Marine connections of Amazonia: Evidence from foraminifera and dinoflag-
ellate cysts (early to middle Miocene, Colombia/Peru)

M. Boonstra, M.I.F. Ramos, E.I. Lammertsma, P.O. Antoine, C. Hoorn

PII: S0031-0182(14)00547-1
DOI: doi: 10.1016/j.palaeo.2014.10.032
Reference: PALAEO 7069

To appear in: Palaeogeography, Palaeoclimatology, Palaeoecology

Received date: 9 July 2014

Revised date: 20 October 2014
Accepted date: 22 October 2014

Please cite this article as: Boonstra, M., Ramos, M.I.F., Lammertsma, E.I., Antoine,
P.O., Hoorn, C., Marine connections of Amazonia: Evidence from foraminifera and di-
noflagellate cysts (early to middle Miocene, Colombia/Peru), Palaeogeography, Palaeocli-
matology, Palaeoecology (2014), doi: 10.1016/j.palaeo.2014.10.032

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 ACCEPTED MANUSCRIPT

Marine connections of Amazonia: Evidence from foraminifera and

dinoflagellate cysts (early to middle Miocene, Colombia/Peru)

Boonstra, M. 1, Ramos, M.I.F. 2, Lammertsma, E.I. 1, Antoine, P.-O.3, and Hoorn, C. 1*

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Paleoecology and Landscape Ecology, Institute for Biodiversity and Ecosystem Dynamics

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(IBED), University of Amsterdam, P.O. Box 94248, 1090 GE Amsterdam, The Netherlands
2
Museu Paraense Emílio Goeldi, Campus de Pesquisa, CCTE, Av. Perimetral, 1901, CEP

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66077-830, Belém, PA, Brazil

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Institut des Sciences de l’Évolution, CC064, CNRS, IRD, Université Montpellier 2, Place
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Eugène Bataillon, F-34095 Montpellier, France.
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*Corresponding author e-mail: M.C.Hoorn@uva.nl

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Abstract
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Species composition in the present-day Amazonian heartland has an imprint of past marine

influence. The exact nature, timing and extent of this marine influence, however, are largely

unresolved. Here we use calcareous tests of foraminifera and marine palynomorphs from

Miocene sediments in northwestern Amazonia to extend on current estimates for salinity

ranges, paleoenvironments and paleogeography. Our samples mostly contain tests and/or

organic linings of euryhaline (tolerant to a wide range of salinity) foraminifera of the genera

Ammonia, Trochammina and Elphidium, with Ammonia being by far the dominant genus at

all locations. Organic-walled dinoflagellate cysts (dinocysts), such as Spiniferites spp.,

Polysphaeridium zoharyi and Tuberculodinium vancampoae, are also common at a number of

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sites. This association of foraminifera and dinocyst taxa points at varying salinities, with

aberrant forms of Ammonia indicating lower limits of 0 -10 psu (practical salinity units)

whereas dinocyst associations suggest more marine conditions. Such regional heterogeneity is

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common at the interface of shallow marine to freshwater environments, like estuaries. We

conclude that during the early and middle Miocene marginal marine conditions reached at

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least 2000 km inland from the Caribbean portal. Global high sea level and fast subsidence in

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the sub-Andean zone are thought to be the controlling mechanism of the marine incursions.

Lowering of global sea level and a change in tectonic regime terminated the incursions in the

course of the Plio-Pleistocene.

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Keywords: Amazonia, Pebas, marginal marine, Neogene, micropaleontology
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1. Introduction
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Amazonia is one of the most species-rich areas on Earth and the origins of this biodiversity
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are widely debated. A relatively little explored speciation driver are past marine incursions
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into the Amazon region, which were recognized based on a wide range of marine evidence

reported in the Neogene Solimões and Pebas formations (laterally equivalent in Brazil and

Peru-Colombia, respectively; Hoorn, 1994a, b). The importance and inland extent of these

marine incursions into South America have been cause for research ever since Ihering first

proposed the “Arm of Tethys”, a marine connection between northern and southern South

America (1927; see Boltovskoy 1991). Although Miocene marine deposits are documented in

both southern and northern continental South America there probably never existed a direct

connection (Hovikoski et al., 2007).

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Nores (1999, 2004) suggested two marine incursions from the Atlantic Ocean created

biological refugia, which triggered speciation bursts after sea level dropped. Furthermore,

Higgins et al. (2011) showed a close relation between high diversity and the Neogene

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Solimões and Pebas formations. Marine incursions may also have affected the composition

and evolution of the fish fauna (Monsch, 1998; Hubert and Renno, 2006; Lovejoy et al., 1998;

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2006; Cooke et al., 2012) and led to allopatric speciation of invertebrates such as ostracods

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(e.g., within the genus Cyprideis; Muñoz-Torres et al., 1998; Whatley et al., 1998) and

mollusks (e.g., Wesselingh, 2006; Wesselingh and Ramos, 2010). Accordingly, Amazonian

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river dolphins diversified from the middle-late Miocene (Hamilton et al., 2001; Bianucci et
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al., 2013). This all forms an important motivation to further study the nature of the marine

incursions.
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Paleontological data indicative for marine incursions into Amazonia were first reported by
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Gabb (1869). Subsequent paleontological, palynological, sedimentological, and geochemical

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data extended on these and other findings (e.g. Sheppard and Bate, 1980; Nuttall, 1990;

Hoorn, 1993, 1994a, b; Vonhof et al., 1998; Kaandorp et al., 2003; Hovikoski et al. 2005,
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2007, 2010; Lovejoy et al., 2006, Wesselingh et al., 2002, Wesselingh and Ramos, 2010;

Linhares et al., 2011). Based on this research, the inland aquatic environment was considered

to be brackish and most comparable to an estuarine environment (Hoorn, 1994b; Vonhof et

al., 2003; Wesselingh et al., 2002; Hovikoski et al., 2007).

Marine palynomorphs such as organic linings of the inner wall of foraminifera, organic-

walled dinoflagellate cysts (dinocysts from here onwards) and mangrove pollen were found in

Miocene sediments in Colombia (Hoorn, 1993; 1994a, b; 2006). Linhares et al. (2011) first

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reported calcareous tests of planktonic foraminifera, besides other marine groups such as

bryozoa, mollusks (Melongena woodwardi) and otoliths, in a borehole section in western

Brazil. With this recent finding the question arose whether beds enriched with marine

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organisms represented events or more prolonged marine conditions, and whether salinity was

higher than previously thought (~1-5 psu)(Vonhof et al., 2003).

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In this study we revisit samples collected by Hoorn in 1988-89, Hoorn and Wesselingh in

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1991, and Vonhof et al. (2003),. We document the foraminiferal and dinocyst findings,

provide illustrations of the common taxa, review a selection of sites in northern South
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America (east of the Andes) where marine influence is reported (Fig. 1), and present a revised

interpretation.
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2. Geological background and study sites

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The Neogene history of western Amazonia is strongly influenced by Andes exhumation and

subsidence in the sub-Andean region, while coinciding with global sea level changes. This
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resulted in fluvio-lacustrine conditions with episodic marine influence at the interface of the

Andes and Amazonia (Hoorn, 1993; Wesselingh and Salo, 2006.; Hoorn et al., 2010a)(Fig. 3).

Amazonia’s sedimentary history is, to a large degree, recorded in the Pebas and Solimões

formations, which extend over c. one million square kilometers across the Colombian,

Peruvian and Brazilian Amazon region (Wesselingh et al., 2002). Evidence for marine

conditions was found in sedimentary successions of both Andean origin (e.g. Hoorn, 1993,

1994b; Hovikoski et al., 2005, 2010; Wesselingh et al., 2002; Linhares et al., 2011) and

Amazonian (cratonic) origin (Hoorn, 1994a, 2006; Vonhof et al., 1998). The latter were

deposited prior to, and contemporaneously with the vast extension of the Andes sedimentary

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wedge. Here we will review the most characteristic sites and their lithological properties.

2.1. Site descriptions

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Cratonic fluvial sediments, informally known as Apaporis Sand Unit and Mariñame Sand

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Unit, respectively outcrop along the Apaporis and Caquetá rivers. Marine palynomorphs were

found in samples collected at the key sections of Mariñame and Tres Islas, situated along the

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Caquetá River (sites 1-4; Table 1; Fig. 1-2). The sediments at these locations are composed of

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quartz-rich unconsolidated sands with a stable heavy mineral assemblage, and black organic-

rich clays and lignite. Organic linings of foraminifera and/or dinocysts occur in the sandy
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clays or sandy lignite intervals directly above the organic-rich layers deposited in Mauritia

palm swamps (Hoorn, 1994a)(Fig. 2). In the lower Apaporis River similar sediment
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successions were found, but the dominant pollen types in these organic clays are mangroves,

which alternate with Mauritia-dominated layers. Foraminifera are absent, instead the organic-
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rich sediments are capped by sandy clay layers containing dinocysts (Hoorn, 2006)(sites 5-6;
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Table 1; Fig.1-2).
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The Pebas (Peru-Colombia) and Solimões (Brazil) formations are mostly of Andean origin

and characterized by an alternation of fossil rich, blue silty clays, clays, and organic-rich clays

and lignites. The Solimões Formation has a maximum thickness of 900 m and is composed of

distinct blue siltstones to silty clays and black organic-rich clays (Maia et al., 1977). It has an

unstable heavy mineral association and geochemical signature typical for Andean provenance

(Hoorn, 1993; Roddaz et al., 2006). The sedimentary successions follow a rhythmic

alternation of coarsening upward cycles possibly controlled by Milankovitch cyclicities.

These cycles are expressed by shifts from terrestrial lowstand conditions (swamps) to a

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maximum flooding (marine assemblage) and highstand (shallow lakes)(Vonhof et al., 2003;

Wesselingh et al., 2006b, c; Vonhof and Kaandorp, 2010.

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Marine palynomorphs were found in the c. 30 meter thick Los Chorros and Mocagua sections.

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These outcrops are situated along the Amazon River in Colombia (sites 7-8; Table 1; Fig.1-

2)(Hoorn, 1994b; Vonhof et al., 2003; Wesselingh et al., 2002) and include at least five layers

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composed of a mixture of sand, clay and/or coal with marine palynomorphs. Further north, at

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the locality of Buenos Aires along the Cotuhé River (sites 9-10; Table 1; Fig.1-2, Colombia),

foraminifera tests and marine molluscs were reported by Wesselingh et al. (2002). Based on
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strontium isotope analyses on the molluscs Vonhof et al. (2003) give a ~5 psu salinity

estimate. The best locality for foraminifera is Nuevo Horizonte (site 12; Table 1; Fig.1-2,
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Peru), situated in the proximity of the Iquitos and Itaya Rivers (Wesselingh et al., 2002),

where a rich assemblage of well-preserved foraminiferal tests and ostracods with a ~1 psu
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salinity estimate (Vonhof et al., 2003.

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2.2. Age

The sedimentary successions in western Amazonia were originally dated as c. 16 to 12 Ma,

following the palynological zonation scheme by Lorente (1986)(see Hoorn 1993). These ages

are now adjusted following Jaramillo et al. (2011), who established a new zonation for

northern South America by applying a combination of graphic correlation and constrained

optimization on multiple sections from the Llanos. Rather than single marker species, their

zonation is based on palynological assemblages, ages based on isotopes, magnetostratigraphy,

and foraminifera, and calibrated to the timescale of Gradstein et al. (2005).

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The Mariñame and Tres Islas sections are dominated by Malvacipolloides (previously

Echitricolporites) maristellae and Rhoipites guianensis (Hoorn 1993, 1994a), which are

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characteristic for the early Miocene T13 Echitricolporites maristellae zone (between 17.7 and

16.1 Ma). The Apaporis section presents single occurrences of Grimsdalea magnaclavata and

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none of the characteristic taxa of the other zones. Hoorn (2006) assigned it to the Grimsdalea

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zone (sensu Lorente, 1986), which currently would coincide with the early to middle Miocene

T14 Grimsdalea zone (16.1 to 14.2 Ma). The sediments exposed at Los Chorros, Mocagua,

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and Buenos Aires include Grimsdalea magnaclavata and Crassoretitriletes vanraadshoovenii
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(Hoorn, 1993, 1994b), a combination that according to Jaramillo et al. (2011), is indicative for

a middle Miocene age (T15; 14.2 to 12.7 Ma).

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3. Material and methods

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Here we revisit a selection of samples that were collected between 1988 and 1991 within the

framework of the Tropenbos International research program (Hoorn, 1993, 1994a, b;

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Wesselingh et al., 2002). Organic residues and raw sample material are kept at the University

of Amsterdam. Lithological columns and the position of all samples are shown in Figure 2,

and the counts of all foraminifera and dinocyst specimens found in this study are listed in

Table 2.

3.1. Foraminiferal tests

At the Museu Paraense Emílio Goeldi (MPEG), thirteen samples from Los Chorros, Buenos

Aires, Nuevo Horizonte, and Cotuhé were processed and a semi-quantitative analysis of

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calcareous microfossils was carried out. The samples ranged in weight from 10 to 30 g and

were sieved through a combination of sieves with a mesh of 500 µm, 250 µm, and 125 µm.

The remaining fraction was dried at 60 ºC and the residuals of 250 and 125 µm were picked.

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An exception was sample F70 from Nuevo Horizonte, which was lent to us by F. Wesselingh

and processed previously (see Vonhof et al., 2003). From the initial thirteen samples five

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contained microfossils. The Nuevo Horizonte sample (‘nivel alto’ and F70) were particularly

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rich, with c. 300 counted specimens (Table 2). In addition, three samples from the marine

layer at the Mariñame section were processed at the University of Amsterdam. For each

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sample, 9 to 50 g was sieved through a combination of sieves with 270 µm, 180 µm and 63
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µm meshes. However, no remains of tests were recovered, possibly due to post-depositional

dissolution of calcareous tests.

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A selection of calcareous microfossils was photographed using the Scanning Electronic

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Microscopy (SEM) facility at the MPEG. Type material of the calcareous foraminifera from

Nuevo Horizonte is housed in the micropaleontological collection of MPEG under the

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numbers MPEG-194-M to MPEG-209-M; MPEG-496-M to MPEG-502-M (Plate 1-3).

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3.2 Foraminiferal linings

Palynological processing was performed at the University of Amsterdam (1990-1993 and

2013) following a standard procedure (see Hoorn, 1993). For each sample c. 1 cc was sieved

over a 250 µm-mesh to remove the larger particles. The <250µm fraction was treated with

10% sodium pyrophosphate solution to disperse the clays. Separation of the organic fraction

from the heavier inorganic fraction was done using bromoform with a density of 2.0 g/cm³.

The organic residue is mounted in glycerin on microscope slides and sealed with paraffin.

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In this study sixteen samples from the Los Chorros, Mocagua, Mariñame and Tres Islas

sections were qualitatively analyzed. No taxonomic key exists for organic linings of

foraminifera and so we classified them by assigning morphotypes. Three different categories

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were counted: a) fragments consisting of a single chamber, b) fragments with two to three

chambers and c) fragments from four chambers to complete foraminifera. The latter were

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studied in more detail and, when possible, coupled to recent living taxa. Analysis was

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performed using a Leitz LM microscope and slides are stored at the University of Amsterdam.

The taxa were photographed using the Nomarski Differential Interference Contrast method

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(Allen et al., 1969; Bercovici et al., 2009)(Plate 4).
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3.3 Dinocysts
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Twenty-three sediment samples were processed at RGD (National Geological Survey of the
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Netherlands; now TNO) in 1992 and 1995. The RGD procedure included the removal of
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carbonates using HCl 10%, the removal of silicates using HF 20%, and separation of the

organic fraction using bromoform with a density of 2.1 g/cm³. The organic residue was sieved
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over a 20 µm-mesh to remove the remaining fine (clay) particles. A qualitative analysis of

organic-walled dinocysts was done on, using a Leitz light microscope (x400 magnification).

Identification of the species follows Fensome and Williams (2004).

3.4 Pollen and spores

Three samples collected by Vonhof et al. (2003) at the locality of Nuevo Horizonte were

processed for palynology at the University of Amsterdam following the above-described

method. The sample collected in the lignite layer proved rich in pollen and it was assessed for

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it’s content in biostratigraphic marker specimens.

4. Data and interpretation

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4.1. Age

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The lignite layer at Nuevo Horizonte is dominated by Grimsdalea magnaclavata, but also

includes Crassoretitriletes vanraadshoovenii (rare). Heterocolpites incomptus, H. rotundus,

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H. verrucosus, and Polypodiaceacidites tibui are also common. Other characteristic taxa are

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Mauritiidites franciscoi, Perisyncolporites pokornyi, Psilamonocolpites nanus, P.

amazonicus, Psilatricolporites varius, Retimonocolpites maximus, R. absyae, Retitricolporites

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caputoi, Ranunculacidites operculatus, Verrumonoletes usmensis, and minor amounts of

Malvacipolloides maristellae. The palynological assemblage of Nuevo Horizonte resembles

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that of Los Chorros, Mocagua and Buenos Aires, and following the zonation in Jaramillo et

al. (2011) the age would range between 14.2 and 12.7 Ma (zone T15).
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4.2 Foraminifera
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4.2.1 Calcareous tests

Foraminiferal tests are abundant in the sediment layers exposed at Nuevo Horizonte (Peru).

Specimens are very well preserved, but species diversity is low. In one of the samples more

than 300 specimens were counted, consisting of both Ammonia and Elphidium (Table 2;

Plates 1 and 2). At Buenos Aires Elphidium is more abundant than Ammonia, but both occur

in very low, unrepresentative quantities (Table 2).

In sample F70 (Nuevo Horizonte) Ammonia is dominant and represented by A. tepida and A.

parkinsoniana, as well as aberrant forms that comprise 15% of the total sample. A. tepida is

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the most abundant species with about 5 morphotypes and several types of test abnormalities

such as an abnormal addition of a chamber, abnormally protruding chambers, reduced

chamber size, distorted chamber arrangement, and complex forms (Plate 3). E. granosum is

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also found in this sample, including specimens with deformities, but occurs in lower numbers

than A. tepida. The samples from Buenos Aires form an exception as here Elphidium

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dominates, although in overall lower numbers than the Nuevo Horizonte samples (Table 2).

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The assemblage of the Nuevo Horizonte site is further characterized by charophyte oogonia

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and ostracods. Charophyte oogonia are typical for freshwater conditions but can tolerate

slightly raised salinity levels. The ostracods are mainly represented by the euryaline genus
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Cyprideis, while other genera, typical for brackish (Perissocytheridea and Rhadinocytherura)

and marine (Pellucistoma) waters occur in lower quantities. The paleoenvironmental

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preferences of Pellucistoma –as reconstructed from the Neogene deposits from Solimões

Formation– are still under investigation (Gross et al., 2014, in prep; Ramos et al., in prep.).
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Despite the widespread occurrence and ecological relevance of Ammonia and Elphidium, the

taxonomy of these genera has been the subject of long-standing controversy (see Schweizer et
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al., 2011). These taxa tend to occur in unstable marginal environments with varying

ecological parameters. Consequently they develop intra-specific variations (morphotypes)

with many different forms that constitute an adaption to the physical-chemical variability of

the waters, making taxonomical identifications at species level complex.

In the present study two morphotypes of Ammonia were recognized: A. tepida and A.

parkinsoniana, which are similar to the morphotypes recorded in Europe, Mexico and the

Caribbean. Ammonia tepida has a wide geographical distribution (Hayward et al., 2014) and

is commonly recorded in Europe from Neogene to Present (Schweizer et al, 2011; Pascal et

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al., 2008; Amorozi et al., 2009; Filipescu et al., 2005, 2011), but also in the Americas (Eichler

et al., 1995; Debenay et al., 1998; Bruno, 2013), including the Caribbean (Hayward et al.

2003; Molinares et al., 2012).

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Like Ammonia, the genus Elphidium also presents taxonomical divergences. In the studied

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material the specimens of Elphidium are very similar to E. granosum (d´Orbygny, 1826) from

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the western Mediterranean (Milker & Schmiedl, 2012: fig. 27, 17-18). In Europe the species

is also referred to as Porosononion granosum (Poignant et al., 2000; Debenay et al., 2005;

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Felipescu et al., 2011) and Criboelphidium granosum (Amorosi et al., 2014). E. granosum
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differs from other species in this genus because of the absence of the typical ponticuli and the

openings of the associated channel system (retral process of elphidiids). Instead, this species

has an ornamented test surface with pustules and granules in the umbilical region; sutures are
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incised and curved backward with granules or pustules extending from the umbilical region.
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4.2.2 Organic linings

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Not all foraminifera have organic linings, but their presence (together with calcareous tests) is

particularly useful when reconstructing paleoenvironments at the interface of the terrestrial

and marine realms (Mamo et al., 2009). In spite of this, there is no taxonomic key for their

classification and because of this we have divided them into three morphotypes.

- Morphotype 1 (Plate 4, type A-G) is characterized by slowly evolving, pitted

chambers, and with a distinct wide umbilicus. Within this morphotype there is some

degree of variation. This morphotype has affinity to the Rotaliaceae (Ammonia or

Elphidium) because the calcareous tests of these taxa are pitted, which can be

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replicated in their organic lining.

- Morphotype 2 (Plate 4, type H). Individuals of this type are relatively small, with a
wide umbilicus; all observed specimens had no more than four chambers. This

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morphotype is assigned to juveniles that probably belonged to the Rotaliaceae

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(Ammonia or Elphidium).

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- Morphotype 3 (Plate 4, type I & J) is characterized by large, inflated chambers with

solid and thick walls and no visible pits. We tentatively assign this form to

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Trochammina, an agglutinated species with a wall consisting of clay minerals and

small quartz grains and a thicker lining than Ammonia.

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In all sections morphotype 1 (Ammonia-Elphidium) is by far the most abundant taxon whereas
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specimens of other morphotypes are rare (Table 2). Sometimes this type is found in
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combination only with indeterminate juveniles (C31) or Trochammina (C23b). In C108 and

C54, the ratio between Ammonia-Elphidium, Trochammina and juveniles is (almost) equal,
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but in Mocagua Ammonia-Elphidium are the most abundant forms, with respectively seven
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and twelve individuals. In one sample from Mocagua, both Trochammina and juveniles are

present (Moc20) and in the other sample one individual of Trochammina is present (Moc24).

In the sample ‘bonebed 15-20’ (Los Chorros section) all morphotypes had resemblance to

Ammonia-Elphidium.

The organic linings of foraminifera are very fragile and after dissolution of the calcareous

and/or agglutinated test, they can easily get damaged or partly folded. This explains why in

our samples the organic linings of foraminifera present various degrees of preservation.

4.2.3 Ecology

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Benthic foraminifera such as Ammonia and Elphidium have great potential as indicators of

environmental conditions, as they are sensitive and expressive markers of both naturally and

anthropogenically stressed marine environments (Coccioni, 2000). These genera are common

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in paralic environments, where few highly adaptable and widespread foraminiferal species

occupy most ecological niches. These paralic taxa are salinity tolerant (ranging from

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hyposaline to hypersaline waters) and can be used as salinity indicators in the standard

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estuarine classifications based on the spatial distribution of salinity (Debenay et al., 2000). An

overview of the salinity range for different genera (our study and other sites in Amazonia) is

shown in the Appendix.

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The Miocene assemblage in Amazonia is composed of euryhaline foraminifera such as

Ammonia, Trochammina and Elphidium, which are common in shallow ponds, coastal
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lagoons and mangrove environments in the Caribbean region (Javaux and Scott, 2003;
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Molinares et al, 2012), but also in Europe (Poignant et al., 2000; Chendes et al., 2004,

Filipescu et al., 2011, Bella et al., 2011), in the Mediterranean (Carboni et al., 2009), and in
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the Americas (Eichler et al., 1995; Debenay et al., 1998; Bruno, 2013).
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The genus Ammonia is an infaunal/littoral taxon and can occur from fully marine to brackish

environments, but is particularly common in marginal marine settings. Under stressful

environmental conditions (i.e. hypo- or hypersalinity) it can develop aberrant forms (Geslin et

al., 2000). Ammonia is also commonly found in marshes and subtidal environments and in

sediments with a high variability in mud and total organic carbon (TOC) content (Hayward et

al. 2004; Murray, 2006) and tolerates a wide range of salinity (<5 to >35 psu)(Melis and

Covelli, 2013; Dissarda et al., 2010). There are around 25 to 30 species of Ammonia and

species referred to this genus that are known to withstand low oxygen conditions <0.1 ml/L

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for several days (Moodley and Hess, 1992; Murray, 2006), but with the oxygen levels <1.2

ml/L the metabolic rate can slow down as the organism struggles to cope with the poor

conditions (Bradshaw, 1968; Murray, 2006).

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A. tepida is an opportunistic species which colonizes lakes or lagoons shortly after their

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formation, but can disappear with salinity changes from mesohaline to freshwater conditions

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or due to the maturation of the lacustrine community, predation or competition causing its

extinction (Almogin-Labin et al., 1995). The taxon occurs muddy sediment with a high total

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organic carbon (TOC), and is characteristic for outer paralic systems with thermohaline
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stratification. Furthermore, A. tepida is particularly sensitive to the latter because it relies on

organic matter as food resource. In Mediterranean lagoons the highest abundances of A.

parkinsoniana and A. tepida occur in intertidal and subtidal areas of the outer lagoon with
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modest hydrodynamics. However, A. parkinsoniana, prefers sandier substrates and

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oxygenated waters, and is reported as coastal taxon (Jorissen, 1988; Mellis and Covelli,

2013).
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Elphidium is a shallow-water and infralittoral taxon while E. granosum is reported from

lagoonal and shallow-marine settings (Zampi and D’Onofrio, 1987; Albani and Barbero,

1990; Bellotti et al., 1994). It inhabits sediments with highly variable sand, mud and TOC

contents and a salinity range of 15-31 psu (Alve and Murray; 1999; Murray, 2006). Ammonia

beccarii and E. granosum are considered by Jorissen (1988) as inhabitants of a near-shore

zone (7.5–25 m water depth).

According to Bella et al. (2011), the occurrence of taxa such as A. parkinsoniana, A. tepida,

E. poeyanum and E. granosum may indicate a brackish environment, such as a lagoon, or a

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slightly confined infralittoral environment affected by freshwater influence. E. granosum is a

hyposaline taxon characteristic of low salinity areas (Melis and Covelli, 2013) and does not

tolerate oxygen stress (Jorissen, 1988). Bella et al. (2008) further point out that E. granosum

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has a positive correlation with clayey substrates and high concentrations of organic matter.

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The genus Trochammina is common in lower shore face deposits, on tidal flats, channels and

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mangroves (Simmons et al., 1999), and in supratidal and landwards zones (Debenay et al.,

2000). Trochammina inflata is an agglutinated foraminifera characteristic for salt marshes and

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associated with marsh plants, but also occurs outside the marshes (Alve and Murray, 1999).
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The agglutinated test protects this species from dissolution by the low pH in marshes, and it

can live in almost fresh to brackish environments (0.5-30 psu)(Murray et al., 2011).
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According to Melis and Covelli (2013) foraminifera react to stressed and/or polluted en-

vironments through variations in abundance and species richness (community structure), but
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also through morphological abnormalities which affect shape, size or disposition of one or
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more chambers of the test. Test abnormalities in foraminifera are frequently observed in

naturally stressed marine environments – possibly induced by a deviation from optimum

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chemical and physical parameters, such as temperature, salinity, dissolved oxygen, organic

matter etc. (Boltovskoy et al., 1991; Almogi-Labin et al., 1992; Yanko et al., 1998; Geslin et

al., 2000; Coccioni, 2000). In the Dead Sea, a hypersaline inland pool, the highest relative

abundances (7-57%) of abnormal test of Ammonia tepida were recorded (Almogi-Labin et al.,

2002), whereas under normal conditions, or in a non-stressed population this is about 1%

(Geslin et al., 2002).

Based on the above we interpret the early and middle Miocene strata in the western Amazonia

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as formed in a complex system typical of paralic environments. This environment that was

characterized by cyclical and abrupt changes typical for a marginal marine to estuarine setting

which includes lagoons, salt marshes and mangroves, with low, near-bottom oxygen levels

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where sediments varied from muddy to sandy. The composition, low diversity, relatively high

number of specimens and chamber abnormalities (15%) of the foraminiferal assemblage in

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our samples, points at a sedimentary system at the interface of brackish and freshwater. The

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very rapid changes in ecological parameters such as the oxygen, salinity levels and

hydrodynamics caused stressful conditions that triggered aberrant forms in the foraminifera.

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4.3 Dinocysts

4.3.1 Results
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Dinocysts have been observed in ten of the total twenty-three analyzed samples, and are
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relatively abundant (>20 specimens) in two samples (Table 2, Plate 4, K-S). The associations
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seem to vary between samples and sites, but this could be an artifact of the overall low

dinocyst counts. The phototrophic taxon Spiniferites is the most frequently observed dinocyst,
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but Polysphaeridium zoharyi and Tuberculodinium vancampoae are common as well. Also

frequently observed are so-called ‘round brown’ cysts with an indication of an archeopyle

likely belonging to the heterotrophic type Brigantedinium. Some other Protoperidiniaceae,

such as Lejeunecysta sp., were also observed, however most of them could not be identified to

species level due to folding and/ or damage. Interesting is the presence of Quadrina? condita

in one sample (Los Chorros 44), a palynomorph seldom observed so far and for which the

status ‘dinocyst’ is yet to be confirmed. A few specimens of typical late Cretaceous to

Paleogene genera have also been identified in a number of samples (e.g. Dinogymnium in

Cotuhé 82), indicating the presence of reworked older marine material in these deposits.

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4.3.2 Ecology

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The common Spiniferites is generally considered a cosmopolitan: it is found in shallow

coastal as well as in open oceanic sites of polar to equatorial regions (Zonneveld et al., 2013).

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Due to poor preservation, Spiniferites could not be identified to the species level. However,

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specimens of Spiniferites with a distinct ‘cruciform’ shape, which is associated with fresh to

brackish water conditions in the central Asian and southeastern European seas (Wall et al.,

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1973; Kouli et al., 2001; Mudie et al., 2001; Marret et al., 2004), have not been observed in
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the Amazonian samples. Both Polysphaeridium zoharyi and Tuberculodinium vancampoae

are observed in (sub-)tropical to equatorial coastal sites (Zonneveld et al., 2013). Particularly

P. zoharyi is associated with enhanced or reduced surface water salinity, like lagoons and
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estuaries, (Wall et al., 1977, Morzadec-Kerfourn, 1979; Bradford and Wall, 1985; Marret and
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Zonneveld, 2003), but is not restricted to these environments (Zonneveld et al., 2013).

Similarly, Tuberculodinium vancampoae is found at sites within a broad salinity range of

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27.8-39.2 psu, but the highest abundances occur where salinity is >32.9 psu (Wall et al., 1977;
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Zonneveld et al., 2013). The heterotrophic type Brigantedinium is a cosmopolitan which can

be abundant in inner neritic as well as open oceanic sites at polar to equatorial latitudes. High

abundances are, like for most heterotrophic Protoperidiniaceae, related to high surface water

productivity due to nutrient availability (Zonneveld et al., 2013). Although the present-day

ecology of Quadrina? condita is unknown, it is often found in association with thermophilous

taxa in the paleorecords (Soliman et al., 2012), which is consistent with our current finding.

Although generally the highest abundances of most identified dinocyst taxa nowadays occur

at sites with an average sea surface salinity of 30-38 psu, occasionally they are also found in

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areas with low salinities, related to seasonally enhanced runoff (Zonneveld et al., 2013). For

example, P. zoharyi dominates the assemblages in the shallow Mississippi Sound where the

sea surface salinity seasonally drops below 30 psu (Edwards and Willard, 2001). Spiniferites

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and P. zoharyi are commonly found together at various other sites along the Gulf of Mexico

coast, for example in Mexican lagoons (Limoges et al., 2013), the Florida Bay (e.g. Wingard

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et al., 1995; Ishman et al., 1996; Brewster-Wingard et al., 1998), and Florida estuaries

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(Cremer et al., 2007). Surface water salinity differs between these sites; in the lagoons it

shows little annual variation (35-37 psu; Limoges et al., 2013), whereas in the estuaries

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salinity can drop below 14 psu during the wet season (Stoker, 1992). These sites are generally
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shallow (< 5 m deep). Small proportions of T. vancampoae have been observed together with

P. zoharyi and Spiniferites in brackish to highly saline estuaries of various Caribbean islands

(Wall et al., 1977), as well as in Florida estuaries like Rookery Bay (Cremer et al., 2007).
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High relative abundances of Brigantedinium are found in the nutrient-rich Amazon River
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plume along the northern South America coast (Vink et al., 2000; Zonneveld et al., 2013).

Moreover, Brigantedinium is found in association with Spiniferites and P. zoharyi along the
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Florida coast at sites up to 26 m deep and with annual salinity of ~35 psu (Limoges et al.,
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2013), but also in the shallow estuarine system of Rookery Bay (FL) where salinity seasonally

varies between 0-38.4 psu (Cremer et al., 2007 and references therein). Outside the Gulf of

Mexico region, similar dinocyst associations are observed in large estuarine inlets like

Chesapeake Bay (VI, USA)(Willard et al., 2003).

Despite the presence of some Cretaceous and Paleogene dinocyst taxa, the overall association

in the Amazonian samples appears to be more typical of the Neogene. Particularly

Tuberculodinium vancampoae and Quadrina? condita are important biostratigraphic

indicators, with first occurrences in the latest Oligocene (Powell, 1992) and middle Miocene

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(de Verteuil and Norris, 1992) respectively. This supports our inference that although some

marine material is present, most observed marine palynomorphs are of Miocene age.

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In surface samples of Florida Bay locations, particularly Spiniferites and P. zoharyi are

reported in association with foraminifera Ammonia and Elphidium, amongst others (Wingard

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et al., 1995; Ishman et al., 1996; Brewster-Wingard, 1998). Considering the present-day

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distribution of the observed dinocyst taxa, the Pebas dinocyst assemblage likely reflects an

estuarine to shallow marine environment, which is consistent with the interpretation of other

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proxies. Important to note is that high abundances of foraminiferal linings and high
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abundances of dinocysts do not occur in the same samples (Table 2). This further illustrates

the spatial variability within the wetland, with foraminifers likely occurring in the shallow

parts whereas the dinoflagellates likely thrived in the deeper channels. Additional
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sedimentological research could confirm different depositional conditions for the different
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marine (micro)fossils found throughout Amazonia.

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5. Discussion
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5.1 Paleoenvironmental implications of our data and regional context

Miocene marine palynomorphs from Colombian Amazonia and an exceptionally well-

preserved calcareous foraminifera assemblage from Nuevo Horizonte (Peru) characterize

selected strata in a sedimentary succession otherwise deposited in a terrestrial environment.

Our findings suggest that during the Miocene the Amazonian wetlands were affected by

marginal marine conditions of varying salinities. Elsewhere in northern South America,

marine evidence from lower to middle Miocene strata further confirm the regional nature of

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these marine ingressions which linked Amazonia through the Llanos Basin with the

Caribbean (Fig. 1). This is in agreement with Nuttall (1990), Hoorn (1993), Lundberg et al.

(1998), Lovejoy et al. (1998) and Wesselingh and Macsotay (2006).

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Regional data further suggest that there were at least four discrete incursion phases into

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Amazonia, one during the early Miocene (zone T13), two during the middle Miocene (zone

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T14? and T15), and one (or more) during the late Miocene (7 Ma; Bolivia)(Uba et al., 2009).

These marine incursions temporarily changed fluvial and fluvio-lacustrine settings into

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marginal marine/estuarine settings while the opposite happened during the sea regression,
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higher precipitation and tectonic stability.

The notably different foraminiferal composition found in middle to upper Miocene strata in
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westermost Amazonia suggests that the marine incursions varied in intensity. Hulka et al.
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(2006) reported a series of marginal marine benthic foraminifer taxa from shallow marine and

lacustrine/alluvial plain deposits in Bolivia (see Appendix). Although Nicolaides & Coimbra
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(2008) question these taxonomical identifications, and the paleoecological interpretation

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should be considered with caution, Uba et al. (2006, 2009) also reported Globigerinacea

(plankton) and Ammonia (benthos) from this same geographic region. In westernmost

Peruvian Amazonia, shallow to marginal marine taxa such as Trochammina cf. pacifica,

Ammonia, Protelphidium (or Haynesina?), and Miliammina fusca are reported, but also

Psammosphaera, Anomalinoides, Karreriella, Buccella, Bathysiphon (P.-O. Antoine, pers.

obs.). This all points at mixed assemblages where both benthic and planktonic forms occur,

and represent both shallow restricted and open marine environments (see e.g. Resig, 1990;

Mallon, 2011).

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It is interesting that the abovementioned assemblages from sub-Andean Peru and Bolivia

resemble Miocene assemblages found in northern Argentina (see Boltovskoy, 1991 for

species list and references). Here too planktonic foraminifera were found together with a

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typical shallow marine assemblage. This all suggests the sub-Andean sites in Peru and Bolivia

might have been more proximal to a marine entrance from the South, i.e. the Paranan Sea

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(Boltovskoy, 1991; Alonso, 2000; Hovikoski et al., 2007) or a marine connection between the

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Northern and Central Andes (the Guayaquil Gap), which is further supported by middle-late

Miocene marine deposits (Steinmann et al., 1999). However, to date no marine fauna has been

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found in the fossil-rich deposits of the intervening basins (e.g. Cuenca and adjacent basins).
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To be sure, this would require further study of the foraminiferal assemblages from a wide

region.
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The pathway for marine incursions and accommodation space for shallow marine to estuarine
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sediment deposition is facilitated by a combination of causes. These are: plate tectonic

changes in the Pacific and flat slab subsidence, Andes uplift, foreland basin formation and
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global high sea level (Marshall and Lundberg, 1997; Roddaz et al., 2010; Shephard et al.,
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2010). As a consequence, Miocene proto-Amazonia was subject to large scale

paleoenvironmental changes that were epitomized by drainage readjustments, wetland

formation and marine incursions (e.g. Hoorn, 1993; Vonhof et al., 2003; Wesselingh et al.,

2002; Hovikoski et al., 2007; Hoorn et al., 2010b; Roddaz et al., 2010).

5.2 Other paleontological and sedimentological evidence for marine incursions

Several other euryhaline taxa in Neogene strata in other areas of Western Amazonia support

the hypothesis of intermittent brackish environments during the Miocene (Table 1; Fig. 1).

Tidal sediment structures (rhythmites and characteristic ichnofacies) are also part of the

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evidence (Hovikoski et al., 2010), in addition to salinity tolerant ray-finned fishes (Lundberg

et al., 1998; Lovejoy et al., 2006), rays and sharks (Monsch, 1998), mollusks (Wesselingh et

al., 2002; Wesselingh and Ramos, 2010; Vonhof et al., 1998, 2003), barnacles, bryozoans,

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ostracods (Sheppard and Bate, 1980; Muñoz-Torres et al., 1998), trace fossils, mangrove

pollen and sedimentological indications of tidal influence in the ancient Pebas system (Hoorn,

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1994a, b; Hovikoski et al., 2007).

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From North to South (Table 1; Fig. 1), such proxies are located nearby Iquitos and

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Contamana (Loreto, Peru), along the Ríos Inuya (Fitzcarrald Arch, Ucayali) and Madre de
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Dios (Cerro Colorado, Madre de Dios, Peru), and in various sections from the Chaco, Bolivia

(Table 1; Fig. 1). These deposits, ranging from the late middle Miocene (c. 13 Ma) up to the

late Miocene (c. 9 Ma), coincide with the end of the “Pebas phase” and the beginning of the
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subsequent “Acre phase” (Hoorn et al., 2010a). They are assigned to the Pebas Formation
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nearby Iquitos (Monsch, 1998; Wesselingh et al., 2010; unpublished data), to the Ipururo

Formation in Contamana (unpublished data) and the Río Inuya (Salas-Gismondi et al., 2006,
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2007; Antoine et al., 2007; Espurt et al., 2010; unpublished data), to the Madre de Dios
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Formation in the Madre de Dios (Campbell et al., 2001, 2010; Hovikoski et al., 2005, 2007;

Antoine et al., 2013), and to the Yecua Formation in the Chaco, Bolivia (Hulka et al., 2006).

Vertebrates of taxa with marine affinities occur in the abovementioned deposits and consist of

rays and sharks (Lovejoy et al., 1998, 2006; Monsch, 1998; Antoine et al., 2007; Lundberg et

al., 2010; unpublished data), but also pristid teeth (sawfishes) were recovered from various

localities (Table 1). Sawfishes are generally fully marine organisms, even if some species are

euryhaline (Lundberg et al., 2010). Palatine teeth of large eagle/bat rays (Myliobatis spp.) and

cownose rays (Rhinoptera sp.) were also reported in latest middle Miocene deposits of the

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Pebas Fm., in Northwestern Amazonia (Monsch, 1998; Lovejoy et al., 2006), and of the

Fitzcarrald local fauna (Bianucci et al., 2013). Such rays have a marine to estuarine habitat

today (Lundberg et al., 2010). A vertebra belonging to a large lamniform shark was found in

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2005 along the Río Inuya (previously misidentified as Carcharhinus sp. (e.g., Bianucci et al.,

2013). This group encompasses obligate marine sharks (including the great white; Cappetta,

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2012). Several fossil teeth of Carcharhinus (bull sharks, Carcharhiniformes) were also

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unearthed in late Miocene deposits of the Acre area, Brazil (Universidad Federal do Acre

collection, Rio Branco, Brazil) but their recent relatives are often reported in the Amazon

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River as far up the river as Iquitos, Peru (Thorson, 1972). Fossil cetacean remains from
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Northwestern Amazonia have been unearthed in a wide array of middle and late Miocene

localities from Brazil, Colombia and Peru (e.g., Negri et al., 2010; Bianucci et al., 2013).

They are mostly referred to as representatives of river dolphin families (for review, see
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Bianucci et al., 2013). Yet, a delphinidan of uncertain affinities was uncovered in the
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Fitzcarrald Miocene fauna (Goillot et al., 2011; Bianucci et al., 2013), within tidally-

influenced deposits c. 13 Ma in age (Espurt et al., 2010).

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A diagram proposed by Hovikoski et al. (2010: Fig. 9.11) illustrates a wide array of

depositional environments ranging from alluvial to marine, and likely to have occurred in

Western Amazonia during the Pebas and Acre phases. Based on molecular clock estimates

and on the available fossil record, Lovejoy et al. (2006) consider that the Pebas megawetland

system provided favorable conditions triggering the adaptation of most “Marine-Derived

Lineages” in Amazonia, such as river stingrays (Potamotrygonidae), drums (Sciaenidae),

anchovies, needlefishes, and mollusks (Wesselingh et al., 2002). However, the recent

discovery of an obligate freshwater potamotrygonid (Potamotrygon ucayalensis) in middle

Eocene deposits of Peruvian Amazonia (Adnet et al., 2014) minimizes the role of Miocene

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marine incursions, at least for the invasion of Amazonian river stingrays.

6. Concluding remarks

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A plethora of different proxies confirm the existence of a Miocene corridor that connected

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Amazonia with the marine realm. In this study, we present calcareous tests and organic

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linings of foraminifera, and organic-walled dinocysts from Miocene sediments in Colombia

and Peru. These findings suggest estuarine conditions with salinities ranging from 0.5 psu

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(freshwater) to well above >5 (brackish) and possibly reaching up to 30 psu (saline) existed.
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The marine corridor allowed an arrangement of paralic environments with a mixture of

marine and brackish water biota to colonize the heartlands of Amazonia, reaching at least

2000 km south from the Caribbean portal of entrance, and 1000 km from the nearest
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coastline.
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The foraminiferal association (Ammonia, Elphidium and Trochammina) and the dinocyst
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assemblage (Spiniferites, Brigantedinium and Tuberculodinium vancampoae) reported in this

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study all point at shallow marine and shallow estuarine/mangrove settings with daily

influence of saline (brackish) water. The aberrant forms of Ammonia found at Nuevo

Horizonte seem to confirm stressful low salinity and oxygen conditions.

Marginal environments such as inferred in the Pebas/Solimões wetland are very susceptible to

physical-chemical changes due to the dynamics of the tidal system and seasonal variation in

precipitation levels and rates of fluvial run-off. The relatively short-lived marine incursions

led to extreme conditions, which allowed taxa such as Ammonia to invade, settle, and

reproduce. However, residence time in these new conditions was short enough for taxa to be

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discontinuous in the studied successions. In the course of the Plio-Pleistocene the return of

stable freshwater conditions created a suitable environment for freshwater biota.

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The edaphic heterogeneity in the Amazon Region is, for an important part, related to

geological bedrock and the Pebas/Solimões formations with the marine strata as an important

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component of these formations. The Miocene salinity fluctuations in space and time created a

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mosaic of ecological niches that played a role in species evolution in this region. This stresses

the importance of these marine incursions for better understanding modern biodiversity and

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proves the need for further study on this aspect of Amazonia’s natural history.
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7. Acknowledgements
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We are very grateful for the help we received from Suzette Flantua (map) and Jan van Arkel
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(photos). We thank Hubert Vonhof for discussion and sharing sample material; Simon

Troelstra and Peter Frenzel for advice on the classification of organic linings; Waldemar
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Herngreen for initial assessment of dinocysts; Roel Verreussel for retrieving samples from the
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TNO repository. Roel Verreussel, Alexander Houben and Francesca Sangiorgi are

acknowledged for sharing their dinocyst expertise. Francisco Parra and Rosa Navarrete have

identified microfossils from Contamana, Peru. We acknowledge CLIM-AMAZON

(www.clim-amazon.eu), a joint Brazilian-European facility for climate and geodynamic

research on the Amazon River Basin sediment, for partially funding this work (E.L. study

grant and C.H. travel funds). Two anonymous reviewers are kindly thanked for their

comments, which helped us to substantially improve the manuscript.

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Ostracoda) and their development over the past 30 million years. In: C. Hoorn & F.P.

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Wesseling (Eds), Amazonia: Landscape and Species Evolution: A look into the past,

Wiley-Blackwell, Oxford, 302-316p.

Wesselingh, F.P., Hoorn, C., Kroonenberg, S.B., Antonelli, A.A., Lundberg, J.G., Vonhof,

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H.B. & Hooghiemstra, H., 2010. On the origin of Amazonian landscapes and biodiversity:

A synthesis. In: C. Hoorn & F.P. Wesseling (Eds), Amazonia, landscape and species

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evolution, Wiley-Blackwell, Oxford, 421-432p.

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Whatley, R.C., Muñoz-Torres, F. & van Harten, D., 1998. The Ostracoda of an isolated

Neogene saline lake in the western Amazon Basin, Peru. Bulletin du Centres de

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Recherches Elf Exploration-Production. Memoires 20: 231-245.
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Willard, D.A., Cronin, T.M. & Verardo, S., 2003. Late-Holocene climate and ecosystem

history from Chesapeake Bay sediment cores, USA. Holocene 13: 201-214.

Wingard, G.L., Ishman, S., Cronin, T., Edwards, L.E., Willard, D.A. & Halley, R.B., 1995.
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Preliminary analysis of down-core biotic assemblages: Bob Allen Keys, Everglades

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National Park, Florida Bay. US Geological Survey Open-File Report: 98-122.

Yanko, V., Ahmad, M. & Kaminsky, M., 1998. Morphological deformities of benthic
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foraminiferal tests in response to pollution by heavy metals: implications for pollution

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monitoring. Journal of Foraminiferal Research 28(3): 177-200.

Zampi, M. & D’Onofrio, S., 1987. I foraminiferi della laguna di Levante (Orbotello,

Grosetto), Atti Societa Toscana Scienze Naturali Memorie 93: 101-127.

Zonneveld, K.A.F., Marret, F., Versteegh, G.J.M., Bogus, K., Bonnet, S., Bouimetarhan, I.,

Crouch, E.; de Vernal, A., Elshanawany, R., Edwards, L., Esper, O., Forke, S., Grøsfjeld,

K., Henry, M., Holzwarth, U., Kielt, J.-F., Kim, S.-Y., Ladouceur, S., Ledu, D., Chen, L.,

Limoges, A., Londeix, L., Lu, S.-H., Mahmoud, M.S., Marino, G., Matsouka, K.,

Matthiessen, J., Mildenhal, D.C., Mudie, P., Neil, H.L., Pospelova, V., Qi, Y., Radi, T.,

Richerol, T., Rochon, A., Sangiorgi, F., Solignac, S., Turon, J.-L., Verleye, T., Wang, Y.,

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Wang, Z., Young, M., 2013. Atlas of modern organic dinoflagellate cyst distribution

based on 2405 data points. Review of Palaeobotany and Palynology 191: 1-197.

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Plate 3

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CAPTIONS

Figure 1 Localities with reported marine influence during the Miocene(see for coordinates

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and information Table 1). The study sites are the red dots: 1, Mariñame; 2, Tres Islas 3; 5,

Apaporis 4; 7, Los Chorros; 8, Mocagua; 9, El Salado (Cotuhé); 10, Buenos Aires; 11. Puerto

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Caimán; 12, Nuevo Horizonte. Other localities are in black dots.

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Figure 2 Lithological columns of study sites, and the sampled levels where marine

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palynomorphs and calcareous microfossils were found.
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Figure 3 Palaeogeographic map of northern South America in Miocene times, constrained by

our new data and from literature. See Table 1 for further details on the concerned localities.
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Coastlines, shields, and Andean range outlines were modified from Uba et al. (2009), Hoorn
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et al. (2010a), Roddaz et al. (2010), Montes et al. (2012), Antoine et al. (2013), and references

therein.
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Plate 1 SEM microphotography of Elphidium granosum? from Nuevo Horizonte (Peru)

(magnification bar = 50 µm): a – l) Elphidium granosum? ; a) side view (MPEG-194-M); b)

side view (MPEG-195-M); c) side view; d) fully developed stage; side view (MPEG-196-

M);e) detail of the sutures and granules (MPEG-196-M); f) side view (MPEG-197-M); g) side

view (MPEG-198-M); h) side view (MPEG-199-M); i) detail of the sutures (MPEG-199-M);

j) side view, small form (MPEG-200-M); k) deformed chambers with reduced size (MPEG-

201-M); l) deformed chambers with reduced chamber (MPEG-202-M).

Plate 2 SEM microphotography of Ammonia, Nuevo Horizonte (Peru)(magnification bar = 50

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µm): a) Ammonia parkinsoniana, umbilical side (MPEG-203-M); b-l) Different morphotypes

of Ammonia tepida. b) morphotype 4, spiral side (MPEG-204-M); c) morphotype 5, spiral

side (MPEG-205-M); d-f) morphotype 1 (according to Coccioni, 2000, fig.2/1-3), (MPEG-

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206-M) d) spiral side, e) umbilical side, f) apertural view; g – i) morphotype 2 (MPEG-207-

M), g) spiral side, h) umbilical side, i) apertural side; j – l) morphotype 3 (according to

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Coccioni, 2000, fig.2/16-19), (MPEG-208-M) j) spiral side, k) umbilical side, l) apertural

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side..

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Plate 3 SEM microphotography of different kinds of abnormal tests of Ammonia tepida,
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Nuevo Horizonte (Peru)(magnification bar = 50 µm): a) distorted chamber position, umbilical

side (MPEG-209-M); b) complex form (MPEG-496-M); c) abnormal addition of chamber,

spiral side (MPEG-497-M); d) distorted chambers arrangement, umbilical side (MPEG-498-

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M); e) reduced chamber size, umbilical side (MPEG-499-M); f) reduced chamber size, spiral
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side (MPEG-500-M); g –h) abnormal addition of chambers, spiral side (MPEG-501-M;

MPEG-502-M).
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Plate 4 Selection of dinocysts and organic linings of foraminifera from Miocene sediments of

various sites in western Amazonia, photographed using a light microscope with Nomarski

Differential Interference Contrast method.

Foraminifera: Types A-G are assigned to Rotaliacea (Ammonia-Elphidium), type H possibly

is a juvenile of Ammonia-Elphidium and types I & J are assigned to Trochammina (Type #)

sample code, England Finder coordinates: (A) Chorros 31, D36-3; (B) c23b, S24; (C) C31.3,

N25-1; (D) c23b, N13-N14; (E) 3Is53, G19-2;(F) C55, N30-2; (G) 3Is138, L25; (H) C108,

P23; (I) c23b, Q21-R21; (J) c23b, M14.

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Dinoflagellate cysts: (K) Dinogymnium sp., 1-2 different focus, Cotuhé 82 V44/2. (L):

Polysphaeridium zoharyi, Mariñame 5-91, M43/3. (M): Quadrina? condita, Los Chorros 44,

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W49/4 orientation uncertain. (N-O): Spiniferites sp., (N) Apaporis 161, T64/4, (O) Mocagua

3 K44/2 orientation uncertain. (P): Tuberculodinium vancampoae, Apaporis 161, F62/3. (Q-

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R): Brigantedinium sp., (Q) Los Chorros 44, X48/1, (R) Apaporis 161, T55/2. (S): cf.

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Lejeunecysta sp., Apaporis 161, T55/2.

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Table 1

Sources (age) Fossil taxa/Sedimentology Sources (Marine proxies)

Zonocostites pollen, foraminifera (linings),

Hoorn (1993, 1994a) Hoorn (1993, 1994a)
dinocysts (see table 2)

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Zonocostites pollen, foraminifera
Hoorn (1990, 1994a) Hoorn (1990, 1994a)
(linings)(see table 2)
Hoorn (1990) Zonocostites pollen, foraminifera (linings) Hoorn (1990)
Hoorn (1990, 1993, 1994a) Zonocostites pollen Hoorn 1990, 1993, 1994a)

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Hoorn (2006) Zonocostites pollen, dinocysts (see table 2) Hoorn (2006)
Hoorn (2006) Zonocostites pollen Hoorn (2006)

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Sr isotopes (0.708113-0.708548),
Hoorn (1990, 1993,
Hoorn (1990, 1993, 1994b) Zonocostites pollen, foraminifera (linings),
1994b)
dinocysts (see table 2)
Zonocostites pollen, foraminifera (linings),
Hoorn (1993, 1994b) Hoorn (1993, 1994b)

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dinocysts (see table 2)
Zonocostites pollen (rare), dinocyst
Hoorn (1990, 1994b) Hoorn (1990, 1994b)
(reworked/rare)
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Sr isotopes (0.708894-0.7090267), Hoorn (1993, 1994b);
Hoorn (1993, 1994b) molluscs, dinocysts, foraminifera (linings Wesselingh et al. (2002);
and calcareous tests)(see table 2) Vonhof et al. (2003)

Hoorn (1990, 1993,

Hoorn (1990, 1993, 1994b) Zonocostites pollen (common)
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1994b)
Sr isotopes (0.708134-0.708617),
Wesselingh et al. (2002);
this work ostracods (Cyprideis), molluscs, calcareous
Vonhof et al. (2003)
foraminifera (see table 2).
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Sr isotopes (0.707993-0.709299), Hoorn (1993), Vonhof et

Hoorn (1993)
Zonocostites pollen, dinocysts al. (2003)
Ostracods (Cyprideis), planktonic
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foraminifera (Globigerinoides, Globorotalia

and Globigerina) and benthic foraminifera
Linhares et al. (2011) Linhares et al. (2011)
(Amphistegina, Planorbulina and
Quinqueloculina), molluscs, fishes teeth,
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decapoda & bryozoans

Vertebrata: Pristis sp. (sawfish, oral teeth);
Monsch (1998);
unpublished data Myliobatis sp. (eagle/bat ray, dozens of
unpublished data
teeth)
Foraminifera: Karreriella sp., Trochammina
unpublished data
cf. pacifica, Miliammina fusca,
(identification of forams by
unpublished data Psammosphaera sp.
Rosa Navarrete &
Vertebrata: Pristis sp. (sawfish, rostral and
Francisco Parra)
oral teeth)
Foraminifera: Ammonia spp.,
unpublished data
Anomalinoides spp.,
(identification of forams
Protelphidium/Haynesina(?) sp., Karreriella
unpublished data and sea urchins by Rosa
sp., Buccella spp., Bathysiphon sp.
Navarrete & Francisco
Echinodermata: Echinoidea indet. (sea
Parra)
urchin spines)

Vertebrata: Pristis sp. (sawfish, oral teeth); Salas Gismondi et al.

Salas Gismondi et al.
Rhinoptera sp. (cownose ray, palatine (2006); Antoine et al.
(2006, 2007); Antoine et
teeth); Delphinoidea indet. (dolphin); (2007); Bianucci et al.
al. (2007)
Rhythmites, tidalites (2013, unpublished data)

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Salas Gismondi et al.

Vertebrata: Large lamniform shark Salas Gismondi et al.
(2006); Antoine et al.
(vertebra) -- surface collect; Rhythmites, (2006); Antoine et al.
(2007); Espurt et al.
tidalites (2007); unpublished data
(2010); Goillot et al. (2011)
Salas Gismondi et al.
unpublished data (S.
(2006); Antoine et al.
Rhythmites, tidalites Brusset, pers. com;,
(2007); Espurt et al.

(2010); Goillot et al. (2011)
Campbell et al. (2001,
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2005)
Ophiomorpha, Thalassinoides, Gyrolithes,
Hovikoski et al. (2005,
and Asterosoma; Rhythmites, cyclic
2007, 2010). See 2007 for

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2010) tidalites (no microfossil indicators for marine
additional locations.
conditions)
Foraminifera: Ammonia tepida,

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Anomalinoides cf. salinensis, Cibicidoides
cf. cushmani, C. aff. mckennai, Gyroidina
rosaformis, Hansenisca multicamerata,
Holmanella baggi, H. valmonteensis,

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Nonionella miocenica, Pseudoparella
Hulka et al. (2006); Uba et
californica;
Uba et al. (2009) al. (2009); Nicolaides and
Ostracoda: Cyprideis spp., Darwinulla,
Coimbra (2008)
Iliocypris, Cypria, Perissocytheridea,
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Cyprideis aff. amazonica, C. aff. torosa, C.
aff. truncata, C. sp., Heterocypris sp.,
Bivalvia: Neocorbicula sp., Tellina sp.;
Gastropoda: Heleobia; Tidal and shoreline
facies; Ophiomorpha; marine barnacles;
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Miocene: dinocysts, foraminifera (linings) &

Bayona et al., 2007 Bayona et al. (2007)
marine acritarcha
idem idem idem
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idem idem idem

idem idem idem
idem idem idem
idem idem idem
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idem idem idem

idem idem idem
idem idem idem
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idem idem idem

idem idem idem
idem Zonocostites pollen. Dueñas (1980)

Zonocostites, Psilatricolporites crassus

Rull (2001) Rull (2001)
pollen, dinocysts, foraminifera (linings)
early-middle Miocene: Zonocostites,
Psilatricolporites crassus, Avicennia type
Lorente (1986) pollen, dinocysts, foraminifer linings; Lorente (1986)
calcareous foraminifera; late Miocene-
Pleistocene: alluvial plain
idem see below Lorente (1986)

Miocene: Zonocostites, microplankton and

idem Lorente (1986)
foraminifera; Plio-Pleistocene: alluvial plain

idem see above Lorente (1986)

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idem Late Miocene, Zonocostites Lorente (1986)

idem Miocene, Zonocostites throughout Lorente (1986)

Zonocostites; sedimentary structures

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Hoorn (1994b) Hoorn (1994b)
indicative of tidal deposition

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Table 2

AGE

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M Proc (Bio FORA FORAMINIF
A SAM esse zon M. ERAL
P SITES PLE d e) TESTS LININGS ORGANIC WALLED DINOCYSTS

Lejeunecysta sp.

Indeterminate
Spiniferites

Polysphaeridium zoharyi (epicyst)

cf. Dinogymnium (reworked)

Protoperidinoid indet brown spiny

Cretaceous forms (reworked)

cf. Quadrina? condita
cf. Brigantedinium

Protoperidinioid (indet smooth)

Tuberculodinium vancampoae
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Juveniles (Indeterminate)
Elphidium granosum?

Ammonia-Elphidium

Indeterminate
Trochammina
ED Ammonia
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Mar 1990 T13

Mariñ 05-91 ; 1
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1 ame 1992 8 2 5 2 3
Mar 1990 T13
Mariñ 09-91 ;
1 ame 1992 3 5
Mariñ 3Is13 T13 2
1 ame 8 1990 1 4 0
Mariñ Mar1 T13
1 ame -91 1990 2
1990 T13
Mariñ Mar2 ;
1 ame -91 1992 4 4
1990 T13
Mariñ Mar3 ;
1 ame -91 1992 5 3
Tres T13 1 3
2 Islas 3Is53 1990 7 2

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1990
Apapo ; 1 1
5 ris 4 161 1992 T14 5 5 1 2 1 1
1990
Apapo ; W
5 ris 4 162 1992 T14 H 3 6

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1990
Bueno ;
1
s Aires 1992

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0
(Cotuh ; W W
e) 53 2014 T15 0 0 H H

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Bueno
1 s Aires 1990
0 (Cotuh ; 1

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e) 52(1) 2014 T15 1 0
El
Salado
1
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de 1990
0
Cotuh ; 1 W
e 82 2014 T15 0 1 H
Los 1990
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7 Chorro ; 1 2
s C31 1992 T15 3 3 4 3
Los 1990
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7 Chorro ;
s C44 1992 T15 2 4
Los
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7 Chorro C23b 2 3
s (1) 1990 T15 5 2 6
Los
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7 Chorro C108 1
s (1) 1990 T15 1 2 2 0
bone
Los bed
7
Chorro 15-20
s (1) 1990 T15 6
Los
7 Chorro C54
s (1) 1990 T15 1 1 1
Los
7 Chorro 1 1
s C55 1990 T15 4 3 2 9
1990
8 Mocag ;
ua Moc3 1992 T15 1
Mocag 1990 W W W
8
ua Moc7 ; T15 H H H

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1992
1990
8 Mocag Moc2 ;
ua 0 1992 T15 1 7 1 6
Mocag Moc2 1
8
ua 4 1990 T15 1 2 9

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Nuevo 2
1
Horizo nivel 2 7
2
nte alto 2014 T15 5 5

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Nuevo 1
1
Horizo nivel 0 1

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2
nte bajo 2014 T15 3 7
Nuevo 1 1
1
Horizo F70= 8 1

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2
nte F141 2003 T15 7 3
Nuevo lignit
1
Horizo e
2
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nte layer 2014 T15 0 0

Table 1 Overview of localities in northern South America (east of the Eastern Cordillera)
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where Miocene marine evidence was recorded. The coordinates in this table form the basis for
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Figure 1.
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Table 2 Foraminifera and dinocyst counted in samples collected in Colombia and Peru.‘WH’

indicates observations made by Waldemar Herngreen (Dutch Geological Survey) in 1992 and
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1995.

Appendix Salinity ranges for foraminifera and dinocysts found in this study (grey shade) and

from previously documented, and on-going, studies in the sub-Andean Bolivia and Peru.

Identifications by Hulka et al. (2006) are questioned by Nicolaidis and Coimbra (2008).

Salinity ranges for the listed foraminifera are copied from the Encyclopedia of Life (EOL)

and for dinocysts from Marret and Zonneveld (2003) and Zonneveld et al. (2013).

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HIGHLIGHTS

 During the Miocene marine protists lived in Amazonia.

 Protists in Miocene Amazonia point at marginal marine conditions.

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 Lower salinity in Miocene Amazonia ranged between 0 - 10 practical salinity
units.

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 Alternating marginal marine and freshwater deposits indicate estuarine
conditions.

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 Regional data show that in the Miocene Amazonia and the Caribbean were
connected.

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