaud et al., 1997). The first type is rare and until now
has always been considered as dinosaurian. Lately described
as “dinosauroid spherulitic type, tubospherulitic morpho-
780 B. Sigé et al. / Geobios 37 (2004) 771–794

Table 1 Table 1 (suite)

Updated lists of LU-3 and Chulpas local faunas (levels) from the red muds- Class Mammalia
tone unit (Lower Muñani Formation) at Laguna Umayo Subclass Metatheria
Mise à jour des listes fauniques de LU-3 et Chulpas de l’affleurement des Order Peradectia
pélites rouges (Formation inférieure Muñani) de Laguna Umayo
Family Peradectidae
LU-3 local fauna = lower level Peradectes (nec Alphadon) austrinum (Sigé, 1971, 1972;
Phylum Charophyta Crochet, 1980)
Order Porocharacea Family ?Pediomyidae or ?Microbiotheriidae (Sigé, 1972: 392;
Amblyochara peruviana (Grambast et al., 1967; Feist and this paper)
Grambast-Fessard in Jaillard et al., 1993:646) indet.
Feistiella (= Porochara) gildemeisteri (idem) Order Didelphimorphia
Feistiella gildemeisteri forma minor (Feist and Grambast-Fessard Family Didelphidae
in Jaillard et al., 1993) indet. (= Didelphidae indet. 1 in Sigé, 1972, = Didelphini
indet. in Crochet, 1980)
Class Osteichthyi Didelphimorphia indet. (Sigé, 1972)
Order Semionotiformes Subclass Eutheria
Family Semionotidae Order ?Notoungulata
Lepidotyle enigmatica (Meunier et Gayet, 1992 *) Family Perutheriidae
Order Characiformes Perutherium altiplanense (Thaler in Grambast et al., 1967;
Family indet. Marshall et al., 1983)
Tiupampichthys intermedius Gayet, Jégu et al., 2003 * Order Condylarthra
Family Characidae Family ?Didolodontidae
Subfamily Serrasalminae indet. (first quoted in Kerourio and Sigé, 1984: 140, n3)
indet. (Gayet and Meunier, 1998: 95 *)
Subfamily Tetragonopterinae CHULPAS local fauna = upper level
Indet. (Gayet and Meunier, 1998: 95 *) Phylum Charophyta
Order Siluriformes Order ?Porocharacea
“Doradoida” indet. indet.
Nov. Gen. (Gayet and Meunier, 1998: 99 *) Class Osteichthyi
Order Perciformes Order Characiformes
Family Latidae Family Characidae
Nov. Gen. (Gayet and Meunier, 1998: 101 *) Subfamily Serrasalminae
N.B. Les travaux signalés * fournissent les synonymies des citations indet. (Gayet pers. comm.)
antérieures pour LU3 Subamily Tetragonopterinae
Order Dipnoi indet. (Gayet pers. comm.)
Family Ceratodontidae Order Elopiformes
Ceratodus sp. (Schultze, 1991: Pl. 1, Fig. 4) Family cf. Albulidae
Family Lepidosirenidae indet. (Gayet pers. comm.)
Lepidosiren cf. L. paradoxa (Schultze, 1991: 444) Order Dipnoi
Class Amphibia Family Lepidosirenidae
Order Anura Lepidosiren cf. L. paradoxa (this paper)
Family cf. Leptodactylidae Class Reptilia
indet. (fide Vergnaud-Grazzini in Sigé, 1968) Order Crocodilia
Family indet.
Class Reptilia
Order Chelonia Eggshell material, rare tiny fragments; indet. (Vianey-Liaud et al., 1997,
Family Podocnemididae 86, Figs. 6-4)
?Roxochelis vilavilensis (Bonaparte and Powell, 1980; Class Mammalia
De Broin, 1991: 513): the collection level is unknown, that Subclass Metatheria
of Chulpas is reasonably excluded (outcrop not exposed in Order Didelphimorphia
the road cut previous 1978–1979) Family Didelphidae, indet.; at least three forms (Crochet and
Order Squamata Sigé, 1993: 101)
Family Aniliidae Order Polydolopimorphia
cf. Coniophys (Rage, 1981) Family indet.
Order Crocodilia Subfamily n. subfam. (Sigé et al., in progress)
Family indet. (Sigé, 1968; S. Salisbury in litt. 1997) Chulpasia mattaueri (Crochet and Sigé, 1993: 99)
aff. Chulpasia indet. (Crochet and Sigé, 1993: 100)
Eggshell material Family ?indet.
Dinosaurians, ?ornithopods (fide Dughi and Sirugue in Sigé, Sillustania quechuense (Crochet and Sigé, 1996)
1968), ?sauropods (fide Chiappe et al., 1998), Megaloolithus sp.
(Vianey-Liaud et al., 1997). N.B. The unreproduced find of this rare shell Subclass Eutheria
type at LU-3 actually makes its origin dubious (see text, p. 12) Order Proteutheria indet. (Crochet and Sigé, 1993: 102)
Birds or dinosaurians: ornithoid, ratite or dinosauroid angustispherulitic Order Notoungulata indet.; at least three forms (Crochet and Sigé,
type (Vianey-Liaud et al., 1997) 1993; 103)
(suite)
 B. Sigé et al. / Geobios 37 (2004) 771–794
type, tubocanaliculate pore system”, it was assigned to the
genus Megaloolithus of the Megaloolithidae family
(Vianey-Liaud et al., o.c.). The more common second type
has been interpreted as either avian (Dughi and Sirugue in
Sigé, 1968) or possibly ornithoid dinosaurian (Kerourio and
Sigé, 1984). Its description as “ornithoid type, ratite morpho-
type or dinosauroid type, angustispherulitic morphotype”
allows interpretation as either ratite-avian, or dinosaurian
with a ratite morphotype (Vianey-Liaud et al., o.c.). These
eggshell records for dinosaur occurrence were used to
strengthen the alleged Late Cretaceous age of the rock unit
(Sigé, 1968; Kerourio and Sigé, 1984). Both Laguna Umayo
1 and 2 eggshell types are now recognized, separately, in
other localities (Vianey-Liaud et al., o.c.). These results are
discussed below.
The LURMU outcrops were restudied in situ and sampled
in 1980 and 1985. A few years prior to 1980, the old access
track to Hacienda Umayo was widened and its route locally
straightened in order to improve accessibility to the archeo-
logical and tourist site of Sillustani (Sigé, 1972: Fig. 2; here
Fig. 1). Ditches and a 1–2 m high embankment were con-
structed along the new road. These excavations provided
fresh exposures of the red mudstones, revealing in particular
a diffuse tuffaceous component in the upper part of the
exposed section. A new microconglomerate level also was
uncovered. Like similar cross-bedded channel deposits in the
section, it is lenticular. Being the last microconglomerate
level well exposed in the red mudstone sequence, it was first
named “Upper/ or Top Level”. It occurs ~90 m below the
boundary between the red mudstones and the pyroclastic
conglomerate and ~200 m above the LU-3 level. It proved to
be as fossiliferous as or more than the LU-3 level and it also
yielded mammals among other vertebrate remains. It was
formally named “Chulpas level” by Crochet and Sigé (1993),
from the Quechua name of the neighbouring funeral towers
at Sillustani. Its mammal fauna was reported with description
of a marsupial, Chulpasia mattaueri (Crochet and Sigé,
1993). Another marsupial from Chulpas was described as
Sillustania quechuense (Crochet and Sigé, 1996), and vari-
ous other taxa from the Chulpas and LU-3 local faunas, lower
vertebrates and mammals, remain under study (Table 1).
4.2. Taphonomic process
For LU-3 and Chulpas levels among similar LURMU
channel deposits (see Geological setting), the fossil content
consists of generally disassociated small (millimeter-length)
material, with exceptional centimeter-length fragments such
as the Perutherium type specimen, or the turtle plastron from
an unknown location in the section (Table 1). All the material
has its origin in living organisms of the lacustrine local
environment, within which some of their hard remains were
collected, carried, then deposited by channel currents of low
energy. This process affected not only purely aquatic animals
such as fish, but also organisms linked to banks and shallow
waters such as charophytes, amphibians, crocodiles, turtles,
781
eggs, as well as disassociated remains of terrestrial animals
from the peripheral zone such as mammals. The lack of
typical material from similar areas, such as gastropods, could
imply difficult conditions for preservation of more fragile
calcified structures.
4.3. Discrepant data
Following the description of a didelphimorph marsupial
assemblage (Sigé, 1971, 1972), and despite the acknowl-
edged progressive dental state of Perutherium altiplanense,
the proposed Cretaceous age of the LU-3 local fauna was
initially accepted (e.g. Clemens et al., 1979). The situation
changed with the increasing amount of new data independent
from the locality itself. Van Valen (1988) still remains the
only author who has provided arguments to challenge the
alleged Cretaceous age. Explicitly or more often not, subse-
quent statements retaining an Early Tertiary age of the LU-
3 local fauna refer to Van Valen’s (1988) assessment. Van
Valen took into account data of all the numerous fossil
groups. Some data were subsequently shown to be inaccurate
(Cappetta, 1990). Van Valen (o.c.) also constantly combined
two geographically distant local faunas (Tiupampa, Bolivia,
and Laguna Umayo, Peru) and two geological formations to
which these localities had been initially referred (El Molino
and Vilquechico formations, respectively). Both of these
stratigraphic attributions have now proven to be inaccurate.
On this biased basis, Van Valen’s (1988) analysis remained
unconclusive (contra Bonaparte, 1994, 131), as expressed by
the alternative in the title of his monograph. Nevertheless
most subsequent authors preferred a Tertiary age interpreta-
tion of Laguna Umayo over a Cretaceous one, whereas this
alternative is adequately considered as still unresolved for
some authors (e.g. Clemens, 2001).
The locality of Tiupampa has become increasingly well
known paleontologically and more precisely located strati-
graphically. Its current attribution to the Middle Santa Lucía
Formation and to the early Late Paleocene portion of Chron
26r (~59.0 Ma) is supported by congruent faunal and geo-
chronologic data (Marshall et al., 1997; Sempere et al.,
1997). On this point, the age allocation favored by Van Valen
(1988) for Tiupampa is verified and constrained (Marshall et
al., 1997). This was not the same for Laguna Umayo for
which the paleontological data remained apparently conflict-
ing until now. The hypothesis of correlation between Tiu-
pampa and Laguna Umayo is still not supported by undis-
puted paleontological data, i.e. shared taxa except those of
high rank or with stratigraphic ranges not limited to the K/T
boundary. The Tiupampa record (Marshall and Muizon,
1988) of the LU-3 fauna Peradectes austrinum has been
challenged (Sigé in Jaillard et al., 1993; Muizon, 1991). This
issue is discussed below.
This situation highlights the need to critically reconsider
the current available data concerning the Umayo Formation
as well as the fossil record of its two main levels (LU-3 and
Chulpas). The present aim is to evaluate, without exclusive
782 B. Sigé et al. / Geobios 37 (2004) 771–794
statements as still seen in recent papers, the possible age
constraining value of the Laguna Umayo fossil record; and
consequently to consider to what extent this fossil record fits
the newly available data from stratigraphic correlations and
magnetostratigraphy.
4.4. Constraining age value of the Laguna Umayo fossil
record
4.4.1. Charophytes
With at least one undisputed occurrence, a form “cf. Feist-
iella gildemeisteri” is reported as common in the basal Upper
Santa Lucía Formation in Bolivia (Dejax, 1989). Based on
very scanty sampling, the species Amblyochara peruviana is
cited in the Middle Santa Lucía Formation in the same paper,
Amblyochara being previously recorded from the Paleocene
in Europe (Grambast-Fessard, 1980). Dejax’s identifications
were confirmed through examination of the material by N.
Grambast-Fessard (pers. com. to B.S. and T.S.). Numerous,
securely unreworked, Feistiella gildemeisteri specimens
identified by Dejax, Feist and Grambast-Fessard were subse-
quently recovered from the basal Upper Santa Lucía at Tiu-
pampa (Sempere et al., 1997).
Until then, F. gildemeisteri and A. peruviana were known
by various records in securely determined Late Cretaceous
levels, and their association was stated to indicate the upper
part of the Maastrichtian (Feist and Grambast-Fessard in
Jaillard et al., 1993, 1994). However, new data including
those summarized below (p. 18), firmly substantiate the sur-
vival of F. gildemeisteri into at least the Late Paleocene and
the base of the Eocene, whereas a presumably similar situa-
tion for A. peruviana needs further documentation. The pre-
vious stratigraphic indication of their association has now
lost its constraining value and a reconsideration of charo-
phyte biostratigraphy in the Central Andes is warranted.
4.4.2. Eggshells
The recent study (Vianey-Liaud et al., 1997) of eggshell
material from the Fundo el Triunfo Formation (Late Creta-
ceous, northern Peru) has provided the opportunity to recon-
sider the material described from the LU-3 level at Laguna
Umayo (Sigé, 1968; Kerourio and Sigé, 1984). As most of
the LU eggshell material that was loaned to specialists is lost,
this re-evaluation is principally based on the published illus-
trations. This re-examination benefits, however, from current
progress in the typology of eggshell structures.
The type 1 from Laguna Umayo (“dinosauroid spherulitic
type, tubospherulitic morphotype, tubocanaliculate pore sys-
tem”) is recognized as belonging to the Megaloolithus ooge-
nus as an indeterminate species (Vianey-Liaud et al., 1997:
86). This type is clearly dinosaurian for these authors as it
was for previous authors. From associated egg and bone
embryo evidence, some megaloolithid eggs have been shown
to correspond to sauropod dinosaurs (Chiappe et al., 1998).
Megaloolithus’ known spatial distribution includes different
continental areas (Central India, Northern Peru, SW Europe)
and its stratigraphic range spans the Late Cretaceous, more
precisely the Late Maastrichtian. The LU type 1 is docu-
mented by scarce (and now lost) material resulting from only
the first B.S.’s field trip (1967). It has never been found since
then (1980 and 1985 field work), whereas type 2 was. We
thus have to consider the possibility that technical confusion
occurred during the earliest stage of the research (1967–
1968) when eggshells from various Late Cretaceous locali-
ties as well as different continental areas were under study at
the Aix-en-Provence Museum. At least the type 1 occurrence
needs to be firmly confirmed from new sampling at Laguna
Umayo, prior to be considered as having biostratigraphic
value.
The type 2 from Laguna Umayo (avian type for Dughi and
Sirugue in Sigé, 1968; ornithoid dinosaurian type sensu Ker-
ourio and Sigé, 1984) is recognized as an “ornithoid type,
ratite morphotype or dinosauroid spherulitic type, angusti-
spherulitic morphotype” by Vianey-Liaud et al. (1997: 86). It
has not been possible to decide if it is a ratite or a spherulitic
dinosauroid type. The same ornithoid eggshell type has been
identified in the LU-3 level of the LURMU of southern Peru
and the Fundo El Triunfo Formation of northern Peru (ibid.).
Based on its biostratigraphic record and stratigraphic con-
text, the age of the latter is well established as Late Campa-
nian–Maastrichtian (Mourier et al., 1986, 1988).
It is tempting to suspect relationships between the Umayo
eggshell type 2 and those of flightless birds believed to have
differentiated during the Late Cretaceous in South America
(C. Mourer-Chauviré in litt. 1994; Marshall, 1994). Flight-
less birds are documented as early as the Early Tertiary
(Paleocene, Eocene) on this continent as well as in Antarctica
(Case et al., 1987). Among these birds are the phororhacids
(fossil phororhacid eggshells have not yet been, or know-
ingly, described) which have Eocene representatives in Eu-
rope (Peters, 1987) and persist in the recent Neotropical
fauna with the Cariama and Chunga genera. From available
data (Mikhailov, 1997: 47, 49), the Cariama-type eggshell
differs from both the Umayo types 1 and 2 in its highly
derived condition, and does not solve the relationship alter-
native regarding the Umayo type 2. Furthermore, the mostly
small-sized bone and dental material from LU-3 does not
include specimens clearly referable to dinosaurs or birds.
There is the exception of a larger-size bone fragment, the
histology of which is compatible with that of dinosaurs (de
Riqlès cited in Sigé, 1968: n10).
Based on present knowledge, however, the occurrence of
at least one dinosaur trace (eggshell type 1) at Laguna Umayo
(LU-3 site and level) cannot be entirely ruled out. As dis-
cussed above, the point is actually open to question and needs
to be tested by re-collecting. If presence of eggshell type 1 at
Laguna Umayo is confirmed by new material, its dinosaurian
reference would be strengthened by recent reassessment
(Vianey-Liaud et al., 1997).
It has been suggested in oral discussions that the eggshell
fragments, charophytes, and vertebrate remain need not have
the same origin in the red mudstone unit at Laguna Umayo.
 B. Sigé et al. / Geobios 37 (2004) 771–794
However, the collector (B.S.) has obtained these different
fossils in the same level (LU-3), a 30–40 cm-thick microcon-
glomerate. Their collection has involved the extraction and
examination of the rocks, underwater disaggregation and
screening of the matrix, then acid processing concentration
of the residues. Another often stated query is that eggshells
could be reworked from an older formation. This objection is
not consistent with the lithological observations and inferred
taphonomic processes, noted above.
4.4.3. Didelphimorph marsupials
The rare forms from LU-3 assigned to peradectids and
tentatively to pediomyids (Sigé, 1972) show closest affinities
to forms known from the Late Cretaceous of North America.
In contrast, the varied didelphimorph fauna from the Late
Paleocene of Itaboraí (Marshall, 1987; Marshall et al., 1997)
includes mostly representatives of more derived types, re-
ferred to the Didelphoidea (sensu Marshall, 1987).
In addition to these general aspects, the LU-3 local fauna
includes: (1) Peradectes (nec Alphadon fide Crochet, 1980),
the oldest representative of which (except P. austrinum) is
reported from the Early Paleocene of North America (Mar-
shall and Muizon, 1988); (2) a possible pediomyid, the fam-
ily being well documented in the Late Cretaceous of North
America, with also a Late Paleocene genus reported from
Itaboraí (Marshall, 1987); nevertheless the scarce LU-
3 record requires caution since these alleged pediomyids may
also be regarded as microbiotheriids (Marshall et al., 1989)
which are securely known at Tiupampa (Marshall and
Muizon, 1988); and (3) possibly a didelphid (fide Crochet,
1980), the oldest representatives of which are reported from
the early Late Paleocene of South America (Marshall, 1987;
Marshall et al., 1997).
The LU-3 species Peradectes austrinum and the referred
specimen from Tiupampa (Marshall and Muizon, 1988) are
distinct species according to Sigé (in Jaillard et al., 1993;
Muizon, 1991). From comparison of the original LU-3 speci-
mens with a cast of the single specimen of P. cf. austrinum
from Tiupampa, these species evidently share peradectine
upper molar pattern and main proportions, which support
their generic attribution. Nevertheless P. cf. austrinum shows
differences in size and morphological aspects (larger size
beyond individual or tooth rank variation; longer, shallower,
and less defined median ectoflexus of M3/; StC (stylar cusp
C) more extended; StE poorly differentiated, whereas it is
clearly separated by a groove from StD in P. austrinum, on
M1-2/ as well as on M3/). These differences clearly argue
against specific identity. Two possibilities arise: (1) these
species represent different stages of a single evolutionary
lineage; in this case, the smaller austrinum species would be
ancestral and somewhat older in age, and some evolutionary
trends in the buccal upper molar would be suggested. From
the study of didelphimorph evolutionary lineages among
European Tertiary marsupials (Crochet, 1980), a difference
from 1 to 3 m.y. could be expected. Or (2) these species
belong to two parallel lineages of the Peradectes genus. In
783
this case, the rather close evolutionary stages of P. austrinum
and P. cf. austrinum would only suggest a short time span
(some m.y. as a mean value) between them, without relative
age indication. Neither of these possibilities can be dis-
carded.
Finally, in the present state of knowledge, no clear bios-
tratigraphical constraints arise from this didelphimorph
group, except the suggestion, from Peradectes representa-
tives, of some closeness in time of the Laguna Umayo (LU-3)
and Tiupampa faunas. The later is recognized early Late
Paleocene in age on other evidence (e.g. Sempere et al.,
1997), whereas a different, Early Paleocene age, is assumed
elsewhere for Tiupampa (e.g. Muizon and Cifelli, 2001).
4.4.4. Perutherium altiplanense
Represented by a single fragmentary lower jaw (Thaler in
Grambast et al., 1967), this mammal possesses brachyodont
molars, the cusps of which are arranged in a pronounced
crestiform structure. The resulting brachylophodont mor-
phology denotes an advanced adaptive state, related to some
extent of herbivorous diet. The Perutherium molar pattern is
unquestionably progressive relative to what is known about
other mammals from near the K/T boundary elsewhere in the
world. This point has been stressed by various authors (v
Sigé, 1972, 379).
The statement that Perutherium is too advanced for a Late
Cretaceous age should imply that we know securely to which
group this fossil is referred, in order to analyze its morphol-
ogy in a relevant evolutionary time frame. However, there is
no absolute consensus about the higher systematic relation-
ships of Perutherium. It has been considered to be a condy-
larth or a bunodont marsupial by Thaler (in Grambast et al.,
1967), probably a condylarth without exclusion of bunodont
marsupial affinities by Hoffstetter (1970: n 6), a peculiar
(“perutheriine”) periptychtid condylarth by Van Valen
(1978), a doubtful condylarth by Kielan-Jaworowska et al.
(1979), an ancestral (perutheriid) notoungulate by Marshall
et al. (1983) and accepted as such by Sloan (1987) and Van
Valen (1988), most recently regarded as a South American
ungulate, Perutheriidae, “possibly a didolodontid or a primi-
tive notoungulate”, by McKenna and Bell (1998: 450).
Following the accepted interpretation of Marshall et al.
(1983), Perutherium could appear as a primitive form related
to notoungulates, which were already diversified in the Late
Paleocene elsewhere in South America (Marshall et al.,
1997).
The once classic evolutionary view (e.g. Van Valen, 1978)
in which all ungulates are derived from a latest Cretaceous-
early Paleocene North American common ancestor (namely
Protungulatum) is challenged by modern data. First, Protun-
gulatum appears rather recent for having generated the spec-
tacular radiation implied, already important by Late Pale-
ocene times. Second, evolution from a generalized
tribosphenic pattern of “insectivorous” type with high coni-
cal cusps, to a more bunodont condition, allowing the attri-
tion movements to progressively take a more tangential di-
784 B. Sigé et al. / Geobios 37 (2004) 771–794
rection leading to a lophodont morphology, seems to have
occurred more than once in eutherian history. An incipient
eutherian stage in such a trend is shown in the early Late
Cretaceous (Coniacian) of Central Asia (Nessov, 1987;
Nessov and Kielan-Jaworowska, 1991). In this new appraisal
of the eutherian radiation and ungulate origins, is Peruth-
erium really too advanced for an alleged Late Cretaceous
ungulate? Conversely, Perutherium is plesiomorphic enough
to receive consideration as a primitive South American ungu-
late, such as a possible ancestral notoungulate as proposed
(Marshall et al., 1983).
4.4.5. “Didolodontid indet”
This form from LU-3 (quoted in Kerourio and Sigé, 1984)
is an isolated lower M/3, the morphology and relatively large
size of which resemble those seen in classic didolodontids
from the Late Paleocene and Eocene of Patagonia, southern
Argentina. Although not identical, the LU-3 form is likely
related to Didolodus, and minor differences in evolutionary
stage of their dental morphology suggest a short time span
between both forms. The LU-3 specimen also resembles
Escribania chubutensis, described by Bonaparte et al. (1993)
as a mioclaenid condylarth from the early Late Paleocene
(sensu Marshall et al., 1997) of southern Argentina (Punta
Peligro, Chubut Province).
4.4.6. The Chulpas local fauna
The fossiliferous bed which has yielded this local fauna is
located ~200 m above level LU-3. While the Laguna Umayo
red mudstone succession seems to form a continuous and
homogeneous sedimentary unit below the pyroclastic forma-
tion, the fossils which have indicated (the charophyte asso-
ciation), or have been used to support (eggshell material of
tuberculate type) the Late Cretaceous age of LU-3, have not
been recognized yet among the scanty Chulpas sample
(Vianey-Liaud et al., 1997; Table 1). As in LU-3, the material
from Chulpas is very fragmentary. The lower-rank vertebrate
Chulpas fauna is not significantly different from that of LU-3
(M. Gayet pers. com.; Table 1). The Chulpas mammals
include marsupials and placentals (Crochet and Sigé, 1993)
(Table 1).
The placental taxa include small generalized proteutheri-
ans and relatively hypsodont notoungulates. The proteutheri-
ans, as well as the didelphimorph marsupials, have no con-
straining biostratigraphic value, whereas the diversity and
the evolutionary stage of the Chulpas notoungulates (mainly
tooth fragments) display rather progressive features, e.g.
some degree of hypsodonty.
A bunodont marsupial was described by Crochet and Sigé
(1993) under the name Chulpasia mattaueri. This taxon as
well as a presumably related smaller species were first as-
signed to the caroloameghiniines. Within these, Chulpasia
more closely resembles the Late Cretaceous North American
genus Glasbius than the Paleogene South American repre-
sentatives. More recently, Chulpasia appeared to strikingly
resemble bunodont marsupials that were almost simulta-
neously reported from the Early Eocene Tingamarra Local
Fauna of Murgon, S. Queensland, Australia, viz. first Thyla-
cotinga bartholomaii (Archer et al., 1993), then another new
species being referred to the Chulpasia genus itself (Sigé et
al., in progress). These taxa share derived and plesiomorphic
features which allow recognition of them as members of a
monophyletic subfamilial unit of still unclear familial rela-
tionships among polydolopimorphian marsupials (ibid.). The
dispersal of this therian Gondwanan clade spans the Southern
Hemisphere, similarly to that of the ornithorhynchid
monotremes (Pascual et al., 1992). The closeness of these
Chulpas and Murgon marsupials favors a short time span
between both forms, and so would favor an Early Tertiary age
for Chulpas.
Crochet and Sigé (1996) described another Chulpas mar-
supial, Sillustania quechuense, and assigned it to the order
Polydolopimorphia, whereas it was formerly recognized as
an undetermined caenolestoid (Crochet and Sigé, 1993). It is
characterized by progressive dental features such as a hypo-
cone on the upper molars. So far, the oldest known marsupi-
als with a hypocone were other polydolopimorphians, Epi-
dolops ameghinoi from the Late Paleocene of Itaboraí and
Epidolops sp. from the Late Paleocene of Patagonia (Mar-
shall et al., 1997). The evolutionary stage of marsupial dental
morphology suggests a relatively brief time span between the
Chulpas form and Epidolops.
4.4.7. Appraisal of the fossil record
The biochronologic value of the Laguna Umayo fossil
record is now reduced to a few critical arguments. Indepen-
dently they do not resolve the discrepancy formerly identi-
fied, but as a whole they are generally consistent with the
revised regional stratigraphy.
Until now the main argument in favor of a Late Cretaceous
age was the LU-3 eggshell Type 1 referred to the oogenus
Megaloolithus, which is considered as dinosaurian by differ-
ent specialists. As stated above, the lack of confirmed record
from Laguna Umayo fossil sites now lead us to question this
argument. On the other hand, the charophyte association
claimed as a precise biostratigraphic Late Cretaceous indica-
tor is not restricted to the K/T boundary, or at least warrants
further examinations from specialists. The remaining and
weak arguments allowing a Cretaceous age are: (1) the mor-
phological similarities between Chulpasia and North Ameri-
can Late Cretaceous Glasbius, now challenged by more con-
sistent relationships with Early Eocene Australian forms; (2)
the familial affinities of the didelphimorphian marsupials
from LU-3; and (3) the ambiguous (at once primitive and
progressive) Perutherium ungulate, which thus fails to be
biostratigraphically informative.
The arguments for an Early Tertiary age are more numer-
ous and they all concern mammals: (1) Peradectes austrinum
from LU-3 and P. cf. austrinum from Tiupampa are either
ancestor-descendant related stages in a single evolutionary
lineage, or close evolutionary stages of two parallel lineages;
in both cases a short time span between them is suggested;
 B. Sigé et al. / Geobios 37 (2004) 771–794 785

(2) Perutherium altiplanense from LU-3 is identified as a

primitive notoungulate, which favors its age preceeding the
incipient Late Paleocene notoungulate radiation, neverthe-
less close to it; (3) the “didolodontid indet.” from LU-
3 resembles the early Late Paleocene Escribania and the Late
Paleocene–Eocene Didolodus, both from Patagonia; (4) the
strong resemblance between Chulpasia and Australian Early
Eocene bunodont marsupials is more consistent in the con-
text of a transantarctic monophyletic group of Early Tertiary
age (Paleocene-Early Eocene); (5) the presence of a hypo-
cone on upper molars in another marsupial (Sillustania) from
Chulpas would be an early occurrence of this cusp for a Late
Cretaceous form, whereas it is found in some Late Paleocene
South American marsupials; and (6) the notoungulate re-
mains from Chulpas are both diversified and relatively ad-
vanced in hypsodonty.
Although collectively strongly suggestive of a Paleocene
age, more probably Late Paleocene, the biotic, mostly mam-
malian, arguments are not yet absolutely conclusive. The
rather general precocity of the South American mammal
fauna in the latest Cretaceous cannot be definitely ruled out,
since undisputed diversified local faunas from this time inter-
val remain almost unknown.

5. Paleomagnetism

A ~400 m-thick stratigraphic section of the LURMU was

measured and paleomagnetic samples were collected by C.
Winker in 1985. Within this succession, 43 paleomagnetic
sites (three or four samples/site) were collected, the vast
majority (40 sites) from the lower ~300 m of the section.
Correspondance of the LURMU lithologic section and the
sampled section for paleomagnetism is shown in Fig. 7.
Fig. 8 illustrates the range of thermal demagnetization
behaviors observed for samples from this locality. For some
samples (Fig. 8(a)), the natural remanent magnetism (NRM)
is univectorial with unblocking temperatures ≤580 °C, indi-
cating that magnetite is the carrier of the characteristic com-
ponent of NRM (ChRM) interpreted to be of depositional
origin. Many samples show evidence that a significant por-
tion of the NRM is carried by hematite. The demagnetization
behavior illustrated in Fig. 8(b) indicates that components of
NRM are carried by both magnetite and hematite. These
components of NRM are both of reversed polarity but are not
collinear.
Dramatic examples of multicomponent NRM are encoun-
tered in some samples, with the components of NRM often
nearly antiparallel (Fig. 8(c, d)). In these cases, the highest
unblocking temperature (TUB) component is invariably of
normal polarity, has TUB up to 680 °C, and is carried by
hematite. A reversed polarity NRM component with un-
blocking temperatures in the range ~300 °C ≤ TUB ≤ ~550 °C
is usually observed from sites with multicomponent NRM.
The directions of these NRM components were determined
Fig. 7. Correlated lithologic sections A–B of the Laguna Umayo red clay
unit and sampled section for paleomagnetic studies. Fossil-bearing levels
within the red mudstone unit are numbered from 0 up to 8; among them
LU-3 and Chulpas (LU-8) provided the fossil assemblages which have been
studied. For more details on lithology, see Sigé (1972), Fig. 1.
Corrélation de la coupe lithologique A–B de l’unité pélitique rouge de
Laguna Umayo et de la coupe échantillonnée pour l’étude du paléomagné-
tisme. Les niveaux fossilifères sont dénombrés de 0 à 8 ; parmi eux les
niveaux LU-3 et Chulpas (LU-8) ont fourni les assemblages fossiles étudiés
à ce jour. Pour plus de précisions lithologiques, voir Sigé, 1972, Fig. 1
by free line fits to the progressions of vector end points
observed in the temperature range through which the compo-
nent was unblocked. We interpret this behavior to indicate
that a reversed polarity depositional component of NRM
carried by magnetite is overprinted by a normal polarity
component carried by hematite. This interpretation is sup-
ported by the observation that the normal polarity overprint is
not confined to particular stratigraphic intervals but instead
occurs randomly throughout the sampled stratigraphic inter-
val. We do not regard any of the normal polarity components
as valid recordings of normal geomagnetic polarity during
deposition of the Laguna Umayo section. Accordingly, the
site-mean directions of the normal polarity components are
786 B. Sigé et al. / Geobios 37 (2004) 771–794
Fig. 8. Vector end-point diagrams showing representative thermal demagne-
tization behaviors. Filled circles are projections onto the horizontal plane.
Open circles are projections onto the vertical plane. Numbers adjacent to
data points indicate temperature of thermal demagnetization in °C. a, uni-
vectorial behavior of sample with TUB restricted to ≤580 °C. b, more
complex demagnetization behavior of sample with multiple reversed pola-
rity components of NRM. c, d, complex demagnetization behaviors of
samples with reversed polarity lower TUB component and superimposed
high TUB component of normal polarity.
Diagrammes de désaimentation thermique des échantillons représentatifs.
Cercles pleins : projection dans le plan horizontal ; cercles vides : projection
dans le plan vertical. Les nombres à côté des points donnent la température
de désaimantation en °C. a, exemple d’un échantillon à une seule compo-
sante et dont la température de déblocage est ≤ 580 °C ; b, comportement
complexe d’échantillon montrant plusieurs composantes de polarité in-
verse ; c, d, comportements plus complexes, avec une composante inverse à
faible température de déblocage et une composante normale à haute tempé-
rature de déblocage.
not used in determining the magnetostratigraphy and are not
shown in Fig. 9.
Interestingly, the normal polarity overprint direction is not
aligned with the present geomagnetic field direction at the
sampling location nor with the axial dipole field direction.
This indicates that the overprint direction is not of recent
origin. Fig. 9(a) illustrates site-mean directions of the highest
TUB components of NRM for the Laguna Umayo section;
these site-mean directions are listed in Table 2. The set of
normal polarity site-mean directions carried by hematite are
roughly antiparallel to the set of reversed polarity site-mean
directions many of which are carried by magnetite. These
normal polarity high TUB components have directions which
are distinct from both the present geomagnetic field direction
and the recent axial dipole direction at the collecting site.
Furthermore, these high TUB components have experienced
the same vertical-axis tectonic rotations as the lower TUB
(depositional) components (Butler et al., 1995). These obser-
Fig. 9. Equal-area projections of site-mean directions from Umayo Forma-
tion. Open circles indicate directions in the upper hemisphere while filled
circles indicate directions in the lower hemisphere of the projection. a,
site-mean directions from high TUB components. Open star indicates axial
dipole field direction at sampling site; open square is present geomagnetic
field direction at sampling site. b, site-mean directions from lower TUB
components (TUB r ≤ 580 °C) interpreted as depositional remanence carried
by magnetite. See text for discussion.
Projection stéréographique des directions moyennes de la LURMU. Les
cercles vides indiquent la direction dans l’hémisphère supérieur alors que les
cercles pleins donnent la direction dans l’hémisphère inférieur. a, direction
moyenne des composantes de haute température de déblocage ; l’étoile vide
indique la direction du champ dipolaire au site échantillonné ; le carré blanc
donne la direction du champ géomagnétique ; b, directions moyennes de la
composante de faible température de déblocage (≤580 °C) interprétée
comme une aimantation rémanente portée par la magnétite. Voir le texte.
vations suggest that the normal polarity hematite overprint
was acquired as a chemical remanent magnetism perhaps a
few m.y. after deposition.
Site-mean directions of the reversed polarity “magnetite”
component of NRM (~300 °C ≤ TUB ≤ ~550 °C) are illus-
trated in Fig. 9(b) and are listed in Table 2. The site-mean
directions for this component of NRM are indistinguishable
from the reversed polarity site-mean directions for sites with
univectorial NRM carried by magnetite and for sites with
univectorial NRM carried by hematite. These reversed polar-
ity “magnetite” component directions thus appear to be reli-
able records of the paleomagnetic field at the time of deposi-
tion, despite having a superimposed overprinting component
carried by hematite. The site-mean directions of these NRM
components are used along with the ChRM components from
the univectorial sites to determine the magnetic polarity
sequence at Laguna Umayo.
 B. Sigé et al. / Geobios 37 (2004) 771–794 787

Table 2
Site-mean characteristic magnetization parameters for the red mudstone unit (Lower Muñani Formation) at Laguna Umayo
Direction moyenne des paramètres d’aimantation pour l’affleurement des pélites rouges (Formation Muñani inférieure) à Laguna Umayo
In situ Corrected
Site Level N/N0 k ␣95 I D I′ D′ VGP Lat VGP Long
(m) (°) (°) (°) (°) (°) (°N) (°E)
LU001 a 4.0 3/3 397.6 6.2 18.3 96.0 22.7 107.4 –19.7 12.7
LU002 a 6.25 3/3 120.9 11.3 19.9 100.5 21.9 112.2 –24.2 14.2
LU003 22.75 3/3 21.1 27.5 42.6 130.3 27.3 149.1 –60.1 18.1
b b
LU004 36.75 2/2 –30.1 262.9 –39.1 283.3 17.9 181.2
LU005 52.75 3/3 18.4 29.6 36.5 167.7 10.1 174.2 –77.9 81.0
LU006 58.25 3/3 383.0 6.3 –16.1 296.8 –10.6 304.0 34.1 204.1
LU007 64.75 3/3 12.5 36.5 70.9 163.8 43.3 184.6 –79.6 266.2
LU008 71.75 3/3 6.3 53.7 –37.8 311.9 –25.2 331.3 62.1 201.3
LU008 La 71.75 3/3 250.1 7.8 47.6 146.4 27.2 167.2 –77.6 24.5
b b
LU010L 85.75 2/2 29.4 138.2 15.3 151.0 –60.5 32.1
LU011 L a 96.0 3/3 63.6 15.6 36.1 123.2 28.0 144.0 –55.3 16.3
LU012 a 107.25 3/3 79.0 14.0 39.6 144.6 21.1 161.3 –71.2 32.4
LU013 125.0 3/3 5.9 56.0 42.8 149.5 21.9 166.5 –76.2 36.6
LU014 132.25 3/3 7.2 50.0 61.0 151.9 36.5 179.3 –85.4 298.7
LU015 120.0 3/3 23.6 25.9 40.0 99.5 43.2 129.9 –42.4 0.0
LU016 137.5 3/3 26.8 24.3 27.0 134.1 15.2 146.4 –56.2 29.6
LU017 143.5 3/3 36.8 20.6 28.1 95.4 36.5 116.3 –29.6 5.1
b b
LU018 165.5 2/2 40.1 146.9 21.2 163.6 –73.3 34.5
b b
LU019L 172.75 2/2 17.5 113.5 18.1 124.4 –35.5 19.6
LU020 178.0 3/3 35.1 21.1 –14.4 283.3 –21.1 294.0 25.8 195.2
LU020 L a 178.0 3/3 127.4 11.0 44.7 140.0 27.3 161.3 –71.9 21.5
LU021 a 191.5 3/3 63.9 15.6 38.1 135.7 23.6 154.2 –64.7 24.3
LU022 196.0 3/3 16.9 31.0 –7.5 290.5 –11.5 296.1 26.7 201.3
LU022 L a 196.0 3/3 275.6 7.4 36.6 133.6 23.5 151.8 –62.4 23.5
LU023 200.5 3/3 313.9 7.0 –8.6 269.5 –23.5 278.3 11.1 190.1
LU023 L a 200.5 3/3 206.3 8.6 30.6 128.5 21.0 144.2 –54.9 23.4
LU024 205.5 3/3 15.6 32.4 26.4 142.1 10.9 152.3 –61.0 37.5
b b
LU025 211.75 2/2 28.1 112.4 27.3 130.2 –42.1 14.4
b b
LU026L 219.25 2/2 29.7 133.9 17.5 147.8 –57.9 28.2
LU027 225.25 3/3 44.8 18.6 –6.1 262.3 –24.9 270.1 3.6 187.3
LU027 L a 225.25 3/3 45.4 18.5 36.2 129.3 25.1 148.4 –59.3 20.5
b b
LU028 231.5 2/2 –34.8 316.0 –20.8 332.4 62.7 207.0
LU029 236.5 3/3 13.3 35.2 27.7 124.5 20.6 139.4 –50.2 22.1
LU031 246.0 3/3 109.4 11.8 –38.8 297.6 –32.9 322.0 53.6 190.9
LU031L 246.0 3/3 13.9 34.4 14.8 142.0 1.0 146.3 –53.5 41.3
LU033 a 259.5 3/3 64.5 15.5 7.1 105.1 14.0 111.3 –22.4 18.5
LU034 a 265.75 3/3 67.6 15.1 19.9 120.6 16.3 131.6 –42.2 22.9
LU035 a 270.75 3/3 152.9 10.0 17.7 104.0 23.5 116.6 –28.6 14.3
LU036 275.5 3/3 17.7 30.2 31.4 125.1 23.3 142.3 –53.2 20.5
LU038 284.25 3/3 6.9 51.2 36.9 115.8 32.4 139.1 –50.9 11.2
LU040 297.75 3/3 30.6 22.7 17.2 99.1 25.6 112.0 –24.5 11.9
LU042 381.5 3/3 7.6 48.4 –33.8 291.1 –32.5 313.4 45.4 190.5
Notes: Site: identification number for paleomagnetic site; L following site number indicates lower-temperature component of NRM. Level: stratigraphic level
of collecting site. N/N0: number of samples used to determine site-mean direction divided by number of samples analyzed. k: concentration parameter. ␣95:
radius of cone of 95% confidence about site-mean direction. I and D: inclination and declination of in situ site-mean direction. I′, D′: inclination and declination
of the site-mean direction corrected for bedding tilt. Lat and Long: latitude and longitude of the site-mean virtual geomagnetic pole (VGP) computed from the
corrected direction. a Indicates site-mean directions used in tectonic analysis of Butler et al. (1995). Site numbers in italics indicate site-mean directions not used
in magnetostratigraphic analysis. b Indicates undefined values.

The LURMU magnetostratigraphy is illustrated in
Fig. 10. The ~300 m section which was densely sampled is
entirely of reversed magnetic polarity. The upper ~100 m of
the unit does not have available outcrop for sufficient sam-
pling, so the upper three paleomagnetic sites (all of reversed
polarity) within this portion of the stratigraphic section are
not included in Fig. 10. The fossil vertebrate level LU-
3 occurs ~200 m above the base of the section, while the
Chulpas level occurs at the top of the ~400 m sampled
section.
788 B. Sigé et al. / Geobios 37 (2004) 771–794
Fig. 10. Polarity stratigraphy of Laguna Umayo section. VGP latitudes
determined from site-mean ChRM directions are plotted against stratigra-
phic level. Sites with clustering of sample ChRM directions significant from
random at the 5% significance level are shown by solid circles; VGP
latitudes for sites with poorer within-site clustering are shown by open
circles. Interpreted magnetic polarity zonation is shown at the right; the
entire stratigraphic section is interpreted as reversed polarity.
Magnétostratigraphie de la LURMU. Les latitudes du GVP déterminées à
partir des directions caractéristiques moyennes sont projetées en fonction du
niveau stratigraphique. Les cercles pleins montrent le groupement des direc-
tions caractéristiques des échantillons avec une erreur de 5 % ; les latitudes
du GVP des échantillons avec un plus faible groupement sont représentées
par les cercles vides. A droite figure l’interprétation des zones de polarité
magnétique ; la section stratigraphique est interprétée comme de polarité
inverse dans son ensemble.
6. Discussion
6.1. LURMU age range based on current data
Because the LURMU magnetostratigraphy shows a con-
tinuous reversed polarity over a ~300 m-thick section, the
possible correlations (as constrained by the magnetostrati-
graphic data) and potential age range (latest Paleocene–Early
Eocene as constrained by stratigraphy, and Late Cretaceous–
Early Tertiary, more probably Late Paleocene, as suggested
by the fossil record) are restricted to reverse chrons 29r
through 24r of the Geomagnetic Polarity Time Scale (GPTS)
(sensu Cande and Kent, 1992, 1995).
6.1.1. Late Cretaceous
If the LURMU age is, at least in part, Late Cretaceous as
previously assumed (up to Vianey-Liaud et al., 1997), this
unit could correlate within Chron 29r (latest Maastrichtian–
earliest Paleocene). Such a correlation is contrary to the
available regional stratigraphic observations, and the previ-
ous paleontological arguments used for this age assignment
are now shown to be weak.
6.1.1.1. Distribution of charophytes and correlations. Rely-
ing on its charophyte assemblage, the LURMU was first
considered equivalent to the then unsubdivided Vilquechico
Formation, which was regarded as Late Cretaceous in age
(Newell, 1949; Grambast et al., 1967). A later sedimento-
logic and paleontologic study (Jaillard et al., 1993) of the
type section at Vilquechico, 75 km NNE of Laguna Umayo,
proposed a refined age on the basis of regional correlations:
the LURMU was interpreted as equivalent to the “third minor
sequence” of the Upper Vilquechico Formation (UVF)
mainly on the basis of an updated charophyte biochronology
(Feist and Grambast-Fessard in Jaillard et al., 1993, 1994).
Subsequent correlation of the UVF with the Middle (basal
part excepted) and Upper El Molino Formation of Bolivia
(Sempere et al., 2000a; contra Jaillard et al., 1993) now
implies that it was deposited during the ~66.5–60 Ma interval
(Sempere et al., 1997), and therefore that its “third (=upper)
minor sequence” is Paleocene in age, as are the charophytes
it yielded.
The charophyte assemblage shared by the UVF, LURMU,
and the Santa Lucía Formation are now also known from the
Lower Potoco Formation, a Bolivian equivalent of the Lower
Muñani Formation (det. P. Meza, Universidad Nacional San
Antonio Abad del Cusco, on samples collected by V. Carlotto
and T. Sempere in the Camargo syncline, southern Bolivia, in
1998). Occurrence of these charophytes in LURMU (here
identified as the Lower Muñani Formation according to up-
dated regional stratigraphy) cannot exclusively support a
correlation with the UVF but is consistent with the correla-
tion of the Lower Muñani and Lower Potoco formations
(Fig. 4). It can be concluded that the charophyte fossil record
in the Central Andes is in dire need of absolute age calibra-
tion.
6.1.1.2. Dinosaur occurrence. Dinosaur occurrence in
LURMU is recognized to be doubtful and requiring new field
sampling and checking. Furthermore such an occurrence, if
confirmed, does not necessary retain a diagnostic value for a
Cretaceous age. The survival of dinosaurs in restricted geo-
graphic areas has been invoked and received support from
records assumed to post-date the K/T boundary (Sloan et al.,
1986; Van Valen, 1988; Antunes and Sigogneau-Russell,
1992; more recently Fassett et al., 2000). Based on dinosaur
occurrence, the LURMU age could possibly be on either side
of the K/T boundary, but anyway very close to it, or even
encompassing it (LU-3 on the Cretaceous side and Chulpas
on the Tertiary), and in any case consistent within Chron 29r.
Whereas the hypothesis for a much longer dinosaur survival
is highly doubtful, keeping in mind that the whole Paleocene
fossil record worldwide is relatively well known and does not
include dinosaurs so far (except such restricted transbound-
ary lineages in the earliest Paleocene beds).
In summary, the available evidence calls into question the
Late Cretaceous age previously assumed for the Laguna
Umayo redbeds, including both, the LU-3 and Chulpas fos-
siliferous levels.
6.1.2. Early Tertiary
An Early Tertiary age for the LURMU seems to corre-
spond better with the evidence from both the updated stratig-
raphy and the complete fossil record, mainly the mammal
fauna. In this case, our thick magnetostratigraphic section
may best correlate with the long reversed polarity chrons 26r
 B. Sigé et al. / Geobios 37 (2004) 771–794
(~60.9–57.9 Ma; early Late Paleocene, Selandian) or 24r
(~55.9–53.4 Ma; late Thanetian–early Ypresian).
The regional stratigraphic observations (Fig. 6) would
lead us to favor correlation with Chron 24r. In the latter, the
undecompacted depositional rate for the measured section is
≥120 m/m.y. Although an upper boundary for this rate is
impossible to assess, this order of magnitude is consistent
with depositional rates ≥500 m/m.y. commonly found in
foreland basins.
The fossil record, particularly the mammalian, fits well
with chron 26r, as argued in the “Fossil record appraisal”
paragraph. Furthermore chron 24r fits with some biotic
events such as the chulpasiine marsupials’ transantarctic dis-
persal, close to the Paleocene–Eocene boundary (Sigé et al.,
in progress). Placement of the LURMU within much shorter,
chron 25r (~57.5–56.3 Ma; Thanetian) cannot be definitely
ruled out because a high depositional rate can be expected to
have occurred in this foreland basin.
6.2. Evaluation of the three major hypotheses
The current data on stratigraphy, basin dynamics and
correlations, as well as age indications from the improved
Laguna Umayo fossil record, support tentative correlation of
the reversed polarity magnetozone of the LURMU and the
Geomagnetic Polarity time scale (GPTS). In its lower part,
the LURMU overlies a sandstone unit of several hundred
meters thickness. Potential gaps below the red mudstone unit
(see stratigraphy) could restrict reliable downward extension
of stratigraphic analyses. At the top, the LURMU is discon-
formably overlain by the volcaniclastic conglomerates and
sandstones of the Upper Muñani Formation.
The Laguna Umayo red bed succession is composed
mainly of mudstones with sandstone intercalations and mi-
croconglomerate channel lenses. A diffuse tuffaceous com-
ponent is recorded in the ~50 m underlying the fossiliferous
Chulpas level. Considering the continuous reversed sequence
of the LURMU paleomagnetic section, the available data
from stratigraphy and paleontology favor three main hypoth-
eses concerning correlation of the studied section with the
GPTS (Fig. 11).
6.2.1. Hypothesis 1
The LURMU long reversed polarity sequence could cor-
relate within Chron 29r, between 65.58 and 64.75 Ma (Cande
and Kent, 1995). In this case, these rocks and their fossils
were deposited during a latest Cretaceous–earliest Paleocene
interval (encompassing the K/T boundary). This hypothesis
would partly overlap the age evaluation previously assumed
for the Laguna Umayo fossil-bearing levels. It would support
the possibility that the LU-3 level (with dinosaurian egg-
shells) could be close to K/T boundary, maybe latest Creta-
ceous, while the Chulpas level (with no dinosaurian egg-
shells so far) could be earliest Paleocene. Also according to
this hypothesis, the Laguna Umayo (LU-3 and Chulpas)
local faunas would have few equivalent mammals (as well as
789
other vertebrates, among which dinosaurs) in other faunas
described from South America. These are restricted to rare
occurrences: e.g. the tooth fragment from the Fundo el Tri-
unfo Formation, northern Peru (Sigé in Mourier et al., 1986),
Late Campanian–Maastrichtian (Naeser et al., 1991; Jaillard
et al., 1993) or Late Maastrichtian in age (Vianey-Liaud et
al., 1997); and some remains of tribosphenic and non-
tribosphenic mammalian teeth from the El Molino Formation
of Bolivia, Middle Maastrichtian (Gayet et al., 2001). Such
scarce data remain very far from filling the several million
year gap during which occurred the major faunal change
separating the Alamitian non-therian mammal faunas from
Patagonia (e.g. Pascual, 1998), unprecisely dated within a
Campanian–Maastrichtian interval, from the first Paleocene
eutherian and metatherian faunas of Peligrian sub-age (Sem-
pere et al., 1997; Marshall et al., 1997). A converging view is
found in Muizon and Cifelli (2001).
Nevertheless, the argued fossils from the LURMU are
now understood to have limited value in constraining its age
(i.e. no longer diagnostic Late Cretaceous significance of the
charophyte association; unreproduced sampling and thus
doubted reliability of the dinosaurain eggshell material; rela-
tive primitiveness of some LU-3 mammalian taxa (didelphi-
morph marsupials, Perutherium) previously interpreted as
supporting a Late Cretaceous age; and rather derived condi-
tion of most mammals from Chulpas level). Hypothesis 1 is
now challenged by the new data from regional stratigraphy
and correlations, which lead us to identify the LURMU as
representing locally the Lower Muñani Formation and not
the Upper Vilquechico Formation (UVF) as recently as-
sumed (Jaillard et al., 1993).
In total, Hypothesis 1 cannot be retained here as a consis-
tently sustained interpretation.
6.2.2. Hypothesis 2
The LURMU long reversed polarity sequence may well
correlate with a time interval within Chron 26r, between
60.9 and 57.9 Ma (Cande and Kent, 1995). If so, the strata
and the fossils they bear would have been deposited during
an early Late Paleocene interval, and thus the Laguna Umayo
fossil assemblages would be semicontemporaneous with
those known from Tiupampa (Bolivia) and some localities in
northwest (Salta basin) and southern (San Jorge basin) Ar-
gentina (Bertini et al., 1993).
On the stratigraphic aspect, Hypothesis 2 is consistent
with correlation of the LURMU with the lower part of the
Lower Muñani Formation (Fig. 4). As regards the fossil data,
Hypothesis 2 (early Late Paleocene age) is consistent with
the evolutionary stage of most studied mammals, the older
ones from LU-3 as well as the younger ones from Chulpas:
Perutherium is appropriate as a primitive form with a pos-
sible ancestral role among South-American ungulates, more
precisely within notoungulates, already varied in the Late
Paleocene; the didelphimorph marsupials are primitive
enough, Peradectes austrinum as well as the presumed pedi-
omyids versus microbiotheriids; the didolodontid is rather
790 B. Sigé et al. / Geobios 37 (2004) 771–794

Fig. 11. Preferred (Hypothesis 3) and alternative correlations of Umayo paleomag section with GPTS and vertebrate-bearing units in the Andean basin of
southern Bolivia and northwest Argentina (modified from Fig. 10 in Sempere et al., 1997).
Corrélations privilégiée (Hypothèse 3) et alternatives de la coupe paléomagnétique de Laguna Umayo avec le standard GPTS et les unités fossilifères du bassin
andin de Sud-Bolivie et du Nord-Ouest d’Argentine (repris et modifié de Sempere et al., 1997, Fig. 10).

close to Paleocene-Eocene Patagonian relatives; and Chulpa-
sia has relatives, even more derived, among the heteroge-
neous bunodont marsupials recorded from the Late Pale-
ocene sediments in Brazil and Argentina. The derived
condition of Sillustania finds Late Paleocene equivalents in
Brazil and Argentina. Lastly, the notoungulate tooth material
from Chulpas already suggests an advanced condition in
hypsodonty.
In total the stratigraphy and the mammal record support
the correlation of the LURMU within Chron 26r, and so
sustain an early Late Paleocene age for these sediments and
their fossils.
Nevertheless Hypothesis 2 introduces a problem that
arises from the striking disparity between the Laguna Umayo
local faunas and the ~450 km ESE-ward distant Tiupampa
local fauna in Bolivia. The little diversified Laguna Umayo
 B. Sigé et al. / Geobios 37 (2004) 771–794
samples are biased and obviously incomplete. Conversely,
the Tiupampa local fauna, although taxonomically rich and
varied, does not share species with the Laguna Umayo fau-
nas, with the exception of possibly ancestor-descendant re-
lated Peradectes austrinum and P. cf. austrinum. The best
characterized Laguna Umayo taxa (e.g. Perutherium alti-
planense, didolodont sp., Chulpasia mattaueri, Sillustania
quechuense) are lacking at Tiupampa. Does this difference
result from a biased record at Tiupampa, or from peculiar
geographic, ecological, or chronological features?
6.2.3. Hypothesis 3
The LURMU reversed polarity zone may correlate to a
time interval within Chron 24r. These strata and the fossils
they bear would thus have been deposited during a latest
Paleocene–earliest Eocene interval (late Thanetian–early
Ypresian), between ~55.9 and 53.4 Ma (Cande and Kent,
1995). This age attribution would involve both LU-3 and
Chulpas fossil-bearing levels.
Hypothesis 3 is the one preferred from the stratigraphic
point of view, for reliable regional correlation of the LURMU
and the local sequence including the Lower Muñani Forma-
tion unconformably overlain by the Upper Muñani Forma-
tion. The LURMU fossil (mammal) record appears less con-
sistent with Hypothesis 3 than Hypothesis 2, although
alternative interpretations of main taxa are possibly consis-
tent with the younger age (in fact a rather minor difference in
respect to the whole Late Paleocene ca 5 m.y. in duration).
Nevertheless, new data require consideration of Hypoth-
esis 3. The bunodont marsupials from the Chulpas level,
Chulpasia itself as a scarce, related smaller form (Table 1),
share close relationships with an Australian fossil species
from the Early Tertiary Murgon locality in Queensland, Aus-
tralia, originally described by Archer et al. (1993) as Thyla-
cotinga bartholomaii. This closeness was noted by Sigé et al.
(1995). A bit later another smaller bunodont marsupial was
found at Murgon, closer in size and morphology to Chulpa-
sia (Sigé et al., in progress), so making this genus shared by
two austral continental areas, South America and Australia.
Together with Thylacotinga, they are interpreted to represent
a distinct group, the chulpasiines, among the many and het-
erogeneous Late Cretaceous and Early Tertiary polydolopi-
formian bunodont marsupials reported so far, mostly from
the Americas and Antarctica. If this interpretation is sus-
tained, Chulpas and Murgon chulpasiines have significant
bearing on the relative age of these faunas. The Murgon beds
were first assigned an Early Eocene age of 54.6 Ma from
K/Ar illite radiometric dating (Godthelp et al., 1992). Al-
though questioned (Woodburne and Case, 1996), this age is
consistent with general Murgon faunal data (e.g. Archer et
al., 1999).
The chulpasiines’ geographic range reveals a transantarc-
tic dispersal of these animals across an extended East-
Gondwanan faunal province (Weddellian Province). As re-
gards marsupial and placental taxa, dispersal is accepted as
having selectively occurred from South America to Australia
791
via a western Antarctic way, during a Latest Cretaceous–
Early Paleocene interval (Woodburne and Case, 1996; Pas-
cual, 1998). There are now arguments, such as a typical
archeonycteridid bat at Murgon (Hand et al., 1994), for a
mammal dispersal lasting until Early Eocene times. Another
piece of evidence of mammal transantarctic interchange is
the Patagonian Late Paleocene Monotrematum sudameri-
canum, an ornithorhynchid monotreme from presumed Aus-
tralian migrant ancestors (Pascual et al., 1992).
At least West-Antarctica was “part of the territory where
the peculiar evolution of “South American” land mammals
occurred [during] the Late Cretaceous?—Early Paleogene”,
as shown by continental vertebrates, among which various
typical South-American land mammals (marsupials, xenar-
thrans, ungulates) recovered from Early Tertiary beds (e.g.
Viscaino et al., 1997, 1998). Prior to opening of the Drake
Passage, the terrestrial connection was formed by Patagonia
and Antarctic Peninsula, and is thought to have lasted up to
ca. 36 Ma, Late Eocene (Woodburne and Zinsmeister, 1984).
On the opposite transantarctic side, reconstructions from
geophysical data imply an early Antarctic-Australia separa-
tion (see Viscaino et al. o.c. for extended review): either
shallow marine ingressions within a long interval from 70 to
50 Ma (McGowran et al., 2000), or a wide Southern Ocean
that opened as early as 64 Ma (Lawver et al., 1992), or at the
other extreme late Early Oligocene age for the final separa-
tion (Veevers, 1991).
The closeness of Chulpas and Murgon Chulpasia species
in tooth size and morphology can be interpreted as reflecting
a very short time span, from close to 0 to 1.5 m.y. in evolution
between them, that is presumably after the common genetic
pool was interrupted. This perhaps represents the breaking
away of two regions of the Weddelian Province: South
America and Antarctica on one side, and Australia on the
other. In this scholastic hypothesis, the age of Chulpas
bearing-fossils sediments is either pre- or post-separation. In
the first case Chulpasia mattaueri could have been ancestral
to the Murgon Chulpasia; in the second both species could
have been sister species from a common, South-American
ancestor lineage. Another hypothesis would be that this ma-
jor paleogeographic break occurred later and the chulpasi-
ines had sympatric differentiation.
In any situation the age of Chulpas and Murgon must be
rather close. As stated above, the general Chulpas mamma-
lian data suggest Late Paleocene affinities, so a possible
pre-Murgon age.
7. Conclusion
If true, the close relationship between both Chulpas
(Southern Peru) and Murgon (SW Queensland) Chulpasia
species provides reliable congruency of fossil information
and regional and correlational stratigraphic data, sustaining
Hypothesis 3. The stratigraphic and age reassessment of the
LURMU (Laguna Umayo fossil-bearing red mudstone unit)
792 B. Sigé et al. / Geobios 37 (2004) 771–794
and its attribution to the Lower Muñani Formation and not to
the Upper Vilquechico Formation, and correlation to a time
interval within Chron 24r, thus implies deposition during a
latest Paleocene–earliest Eocene (late Thanetian–early Ypre-
sian) interval between ~55.9 and 53.4 Ma.
If the chulpasiine interpretation is shown to be wrong,
then the fossil mammal record, as it stands, would be equally
consistent with Hypothesis 2 (mostly Chron 26r; early Late
Paleocene age, between 60.9 and 57.9 Ma), while strati-
graphic data would still favor Hypothesis 3. The extra-
mammalian arguments to retain Hypothesis 1 (Chron 29r,
latest Cretaceous–earliest Paleocene, between 65.58 and
64.75 Ma), which would overlap the Late Cretaceous age
assignment assumed until recently for the fossil-bearing La-
guna Umayo mudstones, have now lost credibility. The
present state of knowledge leads us to favor an Early Tertiary
age, more probably latest Paleocene–earliest Eocene for the
biotas and the fossil-bearing red mudstone unit at Laguna
Umayo.
Acknowledgments
Aspects of this study were supported by grants from the
following: the National Geographic Society (2467-82, 2908-
84, 3381-86); the Gordon Barbour Fund, Department of
Geological and Geophysical Sciences, Princeton University;
the National Science Foundation (grants EAR-88-04423 and
EAR-90-17382); the CNRS through URA 327 (1984) and
ATP Evolution (1985); the Ministère de la Recherche, France
(1994); an Action Spécifique (1983–1989) from the Museum
National d’Histoire Naturelle, Paris; the Institut Français des
Études Andines, Lima (1967–1985); the Institut de Recher-
che pour le Développement (IRD, ex-ORSTOM), Lima; the
Universities of Lima and Arequipa and administrative ser-
vices in Peru. For provided information, cast or paper docu-
mentation, technical assistance and/or collaboration we
thank Mouloud Benammi, France de Broin, Christine Bibal,
Gabriel Carlier, Víctor Carlotto, Géraldine Garcia, Mireille
Gayet, Catherine Girard, Nicole Grambast-Fessard, Sue
Hand, Jean-Louis Hartenberger, Serge Legendre, Jean-
Michel Liotard, Bernard Marandat, Laurence Meslin, Claire
Millerand, Cécile Mourer-Chauviré, Christian de Muizon,
Édison V. Oliveira, Rosendo Pascual, Jean-Claude Rage,
Steve Salisbury, Régine Simon-Coinçon, Thierry Smith, C.
Winker. Professors William A. Clemens and Michael Archer
sustained and kindly improved a previous draft of this paper.
For his initiating role when discovering Perutherium at La-
guna Umayo, as well his maintained interest and stimulating
discussions about this matter, Professor Maurice Mattauer is
saluted as a master in geology, a science he teaches to begin
in the field with solid data, fossils among them, with full care
for their significant value.
References
Antunes, M.T., Sigogneau-Russell, D., 1992. La faune de petits dinosaures
du Crétacé terminal portugais. Comunicaçöes dos Serviços Geológicos
de Portugal 70, 49–62.
Archer, M., Godthelp, H., Hand, S.J., 1993. Early Eocene Marsupial from
Australia. Kaupia, Darmstädter Beiträge zur Naturgeschichte 3, 193–
200.
Archer, M., Arena, R., Bassarova, M., Black, K., Brammall, J., Cooke, B.N.,
et al., 1999. The evolutionary history and diversity of Australia’s mam-
mals. Australian Mammalogy 21, 1–45.
Audebaud, E., Laubacher, G., Marocco, R., 1976. Coupe géologique des
Andes du Sud du Pérou de l’Océan Pacifique au bouclier brésilien.
Geologische Rundschau 65, 223–264.
Bains, S., Corfield, R.M., Norris, R.D., 1999. Mechanisms of climate warm-
ing at the end of the Paleocene. Science 285, 724–727.
Bellon, H., Lefèvre, R., 1976. Données géochronométriques sur le volcan-
isme andin dans le Sud du Pérou. Implications volcano-tectoniques.
Comptes Rendus de l’Académie des Sciences de Paris (D) 283, 1–4.
Bellon, H., Lefèvre, R., 1977. Spectre d’âges radiométriques du volcanisme
cénozoïque du Pérou central (région de Castrovirreyna-Ayacucho-
Nazca). Réunion Annuelle des Sciences de la Terre 5, 58.
Bertini, R.J., Marshall, L.G., Gayet, M., Brito, P., 1993. Vertebrate faunas
from the Adamantina and Marília formations (Upper Baurú Group, late
Cretaceous, Brazil) in their stratigraphic and paleobiogeographic con-
text. Neues Jahrbuch för Geologie Paläontologie Abhandlungen 188,
71–101.
Bonaparte, J.F., 1994. Approach to the Significance of the Late Cretaceous
Mammals of South America. Berliner Geowissenschaften Abhandlun-
gen E 13, 31–44.
Bonaparte, J.F., Powell, J.E., 1980. A continental assemblage of tetrapods
from the Upper Cretaceous beds of El Brete, northwestern Argentina
(Sauropoda–Coelurosauria–Carnosauria–Aves). Mémoire de la Société
géologique de France. NS 139, 19–28.
Bonaparte, J.F., Van Valen, L.M., Kramartz, A., 1993. La fauna local de
Punta Peligro, Paleoceno inferior, de la Provincia del Chubut, Patagonia,
Argentina. Evolutionary Monographs, 1–61.
de Broin, F., 1991. Fossil turtles from Bolivia. Suarez-Soruco, R. (Ed.).
Fosiles y facies de Bolivia. 1 Vertebrados. Revista Técnica de los
Yacimientos Petroleros Fiscales de Bolivia 12, 527–909.
Butler, R.F., Richards, D.R., Sempere, T., Marshall, L.G., 1995. Paleomag-
netic determinations of vertical-axis tectonic rotations from Late Creta-
ceous and Paleocene strata of Bolivia. Geology 23, 799–802.
Cande, S.C., Kent, D.V., 1992. A new geomagnetic polarity time scale for
the late Cretaceous and Cenozoic. Journal of Geophysical Research 97,
13917–13951.
Cande, S.C., Kent, D.V., 1995. Revised calibration of the geomagnetic
polarity timescale for the Late Cretaceous and Cenozoic. Journal of
Geophysical Research 100, 6093–6095.
Cappetta, H., 1990. Tertiary dinosaurs in South America? A reply to some of
Van Valen’s assertions. Historical Biology 3, 265–268.
Carlotto, V., 1998. Evolution andine et raccourcissement au niveau de Cusco
(13°–16°S, Pérou). Thèse de Doctorat, Université de Grenoble, France,
1–159.
Case, J.A., Woodburne, M.O., Chaney, D.S., 1987. A gigantic phororhacoid
(?) bird from Antarctica. Journal of Paleontology 61, 1280–1284.
Chiappe, L.M., Coria, R.A., Dingus, L., Jackson, F., Chinsamy, A., Fox, M.,
1998. Sauropod dinosaur embryos from the Late Cretaceous of Patago-
nia. Nature 396, 258–261.
Clemens, W.A., 2001. Patterns of mammalian evolution across the Creta-
ceous–Tertiary boundary. Mitteilungen aus dem Museum für
Naturkunde in Berlin. Zoologische Reihe 77, 175–182.
Clemens, W.A., Lillegraven, J.A., Lindsay, E.H., Simpson, G.G., 1979.
Where, when and what—a survey of known Mesozoic mammal distri-
bution. In: Lillegraven, J.A., Kielan-Jaworowska, Z., Clemens, W.A.
(Eds.), Mesozoic Mammals: the First Two-thirds of Mammalian History.
University of California Press, Berkeley, pp. 7–58.
 B. Sigé et al. / Geobios 37 (2004) 771–794
Crochet, J.-Y., 1980. Les Marsupiaux du Tertiaire d’Europe. Éditions Fon-
dation Singer-Polignac, Paris, 279.
Crochet, J.-Y., Sigé, B., 1993. Les mammifères de Chulpas (Formation
Umayo, transition Crétacé-Tertiaire, Pérou): données préliminaires.
Documents des Laboratoires de Géologie de la Faculté des Sciences de
Lyon 125, 97–107.
Crochet, J.-Y., Sigé, B., 1996. Un marsupial ancien (transition Crétacé–
Tertiaire) à denture évoluée en Amérique du Sud (Chulpas, Formation
Umayo, Pérou). Neues Jarhbuch für Geologie und Paläontologie
Monathefte 1996 (10), 622–634.
De Jong, K.A., 1974. Melange (Olistostrome) near Lake Tioticaca, Peru.
American Association of Petroleum Geologists Bulletin 58, 729–741.
Dejax, J., 1989. Sur la présence de gyrogonites de charophytes dans les
sédiments de la formation El Molino affleurant sur le site de Tiupampa
(près de Vila-Vila, département de Cochabamba, Bolivie). 1er Colloque
International sur les charophytes actuelles et fossiles, Montpellier
4–8 juillet 1989, Résumés, 16.
Fassett, J.E., Lucas, S.G., Zielinski, R.A., Budhan, J.R., 2000. Compelling
new evidence for Paleocene dinosaurs in the Ojo Alamo Sandstone. In:
Catastrophic Events and Mass Extinctions: Impacts and Beyond. Lunar
and Planetary Institute Contributions, Houston, Texas 1953, pp. 45–46.
Gayet, M., Meunier, F.J., 1998. Maastrichtian and Early Late Paleocene
Freshwater Osteichthyes of Bolivia: additions and comments. In: Mala-
barba, L.R., Reis, R.E., Vari, R.P., Lucena, Z.M., Lucena, C.A.S. (Eds.),
Phylogeny and Classification of Neotropical Fishes. Ediprucs, Porto
Alegre, pp. 85–110.
Gayet, M., Marshall, L.G., Sempere, T., Meunier, F.J., Cappetta, H.,
Rage, J.-C., 2001. Middle Maastrichtian vertebrates (fishes, amphibians,
dinosaurs and other reptiles, mammals) from Pajcha Pata (Bolivia).
Biostratigraphic, palaeoecologic and palaeobiogeographic implications.
Palaeogeography, Palaeoclimatology, Palaeoecology 169, 39–68.
Gayet, M., Jégu, M., Bocquentin, J., Negri, R., 2003. New characoids from
Upper Cretaceous and Paleocene of Bolivia and Mio-Pliocene of Brazil.
Phylogenetic and palaeobiogeographic implications. Journal of Verte-
brate Paleontology 23, 28–46.
Godthelp, H., Archer, M., Cifelli, R., Hand, S., Gilkeson, C.F., 1992. Earliest
known Australian Tertiary mammal fauna. Nature 356, 514–516.
Grambast, L., Martinez, M., Mattauer, M., Thaler, L., 1967. Perutherium
altiplanense nov. gen., nov. sp., premier mammifère mésozoïque
d’Amérique du Sud. Comptes Rendus de l’Académie des Sciences de
Paris (D) 264, 707–710.
Grambast-Fessard, N., 1980. Les charophytes du Montien de Mons
(Belgique). Revue de Paléobotanique et Palynologie 30, 67–88.
Hand, S., Novacek, M., Godthelp, H., Archer, M., 1994. First Eocene Bat
from Australia. Journal of Vertebrate Paleontology 14, 375–381.
Hardenbol, J., Thierry, J., Farley, M.B., Jacquin, T., de Graciansky, P.-C.,
Vail, P.R., 1998. Mesozoic and Cenozoic sequence chronostratigraphic
framework of European basins, chart 1. In: de Graciansky, P.-C., Hard-
enbol, J., Jacquin, T., Vail, P.R. (Eds.), Mesozoic and Cenozoic Sequence
Stratigraphy of European basins. SEPM Special Publication 60.
Hoffstetter, R., 1970. Radiation initiale des Mammifères Placentaires et
biogéographie. Comptes Rendus de l’Académie des Sciences de Paris
270, 3027–3030.
Jaillard, E., 1995. La sedimentación albiana-turoniana en el sur del Perú
(Arequipa–Puno–Putina). Sociedad Geológica del Perú, Volumen Jubi-
lar Alberto Benavides, 135–157.
Jaillard, E., Cappetta, H., Ellenberger, P., Feist, M., Grambast-Fessard, N.,
Lefranc, J.P., Sigé, B., 1993. Sedimentology, palaeontology, biostratig-
raphy and correlation of the Late Cretaceous Vilquechico Group of
southern Peru. Cretaceous Research 14, 623–661.
Jaillard, E., Feist, M., Grambast-Fessard, N., Carlotto, V., 1994. Senonian–
Paleocene charophyte succession of the Peruvian Andes. Cretaceous
Research 15, 445–456.
Kaneoka, I., Guevara, C., 1984. K-Ar age determinations of late Tertiary and
Quaternary Andean volcanic rocks, southern Peru. Geochemical Journal
18, 233–239.
793
Kerourio, P., Sigé, B., 1984. L’apport des coquilles d’œufs de dinosaures de
Laguna Umayo à l’âge de la Formation Vilquechico et à la compréhen-
sion de Perutherium altiplanense. Newsletters on Stratigraphy 13, 133–
142.
Kielan-Jaworowska, Z., Bown, T., Lillegraven, J.A., 1979. Eutheria. In:
Lillegraven, J.A., Kielan-Jaworowska, Z., Clemens, W.A. (Eds.), Meso-
zoic Mammals: the first two-thirds of mammalian history. University of
California Press, Berkeley, pp. 221–258.
Klinck, B.A., Allison, R.A., Hawkins, M.P., 1986. The geology of the
Cordillera Occidental and Altiplano west of Lake Titicaca, southern
Peru. British Geological Survey, Nottingham, and INGEMMET, Lima,
1–353.
Laubacher, G., Marocco, R., 1990. La cuenca cretácica del Altiplano
Peruano. Litoestratigrafia e integración secuencial. Boletín de la
Sociedad Geológica del Perú 81, 33–46.
Lawver, L.A., Gahagan, L.M., Coffin, F.M., 1992. The development of
paleoseaways around Antarctica. American Geophysical Union Antarc-
tic Research Series 65, 7–30.
Marshall, L.G., 1987. Systematics of Itaboraian (Middle Paleocene) age
“opossum-like” marsupials from the limestone quarry at São Jose de
Itaboraí, Brazil. In: Archer, M. (Ed.), Possums and Opossums. Surrey
Beatty and Sons and the Royal Zoological Society of New South Wales,
Sydney, pp. 91–160.
Marshall, L.G., 1994. The Terror Birds of South America. Scientific Ameri-
can 270, 64–69.
Marshall, L.G., de Muizon, C., 1988. The Dawn of the Age of Mammals in
South America. National Geographic Research 4, 23–55.
Marshall, L.G., de Muizon, C., Sigé, B., 1983. Perutherium altiplanense, un
notongulé du Crétacé supérieur du Pérou. Palaeovertebrata 13, 145–155.
Marshall, L.G., Case, J.A., Woodburne, M.O., 1989. Phylogenetic relation-
ships of the families of marsupials. Current Mammalogy 2, 433–502.
Marshall, L.G., Sempere, T., Butler, R.F., 1997. Chronostratigraphy of the
Mammal–Bearing Paleocene of South America. Journal of South Ameri-
can Earth Sciences 10, 49–70.
McGowran, B., Archer, M., Bock, P., Darragh, T.A., Godthelp, H., Hage-
man, S., et al., 2000. Australasian palaeobiogeography: the Palaeogene
and Neogene record. Memoir of the Association of Australasian Palae-
ontologists 23, 405–470.
McKenna, M.C., Bell, S.K., 1998. Classification of Mammals above the
species level. Columbia University Press, New York, Chichester, West
Sussex, xi-xii, 1–631.
Meunier, F.J., Gayet, M., 1992. Remaniement de la ganoïne chez un Semi-
onotidae nouveau du Crétacé supérieur de Bolivie : intérêt paléobi-
ologique. Geobios 25, 767–774.
Mikhailov, K.E., 1997. Avian Eggshells: an Atlas of Scanning Electron
Micrographs. British Ornithologists’. Club Occasional Publications 3,
1–88.
Mourier, T., Jaillard, E., Laubacher, G., Noblet, C., Pardo, A., Sigé, B., et al.,
1986. Découverte de restes dinosauriens et mammalien d’âge crétacé
supérieur à la base des couches rouges du synclinal de Bagua (Andes
nord-péruviennes): aspects stratigraphiques, sédimentologiques et palé-
ontologiques concernant la régression fini-crétacée. Bulletin de la
Société géologique de France 2 (8), 171–175.
Mourier, T., Bengtson, P., Bonhomme, M., Buge, E., Cappetta, H., Cro-
chet, J.-Y., et al., 1988. The Upper Cretaceous—lower Tertiary marine to
continental transition in the Bagua basin, Northern Peru. Paleontology,
biostratigraphy, radiometry, correlations. Newsletters on Stratigraphy
19, 143–177.
de Muizon, C., 1991. La fauna de mamíferos de Tiupampa (Paleoceno
inferior, Formacion Santa Lucia), Bolivia. In: Suarez-Soruco, R. (Ed.).
Fosiles y facies de Bolivia. 1 Vertebrados. Revista Técnica de Yacimien-
tos Petroleros Fiscales de Bolivia, 12. pp. 575–624.
de Muizon, C., Cifelli, R.C., 2001. A new basal “Didelphoid” (Marsupialia,
Mammalia) from the Early Paleocene of Tiupampa (Bolivia). Journal of
Vertebrate Paleontology 21, 87–97.
Naeser, C.W., Crochet, J.-Y., Jaillard, E., Laubacher, G., Mourier, T.,
Sigé, B., 1991. Tertiary Fission-track ages from the Bagua syncline
(Northern Peru). Journal of South American Earth Sciences 4, 61–71.
794 B. Sigé et al. / Geobios 37 (2004) 771–794
Nessov, L.A., 1987. Results of search and study of Cretaceous and Early
Paleocene mammals on the territory of the USSR. Ezhgodnik Vse-
soyuznog Paleontologicheskogo Obshchestva 30, 199–219.
Nessov, L.A., Kielan-Jaworowska, Z., 1991. Evolution of the Cretaceous
Asian Therian Mammals. Fifth Symposium on Mesozoic. Terrestrial
Ecosystems and Biota. Extended Abstracts. Contributions from the Pale-
ontological Museum University of Oslo, 364, 51–52.
Newell, N.D., 1949. Geology of the Titicaca region, Peru and Bolivia.
Geological Society of America 36, 1–111.
Norris, R.D., Röhl, U., 1999. Carbon cycling and chronology of climate
warming during the Palaeocene/Eocene transition. Nature 401, 775–778.
Palacios, O., De La Cruz, J., De La Cruz, N., Klinck, B.A., Allison, R.A.,
Hawkins, M.P., 1993. Geología de la Cordillera Occidental y Altiplano al
oeste del Lago Titicaca, Sur del Perú. Boletín del INGEMMET A 42,
1–257.
Pascual, R., 1998. The history of South American land mammals: the
seminal Cretaceous–Paleocene transition. Asociación Paleontólogica
Argentina, Publicación Especial 5, Paleógeno de América del Sur y de la
Península Antárctica, 9–18.
Pascual, R., Archer, M., Ortiz-Jaureguizar, E., Prado, J.L., Godthelp, H.,
Hand, S., 1992. The first discovery of monotremes in South America.
Nature 356, 704–705.
Peck, R.E., Reker, C.C., 1947. Cretaceous and lower Cenozoic Charophyta
from Peru. American Museum Novitates 1369, 1–6.
Peters, D.S., 1987. Ein “Phorusrhacide” aus dem Mittel–Eozän von Messel
(Aves: Gruiformes, Cariamae). Mourer-Chauviré, C. (Ed.). L’Évolution
des Oiseaux d’après le témoignage des fossiles. Table Ronde internatio-
nale du CNRS, Lyon-Villeurbanne, Septembre 1985. Documents des
Laboratoires de Géologie de la Faculté des Sciences de Lyon 99, 71–87.
Portugal, J.A., 1974. Mesozoic and Cenozoic Stratigraphy and Tectonic
Events of Puno-Santa Lucia Area. Department of Puno, Peru. American
Association of Petroleum Geologists Bulletin 58, 982–999.
Rage, C., 1981. Les continents péri-atlantiques au Crétacé supérieur : migra-
tions des faunes continentales et problèmes paléogéographiques. Creta-
ceous Research 2, 65–84.
Röhl, U., Bralower, T.J., Norris, R.D., Wefer, G., 2000. A new chronology
for the late Paleocene Thermal Maximum and its environmental impli-
cations. Geology 28, 927–930.
Schultze, H.-P., 1991. Lungfish from the El Molino (Late Cretaceous) and
Santa Lucia (Early Paleocene) Formations in southcentral Bolivia.
Suarez-Soruco, R. (Ed.). Fosiles y facies de Bolivia. 1 Vertebrados.
Revista Técnica de Yacimientos Petroleros Fiscales de Bolivia 12, 441–
448.
Sempere, T., Butler, R.F., Richards, D.R., Marshall, L.G., Sharp, W.,
Swisher III, C.C., 1997. Stratigraphy and chronology of Upper Creta-
ceous–lower Paleogene strata in Bolivia and northwest Argentina. Geo-
logical Society of America Bulletin 109, 709–727.
Sempere, T., Acosta, H., Carlotto, V., 2000a. Estratigrafía del Mesozoico y
Paleogeno en la region del Lago Titicaca: hacia una solución? X Con-
greso Peruano de Geología, Lima, CD-ROM file GR50, 1–41.
Sempere, T., Jacay, J., Carrillo, M.-A., Gómez, P., Odonne, F., Biraben, V.,
2000b. Características y génesis de la Formación Ayabacas (departamen-
tos de Puno y Cusco). Boletín de la Sociedad Geológica del Perú Lima
90, 69–76.
Sigé, B., 1968. Dents de micromammifères et fragments de coquilles d’œufs
de dinosauriens dans la faune de vertébrés du Crétacé supérieur de
Laguna Umayo (Andes péruviennes). Comptes Rendus de l’Académie
des Sciences de Paris (D) 267, 1495–1498.
Sigé, B., 1971. Les Didelphoidea de Laguna Umayo (formation
Vilquechico, Crétacé supérieur, Pérou), et le peuplement marsupial
d’Amérique du Sud. Comptes Rendus de l’Académie des Sciences de
Paris 273, 2479–2481.
Sigé, B., 1972. La faunule de mammifères du Crétacé supérieur de Laguna
Umayo (Andes péruviennes). Bulletin du Muséum National d’Histoire
Naturelle de Paris 99 (3), 375–405.
Sigé, B., Archer, M., Godthelp, H., Hand, S., Crochet, J.-Y., 1995. Peruvian–
Australian Paleogene mammal connection. Cavep’s 95, Fifth Confer-
ence on Australian Vertebrate Evolution, Palaeontology and Systematics,
Canberra, Programme and Abstracts, 2.
Sloan, R.E., 1987. Paleocene and latest Cretaceous mammal ages, biozones,
magnetozones, rates of sedimentation, and evolution. Geological Soci-
ety of America. Special Paper 209, 165–200.
Sloan, R.E., Rigby, J.K., Van Valen, L.M., Gabriel, D., 1986. Dinosaur
extinction and simultaneous ungulate radiation in the Hell Creek Forma-
tion. Science 232, 629–633.
Spence, G.H., Tucker, M.E., 1997. Genesis of limestone megabreccias and
their significance in carbonate sequence stratigraphic models: a review.
Sedimentary Geology 112, 163–193.
Van Valen, L.M., 1978. The beginning of the Age of Mammals. Evolutionary
Theory 4, 45–80.
Van Valen, L.M., 1988. Paleocene dinosaurs or Cretaceous ungulates in
South America. Evolutionary Monographs 10, 1–79.
Veevers, J.J., 1991. Phanerozoic Australia in the changing configuration of
Proto-Pangea through Gondwanaland and Pangea to the present dis-
persed continents. Australian Systematic Botany 4, 1–11.
Vianey-Liaud, M., Hirsch, K., Sahni, A., Sigé, B., 1997. Late Cretaceous
Peruvian eggshells and their relationships with Laurasian and Eastern
Gondwanian material. Geobios 30, 75–90.
Viscaino, S.F., Bond, M., Reguero, M.A., Pascual, R., 1997. The youngest
record of fossil land mammals from Antarctica; its significance on the
evolution of the terrestrial environment of the Antarctic Peninsula during
the Late Eocene. Journal of Paleontology 71, 348–350.
Viscaino, S.F., Pascual, R., Reguero, M.A., Goin, F.J., 1998. Antarctica as
background for mammalian evolution. Asociación Paleontólogica
Argentina, Publicación Especial 5, Paleógeno de América del Sur y de la
Península Antárctica, 199–209.
Woodburne, M.O., Case, J.A., 1996. Dispersal, vicariance, and the post-
Gondwana Late Cretaceous to Early Tertiary biogeography from South
America to Australia. Journal of Mammalian Evolution 3, 121–161.
Woodburne, M.O., Zinsmeister, W.J., 1984. The first land mammal from
Antarctica. Science 218, 284–286.