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Sedimentary Geology 191 (2006) 227 – 254

Sedimentology and paleoecology of an Eocene–Oligocene
alluvial–lacustrine arid system, Southern Mexico
Hugo Beraldi-Campesi a,⁎, Sergio R.S. Cevallos-Ferriz a,1 , Elena Centeno-García a,2 ,
Concepción Arenas-Abad b,3 , Luis Pedro Fernández c,4

Institute of Geology, UNAM, Ciudad Universitaria, Coyoacán, 04510, DF México
Area of Stratigraphy, Department of Earth Sciences, University of Zaragoza, E-50009 Zaragoza, Spain
Area of Stratigraphy, Department of Geology, University of Oviedo, C/ Jesús Arias de Velasco s/n, E-33005 Oviedo, Spain


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

A depositional model of the Eocene–Oligocene Coatzingo Formation in Tepexi de Rodríguez (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 sedimentological–paleontological 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: Tepexi–Coatzingo; Eocene–Oligocene; 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: (H. Beraldi-Campesi), (S.R.S. Cevallos-Ferriz), (E. Centeno-García),
(C. Arenas-Abad), (L.P. Fernández).
Tel.: +52 55 5622 4312.
Tel.: +52 55 5622 4310.
Tel.: +34 976 762 129.
Fax: +34 98 510 3103.
0037-0738/$ - see front matter © 2006 Elsevier B.V. All rights reserved.

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; Morán-Zenteno
et al., 1999). Paleontological studies have provided
references for dating rocks and have contributed to a
more comprehensive understanding of the biotic


H. Beraldi-Campesi et al. / Sedimentary Geology 191 (2006) 227–254

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 Eocene–Oligocene Coatzingo Formation, located in the Central–South 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 Tepexi–Coatzingo basin. (B) Local geology (solid line-box in map A) of the
Tepexi de Rodriguez area. Pz = Paleozoic; Mz = Mesozoic; K = Cretaceous; E = Eocene; E–O = Eocene–Oligocene; P = Pliocene–Pleistocene;
Q = Quaternary.

An overall integration of the sedimentological and paleontological data from the region is taken into consideration to further discuss relevant biotic events. 2). In general. the Cenozoic succession is covered with the Plio-Pleistocene Agua de Luna Formation (Pantoja-Alor et al. 1996). 18°36′42ʺN. 1987). biostratigraphical correlations and floral descriptions. extending its area further to the west and south of the study site (Fig. 1680 masl). 1988) based on field observations. and the upper Ahuehuetes Unit. Chigmecatitlán. birds. Local geology The geology of the Tepexi de Rodriguez area (Fig. 2003.g. proboscideans. Initially. PantojaAlor. Beraldi-Campesi et al. close to the Ahuehuetes locality. correspond to Paleogene rocks within the Tepexi–Coatzingo basin.. paleotemperatures and to improve the understanding of biogeographical patterns of North Americas' flora (Calvillo-Canadell and Cevallos-Ferriz. It crops out West of the AS (although their contact is not clearly visible). 2). The Axamilpa Section The Axamilpa Section (AS) is located along the Axamilpa River.. Ramírez-Garduño and Cevallos-Ferriz. being then at least part of the local basement of the Coatzingo Fm. An ever-growing collection of fossil plants from the Ahuehuetes locality. but no sedimentological studies that describe the type of paleolake. extensive palynologic and paleobotanic studies have pointed to an Eocene–Oligocene age for both the Pie de Vaca and Ahuehuetes Units (Calvillo-Canadell and CevallosFerriz.. has been used to estimate ages. 1999). reptiles and the skeleton of a flamingo (Cabral-Perdomo. some of them with profound evolutionary implications. stromatolites. Cabral-Perdomo. Carranza-Sierra and Martínez-Hernández. which represents the northernmost part of the Acatlán Complex (Ortega-Gutierréz et al. although they are geographically separated. which has been informally linked with the Eocene Balsas Conglomerate (Martínez-Hernández and Ramírez-Arriaga. 1999) have served as tools for dating. the metamorphic Tecomate Formation.. and by a Paleozoic basement.g. 1987. Pantoja-Alor et al. Rodríguez de la Rosa et al. the Pie de Vaca Unit was interpreted to be of Miocene to Pliocene–Pleistocene age (Buitrón and Malpica-Cruz. composed mainly of tuff and tuffaceous sandstones (Silva-Romo and GonzálezTorres. In this paper. 2. 1999). such as interchanges and endemisms. and ostracods. 2. both interpreted as fluvial to lacustrine low-energy environments. in the central– southern part of Puebla (97°55′48ʺW. 2005). Outcrops of the Coatzingo Fm in the Pie de Vaca and nearby localities have been interpreted as lacustrine in origin (Buitrón and Malpica-Cruz. Buitrón and Malpica-Cruz. Carranza-Sierra and Martínez-Hernández. The lower rocks of the Cenozoic succession correspond to a white. Later on. 3 km N of the town of Tepexi de Rodríguez.. . That Formation has been locally divided into two units: the lower Pie de Vaca Unit. composed mainly of limestones and cherty and sandy limestones. 1988). several localities (e. 2. e. is thought to underlie the Ahuehuetes Unit. Fig. one of the most wellpreserved Oligocene records in Mexico. which represents the Pie de Vaca Unit (Fig..1. 2002. 1999. that may have occurred in this area. 2001. 2000. Pantoja-Alor et al. Other fossils from the Formation include fossil wood (currently being studied). its evolution or the overall paleoenvironmental conditions that existed have previously been undertaken. The geographical limit of the Coatzingo Fm is at present uncertain. 2002. 1B) has not been studied completely. Zaragoza. 2). 1987. particularly artiodactyl (camel-like and cervid) fossil tracks. This section. 1988) in the SE part of the Tepexi area (Fig. a Cenozoic succession is underlain by Cretaceous rocks. The Coatzingo Formation The Coatzingo Formation.3. however. 1988). termed here the “Axamilpa Section” (AS). previously named Pie de Vaca Fm (Pantoja-Alor et al. and unconformably underlies the eastern part of the Ahuehuetes section. based on facies analysis of one of its most complete sections. the Tlayua Fm (Kashiyama et al. 1A). / Sedimentary Geology 191 (2006) 227–254 ferous localities (e. calcareous. clast-supported conglomerate that 229 crops out in many areas of the Tepexi–Coatzingo basin. 2002). Magallón-Puebla and Cevallos-Ferriz. 2002. in: Calvillo-Canadell and Cevallos-Ferriz. 1994a.g. 2002. Velasco de León and Cevallos-Ferriz. together with those of cat-like mammals. Finally. Martínez-Hernández and Ramírez-Arriaga. Palynomorph assemblages described from several localities in the Formation (Carranza-Sierra. and Cuayuca) have been correlated biostratigraphically (Martínez-Hernández and Ramírez-Arriaga.. we provide a paleoenvironmental reconstruction of part of the Coatzingo Formation. 1995. 1990) and the Rosario Fm.H. based upon fossil mammal tracks from the Pie de Vaca locality.

6. then correlated and combined in a single log (Fig. 1999). Microscopic features of the facies are shown in Fig. minor limestones and marls. dm. coarse sandstones and minor limestone. 2. Stratigraphy and facies of the Axamilpa Section Well-exposed horizontal strata of the AS crop out mainly in a vertical cliff (Fig. Interpretation: These conglomerates represent cohesive debris flow deposits. keeping them floating and favoring a disorganized fabric. 3 to 7 cm in diameter. Colombo. The matrix is a sand–silt mixture with a high percentage of carbonate. 7A) This facies comprises polymictic. Reading. from the base of the section to ∼ 20 m). Log 2 contains only groups b) and c). b) detrital and chemical rocks (sandstones. General stratigraphy of the regional lithologies within the Tepexi–Coatzingo basin and the position of the Coatzingo Formation Units.230 H. although some quartz and limestone clasts are also present. which form irregular beds up to 2 m thick. 4. from ∼ 20 to ∼ 36 m). matrix-supported conglomerates. interpretation also supported by regional geological studies (Silva-Romo. 2002). Beds display planar and trough m-thick. 5. / Sedimentary Geology 191 (2006) 227–254 Fig. with . Fig. Some parts are replaced and cemented with hematite. Ahuehuetes. Three stratigraphic sections were measured at sites A. and are typical of the proximal or mid parts of alluvial fans (e. 1992. and C (Fig. up to 14 cm long. 3. which commonly appear interbedded. Conglomeratic facies Matrix-supported conglomerates (Cgm. claystone and quartz. marls and scarce terrigenous. from ∼ 36 m to the top). 3A) that shows no evidence of major deformation. The localities Axamilpa Section. consisting mainly of schist. Beraldi-Campesi et al. white or reddish limestone.g. Matrix strength and buoyancy prevented large clasts to sink. B. Facies descriptions and interpretations are summarized in Table 1. These conglomerates always form on slopes. 2003. Pore spaces between clasts are commonly filled by calcite cements. Fig. Three lithological groups are recognized in the AS: a) detrital rocks (conglomerates. Clasts are angular to subrounded. 1998 in Calvillo-Canadell and Cevallos-Ferriz. Clast-supported conglomerates (Cgc. 7B–C) These consist of poorly sorted clast-supported conglomerates. Clasts are angular to subrounded. and Pie de Vaca (right) are shown in a stratigraphical context. and c) rocks mainly of chemical origin (limestones and evaporites. strata with erosive bases. Log 1). 1986). Martínez-Hernández and Ramírez-Arriaga. and composed of schists. A second section (Log 2) was measured at site D. The vertical distribution of facies and their associations is shown in Fig. This facies forms tabular. 3B).

B. 5). Log 1 was measured in sites A. (A) Panoramic view.H. of the Axamilpa Section and surroundings. / Sedimentary Geology 191 (2006) 227–254 Fig. 231 . 3. Log 2 was measured in site D (logs shown in Fig. facing NE. Beraldi-Campesi et al. and C. (B) Topographic map of the Axamilpa Section showing the sites where logs were measured.

It is interpreted to have formed from channeled and unconfined flows. Loading structures are observable at the base of the beds. Bhattacharya and Giosan. Sandstone facies Cross-stratified sandstones (Sc. Interpretation: This facies is common in fluvial systems of medium to low energy (Miall. and may contain ‘algal lumps’. 7E) This facies is formed by fine to medium (rarely coarse) sandstones arranged in cm. / Sedimentary Geology 191 (2006) 227–254 normal or inverse grading. and ripple. ostracod remains. cm. arranged in dm to m thick tabular or lenticular strata.g. parallel. 1998. Calcareous cement and hematized zones are common in this facies. separated by pale-green marl layers.g. although turbulent episodes capable of eroding the substratum could have occurred. where gravel deposits may accumulate at the base of the channels (Davis. more rarely. given their close relationship to calcareous lacustrine facies. 1992. Interpretation: The coarsening upward trend and the thinner beds towards the top are common indicators of deltaic progradation (e. show internal horizontal lamination. The common normal grading in the sandstones reflects the waning behavior of these flows. quartz and limestone grains. Fig. 7D) These consist of very coarse to fine-grained sandstones.and low-angle stratification are present in the upper sandstones. as well as gypsum crystals and chalcedony. Beraldi-Campesi et al. it is interpreted as the progradation of small deltaic lobes in distal flat areas. the existence of a water layer. resulting in brecciation and the precipitation of hematite. Fig. Given the thin nature of this facies and its stratigraphical position between rocks of chemical origin. and ripple lamination) that reflect variable energy conditions. the substrate would be exposed to desiccation and oxidation. 1983). which typically form overall coarsening-upward units. 1996. while thickness of sandstones increase. Massive or graded sandstones (Sm. unconfined floods can deposit sand sheets with a variety of structures (e. 1996). 7F–G) This facies is composed of sandstone and marl alternations. The abundance of oolites and ostracods in this facies varies along the section. may also be present. Sandstone and marl alternations (Scs. Some coarsening-upward examples of this facies may record progradation of minor deltaic-type bars. Marl layer thickness decrease toward the top. as implied by the many marly rip-up clasts. Fig. which may display load structures and marl intraclasts in their base. 1979). and may pass laterally into fine-grained conglomerates. Fig. In such settings. 7H) This facies comprises mostly fine. The bioturbation suggests the presence of interstitial organisms and. quartz and carbonate grains extensively cemented with calcite and locally with hematite. Interpretation: This kind of deposits would form in braided channels and bars in middle parts of alluvial systems (Miall. which display normal or inverse grading and cross stratification. bioturbation galleries. therefore. Horizontal-laminated and cross-stratified sandstones (Shc. Fluid escape structures are locally present. but also medium grain-size sandstones. The presence of oolites.g. mainly formed of schist.. or merely reflect migration of the coarser head of longitudinal bars on the finer-grained downcurrent tail. Kowalewska and Cohen. Sorting is . 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.232 H. peloids. flaser. showing N40° paleocurrent directions. During drought periods. alternated clay and carbonate accumulation would account for the formation of marls among the clay-rich layers. which are common in large saline and calcareous lakes fringes ( dm-thick beds. 1996). 1980). Clasts are regularly imbricated. containing clay. which are amalgamated or. Some beds pass laterally into coarse sandstone. oolites. Sandstone beds. Rust. 2003).to dm-thick beds. The separation of both lithologies is not abrupt but continuous. Sandstones contain large amounts of schist fragments and also scattered angular quartz and other rock fragments (9–30 mm in diameter). Swirydezuk et al. These beds can contain conglomeratic lenses. sometimes with cross and flaser stratification and ripple lamination. Miall. in the middle-to-distal parts of alluvial fans. Brecciation and nodulation. indicates currents or oscillation produced by wind and swell during periods of absence of terrigenous. Fluid escape structures may be evidence of contemporary seismic movements or other factors that led to sediment disturbance. During quiet periods or the weakest flows. The marls are porous and poorly indurated.

Stratigraphic logs from the Axamilpa Section. / Sedimentary Geology 191 (2006) 227–254 Fig. Beraldi-Campesi et al.H. 233–236 . 4. pp.

Fossil plants found at one level. Marzo. Coarse. ostracod remains. Deposits on floodplains of distal alluvial-fan or lacustrine delta systems. Channelled and unconfined flows in the middle to distal parts of alluvial fans. Interpretation: This facies suggest direct sedimentation of sand from waning. bioturbation and algal lumps are present. turbulent sediment-laden flows with no (or little) traction at the bed. and ostracods. Fig. with reduced terrigenous input. The presence of fluid escape structures also indicates liquefied flows that resulted from rapid flow deceleration and quick deposition. silicified oncolites.g. Nodular gypsum. nodulation. in a wide and flattened alluvial or deltaic setting. brecciation. Gypsum and calcite laminae. also present. desiccation cracks. Debris-flow deposits in proximal or middle parts of alluvial fans. Chert bands and gypsum nodules. and are either massive or have horizontal lamination. Rhythmic carbonate deposition in relatively deep. vertebrate ichnofossils. near lake margins. Palustrine setting with soil formation and carbonate deposition. Varve-like laminated limestones with flaser stratification. Planar cross-stratification. Beraldi-Campesi et al. 1979). / Sedimentary Geology 191 (2006) 227–254 237 Table 1 General description and interpretation of the Axamilpa Section facies (see text for details) Facies Description Interpretation Conglomeratic Cgm Polymictic. peloids and bioturbation galleries are found. they show horizontal lamination and scarce fluid escape structures. bacterial or algal dm-thick. Progradation of small deltaic lobes in flat marginal lacustrine areas. matrix-supported conglomerates. ripples. oolites. microbial-like lamination. internal horizontal lamination and rare fluid-escape medium-grained sandstones with horizontal and cross-stratification. pores. Subaerial exposure events. Load structures. Marginal shallow and low-energy lacustrine environments. chert and gypsum nodules. These probably arose from flow expansion at the mouth of confined conduits (channel mouths?). Fine. Net coarsening-upward intervals. although the basal part of some beds contains scattered grains up to 8 mm. Limestones in massive or stratified layers containing desiccation cracks. Marl facies Massive or laminated marls (Mm. load structures. with prolonged evaporation and depletion of water. inverse or normally fine-grained sandstones in tabular or lenticular strata. in some cases. Neighboring carbonate strata suggest deposition in a distal floodplain to marginal lacustrine setting. Alternation of calcareous marls and clay-rich layers. Braided channel and bar deposition in middle to distal parts of alluvial systems. 1992. Massive oolitic limestones with gypsum nodules and chert bands. chalcedony. where herbaceous vegetation became established. Marl layers. massive or with horizontal lamination. although. Suspended deposition (flash floods) in flat alluvial or deltaic settings. The differences in grading would reflect sedimentation from high to low density fluids (e. Sandstone and marl alternations. Periodic or continuous settling of carbonate muds in offshore. Mutti et al. chert bands and nodules. Coated grains and pisoids. low-energy lacustrine areas. ripple and low-angle cross stratification. Carbonate deposition in shallow lacustrine to palustrine settings. Beds are massive or inversely or normally graded. Stromatolites capped with gypsum. good. brecciation and bioturbation are present.g. perhaps with an eolian contribution.H. Chert bands and microcrystalline gypsum nodules up . bioturbation. and peloids. microgranular and nodular gypsum. Cgc Sandstone Sc Shc Scs Sm Marl Mm Mh Calcareous Lm Lo Ll Ls Evaporitic Gy Clast-supported conglomerates with erosive bases. Halite Intense and prolonged lake evaporation events.. Margins of saline carbonate lakes with agitated shallow waters. normal or inverse grading. rhizocretions. Load structures. peloids. Fine to medium massive sandstone. 1996). low-energy offshore lacustrine areas. fluid escape structures. The flows loaded with sand were perhaps generated by sudden floods during heavy rainstorms (e. cm. and cm-thick interbedded marl layers. vertebrate tracks. Lowe. Cross-stratification and conglomeratic lenses. 7I) These marls form tabular layers.

small anhedral gypsum crystals. 1997) and in paleosols (Davies and Gibling. Beraldi-Campesi et al. Laterally they have mm-thick varve-like horizontal and wavy laminae and no bioturbation is observable. 7J–K) Alternation of cm. mm to cm in diameter. bioturbated. 1998. radial calcite crystals. in strata varying from 60 to 70 cm in thickness. Oolitic limestones (Lo. load structures. they show partial dolomitization. Some horizons are brecciated. possibly grass-like plants adapted to basic conditions. columnar and domal stromatolites. bioturbation and hematite replacements. and show microbial-like lamination (with bacterial or algal remains preserved. vertebrate fossil tracks. ostracods. The varve-like appearance suggests that the carbonate versus terrigenous sedimentation was influenced by seasonal factors. hemispherical stromatolites. Interpretation: These marls represent periodic or continuous settling of carbonate mud in offshore. as well as peloids and bioturbation. Laminated limestones (Ll. 8A) Beige oolitic grainstones to packstones represent this facies. The scarcity of oncolites suggests that they were formed in shallower zones and transported to deeper zones. Fig. Two types of stromatolites were found at two different levels: a) large. Some beds contain microgranular and microcrystalline nodular gypsum. and pores (1 to 3 mm in diameter) are present. Likewise. Desiccation cracks. In a few places. Fungal or algal filaments are also present. Locally they show flaser stratification and contain scarce silicified oncolites. Vertebrate fossil tracks were observed at one level. 30 to 40 cm high. and b) small. Load structures. 2003. fluid escape structures. Marls and claystones (Mh. with large amounts of oolites. Fig. mm-thick laminae of carbonatecemented sandstone may be present in the marl levels. although rarely). Porosity is conspicuous with pores up to 4 mm in diameter. rhizocretions. microgranular chert. and desiccation cracks are very common (Fig. scattered terrigenous suggest eolian inputs during high level stages. 1979). low-energy lacustrine areas with little terrigenous input. as well as algal-like aggregates. Schroeder et al. peloids.5 mm to 10 cm in diameter. clays and quartz grains are seen. The poor preservation of the plant fragments implies an early diagenetic degradation. and chert bands. Kowalewska and Cohen. Some are porous and contain microcrystalline gypsum nodules. coated grains and pisoids. Limestone facies Massive limestones (Lm. Fig. and chalcedony. beige or green mudstones that form cm. continuous laterally coalescent bioherms. In thin section. Some marls show brecciation and bioturbation. mm to cm thick. 7K). which occur alone or in clusters of two or . 10 to 12 cm high. Fig. bands and nodules of chert. Interpretation: Vermiculite can be found in soils (McCarthy and Plint. / Sedimentary Geology 191 (2006) 227–254 to 1 cm are present. 0. chert and gypsum nodules.238 dm-thick layers of calcareous marls with cm-thick greenish clay-rich vermiculite layers. 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. 8E–F) Stromatolites appear associated with facies Ll and Lm. A shallow or fringing lacustrine environment with agitated water that promoted the formation of oolites is inferred for this dm-thick tabular layers. This may imply also changes in the lake level. Gypsum nodules may have formed during periods of evaporation. Fossil plants were found at one level. They are typically partially silicified. Stromatolitic limestones (Ls.g. Thin. Ripples may be present locally.. Fig. which form abundant. Replaced and fragmentary ostracod valves. The vermiculite forms layers 1–3 cm thick that rhythmically interrupt the marls. The surfaces of these rocks are commonly replaced by hematite. complete and broken oolites. Interpretation: These deposits are typical of those that form along the edges of saline carbonate lakes (e. 2003). 7L) This facies consists of white-yellowish. Swirydezuk et al. and thus the lake area. Interpretation: These deposits may have formed in relatively deep and low-energy lacustrine offshore areas by carbonate deposition and variable terrigenous inputs. and peloids occur but are uncommon. indicating that soils may have formed in a palustrine setting where carbonate deposition occurred during rainy seasons with high lake levels. 8B–D) This facies is represented by dm-thick tabular strata of mudstones..

Beraldi-Campesi et al. 5. See text for explanation of facies associations. / Sedimentary Geology 191 (2006) 227–254 Fig. pp. Facies distribution in the Axamilpa Section. 239–242 .H.

dark grey). Beraldi-Campesi et al. 6. (H) Bioturbated microbial-like lamination. (G) Brecciated zone with intraclasts (I). (D) Radial growing of diagenetic calcite. black) and nodules (N. (B) Nucleated oolites with chalcedony crystals (C). (I) Partially dolomitized limestone clast. (E) Rhizoidal pores (P. Microscopic features in thin section. (F) Rhizoid pore with inner concentric layers of calcite. (J) Echinoderm fragment (E) along with quartz grains (Q) and pelloidal limestone fragments (P). / Sedimentary Geology 191 (2006) 227–254 243 Fig.H. (C) Large gypsum crystal (G) within a micritic matrix. . (A) Pellets within a spar matrix. with a miliolid foraminifer (arrow).

/ Sedimentary Geology 191 (2006) 227–254 . Beraldi-Campesi et al.244 H.

(E) Fine lamination of facies Shc. (G) Load structures of the facies Scs. (J) Facies Mh showing marl (M. / Sedimentary Geology 191 (2006) 227–254 245 Fig. (E) Large stromatolites (facies Ls). and the regular morphology of the structures suggest that relatively stable conditions Fig. three domes. showing poor sorted floating clasts. (D) Cross-stratification of the facies Sc. The stromatolitic lamination shows the characteristic dark/light-lamina alternation (Reid et al. dark). (C) Close-up of the facies Ll showing compacted mm lamina. (B) Facies Ll containing gypsum nodules (arrows). and Ll (∼25 m in log 1)... Interpretation: These facies formed in low energy. (I) Facies Mm (arrows) in Log 1 at ∼ 25 m (scale = 1 m). Beraldi-Campesi et al. with the lighter laminae being thicker and dominant. (D) Sequence of the facies Mm. Their continuous and smooth lamination. (K) Flame structures of the facies Mh where thin sandstone layers occur. Lm. Seong-Joo et al. 8. The matrix between the stromatolites is massive or laminated carbonate. (H) Partially dissolved and collapsed stratum of gypsum (facies Gy) with calcite laminae. (B) Erosive contact between facies Cgc (above) and facies Cgm (below). 7. Both stromatolitic intervals are noteworthy capped by gypsum in honeycomb-like frameworks. (F) Small isolated stromatolites (facies Ls). hammer – encircled – is scale). (G) Transversal cut of a small stromatolite showing the outside and its internal lamination. Lo. (H) Facies Sm. 2000. (F) Facies Scs lamination. light) and claystone (C. . (C) Facies association A in log 1 (∼ 12 m) showing two cycles (corks). (L) General view of Lm facies in site D (∼11 m. (A) General view of the Facies Lm and Lo next to a rupestrian painting. (I) Intricate growth of gypsum crystals. 2000). Hammer –encircled – is scale. Partial silicification is present toward the centre in both types. Lm. (A) Close-up of the facies Cgm.H. shallow and marginal lake environments.

microcrystalline and macrocrystalline textures are observable. Decreasing terrigenous inputs allowed deposition of massive limestone facies in shallow or even fringing lacustrine conditions. Thus. probably as a complex and recurrent sequence of processes (Arenas et al. Lo.246 H.. Finer sediment reached distal zones of the fan. Diatoms have not been detected and thus they can likely be ruled out as a major source of silica. shallower lacustrine conditions gave place to . overlain by coarser sandstones that grade into conglomerates towards the top..g. Fig. giving rise to small deltaic systems. Marls are evidence of a water body with carbonate deposition. 3. 1999). 2003). Facies that are inferred to be genetically linked have been grouped. causing dissolution and collapse of the evaporite layers. in which sheet-like flows deposited part of the detrital sediments as facies Shc. Sc → Mm → Lm. indicate alluvial fans of small size. with planar contacts and a fining-upward pattern. Morán-Zenteno et al. followed by massive limestones that are up to 35 cm thick. The limited presence of stromatolites in the AS suggests that favorable conditions for their growth were not frequent. 1980. although macrocrystalline gypsum is more common in irregular beds. where silicification takes place in littoral and eulittoral sequences of lacustrine carbonates. Pirajno and Grey. The extensive gypsum on their upper surfaces suggest alkaline conditions with periods of prolonged evaporation. In other cases. Although microbial mats have also been claimed as a source of silica in lacustrine deposits (Bustillo et al. 2002). Eugster. C association: (Shc). Thin and broken carbonate layers and halite casts are also present. Microgranular. They are clearly diagenetic features and are found replacing some of the primary components of the deposits (e. and their close relationship with lacustrine facies. Facies associations The facies described above are associated in vertical sequences that represent the superposition of different subenvironments through time. The sequence represents the retreat of an alluvial system and the establishment of lacustrine and palustrine environments.2. B association: Scs → Shc. In some cases. either massive or interbedded with marls. beginning at the base with sandstone facies. Features of the alluvial-fan conglomerates. can be debatable. Sm. Silicification processes Many of the facies above described contain chert nodules and bands and chalcedony. It comprises mainly clastic facies with thicknesses from 30 to 250 cm. 6) can be interpreted as a sequence of events that describe the sedimentary evolution of the section.1. 2000. and caused offshore marl deposition (facies Mm). This association records the progradation of the alluvial system by channeled flows that entered a shallow lake area. Dunagan and Turner. Interpretation: This facies represents increasing solute concentration in the lake and sediment pore water due to lowering of the water table and intense evaporation. silicification may have occurred during early diagenesis by the mixing of meteoric waters with the evaporite pore waters. varying from 4 to 70 cm thick. Sm → Shc. Evaporitic facies Gypsum (Gy. such as the clast size (8–14 cm). Sm → Sc →→ Cgc This comprises a clastic facies with thickness from 50 to 250 cm and coarsening upward evolution.g. tabular to very irregular beds of dissolved and collapsed gypsum. as it has been seen in other basins (e. stromatolites. 3. / Sedimentary Geology 191 (2006) 227–254 prevailed during their formation. oncolites.. 8G–H) This facies consists of dm-thick. It begins with fine detrital inputs that induced a subsequent rise of the water table. A association: Cgc → Sc. This association occurs cyclically and suggests gravel and sand deposition in braided bar and channel systems of alluvial fans that reached lacustrine areas. calcite cementation and silicification. Ll This consists of clastic and carbonate facies. 1999) would have accounted for the silica input. The source of silica. hydrothermal pulses may have loaded the pores of the preexistent rocks with the silica that would later precipitate as chert. it is more likely that the extensive magmatic activity that prevailed in this region during the Paleogene (Martiny et al. the paucity of microbial deposits in the AS argues against microbial mats being a major source for silica. mostly with active fluvial-dominated sectors in relatively steep areas. Paleocurrents inferred from the conglomerate clast imbrication suggest that the source of alluvial sediments was located to the SW of the study area. Mm → Lm This fining upward sequence begins with erosive or planar contacts. Beraldi-Campesi et al. the local debris-flow events.. as in many silicified deposits elsewhere. 2004. Later. and their vertical associations (Fig. with their apices close to the palustrine and lacustrine areas that existed down stream. limestones). Lo.

which is found in deformed. It continues with carbonate precipitation in either relatively deep (facies Ll) or shallower waters (facies Lo. Fossil tracks with similar characteristics have been described from other localities and related to the Camelidae group (Cabral-Perdomo. Commonly the strata are tabular. 1995). 1) and some 3 km E from . The form of the digits is typical of ungulates (artiodactyls). and would mark the establishment of lacustrine. Ls → Gy This is a complex association formed of cm. although some plant remains resemble legumes and aquatic plants in morphology. Coquinas with silicified ostracods have been observed at the Zaragoza locality (Fig. 9J–K) Scarce ellipsoidal oncolites. These are small (up to 10 mm in length).to dmthick claystone. marl. Leaves: (Fig. 9. The traces are usually oriented in a same direction. relicts of roots (rhizocretions) were observed. as assumed for those of the AS. and mostly appearing as vertical traces. a relative deepening and the beginning of a new sequence. Lm) in which palustrine conditions were present. is constant along the stratigraphic section. 9L) In some strata. Facies Ll could represent a slightly deeper lake setting with episodic carbonate deposition. tracks of at least 4 steps can be easily distinguished. there are many other possible burrowing candidates. Facies Lo and Ls would also imply shallow. probably shallow. Recognized genera include Pseudosmodingium and Pistacia from the Anacardiaceae and Cedrelospermum from the Ulmaceae. both fragmental and complete specimens. 3. and Mh → Lm → (Ll) → Lo → Gy. carbonate lacustrine conditions in marginal areas affected by waves (Lo) or in calm areas (Ls). for example. Ostracods are present either as conjoined or disarticulated valves. were found in limestones (Facies Ll). Fossil record of the Axamilpa Section Stratigraphic appearance of the Axamilpa Section fossils is indicated in Fig. The shape and size of the valves. 1990). and gypsum nodules to form in earlier carbonate facies. may have caused dilution of the lake waters. in several levels of the section. but probably more saline. However. Oncolites: (Fig. Beraldi-Campesi et al.3. They are moderately well preserved carbonaceous imprints. They do not display ornamentation. which indicates cyclic water level changes attributed to deepening–shallowing events.H. suggesting different consistency and water saturation of the substrate at the time of the imprint. The sequence represents an overall shallowing that implies the change from lacustrine–palustrine to evaporitic conditions. nodulation. followed by deepening–shallowing lacustrine events. whereas the external surface remains unsilicified. and it is possible that they contributed to the bioturbation. / Sedimentary Geology 191 (2006) 227–254 carbonate deposition in marginal areas with or without agitation (facies Lo and Lm. respectively). Lm → Lo. Two of the associations that reflect that evolution are: Mm → Ll. 9D–I) At least six different types of leaves were recognized in the facies Mh. 6). oxidized (inferred by the color). Sometimes ostracods are found in the same sediments. Rhizocretions: (Fig. Mh → Ll. 1990). The remaining material has not been identified due to the absence of diagnostic characters. although they may appear randomly arranged. Two of the most common sequences of this shallowing process are D1: Mm → Ll → Lm. which is common in non-marine ostracods (Henderson. and D2: Mh → Ll. They are abundant regionally. 9A–B) All the tracks observed in the AS were found on top of exposed bedding planes at various levels of the section. being quite smooth. 9M–N) These are present in several horizons of the succession. 247 Although footprints are not always well marked. Their concentric lamination may be obliterated by silicification. This association reflects the end of the alluvial– deltaic dominance. Vertebrate fossil tracks: (Fig. Foot lengths vary from 10 to 18 cm. It begins with deposition in low-energy offshore areas as a consequence of a lake deepening and expansion. D3: Mm. Bioturbation ‘galleries’: (Fig. the tracks are better defined than in others. D association: Mm. followed by high rates of evaporation and desiccation that formed facies Gy in very shallow ponds. between 400 and 550 μm in length. always in low-energy lacustrine marls and limestones. Ls → Gy. except for the gypsum. due to supersaturation of the lake by increased evaporation that caused sulfate deposition. Ostracods: (Fig. collapsed and partially dissolved strata. Representative specimens are shown in Fig. calcareous and evaporitic facies. given the benthic and excavator habits of some species (Henderson. carbonate depositional conditions after an initial deepening event. Rises in the lake water level due to freshwater inputs. 9C) Galleries are observed in thin laminae. Facies Lm and Ll can be found alternating through time. 3 to 5 cm in diameter. They are usually found displaying alternated right and left steps. 4. suggesting stability in diversity of species. Mh → Lm → Ll → Lm (Fig. complete or fragmented. which is most intense toward the center. In some strata. Lm. and also caused brecciation.

Beraldi-Campesi et al. / Sedimentary Geology 191 (2006) 227–254 .248 H.

the similarity between faunas may indicate a relationship between the water bodies. 10.H. (B) Shape of an ichnite delimited by a draw. (F) Poorly preserved fossil leaf of Pistacia sp. (G) Fossil leaf of Pseudosmodingium sp. The transitional stage records the influence of both the alluvial and the lacustrine depositional environments. (C) Lacustrine stage. Microorganisms: (Fig. and finally 4) evaporitic (Fig. In the studied succession this can be conceived as four main depositional stages: 1) alluvial–fluvial (Fig. 4. 2) transitional (Fig. intermittent lacustrine settings. 3) lacustrine (Fig. In that case. characterized by braided channel and bar deposits with rare debris flows. . / Sedimentary Geology 191 (2006) 227–254 249 Fig. (K) Transversal cut of a partially silicified oncolite (white calcite coat). 0–20 m). (I) Fossil leaf of a possible aquatic plant. 10A). it. (D) Evaporitic stage. (H) Fossil impression of a legume. characterized mainly by detrital facies (conglomerates. Facies Ls). 10B). followed by expansion of a carbonate deposi- tional lacustrine system that eventually evolved to a more shallower and alkaline. Fig. Sedimentary evolution proposed for the Axamilpa Section (see text for explanation). The alluvial–fluvial stage. (J) Oncolites in the field. characterized by marls. (O) Permineralized microorganisms. wrapped by a mucilage-like sheath. This evolution reflects an overall retrogradation of an alluvial–fluvial system. 9O–P) Permineralized microfossils were observed in thin sections of chert samples. The diameter of each cell varies from 5 to 7 μm. (N) Transversal cut of an ostracod with non-symmetric valves. suggesting that one or more lakes existed at that time. characterized by carbonates. sandstones and mudstones. 36–55 m). The diameter of the groups varies from 10 to 15 μm and they are arranged in small colonies (6 to 20 individuals per colony). 10C). 20–36 m). (L) Rhizoid impressions (scale = 1 cm). 9. Stromatolites: (see Stromatolitic limestones. (D) Fossil leaves of Cedrelospermum sp. (A) Track of vertebrate ichnites. sandstones and limestones (alternating distal alluvial and carbonate lacustrine facies. as indicated by limestones and marls that cap the retrograde cycles. 10D. Evolution of the sedimentary system The Axamilpa Section clearly shows a fining-upward trend from alluvial conglomerates and sandstones to lacustrine carbonates and evaporites. (M) Transversal view of an ostracod showing substituted internal remains. overlapping the alluvial domain. Beraldi-Campesi et al. (E) Carbonaceous impression of an unidentified leaflet. They appear in groups of four cells. (C) Parallel bioturbation galleries. that gave rise to the expansion of palustrine and lake environments northeastward. (A) Alluvial–fluvial stage. experienced a general retrogradation through time (associations A and B). (B) Transitional stage. (P) Individual microbial cells.

B. 1996). inferred as the retrogradation of an alluvial system.. The Salton Sea in southern California (Arnal. is probably the result of the basin extension. 1999). Olson. nodulation. with the cessation of the alluvial influence. Finally. 1983) is a saline lake with scarce peripheral vegetation. it is possible that the discovery of this single flamingo skeleton represents a larger population. extensive faulting and block displacements (e. as inferred from desiccation cracks. and size of the water bodies must have changed substantially over time. 1992.g.250 H. 1996). The decreasing depth. Because extant flamingos are gregarious and migratory (Nager et al. 1961). scarce rainfalls and high evaporation. thus lending support to paleoecological inferences. Beraldi-Campesi et al. but discontinuous. Ls. For example. the influence of meteoric waters may have caused the dissolution and collapse of the evaporitic layers. giving rise to ephemeral and shallow saline lakes. Mm. associations A. Extensive plains are characteristic of these environments. Flamingos (Phoenicopteridae) have been described from many Eocene and Oligocene strata (Martin. 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. rhizocretions. and Mh developed (sequence D). Oolites. gypsum) would have concentrated in the lake and formed early diagenetic features (e. 1996). the geologic setting indicates the existence of shallow lacustrine environments. Lo. could represent the existence of magmatic activity in the subsurface. The common. maybe related to the global climatic changes that occurred in that epoch (Frakes et al. and sedimentary features of the AS rocks. but their presence in the Coatzingo Formation contributes to the understanding of the past habitats and ecological requirements of these birds. the frequency of flow-generated deposits appears to decrease starting from the transitional stage. Hydrothermal pulses. 5.. as shown by the different lithofacies. Similarly. whose intermittency appears to be more frequent as the environment became dryer. and the vegetation is low and open (Kowalewska and Cohen. 2002) prevailed in this area of the Coatzingo Formation. stromatolites. As in modern analogs. The depth.g. changing topography and drainage patterns. such as Cedrelospermum. Further work is needed to correlate precisely all the localities where avian ichnofossils are present. Wolfe. D) was probably related either to tectonic and faulting episodes. / Sedimentary Geology 191 (2006) 227–254 with alluvial inputs in the form of lacustrine deltaic lobes and distal-fan sheet-like deposits (associations B and C). Fossil plants from the Axamilpa Section. This would imply a progressive lowering of the lake level. and ever-smaller water bodies that represent the lacustrine and evaporitic stages. such as rain frequency and intensity. appearance of ostracods along the Axamilpa Section supports the existence of intermittent water bodies. While subjected to intense evaporation. For example.. Ll. It has hydrothermal activity associated with rifting (Harmon. which is reinforced by the presence of a flamingo skeleton in the Pie de Vaca locality (CabralPerdomo. During early diagenesis. fossil content. Although a timeframe for these events is difficult to assess. 1998). This suggests that environmental conditions (shallow and saline waters) similar to those known from where these birds live today (Mascitti and Bonaventura. carbonate and evaporitic deposits.. Morán-Zenteno et al. salinity. such as extrusive and intrusive magmatism. ostracods and migratory birds are common. Gilmore and Castle. sulfates (e.. seem to reflect a change towards more arid conditions. brecciation. as well as their temporal and spatial distribution during the Eocene–Oligocene. Paleoecology The fossil record known from the region correlates well with the sedimentological data. Martiny et al. These lakes have very shallow and even intermittent waters. 1961. gypsum nodules) in the exposed mud flats. 2001).. 1985). Both ancient and present-day hydrothermalism processes have been reported for this area (Carballido-Sánchez and Delgado-Argote. and fenestral porosity present in facies Lm. must have determined sedimentation and biota distribution in the Coatzingo Fm to a great extent. regional events that were taking place simultaneously in central Mexico during the Paleogene. and facies Lm.g.. deduced from the occurrence of magnesite and cherts in the AS and in nearby localities. are . favoring the development of palustrine conditions around a lacustrine water body. Jiménez-Suárez et al. with high slopes or mountains relatively close to the lake environment. along with local factors. or to climatic variations. This notion may also be supported by the fact that the fossil record of aquatic birds in Tepexi de Rodríguez is composed mainly of ichnofossils of their tracks (Cabral-Perdomo. carbonate lacustrine environments became extensive. 1983. The fining-upward evolution of the sequence. 1989. 1966) and combines fluvial and lacustrine sedimentary processes (Arnal. 2000. the cyclical deposition of some facies (e. Great Salt Lake. increasing salinity. Utah. is a shallow calcareous-evaporitic lake (∼ 10 m) influenced along its margins by alluvial systems. 1994).g. Pistacia and Pseudosmodingium.

contrasting lowland xeric and upland mesic vegetations. Given the stratigraphical position of the two localities (Fig. and shelter. Beraldi-Campesi et al. (Carranza-Sierra. Perhaps the Tepexi–Coatzingo basin served as a migratory path for these animals. suggesting a long history of exchange between the North American and Eurasian floras (Ramírez and Cevallos-Ferriz. furthermore. would indicate a grass-like vegetation relatively far from the waterways where the riparian communities became established. shade.. as well as a place to drink. food. so riparian settings would be suitable places for taphonomical processes. and one to South America. may indicate that near-river environments became established more than once in the basin. and serve as spots for endemisms. 2000. and although the Axamilpa Section lacks volcanic components. 2005). 1994b. 2000). however. one to Eurasia and Africa. Stromberg et al. 1991).. along with Cedrelospermum. Cedrelospermum has been referred to subhumid climates in communities of desert scrub (Magallón-Puebla and Cevallos-Ferris. as in modern saline-lake environments (e.. especially in desert springs. This fact agrees with previous assumptions of grass-abundant biomes for this area (Martínez-Hernández and Ramírez-Arriaga. Plant-bearing strata in the Ahuehuetes Unit are composed of volcanic ash. Pollen assemblages also suggest that the mountain ranges that surrounded the basin were populated mostly by conifers (MartínezHernández and Ramírez-Arriaga. 1991). 2000). Camelids originated in North America during the Paleocene–Eocene in North America (Stearn and Carrol. Riparian zones favor plant dispersal and mixing of seeds (Stromberg et al. In the Axamilpa Section. Perhaps near-river plants had a higher potential for fossilization due to their susceptibility of being quickly buried after flooding events (Stromberg et al. 251 2003).e.. which today exists in the Tepexi de Rodríguez area. Velasco de León and Cevallos-Ferriz. .H. where annual precipitation was less than annual evaporation. and may narrow the existence or riparian environments to only a few places. 1999). 2000) might have caused environmental disruption in the surrounding areas. feed or breed. Soria et al. have been proposed for the Miocene and Pliocene–Pleistocene respectively (van der Made et al. 2002). and where there were long dry periods) (Ramírez and Cevallos-Ferriz. which in turn probably influenced the origin and distribution of plants in this region. / Sedimentary Geology 191 (2006) 227–254 also found in the Ahuehuetes Unit. which suggests similar climatic conditions prevailed in this region during the Oligocene.. is considered a pioneer in harsh environments (Ramírez and CevallosFerriz. The distinction between riparian and non-riparian plants may have implications for their distribution and taphonomy. the major volcanism that occurred in this epoch (Martiny et al. The presence of riparian-related plants from the Ahuehuetes Unit... and 2 events of dispersion. 1999. personal communication. These systems may also serve as dispersal ways for plant and animal species (Baker. The Pseudosmodingium species. 2002). so diversity of fossil plants in the Coatzingo Formation may be an indicative of riparian environments. 1A).. 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 Rodríguez region (Fig. 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. 1989). and deciduous scrub or chaparral-like communities are thought to have established there in temperate to semiarid areas (i. In that sense. the presence of Graminidites sp. Their association with desiccation cracks and rhizocretions in the AS suggests animal movement across emergent areas. Populus and some Anacardiaceae (Ramírez-Garduño. would have been an attraction for animals and may have helped as corridors for their dispersal within arid zones. resources such as drinking water. 2002). estimate annual average temperatures of 10–18 °C (Velasco de León. 2000. 1991). 1999).g. 1999). Moreover. higher plant diversity in arid lands is found around the streams (Ali et al. 1998. they are perhaps an indication of their dispersion process. however. 2). Kowalewska and Cohen.. The fact that riparian ecosystems may have existed in this region would have profound paleobiological implications. together with the evidence for high salinity and aridity. Silva-Romo et al. and high number of endemisms can be found there (Stohlgren et al.. thermometric inferences from the Ahuehuetes locality fossil plants. 2003) and fossil roots. these environments could have had an influence on the biodiversity and distribution of plants and animals. the fossil record of this group is incomplete and the starting points of these migrations are unknown. such as Salix. Furthermore. is thought to be endemic to this region and. Furthermore. which are also represented in the AS. 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. although resources from streams would have been their main attraction. 2002). 1986). Morán-Zenteno et al. The tracks indicate that these animals traversed the area regularly.

Facultad de Ciencias. UNAM. 262. Gonzalo Pardo from the University of Zaragoza. Mexico. Ecology of Biological Invasions of North America and Hawaii. B. Conclusions While a wide variety of habitats exist today in this region of the globe.. Scott Bates.E. Ana Luisa Carreño. Mex. Cabral-Perdomo. Dedolomitization and other early diagenetic processes in Miocene lacustrine deposits. California. Drake.g.. Jerjes Pantoja Alor. climáticas y cronoestratigráficas (Municipios de Zacapala and Coatzingo.252 H. Alonso-Zarza. L. where life forms have existed and evolved.. 427–478. Arnal. Estado de Puebla – interpretación preliminar de su emplazamiento. M. Cevallos-Ferriz. 25A. the staff of the Biological Sciences and Earth Sciences Postgraduate Department of UNAM. Sr. or communities.. References Ali. Baker. 24 pp. Geología del cuerpo serpentinítico de Tehuitzingo. 2003. Pardo. and gives organisms. fossils and paleoenvironments from the Axamilpa Section reinforce the importance of understanding the reciprocal biotic and abiotic interactions and influences. 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.M. and microorganisms of the Salton Sea. G. Beraldi-Campesi et al.. Ing. Robin Renaut. sedimentation. Dr. 1986. Revista del Instituto de Geología 8 (2).. C. Carranza-Sierra. and to understand the evolution. H. Madrid Basin. Dr. New York. E. E. and Dr. . 2002..G. M. pp. Spain). M. Geological Society of America Bulletin 72 (3). 1999. una localidad fosilífera famosa de México: I... G. which offers a more dynamic understanding of the natural processes. L. Interpreting past life and the places where they lived opens the opportunity to understand particular biological or geological phenomena.A. 1987. Malpica-Cruz. Ciro Díaz.. M. Sociedad Mexicana de Paleontología..).A. M. México. Blas Valero Garcés. 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. Puebla.E. 1995. Palynology 26. 23–45. J. (Eds. an Oligocene plant from Tepexi de Rodríguez. fossil tracks. Cabral-Perdomo. Claudia Carranza-Sierra. and the influence of physical factors on the local biota and the environment. 1989. Dr. plants) in various levels of the Coatzingo Fm indicate their permanence over time. Caesalpinieae).. Calvillo-Canadell. biogeography and adaptation of organisms in tropical North America. 44–57. Patterns of plant invasion in North America... K. Estado de Puebla).. 2002. L. Puebla State. Félix Aranguti and his family from the Museum of Paleontology (Tepexi de Rodríguez). little is known about these diverse scenarios during the Cenozoic. Arenas.. M. C. Carballido-Sánchez.Sc. 2000. Springer-Verlag..A. While arid and semiarid areas in Mexico are widely distributed and assumed to be of a more recent origin. near Tepexi de Rodríguez. ostracods.. Bhattacharya.. Universidad Nacional Autónoma de México. Bustillo. R. C.A. Bachelor's Thesis. S. Implicaciones paleoambientales. Libreto Guía de la Excursión. Zacapala and Coatzingo municipalities.S. Review of Palaeobotany and Palynology 122 (3–4). et nov.. A flamenco's tafogliph from the Pie de Vaca Formation (late Cenozoic). H. Estado de Puebla. the recurrence of some of the taxa (e. Spain.S. 2003..M. Congreso Nacional de Paleontología. and suggests that either the environmental conditions necessary for their development remained stable or that they adapted to the changing environments. R. Wave-influenced deltas: geomorphological implications for facies reconstruction.. 6. / Sedimentary Geology 191 (2006) 227–254 Finally. their depositional evolution. Tepexi de Rodríguez.. 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. José Carlos García Ramos from the University of Oviedo. reconstruct paleoenvironments. Bruce Sellwood for their comments. Arribas. Puebla. UNAM. Martínez-Hernández.A. 187–210. Sedimentology 50. a context in which they were able to evolve.. Limnology. In: Mooney. Palinoestratigrafía del Grupo Balsas. Diego Aparicio from the Institute of Geology. Cevallos Ferriz. 1961. Moreover. Dr. Acknowledgments We thank Dr. Buitrón. 107–126. Bachelor's Thesis. Bustillo. Predictors of plant diversity in a hyperarid desert wadi ecosystem. Sedimentary Geology 151. J. Balsas Group palynostratigraphy: palaeoenvironmental and chronostratigraphic implications. with leaf architecture similar to Bauhinia and Cercis. Ebro Basin (Spain). 2001. Sedimentary Geology 125. Dr. Journal of Arid Environments 45. A. Ing. Spain.J. 134–148. 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