Sedimentary Geology 191 (2006) 227 254

www.elsevier.com/locate/sedgeo

Sedimentology and paleoecology of an EoceneOligocene

alluviallacustrine arid system, Southern Mexico
Hugo Beraldi-Campesi a,, Sergio R.S. Cevallos-Ferriz a,1 , Elena Centeno-Garca a,2 ,
Concepcin Arenas-Abad b,3 , Luis Pedro Fernndez c,4
a

Institute of Geology, UNAM, Ciudad Universitaria, Coyoacn, 04510, DF Mxico

Area of Stratigraphy, Department of Earth Sciences, University of Zaragoza, E-50009 Zaragoza, Spain
Area of Stratigraphy, Department of Geology, University of Oviedo, C/ Jess Arias de Velasco s/n, E-33005 Oviedo, Spain
b

Received 18 May 2005; received in revised form 24 January 2006; accepted 23 March 2006

Abstract
A depositional model of the EoceneOligocene Coatzingo Formation in Tepexi de Rodrguez (Puebla, Mexico) is proposed,
based on facies analysis of one of the best-preserved sections, the Axamilpa Section. The sedimentary evolution is interpreted as
the retrogradation of an alluvial system, followed by the progressive expansion of an alkaline lake system, with deltaic, palustrine,
and evaporitic environments. The analysis suggests a change towards more arid conditions with time. Fossils from this region, such
as fossil tracks of artiodactyls, aquatic birds and cat-like mammals, suggest that these animals traversed the area, ostracods
populated the lake waters, and plants grew on incipient soils and riparian environments many times throughout the history of the
basin. The inferred habitat for some fossil plants coincides with the sedimentological interpretation of an arid to semiarid climate
for that epoch. This combined sedimentologicalpaleontological study of the Axamilpa Section provides an environmental context
in which fossils can be placed and brings into attention important biotic episodes, like bird and camelid migrations or the origin of
endemic but extinct plants in this area.
2006 Elsevier B.V. All rights reserved.
Keywords: TepexiCoatzingo; EoceneOligocene; Sedimentary evolution; Fossil tracks; Fossil plants; Riparian ambients

1. Introduction
Corresponding author. Present address: School of Life Sciences,
LSE 418, Arizona State University, Tempe, AZ 85287-4601, USA.
Tel.: +1 480 727 7762; fax: +1 480 965 7599.
E-mail addresses: hberaldi@asu.edu (H. Beraldi-Campesi),
scrscfpb@servidor.unam.mx (S.R.S. Cevallos-Ferriz),
centeno@servidor.unam.mx (E. Centeno-Garca), carenas@unizar.es
(C. Arenas-Abad), lpedro@geol.uniovi.es (L.P. Fernndez).
1
Tel.: +52 55 5622 4312.
2
Tel.: +52 55 5622 4310.
3
Tel.: +34 976 762 129.
4
Fax: +34 98 510 3103.
0037-0738/$ - see front matter 2006 Elsevier B.V. All rights reserved.
doi:10.1016/j.sedgeo.2006.03.018

Extensive magmatism and tectonic activity have

contributed to the complex geological history of
central Mexico during the Cenozoic. Paleogene inland
basins there are poorly known, mainly because of the
extensive cover by younger rocks and imprecise
correlations (e.g. Ferrari et al., 1999; Morn-Zenteno
et al., 1999). Paleontological studies have provided
references for dating rocks and have contributed to a
more comprehensive understanding of the biotic

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H. Beraldi-Campesi et al. / Sedimentary Geology 191 (2006) 227254

communities, but yet such studies do not always

provide an environmental context for fossils and they
rarely involve sedimentological studies in their
paleoenvironmental reconstructions. Therefore, Paleogene basins with well-preserved outcrops and fossils

can be extremely valuable components in facilitating

reconstruction studies.
The EoceneOligocene Coatzingo Formation, located in the CentralSouth area of Puebla, Mexico
(Fig. 1A), is known for its plant and animal fossili-

Fig. 1. (A) Map showing the study area and fossil localities from the TepexiCoatzingo basin. (B) Local geology (solid line-box in map A) of the
Tepexi de Rodriguez area. Pz = Paleozoic; Mz = Mesozoic; K = Cretaceous; E = Eocene; EO = EoceneOligocene; P = PliocenePleistocene;
Q = Quaternary.

H. Beraldi-Campesi et al. / Sedimentary Geology 191 (2006) 227254

ferous localities (e.g. Buitrn and Malpica-Cruz,

1987), particularly artiodactyl (camel-like and cervid)
fossil tracks, together with those of cat-like mammals,
proboscideans, birds, reptiles and the skeleton of a
flamingo (Cabral-Perdomo, 1996). Palynomorph
assemblages described from several localities in the
Formation (Carranza-Sierra, 2001; Carranza-Sierra and
Martnez-Hernndez, 2002; Martnez-Hernndez and
Ramrez-Arriaga, 1999) have served as tools for
dating, biostratigraphical correlations and floral descriptions. An ever-growing collection of fossil plants
from the Ahuehuetes locality, one of the most wellpreserved Oligocene records in Mexico, has been used
to estimate ages, paleotemperatures and to improve
the understanding of biogeographical patterns of
North Americas' flora (Calvillo-Canadell and Cevallos-Ferriz, 2002; Magalln-Puebla and Cevallos-Ferriz, 1994a; Velasco de Len and Cevallos-Ferriz,
2000; Ramrez-Garduo and Cevallos-Ferriz, 2002).
Other fossils from the Formation include fossil wood
(currently being studied), stromatolites, and ostracods.
Outcrops of the Coatzingo Fm in the Pie de Vaca and
nearby localities have been interpreted as lacustrine in
origin (Buitrn and Malpica-Cruz, 1987; Pantoja-Alor
et al., 1988) based on field observations, but no
sedimentological studies that describe the type of
paleolake, its evolution or the overall paleoenvironmental conditions that existed have previously been
undertaken.
In this paper, we provide a paleoenvironmental
reconstruction of part of the Coatzingo Formation,
based on facies analysis of one of its most complete
sections, termed here the Axamilpa Section (AS). An
overall integration of the sedimentological and paleontological data from the region is taken into consideration
to further discuss relevant biotic events, such as
interchanges and endemisms, that may have occurred
in this area, some of them with profound evolutionary
implications.
2. Local geology
The geology of the Tepexi de Rodriguez area (Fig.
1B) has not been studied completely. In general, a
Cenozoic succession is underlain by Cretaceous rocks,
e.g. the Tlayua Fm (Kashiyama et al., 2003; PantojaAlor, 1990) and the Rosario Fm, and by a Paleozoic
basement, the metamorphic Tecomate Formation,
which represents the northernmost part of the Acatln
Complex (Ortega-Gutierrz et al., 1999). The lower
rocks of the Cenozoic succession correspond to a
white, calcareous, clast-supported conglomerate that

229

crops out in many areas of the TepexiCoatzingo

basin, which has been informally linked with the
Eocene Balsas Conglomerate (Martnez-Hernndez
and Ramrez-Arriaga, 1999; Pantoja-Alor et al.,
1988). It crops out West of the AS (although their
contact is not clearly visible), and unconformably
underlies the eastern part of the Ahuehuetes section,
being then at least part of the local basement of the
Coatzingo Fm. Finally, the Cenozoic succession is
covered with the Plio-Pleistocene Agua de Luna
Formation (Pantoja-Alor et al., 1988) in the SE part
of the Tepexi area (Fig. 2).
2.1. The Coatzingo Formation
The Coatzingo Formation, previously named Pie de
Vaca Fm (Pantoja-Alor et al., 1988), correspond to
Paleogene rocks within the TepexiCoatzingo basin.
That Formation has been locally divided into two units:
the lower Pie de Vaca Unit, composed mainly of
limestones and cherty and sandy limestones, and the
upper Ahuehuetes Unit, composed mainly of tuff and
tuffaceous sandstones (Silva-Romo and GonzlezTorres, in: Calvillo-Canadell and Cevallos-Ferriz,
2002; Fig. 2), both interpreted as fluvial to lacustrine
low-energy environments.
The geographical limit of the Coatzingo Fm is at
present uncertain; however, several localities (e.g.
Chigmecatitln, Zaragoza, and Cuayuca) have been
correlated biostratigraphically (Martnez-Hernndez and
Ramrez-Arriaga, 1999), extending its area further to the
west and south of the study site (Fig. 1A).
2.3. The Axamilpa Section
The Axamilpa Section (AS) is located along the
Axamilpa River, close to the Ahuehuetes locality, 3 km
N of the town of Tepexi de Rodrguez, in the central
southern part of Puebla (975548W, 183642N;
1680 masl). This section, which represents the Pie
de Vaca Unit (Fig. 2), is thought to underlie the
Ahuehuetes Unit, although they are geographically
separated.
Initially, the Pie de Vaca Unit was interpreted to be of
Miocene to PliocenePleistocene age (Buitrn and
Malpica-Cruz, 1987; Cabral-Perdomo, 1995; Rodrguez
de la Rosa et al., 2005), based upon fossil mammal
tracks from the Pie de Vaca locality. Later on, extensive
palynologic and paleobotanic studies have pointed to an
EoceneOligocene age for both the Pie de Vaca and
Ahuehuetes Units (Calvillo-Canadell and CevallosFerriz, 2002; Carranza-Sierra and Martnez-Hernndez,

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Fig. 2. General stratigraphy of the regional lithologies within the TepexiCoatzingo basin and the position of the Coatzingo Formation Units. The
localities Axamilpa Section, Ahuehuetes, and Pie de Vaca (right) are shown in a stratigraphical context.

2003; Martnez-Hernndez and Ramrez-Arriaga,

1999), interpretation also supported by regional geological studies (Silva-Romo, 1998 in Calvillo-Canadell
and Cevallos-Ferriz, 2002).
3. Stratigraphy and facies of the Axamilpa Section
Well-exposed horizontal strata of the AS crop out
mainly in a vertical cliff (Fig. 3A) that shows no
evidence of major deformation. Three stratigraphic
sections were measured at sites A, B, and C (Fig. 3B),
then correlated and combined in a single log (Fig. 4,
Log 1). A second section (Log 2) was measured
at site D.
Three lithological groups are recognized in the AS: a)
detrital rocks (conglomerates, coarse sandstones and
minor limestone, from the base of the section to 20 m),
b) detrital and chemical rocks (sandstones, minor
limestones and marls, from 20 to 36 m), which
commonly appear interbedded, and c) rocks mainly of
chemical origin (limestones and evaporites, marls and
scarce terrigenous, from 36 m to the top). Log 2
contains only groups b) and c). Facies descriptions and
interpretations are summarized in Table 1. The vertical
distribution of facies and their associations is shown in
Fig. 5. Microscopic features of the facies are shown in
Fig. 6.

Conglomeratic facies
Matrix-supported conglomerates (Cgm, Fig. 7A)
This facies comprises polymictic, matrix-supported
conglomerates, which form irregular beds up to 2 m
thick. The matrix is a sandsilt mixture with a high
percentage of carbonate. Clasts are angular to subrounded, 3 to 7 cm in diameter, and composed of schists,
white or reddish limestone, claystone and quartz. Some
parts are replaced and cemented with hematite.
Interpretation:
These conglomerates represent cohesive debris flow
deposits. Matrix strength and buoyancy prevented
large clasts to sink, keeping them floating and
favoring a disorganized fabric. These conglomerates
always form on slopes, and are typical of the
proximal or mid parts of alluvial fans (e.g. Colombo,
1992; Reading, 1986).
Clast-supported conglomerates (Cgc, Fig. 7BC)
These consist of poorly sorted clast-supported
conglomerates. Clasts are angular to subrounded,
up to 14 cm long, consisting mainly of schist,
although some quartz and limestone clasts are also
present. Pore spaces between clasts are commonly
filled by calcite cements. This facies forms tabular,
dm- to m-thick, strata with erosive bases. Beds
display planar and trough cross-stratification, with

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Fig. 3. (A) Panoramic view, facing NE, of the Axamilpa Section and surroundings. (B) Topographic map of the Axamilpa Section showing the sites where logs were measured. Log 1 was measured in
sites A, B, and C; Log 2 was measured in site D (logs shown in Fig. 5).
231

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H. Beraldi-Campesi et al. / Sedimentary Geology 191 (2006) 227254

normal or inverse grading. Some beds pass laterally

into coarse sandstone. Clasts are regularly imbricated, showing N40 paleocurrent directions.
Interpretation:
This kind of deposits would form in braided channels
and bars in middle parts of alluvial systems (Miall,
1996). Some coarsening-upward examples of this
facies may record progradation of minor deltaic-type
bars, given their close relationship to calcareous
lacustrine facies, or merely reflect migration of the
coarser head of longitudinal bars on the finer-grained
downcurrent tail.
Sandstone facies
Cross-stratified sandstones (Sc, Fig. 7D)
These consist of very coarse to fine-grained sandstones, arranged in dm to m thick tabular or lenticular
strata, which display normal or inverse grading and
cross stratification. These beds can contain conglomeratic lenses, and may pass laterally into fine-grained
conglomerates. Fluid escape structures are locally
present.
Interpretation:
This facies is common in fluvial systems of medium to
low energy (Miall, 1996), where gravel deposits may
accumulate at the base of the channels (Davis, 1983).
It is interpreted to have formed from channeled and
unconfined flows, in the middle-to-distal parts of
alluvial fans. Fluid escape structures may be evidence
of contemporary seismic movements or other factors
that led to sediment disturbance.
Horizontal-laminated and cross-stratified sandstones
(Shc, Fig. 7E)
This facies is formed by fine to medium (rarely
coarse) sandstones arranged in cm- to dm-thick beds,
which are amalgamated or, more rarely, separated by
pale-green marl layers, containing clay, quartz and
limestone grains. Sandstone beds, which may display
load structures and marl intraclasts in their base,
show internal horizontal lamination, sometimes with
cross and flaser stratification and ripple lamination.
Sandstones contain large amounts of schist fragments
and also scattered angular quartz and other rock
fragments (930 mm in diameter). Calcareous
cement and hematized zones are common in this
facies. The marls are porous and poorly indurated,
and may contain algal lumps, peloids, ostracod
remains, oolites, bioturbation galleries. Brecciation
and nodulation, as well as gypsum crystals and
chalcedony, may also be present. The abundance of
oolites and ostracods in this facies varies along the
section.

Interpretation:
This facies may represent deposits laid down on
the floodplains of a distal alluvial-fan setting or
even in the delta plain of a lacustrine delta system
(cf. Miall, 1992, 1996; Rust, 1980). In such
settings, unconfined floods can deposit sand sheets
with a variety of structures (e.g. parallel, flaser, and
ripple lamination) that reflect variable energy
conditions. The common normal grading in the
sandstones reflects the waning behavior of these
flows, although turbulent episodes capable of
eroding the substratum could have occurred, as
implied by the many marly rip-up clasts. During
quiet periods or the weakest flows, alternated clay
and carbonate accumulation would account for the
formation of marls among the clay-rich layers. The
presence of oolites, which are common in large
saline and calcareous lakes fringes (e.g. Kowalewska and Cohen, 1998; Swirydezuk et al., 1979),
indicates currents or oscillation produced by wind
and swell during periods of absence of terrigenous.
The bioturbation suggests the presence of interstitial organisms and, therefore, the existence of a
water layer. During drought periods, the substrate
would be exposed to desiccation and oxidation,
resulting in brecciation and the precipitation of
hematite.
Sandstone and marl alternations (Scs, Fig. 7FG)
This facies is composed of sandstone and marl
alternations, cm- to dm-thick beds, which typically
form overall coarsening-upward units. The separation of both lithologies is not abrupt but continuous.
Marl layer thickness decrease toward the top, while
thickness of sandstones increase. Loading structures
are observable at the base of the beds, and ripple- and
low-angle stratification are present in the upper
sandstones.
Interpretation:
The coarsening upward trend and the thinner beds
towards the top are common indicators of deltaic
progradation (e.g. Bhattacharya and Giosan, 2003).
Given the thin nature of this facies and its
stratigraphical position between rocks of chemical
origin, it is interpreted as the progradation of small
deltaic lobes in distal flat areas.
Massive or graded sandstones (Sm, Fig. 7H)
This facies comprises mostly fine, but also medium
grain-size sandstones, mainly formed of schist,
quartz and carbonate grains extensively cemented
with calcite and locally with hematite. Sorting is

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Fig. 4. Stratigraphic logs from the Axamilpa Section.

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237

Table 1
General description and interpretation of the Axamilpa Section facies (see text for details)
Facies

Description

Interpretation

Conglomeratic Cgm

Polymictic, matrix-supported conglomerates.

Debris-flow deposits in proximal or middle parts of alluvial

fans.
Braided channel and bar deposition in middle to distal parts of
alluvial systems.
Channelled and unconfined flows in the middle to distal parts of
alluvial fans.
Deposits on floodplains of distal alluvial-fan or lacustrine delta
systems.

Cgc
Sandstone

Sc
Shc

Scs

Sm

Marl

Mm

Mh

Calcareous

Lm

Lo

Ll
Ls
Evaporitic

Gy

Clast-supported conglomerates with erosive bases.

Planar cross-stratification, normal or inverse grading.
Coarse- to fine-grained sandstones in tabular or lenticular
strata. Cross-stratification and conglomeratic lenses.
Fine- to medium-grained sandstones with horizontal and
cross-stratification, and cm-thick interbedded marl
layers.
Sandstone and marl alternations. Net coarsening-upward
intervals. Load structures, ripple and low-angle cross
stratification.
Fine to medium massive sandstone; inverse or normally
graded, internal horizontal lamination and rare
fluid-escape structures.
Marl layers, massive or with horizontal lamination. Chert
bands and gypsum nodules, brecciation and bioturbation
are present.
Alternation of calcareous marls and clay-rich layers.
Load structures, fluid escape structures, desiccation
cracks, ripples, peloids and bioturbation galleries are
found. Fossil plants found at one level.
Limestones in massive or stratified layers containing
desiccation cracks, load structures, vertebrate tracks,
rhizocretions, pores, microgranular and nodular gypsum,
chert bands and nodules, chalcedony, brecciation,
bioturbation, microbial-like lamination, nodulation,
ostracod remains, oolites, bacterial or algal
microorganisms, and peloids.
Massive oolitic limestones with gypsum nodules and
chert bands, vertebrate ichnofossils, peloids, and
ostracods. Coated grains and pisoids, bioturbation and
algal lumps are present.
Varve-like laminated limestones with flaser stratification,
silicified oncolites, chert and gypsum nodules.
Stromatolites capped with gypsum.

Progradation of small deltaic lobes in flat marginal lacustrine

areas.
Suspended deposition (flash floods) in flat alluvial or deltaic
settings, near lake margins.
Periodic or continuous settling of carbonate muds in offshore,
low-energy lacustrine areas, with reduced terrigenous input.
Subaerial exposure events.
Palustrine setting with soil formation and carbonate deposition.

Carbonate deposition in shallow lacustrine to palustrine settings,

where herbaceous vegetation became established.

Margins of saline carbonate lakes with agitated shallow waters.

Rhythmic carbonate deposition in relatively deep, low-energy

offshore lacustrine areas.
Marginal shallow and low-energy lacustrine environments, with
prolonged evaporation and depletion of water.
Gypsum and calcite laminae. Nodular gypsum. Halite Intense and prolonged lake evaporation events.
also present.

good, although the basal part of some beds contains

scattered grains up to 8 mm. Beds are massive or
inversely or normally graded, although, in some
cases, they show horizontal lamination and scarce
fluid escape structures.
Interpretation:
This facies suggest direct sedimentation of sand
from waning, turbulent sediment-laden flows with
no (or little) traction at the bed, perhaps with an
eolian contribution. The differences in grading
would reflect sedimentation from high to low
density fluids (e.g. Lowe, 1979). The presence of
fluid escape structures also indicates liquefied
flows that resulted from rapid flow deceleration

and quick deposition. These probably arose from

flow expansion at the mouth of confined conduits
(channel mouths?), in a wide and flattened alluvial
or deltaic setting. The flows loaded with sand
were perhaps generated by sudden floods during
heavy rainstorms (e.g. Marzo, 1992; Mutti et al.,
1996). Neighboring carbonate strata suggest deposition in a distal floodplain to marginal lacustrine
setting.
Marl facies
Massive or laminated marls (Mm, Fig. 7I)
These marls form tabular layers, cm- to dm-thick,
and are either massive or have horizontal lamination.
Chert bands and microcrystalline gypsum nodules up

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H. Beraldi-Campesi et al. / Sedimentary Geology 191 (2006) 227254

to 1 cm are present. Some marls show brecciation and

bioturbation.
Interpretation:
These marls represent periodic or continuous
settling of carbonate mud in offshore, low-energy
lacustrine areas with little terrigenous input. Gypsum nodules may have formed during periods of
evaporation.
Marls and claystones (Mh, Fig. 7JK)
Alternation of cm- to dm-thick layers of calcareous
marls with cm-thick greenish clay-rich vermiculite
layers. Thin, mm-thick laminae of carbonatecemented sandstone may be present in the marl
levels. Ripples may be present locally, as well as
peloids and bioturbation. Fossil plants were found
at one level. Load structures, fluid escape structures, and desiccation cracks are very common (Fig.
7K). The vermiculite forms layers 13 cm thick
that rhythmically interrupt the marls.
Interpretation:
Vermiculite can be found in soils (McCarthy and
Plint, 2003; Schroeder et al., 1997) and in paleosols
(Davies and Gibling, 2003), indicating that soils
may have formed in a palustrine setting where
carbonate deposition occurred during rainy seasons
with high lake levels. The poor preservation of the
plant fragments implies an early diagenetic
degradation.
Limestone facies
Massive limestones (Lm, Fig. 7L)
This facies consists of white-yellowish, beige or green
mudstones that form cm- to dm-thick tabular layers.
Desiccation cracks, load structures, vertebrate fossil
tracks, rhizocretions, and pores (1 to 3 mm in diameter)
are present. Replaced and fragmentary ostracod
valves, complete and broken oolites, and peloids
occur but are uncommon. Fungal or algal filaments are
also present. Some beds contain microgranular and
microcrystalline nodular gypsum, radial calcite crystals, bands and nodules of chert, and chalcedony. Some
horizons are brecciated, bioturbated, and show microbial-like lamination (with bacterial or algal remains
preserved, although rarely). The surfaces of these rocks
are commonly replaced by hematite.
Interpretation:
This facies formed from carbonate deposition in
shallow lacustrine to palustrine areas that underwent
periodic subaerial exposure. The presence of rhizocretions indicates that herbaceous vegetation covered
parts of the surface, possibly grass-like plants
adapted to basic conditions.

Oolitic limestones (Lo, Fig. 8A)

Beige oolitic grainstones to packstones represent
this facies. They are typically partially silicified,
with large amounts of oolites, peloids, ostracods,
coated grains and pisoids, as well as algal-like
aggregates. In a few places, they show partial dolomitization, bioturbation and hematite replacements.
Some are porous and contain microcrystalline
gypsum nodules, mm to cm in diameter, and chert
bands, mm to cm thick. Vertebrate fossil tracks were
observed at one level.
Interpretation:
These deposits are typical of those that form along
the edges of saline carbonate lakes (e.g. Kowalewska
and Cohen, 1998; Swirydezuk et al., 1979). A
shallow or fringing lacustrine environment with
agitated water that promoted the formation of oolites
is inferred for this facies.
Laminated limestones (Ll, Fig. 8BD)
This facies is represented by cm- to dm-thick tabular
strata of mudstones. Laterally they have mm-thick
varve-like horizontal and wavy laminae and no
bioturbation is observable. Locally they show flaser
stratification and contain scarce silicified oncolites,
chert and gypsum nodules, 0.5 mm to 10 cm in
diameter. Porosity is conspicuous with pores up to
4 mm in diameter. In thin section, small anhedral
gypsum crystals, microgranular chert, clays and
quartz grains are seen.
Interpretation:
These deposits may have formed in relatively deep
and low-energy lacustrine offshore areas by carbonate deposition and variable terrigenous inputs. The
varve-like appearance suggests that the carbonate
versus terrigenous sedimentation was influenced by
seasonal factors. This may imply also changes in the
lake level, and thus the lake area. The scarcity of
oncolites suggests that they were formed in shallower
zones and transported to deeper zones. Likewise,
scattered terrigenous suggest eolian inputs during
high level stages.
Stromatolitic limestones (Ls, Fig. 8EF)
Stromatolites appear associated with facies Ll and
Lm, in strata varying from 60 to 70 cm in thickness.
Two types of stromatolites were found at two
different levels: a) large, columnar and domal
stromatolites, 30 to 40 cm high, which form
abundant, continuous laterally coalescent bioherms;
and b) small, hemispherical stromatolites, 10 to
12 cm high, which occur alone or in clusters of two or

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Fig. 5. Facies distribution in the Axamilpa Section. See text for explanation of facies associations.

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243

Fig. 6. Microscopic features in thin section. (A) Pellets within a spar matrix. (B) Nucleated oolites with chalcedony crystals (C). (C) Large gypsum
crystal (G) within a micritic matrix. (D) Radial growing of diagenetic calcite. (E) Rhizoidal pores (P, black) and nodules (N, dark grey). (F) Rhizoid
pore with inner concentric layers of calcite. (G) Brecciated zone with intraclasts (I). (H) Bioturbated microbial-like lamination. (I) Partially
dolomitized limestone clast, with a miliolid foraminifer (arrow). (J) Echinoderm fragment (E) along with quartz grains (Q) and pelloidal limestone
fragments (P).

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245

Fig. 8. (A) General view of the Facies Lm and Lo next to a rupestrian painting. (B) Facies Ll containing gypsum nodules (arrows). (C) Close-up of the
facies Ll showing compacted mm lamina. (D) Sequence of the facies Mm, Lm, Lo, Lm, and Ll (25 m in log 1). (E) Large stromatolites (facies Ls).
(F) Small isolated stromatolites (facies Ls). (G) Transversal cut of a small stromatolite showing the outside and its internal lamination. (H) Partially
dissolved and collapsed stratum of gypsum (facies Gy) with calcite laminae. (I) Intricate growth of gypsum crystals.

three domes. The stromatolitic lamination shows the

characteristic dark/light-lamina alternation (Reid et
al., 2000; Seong-Joo et al., 2000), with the lighter
laminae being thicker and dominant. Partial silicification is present toward the centre in both types. Both
stromatolitic intervals are noteworthy capped by
gypsum in honeycomb-like frameworks. The matrix

between the stromatolites is massive or laminated

carbonate.
Interpretation:
These facies formed in low energy, shallow and
marginal lake environments. Their continuous and
smooth lamination, and the regular morphology of
the structures suggest that relatively stable conditions

Fig. 7. (A) Close-up of the facies Cgm, showing poor sorted floating clasts. (B) Erosive contact between facies Cgc (above) and facies Cgm (below).
(C) Facies association A in log 1 ( 12 m) showing two cycles (corks). Hammer encircled is scale. (D) Cross-stratification of the facies Sc. (E)
Fine lamination of facies Shc. (F) Facies Scs lamination. (G) Load structures of the facies Scs. (H) Facies Sm. (I) Facies Mm (arrows) in Log 1 at
25 m (scale = 1 m). (J) Facies Mh showing marl (M, light) and claystone (C, dark). (K) Flame structures of the facies Mh where thin sandstone layers
occur. (L) General view of Lm facies in site D (11 m; hammer encircled is scale).

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prevailed during their formation. The extensive

gypsum on their upper surfaces suggest alkaline
conditions with periods of prolonged evaporation.
The limited presence of stromatolites in the AS
suggests that favorable conditions for their growth
were not frequent.
Evaporitic facies
Gypsum (Gy, Fig. 8GH)
This facies consists of dm-thick, tabular to very
irregular beds of dissolved and collapsed gypsum.
Thin and broken carbonate layers and halite casts are
also present. Microgranular, microcrystalline and
macrocrystalline textures are observable, although
macrocrystalline gypsum is more common in irregular beds.
Interpretation:
This facies represents increasing solute concentration in the lake and sediment pore water due to
lowering of the water table and intense
evaporation.
3.1. Silicification processes
Many of the facies above described contain chert
nodules and bands and chalcedony. They are clearly
diagenetic features and are found replacing some of the
primary components of the deposits (e.g. oncolites,
stromatolites, limestones). The source of silica, as in
many silicified deposits elsewhere, can be debatable.
Diatoms have not been detected and thus they can
likely be ruled out as a major source of silica.
Although microbial mats have also been claimed as a
source of silica in lacustrine deposits (Bustillo et al.,
2003), where silicification takes place in littoral and
eulittoral sequences of lacustrine carbonates, the
paucity of microbial deposits in the AS argues against
microbial mats being a major source for silica. Thus, it
is more likely that the extensive magmatic activity that
prevailed in this region during the Paleogene (Martiny
et al., 2000; Morn-Zenteno et al., 1999) would have
accounted for the silica input, as it has been seen in
other basins (e.g. Dunagan and Turner, 2004; Eugster,
1980; Pirajno and Grey, 2002). In some cases,
hydrothermal pulses may have loaded the pores of
the preexistent rocks with the silica that would later
precipitate as chert. In other cases, silicification may
have occurred during early diagenesis by the mixing of
meteoric waters with the evaporite pore waters,
causing dissolution and collapse of the evaporite
layers, calcite cementation and silicification, probably
as a complex and recurrent sequence of processes
(Arenas et al., 1999).

3.2. Facies associations

The facies described above are associated in vertical
sequences that represent the superposition of different
subenvironments through time. Facies that are inferred
to be genetically linked have been grouped, and their
vertical associations (Fig. 6) can be interpreted as a
sequence of events that describe the sedimentary
evolution of the section.
A association: Cgc Sc, Sm Shc, Lo, Mm Lm
This fining upward sequence begins with erosive or
planar contacts. It comprises mainly clastic facies with
thicknesses from 30 to 250 cm, followed by massive
limestones that are up to 35 cm thick.
This association occurs cyclically and suggests
gravel and sand deposition in braided bar and channel
systems of alluvial fans that reached lacustrine areas, in
which sheet-like flows deposited part of the detrital
sediments as facies Shc. Finer sediment reached distal
zones of the fan. Decreasing terrigenous inputs allowed
deposition of massive limestone facies in shallow or
even fringing lacustrine conditions. The sequence
represents the retreat of an alluvial system and the
establishment of lacustrine and palustrine environments.
Features of the alluvial-fan conglomerates, such as
the clast size (814 cm), the local debris-flow events,
and their close relationship with lacustrine facies,
indicate alluvial fans of small size, mostly with active
fluvial-dominated sectors in relatively steep areas, with
their apices close to the palustrine and lacustrine areas
that existed down stream. Paleocurrents inferred from
the conglomerate clast imbrication suggest that the
source of alluvial sediments was located to the SW of the
study area.
B association: Scs Shc, Sm Sc Cgc
This comprises a clastic facies with thickness from
50 to 250 cm and coarsening upward evolution,
beginning at the base with sandstone facies, either
massive or interbedded with marls, overlain by coarser
sandstones that grade into conglomerates towards the
top. Marls are evidence of a water body with carbonate
deposition. This association records the progradation
of the alluvial system by channeled flows that entered
a shallow lake area, giving rise to small deltaic
systems.
C association: (Shc), Sm, Sc Mm Lm, Lo, Ll
This consists of clastic and carbonate facies,
varying from 4 to 70 cm thick, with planar contacts
and a fining-upward pattern. It begins with fine detrital
inputs that induced a subsequent rise of the water table,
and caused offshore marl deposition (facies Mm).
Later, shallower lacustrine conditions gave place to

H. Beraldi-Campesi et al. / Sedimentary Geology 191 (2006) 227254

carbonate deposition in marginal areas with or without

agitation (facies Lo and Lm, respectively). Facies Ll
could represent a slightly deeper lake setting with
episodic carbonate deposition, and would mark the
establishment of lacustrine, probably shallow, carbonate depositional conditions after an initial deepening
event. This association reflects the end of the alluvial
deltaic dominance, followed by deepeningshallowing
lacustrine events.
D association: Mm, Mh Ll, Lm Lo, Ls Gy
This is a complex association formed of cm- to dmthick claystone, marl, calcareous and evaporitic facies.
Commonly the strata are tabular, except for the
gypsum, which is found in deformed, collapsed and
partially dissolved strata. It begins with deposition in
low-energy offshore areas as a consequence of a lake
deepening and expansion. It continues with carbonate
precipitation in either relatively deep (facies Ll) or
shallower waters (facies Lo, Lm) in which palustrine
conditions were present. Two of the most common
sequences of this shallowing process are D1:
Mm Ll Lm; and D2: Mh Ll, Lm. Facies Lm
and Ll can be found alternating through time, which
indicates cyclic water level changes attributed to
deepeningshallowing events, for example, D3: Mm,
Mh Lm Ll Lm (Fig. 6).
Facies Lo and Ls would also imply shallow, but
probably more saline, carbonate lacustrine conditions
in marginal areas affected by waves (Lo) or in calm
areas (Ls), followed by high rates of evaporation and
desiccation that formed facies Gy in very shallow
ponds, and also caused brecciation, nodulation, and
gypsum nodules to form in earlier carbonate facies.
Two of the associations that reflect that evolution are:
Mm Ll, Ls Gy, and Mh Lm (Ll) Lo Gy.
The sequence represents an overall shallowing that
implies the change from lacustrinepalustrine to evaporitic conditions, due to supersaturation of the lake by
increased evaporation that caused sulfate deposition.
Rises in the lake water level due to freshwater inputs,
may have caused dilution of the lake waters, a relative
deepening and the beginning of a new sequence.
3.3. Fossil record of the Axamilpa Section
Stratigraphic appearance of the Axamilpa Section
fossils is indicated in Fig. 4. Representative specimens
are shown in Fig. 9.
Vertebrate fossil tracks: (Fig. 9AB) All the tracks
observed in the AS were found on top of exposed
bedding planes at various levels of the section.

247

Although footprints are not always well marked,

tracks of at least 4 steps can be easily distinguished. In
some strata, the tracks are better defined than in
others, suggesting different consistency and water
saturation of the substrate at the time of the imprint.
They are usually found displaying alternated right and
left steps. Foot lengths vary from 10 to 18 cm. The
form of the digits is typical of ungulates (artiodactyls). Fossil tracks with similar characteristics have
been described from other localities and related to the
Camelidae group (Cabral-Perdomo, 1995).
Bioturbation galleries: (Fig. 9C) Galleries are
observed in thin laminae, in several levels of the
section. The traces are usually oriented in a same
direction, although they may appear randomly
arranged. Sometimes ostracods are found in the
same sediments, and it is possible that they contributed
to the bioturbation, given the benthic and excavator
habits of some species (Henderson, 1990). However,
there are many other possible burrowing candidates.
Leaves: (Fig. 9DI) At least six different types of
leaves were recognized in the facies Mh. They are
moderately well preserved carbonaceous imprints,
both fragmental and complete specimens. Recognized genera include Pseudosmodingium and Pistacia from the Anacardiaceae and Cedrelospermum
from the Ulmaceae. The remaining material has not
been identified due to the absence of diagnostic
characters, although some plant remains resemble
legumes and aquatic plants in morphology.
Oncolites: (Fig. 9JK) Scarce ellipsoidal oncolites, 3
to 5 cm in diameter, were found in limestones (Facies
Ll). Their concentric lamination may be obliterated by
silicification, which is most intense toward the center,
whereas the external surface remains unsilicified.
Rhizocretions: (Fig. 9L) In some strata, relicts of
roots (rhizocretions) were observed. These are small
(up to 10 mm in length), oxidized (inferred by the
color), and mostly appearing as vertical traces.
Ostracods: (Fig. 9MN) These are present in several
horizons of the succession, always in low-energy
lacustrine marls and limestones. The shape and size
of the valves, between 400 and 550 m in length, is
constant along the stratigraphic section, suggesting
stability in diversity of species. They do not display
ornamentation, being quite smooth, which is common in non-marine ostracods (Henderson, 1990), as
assumed for those of the AS. Ostracods are present
either as conjoined or disarticulated valves, complete
or fragmented. They are abundant regionally. Coquinas with silicified ostracods have been observed at
the Zaragoza locality (Fig. 1) and some 3 km E from

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249

Fig. 10. Sedimentary evolution proposed for the Axamilpa Section (see text for explanation). (A) Alluvialfluvial stage. (B) Transitional stage. (C)
Lacustrine stage. (D) Evaporitic stage.

it, suggesting that one or more lakes existed at that

time. In that case, the similarity between faunas may
indicate a relationship between the water bodies.
Microorganisms: (Fig. 9OP) Permineralized microfossils were observed in thin sections of chert
samples. They appear in groups of four cells, wrapped
by a mucilage-like sheath. The diameter of each cell
varies from 5 to 7 m. The diameter of the groups
varies from 10 to 15 m and they are arranged in
small colonies (6 to 20 individuals per colony).
Stromatolites: (see Stromatolitic limestones, Facies
Ls).
4. Evolution of the sedimentary system
The Axamilpa Section clearly shows a fining-upward
trend from alluvial conglomerates and sandstones to
lacustrine carbonates and evaporites. This evolution
reflects an overall retrogradation of an alluvialfluvial
system, followed by expansion of a carbonate deposi-

tional lacustrine system that eventually evolved to a

more shallower and alkaline, intermittent lacustrine
settings. In the studied succession this can be conceived
as four main depositional stages: 1) alluvialfluvial (Fig.
10A), characterized mainly by detrital facies (conglomerates, sandstones and mudstones, 020 m); 2) transitional (Fig. 10B), characterized by marls, sandstones and
limestones (alternating distal alluvial and carbonate
lacustrine facies, 2036 m); 3) lacustrine (Fig. 10C),
characterized by carbonates; and finally 4) evaporitic
(Fig. 10D, 3655 m).
The alluvialfluvial stage, characterized by braided
channel and bar deposits with rare debris flows,
experienced a general retrogradation through time
(associations A and B), that gave rise to the expansion
of palustrine and lake environments northeastward,
overlapping the alluvial domain, as indicated by limestones and marls that cap the retrograde cycles. The
transitional stage records the influence of both the
alluvial and the lacustrine depositional environments,

Fig. 9. (A) Track of vertebrate ichnites. (B) Shape of an ichnite delimited by a draw. (C) Parallel bioturbation galleries. (D) Fossil leaves of
Cedrelospermum sp. (E) Carbonaceous impression of an unidentified leaflet. (F) Poorly preserved fossil leaf of Pistacia sp. (G) Fossil leaf of
Pseudosmodingium sp. (H) Fossil impression of a legume. (I) Fossil leaf of a possible aquatic plant. (J) Oncolites in the field. (K) Transversal cut of a
partially silicified oncolite (white calcite coat). (L) Rhizoid impressions (scale = 1 cm). (M) Transversal view of an ostracod showing substituted
internal remains. (N) Transversal cut of an ostracod with non-symmetric valves. (O) Permineralized microorganisms. (P) Individual microbial cells.

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with alluvial inputs in the form of lacustrine deltaic lobes

and distal-fan sheet-like deposits (associations B and C).
Although a timeframe for these events is difficult to
assess, the frequency of flow-generated deposits appears
to decrease starting from the transitional stage, favoring
the development of palustrine conditions around a
lacustrine water body, as inferred from desiccation
cracks, rhizocretions, nodulation, brecciation, and
fenestral porosity present in facies Lm.
Finally, with the cessation of the alluvial influence,
carbonate lacustrine environments became extensive,
and facies Lm, Ll, Lo, Ls, Mm, and Mh developed
(sequence D). The common, but discontinuous, appearance of ostracods along the Axamilpa Section supports
the existence of intermittent water bodies, whose
intermittency appears to be more frequent as the
environment became dryer. This would imply a
progressive lowering of the lake level, giving rise to
ephemeral and shallow saline lakes. While subjected to
intense evaporation, sulfates (e.g. gypsum) would have
concentrated in the lake and formed early diagenetic
features (e.g., gypsum nodules) in the exposed mud
flats. During early diagenesis, the influence of meteoric
waters may have caused the dissolution and collapse of
the evaporitic layers. The depth, salinity, and size of the
water bodies must have changed substantially over time,
as shown by the different lithofacies, fossil content, and
sedimentary features of the AS rocks. The decreasing
depth, increasing salinity, and ever-smaller water bodies
that represent the lacustrine and evaporitic stages, seem
to reflect a change towards more arid conditions.
Some modern hypersaline terminal lakes and playas
show sedimentary processes similar to those inferred for
the Axamilpa Section and could serve as models to
better understand environmental factors. Extensive
plains are characteristic of these environments, with
high slopes or mountains relatively close to the lake
environment. These lakes have very shallow and even
intermittent waters, scarce rainfalls and high evaporation, carbonate and evaporitic deposits. For example,
Great Salt Lake, Utah, is a shallow calcareous-evaporitic
lake ( 10 m) influenced along its margins by alluvial
systems. Oolites, stromatolites, ostracods and migratory
birds are common, and the vegetation is low and open
(Kowalewska and Cohen, 1998). The Salton Sea in
southern California (Arnal, 1961; Gilmore and Castle,
1983) is a saline lake with scarce peripheral vegetation.
It has hydrothermal activity associated with rifting
(Harmon, 1966) and combines fluvial and lacustrine
sedimentary processes (Arnal, 1961).
As in modern analogs, regional events that were
taking place simultaneously in central Mexico during

the Paleogene, such as extrusive and intrusive magmatism, extensive faulting and block displacements (e.g.
Martiny et al., 2000; Morn-Zenteno et al., 1999), must
have determined sedimentation and biota distribution in
the Coatzingo Fm to a great extent, along with local
factors, such as rain frequency and intensity, changing
topography and drainage patterns.
The fining-upward evolution of the sequence,
inferred as the retrogradation of an alluvial system, is
probably the result of the basin extension. Hydrothermal
pulses, deduced from the occurrence of magnesite and
cherts in the AS and in nearby localities, could represent
the existence of magmatic activity in the subsurface.
Both ancient and present-day hydrothermalism processes have been reported for this area (Carballido-Snchez
and Delgado-Argote, 1989; Jimnez-Surez et al.,
2001). Similarly, the cyclical deposition of some facies
(e.g., associations A, B, D) was probably related either
to tectonic and faulting episodes, or to climatic
variations, maybe related to the global climatic changes
that occurred in that epoch (Frakes et al., 1992; Wolfe,
1994).
5. Paleoecology
The fossil record known from the region correlates
well with the sedimentological data, thus lending support
to paleoecological inferences. For example, the geologic
setting indicates the existence of shallow lacustrine
environments, which is reinforced by the presence of a
flamingo skeleton in the Pie de Vaca locality (CabralPerdomo, 1996). This suggests that environmental
conditions (shallow and saline waters) similar to those
known from where these birds live today (Mascitti and
Bonaventura, 2002) prevailed in this area of the Coatzingo Formation. Flamingos (Phoenicopteridae) have
been described from many Eocene and Oligocene strata
(Martin, 1983; Olson, 1985). Because extant flamingos
are gregarious and migratory (Nager et al., 1996), it is
possible that the discovery of this single flamingo
skeleton represents a larger population. This notion may
also be supported by the fact that the fossil record of
aquatic birds in Tepexi de Rodrguez is composed
mainly of ichnofossils of their tracks (Cabral-Perdomo,
1996). Further work is needed to correlate precisely all
the localities where avian ichnofossils are present, but
their presence in the Coatzingo Formation contributes to
the understanding of the past habitats and ecological
requirements of these birds, as well as their temporal and
spatial distribution during the EoceneOligocene.
Fossil plants from the Axamilpa Section, such as
Cedrelospermum, Pistacia and Pseudosmodingium, are

H. Beraldi-Campesi et al. / Sedimentary Geology 191 (2006) 227254

also found in the Ahuehuetes Unit. The Pseudosmodingium species, which today exists in the Tepexi de
Rodrguez area, is thought to be endemic to this region
and, along with Cedrelospermum, is considered a
pioneer in harsh environments (Ramrez and CevallosFerriz, 2002). Cedrelospermum has been referred to
subhumid climates in communities of desert scrub
(Magalln-Puebla and Cevallos-Ferris, 1994b; Velasco
de Len and Cevallos-Ferriz, 2000), which suggests
similar climatic conditions prevailed in this region
during the Oligocene. Furthermore, thermometric inferences from the Ahuehuetes locality fossil plants,
estimate annual average temperatures of 1018 C
(Velasco de Len, 1999), and deciduous scrub or
chaparral-like communities are thought to have established there in temperate to semiarid areas (i.e. where
annual precipitation was less than annual evaporation,
and where there were long dry periods) (Ramrez and
Cevallos-Ferriz, 2002).
Plant-bearing strata in the Ahuehuetes Unit are
composed of volcanic ash, and although the Axamilpa
Section lacks volcanic components, the major volcanism that occurred in this epoch (Martiny et al., 2000;
Morn-Zenteno et al., 1999; Silva-Romo et al., 2000)
might have caused environmental disruption in the
surrounding areas, which in turn probably influenced
the origin and distribution of plants in this region.
Pistacia fossils from the Ahuehuetes locality have close
relatives that are found today only in restricted points of
Asia and one Oligocene locality in Germany, suggesting
a long history of exchange between the North American
and Eurasian floras (Ramrez and Cevallos-Ferriz,
2002). Given the stratigraphical position of the two
localities (Fig. 2), it is possible that some floral elements
found in the Ahuehuetes locality evolved earlier in the
Eocene and lived around the dry and saline environments represented in the Axamilpa Section.
The presence of riparian-related plants from the
Ahuehuetes Unit, such as Salix, Populus and some
Anacardiaceae (Ramrez-Garduo, 1999), which are
also represented in the AS, may indicate that near-river
environments became established more than once in the
basin. In the Axamilpa Section, however, the presence
of Graminidites sp. (Carranza-Sierra, personal communication, 2003) and fossil roots, together with the
evidence for high salinity and aridity, would indicate a
grass-like vegetation relatively far from the waterways
where the riparian communities became established, as
in modern saline-lake environments (e.g. Kowalewska
and Cohen, 1998; Soria et al., 2000). This fact agrees
with previous assumptions of grass-abundant biomes for
this area (Martnez-Hernndez and Ramrez-Arriaga,

251

2003), and may narrow the existence or riparian

environments to only a few places. Pollen assemblages
also suggest that the mountain ranges that surrounded
the basin were populated mostly by conifers (MartnezHernndez and Ramrez-Arriaga, 1999), contrasting
lowland xeric and upland mesic vegetations.
The distinction between riparian and non-riparian
plants may have implications for their distribution and
taphonomy. Perhaps near-river plants had a higher
potential for fossilization due to their susceptibility of
being quickly buried after flooding events (Stromberg et
al., 1991), so riparian settings would be suitable places
for taphonomical processes. Moreover, higher plant
diversity in arid lands is found around the streams (Ali et
al., 2000; Stromberg et al., 1991), so diversity of fossil
plants in the Coatzingo Formation may be an indicative
of riparian environments.
The fact that riparian ecosystems may have existed in
this region would have profound paleobiological
implications. Riparian zones favor plant dispersal and
mixing of seeds (Stromberg et al., 1991), and high
number of endemisms can be found there (Stohlgren et
al., 2005), especially in desert springs. These systems
may also serve as dispersal ways for plant and animal
species (Baker, 1986). Furthermore, resources such as
drinking water, shade, food, and shelter, would have
been an attraction for animals and may have helped as
corridors for their dispersal within arid zones. In that
sense, these environments could have had an influence
on the biodiversity and distribution of plants and
animals, and serve as spots for endemisms.
The presence of this resource rich environment is
further supported by the abundance of fossil tracks
related to camel-like animals at various stratigraphic
levels of the AS and their extensive occurrence in the
Tepexi de Rodrguez region (Fig. 1A). The tracks
indicate that these animals traversed the area regularly;
furthermore, they are perhaps an indication of their
dispersion process. Camelids originated in North
America during the PaleoceneEocene in North America (Stearn and Carrol, 1989), and 2 events of
dispersion, one to Eurasia and Africa, and one to
South America, have been proposed for the Miocene
and PliocenePleistocene respectively (van der Made et
al., 2002); however, the fossil record of this group is
incomplete and the starting points of these migrations
are unknown. Their association with desiccation cracks
and rhizocretions in the AS suggests animal movement
across emergent areas, although resources from streams
would have been their main attraction. Perhaps the
TepexiCoatzingo basin served as a migratory path for
these animals, as well as a place to drink, feed or breed.

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Finally, the recurrence of some of the taxa (e.g. fossil

tracks, ostracods, plants) in various levels of the
Coatzingo Fm indicate their permanence over time,
and suggests that either the environmental conditions
necessary for their development remained stable or that
they adapted to the changing environments.
6. Conclusions
While a wide variety of habitats exist today in this
region of the globe, where life forms have existed and
evolved, little is known about these diverse scenarios
during the Cenozoic. Interpreting past life and the
places where they lived opens the opportunity to
understand particular biological or geological phenomena, and to compare them at different moments, which
offers a more dynamic understanding of the natural
processes.
The depositional model of the Axamilpa Section
provides a conception of the nature and evolution of
a dry paleoenvironment over time, and gives organisms, or communities, a context in which they were
able to evolve. Providing an environmental context
for fossils allows a better understanding of their
ecology and distribution over time and opens the
opportunity for discussions with a timeline vision.
Moreover, fossils and paleoenvironments from the
Axamilpa Section reinforce the importance of understanding the reciprocal biotic and abiotic interactions
and influences.
While arid and semiarid areas in Mexico are widely
distributed and assumed to be of a more recent origin,
the new evidence provided by the well-preserved
sediments of the Axamilpa Section expands our view
on how arid environments may have looked like at
that time, their depositional evolution, and the
influence of physical factors on the local biota and
the environment.
This study may encourage more detailed studies in
the Coatzingo Formation and other Cenozoic basins in
Mexico that can provide data to further precise
timeframes of sediment deposition, reconstruct paleoenvironments, and to understand the evolution, biogeography and adaptation of organisms in tropical North
America.
Acknowledgments
We thank Dr. Ana Luisa Carreo, M.Sc. Claudia
Carranza-Sierra, Ing. Ciro Daz, Dr. Jerjes Pantoja Alor,
Ing. Diego Aparicio from the Institute of Geology,
UNAM. Sr. Flix Aranguti and his family from the

Museum of Paleontology (Tepexi de Rodrguez); Dr.

Gonzalo Pardo from the University of Zaragoza, Spain;
Dr. Jos Carlos Garca Ramos from the University of
Oviedo, Spain; the staff of the Biological Sciences and
Earth Sciences Postgraduate Department of UNAM; Dr.
Scott Bates, Dr. Robin Renaut, Dr. Blas Valero Garcs,
and Dr. Bruce Sellwood for their comments. This
research was supported by Project DGAPA (IN 208500)
to Sergio R.S. Cevallos Ferriz.
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