Cantalamessa, 2005

Palaeogeography, Palaeoclimatology, Palaeoecology 216 (2005) 1 – 25
www.elsevier.com/locate/palaeo
Sequence stratigraphy of the Punta Ballena Member of the Jama

Formation (Early Pleistocene, Ecuador): insights from integrated
sedimentologic, taphonomic and paleoecologic analysis of
molluscan shell concentrations
Gino Cantalamessaa, Claudio Di Celmaa,*, Luca Ragainib
a
Dipartimento di Scienze della Terra, Università di Camerino, Via Gentile III da Varano, 1, I-62032 Camerino (MC), Italy
b
Dipartimento di Scienze della Terra, Università di Pisa, Via S. Maria, 53, I-56126 Pisa, Italy
Received 23 January 2004; received in revised form 3 August 2004; accepted 24 September 2004
Abstract
Early Pleistocene strata are well exposed along the uplifted eastern margin of the Esmeraldas–Caraquez Basin, particularly
on the coastal cliff west of the city of Jama (a type locality for shallow marine Quaternary sedimentary exposures on north-
central Ecuador). Here, at least 180 m of sediments accumulated in inner-shelf to subaerial settings are present (Jama
Formation). This paper is concerned with the middle stratigraphic unit of the Jama Formation (the Punta Ballena Member,
PBM). Integrated sedimentologic, paleontologic, and stratigraphic observations reveal that the PBM is composed of eight,
disconformity-bounded depositional sequences. Hiatal shell concentrations, which formed as a consequence of landward and
seaward stratal attenuation, are associated with sequence bounding and intrasequence surfaces. Together with sedimentologic
characteristics, these fossiliferous levels are important indicators of facies architecture and, therefore, are useful tools for
interpreting sequence stratigraphy. According to their taphonomic features and taxonomic composition, the hiatal shell
concentrations occurring within the PBM may be grouped into two main types, each of which occupies a predictable
stratigraphic position within the sequences: current/wave-winnowed beds at the cycle base and community shell concentrations
in a roughly mid-cycle position.
Current/wave-winnowed beds, commonly underlain by Thalassinoides burrows that penetrate to depths of as much as 1.5 m
below ravinement surfaces, are transported skeletal concentrations and formed through physical processes in shallow, high-
energy environments. They comprise a time-averaged mixture of specimens having different ecologies and states of
preservation and accumulated mainly under conditions of total passing of siliciclastic sediment (i.e. terrigenous sediments are
delivered to the site as suspended load but, owing to the high environmental energy, failed to accumulate).
Community shell concentrations occur in sparsely fossiliferous intervals and result from reduced net sedimentation
within low-energy, relatively deeper settings. They are composed of matrix-supported, loosely clustered, life-oriented and
* Corresponding author. Tel.: +39 737 402 642; fax: +39 737 402 644.
E-mail addresses: [email protected] (G. Cantalamessa)8 [email protected] (C. Di Celma)8 [email protected]
(L. Ragaini).
0031-0182/$ - see front matter D 2004 Elsevier B.V. All rights reserved.
doi:10.1016/j.palaeo.2004.09.012
2 G. Cantalamessa et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 216 (2005) 1–25
little-disturbed fossil assemblages, but are packed in excess of likely living densities and are visually dominated by successive
generations of the infaunal bivalve Raeta undulata, which colonized a slowly aggrading muddy seafloor.
The sedimentological and paleontological evidence presented here not only supports the results of previous studies, which
demonstrate the value of condensed shell concentrations in the identification of stratigraphically significant surfaces, but also
provides adds to the distinctive features of these shell beds.
D 2004 Elsevier B.V. All rights reserved.
Keywords: Early Pleistocene; Shell concentrations; Taphonomy; Mollusk paleoecology; Sequence stratigraphy; North-central Ecuador
1. Introduction Tschopp (1948) the Jama Formation comprises upper

Miocene strata, whereas the geologic explications
An extremely continuous sedimentary succession, available for the published geological maps (Sheet of
mainly composed of cyclic shallow-marine strata Jama at 1:100.000 scale and National Geological Map
variably interbedded with non-marine and marginal- of the Republic of Ecuador at 1:1.000.000 scale), refer
marine sediments, is well exposed along the coastal the sediments to the Onzole Formation (Middle–Late
cliff to the west of the town of Jama (north-central Miocene) and to the Borbon Formation (Late Mio-
Ecuador) (Fig. 1). Between Punta Cabuya to the south cene–Early Pliocene) of the Daule Group.
and Punta Alcatraz to the north, it crops out following The present work is part of a larger research
a gentle, fairly constant, northeast-dipping monocline. project focused on unraveling the paleoenvironmen-
Extensive exposures on the 10-km-long coastal cliff tal history of the marine Plio-Pleistocene strata of
allow the stratigraphy to be traced laterally in great Ecuador (Landini et al., 1991, 2002a,b; Bianucci et
detail at least in the section between Punta Cabuya al., 1993a,b, 1997a,b; Bisconti et al., 2001; Ragaini
and the village of El Matal. On the contrary, cliff faces et al., 2002; Di Celma, 2002; Di Celma et al., 2002;
are poorly exposed southward of Punta Cabuya owing Cantalamessa and Di Celma, 2004). Its aims are to:
to the intense activity of landslides and a luxuriant (i) constrain the middle unit of the Jama Formation
vegetative cover. Here, outcrops are limited to those (the Punta Ballena Member) within a sequence
occurring on the present-day wave-cut platform, and stratigraphic framework by means of integrated
are almost completely lacking south of Punta Pasa paleontological, sedimentological, and stratigraphic
Borracho, where the wave-cut platform is usually observations of mollusk concentrations associated
buried beneath modern beach sands. with key stratal surfaces; (ii) provide adds to the
Pilsbry and Olsson (1941) proposed the name Jama distinctive features of these shell beds. The scarcity
Formation for the entire succession exposed in the sea- of Early Pleistocene shallow-marine deposits pre-
cliffs that extend from the neighbourhood of Jama served onshore in Ecuador further enhances the
southward to Punta Cereza (Cabo Pasado; 80829VW, interest of this locality where strata of this temporal
0822VS) and, on the basis of a very rich marine span are well exposed and easily accessible.
molluscan fossil assemblage, referred these strata to a
generic Pliocene age. To date, apart from Whittaker
(1988) who assigned an early Pliocene age (N18/19) to 2. Geological framework
this formation on the basis of the foraminiferal
assemblage occurring within samples collected south Convergence between the Pacific oceanic plates and
of Punta Pasa Borracho, the biostratigraphy and timing South America has been active at least since Creta-
of deposition have been poorly constrained. Owing to ceous time and has resulted in successive accretions of
the difficulty of obtaining useful biostratigraphic allochthonous terranes onto the previous continental
markers from these shallow-marine deposits (age- margin (Feininger and Bristow, 1980; Feininger, 1987;
diagnostic planktonic foraminiferal fauna are absent Van Thournout et al., 1992; Kerr et al., 2002). West of
or very scarce), many conflicting age determinations the Dolores–Guayaquil Megashear zone (DGM), the
have been proposed. For example, according to basement is a composite and complexly deformed Late
G. Cantalamessa et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 216 (2005) 1–25 3
Fig. 1. (a) Schematic structural and location map of the Ecuadorian coast showing position of the study area (boxed part) with respect to the
place where the Carnegie Ridge impinges on the Ecuador Trench (modified from Lonsdale and Klitgord, 1978; Baldock, 1982; Rosania, 1989).
The Nazca–South America convergence vector from Trenkamp et al. (2002). (b) Map of the Jama area. Locality names used in the text and
figures are those reported on the second edition of the Ecuadorian IGM scale 1:50.000 bJamaQ.
Cretaceous–Early Eocene suite of oceanic terranes, the Plio-Pleistocene siliciclastic successions. The Jama
known as the Piñon–Macuchi Block, which accreted Formation was deposited within the Esmeraldas–
within a dextral shear regime onto the oceanic Caraquez Basin (Rosania, 1989) (Fig. 1a), which
Pallatanga Terrane during the Late Eocene (Jaillard et occupies a forearc position east of the modern
al., 1995; Hughes and Pilatasig, 1999). The Ecuadorian convergent South America–Nazca plate boundary
coastal block of oceanic substratum is the southernmost and separated from the Manabı̀ Basin to east by the
leading area of the North Andes microplate, which is Jama Hills. This roughly north-trending part of the
bounded from the Nazca Plate to the west by the Coastal Range is a structurally complex, Late Mio-
Ecuador trench, and from the South American Plate to cene–Early Pliocene uplifted area located west of the
the east by a system of northeast-trending, right-lateral Jama–Quinindé fault and consists chiefly of a base-
strike-slip faults and north trending thrust faults (the ment-cored series of Paleogene and Miocene sedimen-
DGM) (Trenkamp et al., 2002). The southern extremity tary rocks that were important source terrains for the
of the DGM is commonly thought to intersect the Jama Formation. Although poorly known, the Jama
Ecuador trench in the region of the Gulf of Guayaquil Formation is one of the thickest accumulations of
(Moberly et al., 1982). mainly shallow marine Quaternary strata exposed on
During the last part of Miocene and Early Pliocene, land in Ecuador and its outcrops represent a critical
the western margin of central Ecuador underwent interval and a valuable reference section where
extensive compressive deformation (Whittaker, 1988; environmental changes and tectono-sedimentary evo-
Daly, 1989). This produced the progressive uplifting of lution that affected this part of South America during
the Coastal Range, the shallowing upward trend and the Early Pleistocene can be investigated.
emerging of the Progreso (Benı́tez, 1995) and Manabı̀
basins, and the shifting of the marine sedimentation
into coastal basins sited farther west (Baldock, 1982). 3. Material and methods
At the end of the Early Pliocene, an extensive
planation surface, cutting across the older relief The wide range of facies occurring within the
(Coastal Range and central Ecuadorian Cordillera) studied succession have been distinguished on the
and testifying to the cessation of uplift, was created basis of detailed field observation. For this purpose
throughout Ecuador (Coltorti and Ollier, 1999, 2000). special attention has been paid to physical and
Evidence of Late Pliocene collapse of the forearc biogenic sedimentary structures, grain size, and
accretionary wedge produced by trench-normal exten- paleontological content. Fossils were collected both
sion is provided by the presence of north–south by manual picking at outcrops (for large-bodied and
trending normal faults in northern Ecuador (Aalto rare species) and via bulk-samples (about 30 dm3 each)
and Miller, 1999) and by the sudden eastward marine to obtain quantitative data allowing a comparison
ingression into the continent recorded by the strik- among the different stratigraphic levels. The former
ingly angular juxtaposition of the late Pliocene Canoa are excluded from abundance/dominance computa-
Formation upon the Miocene Tosagua Formation in tions as their presence and interpretation rests on
central Ecuador (Di Celma et al., 2002). Here, the syn- qualitative/semi-quantitative observations carried out
tectonic Late Pliocene–Pleistocene shoaling upward in the field. The bulk samples were washed through a
succession occurring south of Cabo San Lorenzo 1.0 mm-mesh sieve, and coarser bioclasts were picked.
(Canoa and Tablazo Formations), constrains the onset Mollusk specimens were identified taxonomically to
of forearc uplift to the end of the Pliocene (Di Celma species level when possible; the conservative choice to
et al., 2002; Cantalamessa and Di Celma, 2004). leave a fair number of taxa at the generic rank is due to
Pliocene to Pleistocene marine sedimentary rocks in the general shortage of systematic information cover-
Ecuador are chiefly preserved in offshore depocenters ing fossil and modern mollusks of the East Pacific.
(Deniaud et al., 1999). However, while sedimentation Because many taxa remain mainly unrevised, attempts
still continues beneath modern sea level, some margins at a more detailed systematic treatment may result in
of these coastal basins have undergone gentle Pleisto- misidentification rather than useful data. Mollusks
cene uplift, resulting in excellent coastal exposures of were counted following the procedures suggested by
Table 1 Table 1 (continued)

Faunal list of assemblages from current/wave winnowed and Taxa Current/wave Community
community shell beds s.b. s.b.
Taxa Current/wave Community
A D A D
s.b. s.b.
Bivalvia
A D A D Panopea cf. coquimbensis – – 2 0.2
Bivalvia Pholas chiloensis – – 1 0.1
Adrana cf. sowerbyana 3 0.11 – – Pitar paytensis 14 0.5 10 0.96
Adrana sp. 4 0.14 2 0.2 Pitar sp. 3 0.11 7 0.67
Aligena cokeri 1 0.03 – – Psammotreta sp. – – 7 0.67
Anadara aequatorialis 12 0.42 67 6.42 Raeta undulata 2 0.07 247 23.68
Anadara nux 31 1.1 45 4.31 Sanguinolaria bertini X X – –
Anadara obesa 25 0.88 25 2.4 Semele guaymasensis 1 0.03 – –
Anomalocardia callistoides 1 0.03 – – Solecardia sp. 1 0.03 – –
Anomia peruviana 5 0.18 – – Solecurtus broggii 3 0.11 6 0.58
Arcidae indet. 12 0.43 15 1.44 Solen spp. 26 0.92 3 0.29
Bornia cf. venada – – 6 0.58 Strigilla dichotoma 35 1.24 – –
Cardiomya ecuadoriana 1 0.03 – – Strigilla interrupta 3 0.11 – –
Carditamera cf. radiata 1 0.03 – – Tagelus peruvianus X X – –
Carditamera sp. 1 0.03 – – Tagelus spp. 20 0.71 12 1.15
Chione gorgona 1 0.03 10 0.96 Tellina aequizonata 4 0.14 49 4.7
Chione jamaniana 69 2.44 14 1.34 Tellina cf. hertleini – – 30 2.88
Chione traftoni X X – – Tellina laplata X X – –
Chione venadensis X X – – Tellina spp. 75 2.66 14 1.34
Chione spp. 72 2.55 6 0.58 Temnoconcha cognata 17 0.6 38 3.64
Corbula amethystina 2 0.07 – – Trachycardium procerum – – 9 0.86
Corbula cf. nasuta 63 2.23 – – Trachycardium sp. 2 0.07 – –
Corbula cf. nuciformis 82 2.9 – – Trigonocardia granifera X X – –
Corbula porcella 349 12.37 – – Trigonocardia obovalis 191 6.77 4 0.38
Corbula tenuis 60 2.13 9 0.86 Subtotal 1717 60.84 821 78.69
Corbula spp. 85 3.01 8 0.76
Cyathodonta sp. 1 0.03 – – Gastropoda
Cyclinella sp. – – 4 0.38 Acteocina puruha 36 1.28 1 0.1
Galeommatacea indet. 4 0.14 – – Acteocina cf. tabogaensis 1 0.03 6 0.58
Grandiarca grandis X X 8 0.76 Architectonica nobilis 2 0.07 8 0.76
Harvella elegans 7 0.25 X X Asperiscala minuticosta 2 0.07 – –
Linga cancellaris – – X X Asperiscala obtusa 1 0.03 – –
Lucina centrifuga 43 1.52 – – Balcis sp. 20 0.71 – –
Lunarca brevifrons 10 0.35 25 2.4 Calyptraea cf. mamillaris 42 1.49 – –
Macoma cf. grandis – – 45 4.31 Cancellaria metuloides X X 5 0.48
Macoma lamproleuca X X 28 2.68 Cancellaria cf. picta X X 4 0.38
Mactra (Micromactra) sp. – – 8 0.76 Cancellaria solida – – X X
Mactrellona sp. 1 0.03 – – Cancellaria sp. – – 3 0.29
Martesia striata 2 0.07 – – Circulus occidentalis 5 0.18 2 0.2
Montacutidae indet. 1 0.03 – – Circulus sp. 12 0.42 1 0.1
Mulinia pallida 63 2.23 12 1.15 Crepidula sp. 144 5.1 2 0.2
Mysella compressa 32 1.13 7 0.67 Conus cf. cacuminatus – – 4 0.38
Noetia reversa 4 0.14 8 0.76 Conus patricius – – 5 0.48
Nucula exigua 128 4.54 10 0.96 Conus sp. – – 2 0.2
Nuculana eburnea 8 0.28 12 1.15 Cymatophos galerus – – 7 0.67
Nuculana elenensis 48 1.7 – – Depressiscala cf. polita 1 0.03 – –
Nuculana sp. 3 0.11 Epitonium spp. 17 0.6 3 0.29
Orobitella cf. margarita 3 0.11 – – Eulima sp. 10 0.35 2 0.2
Orobitella cf. zorrita 1 0.03 – – Hanetia cf. boggsi X X 4 0.38
Pacipecten tumbezensis 81 2.87 8 0.76 Hanetia cymioides X X 12 1.15
Pandora sp. – – X X (continued on next page)
Table 1 (continued) Di Geronimo and Robba (1976). Faunal compositions

Taxa Current/wave Community are provided in Table 1 with respective abundance
s.b. s.b. (number of specimens per taxon) and dominance
A D A D (proportion pertaining to each taxon). Terms used to
Gastropoda describe bioclastic fabrics and other shell bed features
Hanetia cf. ecuadorensis – – 8 0.76 generally follow Kidwell (1991a) and Kidwell et al.
Hespererato panamensis 1 0.03 – – (1986).
Hirtoscala mitraeforme 2 0.07 – –
Hirtoscala cf. replicata 1 0.03 – –
Marsupina nana – – 7 0.67
Melongena sp. – – 1 0.1 4. Physical stratigraphy
Nassarius gemmulosus 85 3.01 3 0.29
Natica elenae 3 0.11 10 0.96 The Pleistocene siliciclastic fill of the eastern
Natica cf. inespeectans 1 0.03 – –
Natica othello 23 0.82 – –
margin of the Esmeraldas–Caraquez Basin (Jama
Naticidae indet. 136 4.82 – – Formation) is a variable assemblage of marine to
Niso imbricata – – 4 0.38 continental strata. On the basis of sedimentary facies
Niso sp. 13 0.46 1 0.1 associations, it can be subdivided into three main
Nitidiscala cf. durhamiana 3 0.11 1 0.1 formal units that, in stratigraphic order are: (1) the
Northia northiae X X 2 0.2
Oliva incrassata – – 31 2.98
~20-m-thick, marine, Punta Pasa Borracho Member
Oliva sp. 1 0.03 15 1.43 exposed between Punta Pasa Borracho and Punta
Olivella semistriata X X – – Cabuya; (2) the ~100-m-thick, mixed fluvial-marine
Olivella spp. 398 14.1 18 1.73 Punta Ballena Member, exposed extensively from
Polinices intemeratus – – 4 0.38 Punta Cabuya to Punta Alcatraz; (3) the ~60-m-thick,
Polinices otis – – 10 0.96
Polinices cf. panamensis 1 0.03 7 0.67
fluvial El Matal Member, exposed between Punta
Polinices sp. 1 0.03 – – Alcatraz and the village of El Matal. The fluvial strata
Prunum cf. curtum X X – – of the El Matal Member are intercalated with several
Prunum sapotilla 5 0.18 – – tephra horizons up to 50 cm thick. The age of the
Ringicula sp. 4 0.14 – – Jama strata under consideration in the present study
Semicassis centiquadrata – – 8 0.76
Semicassis sp. – – X X
(the Punta Ballena Member) has been determined
Skeneopsis sp. 4 0.14 – – from 40Ar/39Ar dating of the lowermost of these
Sincola ecuadoriana X X – – volcanic layers at Punta Alcatraz, where an ash bed ~8
Strioterebrum cf. camaronense X X 1 0.1 m above the contact between the Punta Ballena and El
Strioterebrum sp. X X 2 0.2 Matal Members yielded a weighted mean radiometric
Strombinophos loripanus – – 5 0.48
Teinostoma sp. 73 2.58 X X
age of 1.16F0.06 Ma.
Terebra elenae – – 1 0.1 The paleontological and sequence stratigraphic
Turbonilla spp. 58 2.05 – – analysis reported in this paper focus on the Punta
Turritella pasada – – 4 0.38 Ballena Member of the Jama Formation (Fig. 2). The
Subtotal 1106 39.13 214 20.57 limited exposure afforded by the modern abrasion
Scaphopoda
platform and by faulting and folding preclude the
Cadulus sp. – – 2 0.2 reconstruction of a comparably well-defined, reliable
Dentalium cf. oerstedi – – 4 0.38 physical stratigraphic architecture for the underlying
Dentalium sp. – – 1 0.1 Punta Pasa Borracho Member.
Subtotal – – 7 0.68
TOTAL 2823 99.97 1042 99.94

4.1. Depositional sequences in the Punta Ballena
Member
A=abundance (number of specimens per taxon); D=dominance
(proportion pertaining to each taxon); X=taxa collected only by
manual picking at outcrop. The Punta Ballena Member consists of eight
superposed, unconformity-bounded depositional
sequences or cyclothems, numbered successively
from PB1 to PB8 (Fig. 2), applying conventional (forestepping shelf wedge, FSW) whose lower contact
sequence stratigraphic concepts developed for pas- on the BSB (downlap surface, DS) marks the turn-
sive-margin settings by researchers at Exxon Produc- around from transgressive to regressive strata (Abbott,
tion Research (e.g. Vail et al., 1977; Wilgus et al., 1997, 2000).
1988; Van Wagoner et al., 1990). This analysis The back-barrier wedge, emplaced within incised-
requires identification of key stratal unconformities valleys and landward of the contemporary ravinement
and paleoenvironmental interpretation of the interven- surface, consists of fluvial to estuarine deposits that, in
ing sediments. Toward this end our fieldwork has many places, pass upward into lagoon/marsh mud-
focused primarily on detailed analysis of physical stone. The incised valleys were infilled during a
sedimentary structures. However, although many relative sea-level rise and so most likely their deposits
environments of deposition and surfaces were obvious (incised-valley fill, IVF) belong genetically to the TST
on the basis of sedimentology alone, we have also (e.g. Allen and Posamentier, 1993). This interpretation
taken advantage of the paleoecological and tapho- is corroborated by the vertical succession of deposi-
nomic attributes of macrobenthic fossil assemblages tional environments within the IVF of sequence PB1
and the ichnology of potential discontinuity surfaces. (fluvial sediments at the base, passing upward into
As largely demonstrated during the last decade by progressively more distal, lagoonal deposits at top),
several works (e.g. Beckvar and Kidwell, 1988; which is transgressive in character. The backstepping
Banerjee and Kidwell, 1991; Kidwell, 1991a,b; shelf wedge above the RS is a deepening- and fining-
Savrda, 1991; Fürsich and Oschmann, 1993; Pember- upward facies succession of shoreface sandstones,
ton and MacEachern, 1995; Brett, 1995, 1998; Abbott including a basal carbonate-rich lithofacies which
and Carter, 1997; Naish and Kamp, 1997; Abbott, grades upward into inner shelf, massive, fine-sandy
1997, 1998; Kondo, 1997; Kondo et al., 1998; Fürsich siltstones topped by a richly fossiliferous interval. In
and Pandey, 1999; Del Rı́o et al., 2001; Dominici, sequences where the strata of the backstepping shelf
2001; Buatois et al., 2002; Di Celma et al., 2002; wedge sit directly upon the truncated sediments of the
Fürsich and Pandey, 2003), these kinds of data subjacent depositional sequence, the RS is superposed
provide a wealth of valuable information on substrate on, and coincides with, the SB. The resulting surfaces
consistency, water depth, shelf depositional processes, are known as E/T surfaces, for the superposition of
and sedimentary dynamics (e.g. sediment starvation erosion and transgression (Plint et al., 1986). Within
versus bypassing versus erosion) at sequence-bound- the Punta Ballena Mbr, ravinement surfaces are
ing and intrasequence surfaces. erosional surfaces displaying small-scale relief and,
The total thickness of individual depositional in some cases, Thalassinoides burrows that indicate
sequences ranges from ~5 to 25 m. An ideal sequence firmground conditions. Although the uppermost por-
of the Punta Ballena Member is composed of a tion of the HST is usually truncated by the superjacent
transgressive systems tract (TST) and a highstand unconformity, depositional sequences PB5 and PB6
systems tract (HST) (Fig. 2). The HST can be divided include TST and HST. The other sequences lack HST
into a back-barrier wedge (BBW), which is interposed sediments because of the erosion at the superjacent
between a basal type-1 sequence boundary (SB) and sequence boundary.
the ravinement surface (RS), and a backstepping shelf
wedge (BSW) above the RS (sensu Thorne and Swift, 4.1.1. Facies architecture
1991). Usually, the BSW consists of an expanded Sedimentary structures, bioturbation, grain size and
siliciclastic core, jacketed by hiatal shell concentra- macrofossil content reveal 11 main sediment facies
tions at the base (onlap shell bed, OSB) and at the top associations, representing inner-shelf to subaerial
(backlap shell bed, BSB) whose accumulation has depositional settings; their principal characteristics
been accompanied by a significant reduction in are summarized in Table 2.
deposition of terrigenous sediment. The BSB lies At Punta Cabuya the PB1 sequence consists of
disconformably on a heavily bioturbated interval trough cross-bedded non-marine conglomerates and
(local flooding surface, LFS) (Abbott, 1997). The minor pebbly sandstones resting in angular uncon-
HST is composed of a shell-poor silty mudstone formity on massive, bioturbated, blue- and light-
Table 2
Description and interpretation of the main facies associations observed in the Punta Ballena Mbr of the Jama Formation
Name Code Description Depositional environment
Non-Marine Nm1 Sharp-based, massive to trough cross-stratified Tidal–fluvial channel and/or
conglomerate composed of rounded extraformational distributary channel.
pebbles and cobbles, and angular, rip-up sandstone pebbles,
overlain by variably bioturbated (Thalassinoides),
pebble-rich, fine- to coarse-grained sandstone and
medium-grained cross-bedded sandstone. Rip-up clasts are
concentrated at the bottom, where they prevail on extra
formational clasts.
Nm2 Well sorted, up to 3-m-thick sets of coarse-grained, trough Stream bars of a braided fluvial
cross-stratified gray sandstone with sparse pebbles. Barren system
of molluscan remains.
Nm3 Medium-scale inclined heterolithic strata composed of Point bars within meandering
coarse- to fine-grained, sandstone and mudstone laminae. channels of a tidally influenced
Internally, this facies association shows multiple, fluvial-estuarine delta
subparallel concave accretion surfaces.
Marginal-Marine Mm1 Massive or centimeter- to decimeter-scale beds of greenish- Lagoon or salt marsh
black mudstone. The lower contact is gradual, whereas the
upper is sharp and erosive. Barren of molluscan remains.
Open-Marine Om1da Erosionally soled alternation of thin shell beds and silty Partially starved, current/wave-
sandstone grading upward into less fossiliferous, dominated upper-shoreface sea-floor
bioturbated, medium-grade sandstone. Shell beds are clast-
to matrix-supported (dispersed current/wave-winnowed
shellconcentration).
Om1sb Erosionally soled, well-sorted, trough cross-stratified and Starved, current/wave-dominated
medium to very coarse-grained shell-rich sands with upper-shoreface sea-floor
occasional pebbles. Typically arranged in 15–45 cm thick
sets (simple current/wave-winnowed shell concentration).
Om2 Low angle, cross-stratified medium siliciclastic sands. Wave-dominated upper-shoreface
Sparsely fossiliferous. Biogenic features are dominated by
the Skolithos ichnofacies and are manifest by the
ichnogenera Skolithos, and by vertical and deep penetrating
Ophiomorpha nodosa.
Om3 Massive, intensely bioturbated very fine to fine siliciclastic Wave-dominated lower shoreface
sandstone. Sparsely fossiliferous.
Om4 Massive, pervasively bioturbated silty fine-sandstone. Shoreface to inner shelf
transition zone
Om5 Massive, fine-sandy silt or silty clay thoroughly Mid-inner shelf
homogenised by biogenic bioturbation. Poorly and sparsely
fossiliferous with rare carbonized wood chips.
Om6b Similar to Om5 but with abundant in situ infaunal bivalves Starved mid-inner shelf
and Thalassinoides burrows.
a
Current/wave-winnowed bed (onlap shell bed).
b
Community bed (backlap shell bed).
brown-colored muddy sandstone and overlain by The entire marine part of the first four depositional
about 2 m of greenish-black mudstone, which are sequences (PB1, PB2, PB3, PB4) is composed almost
marginal-marine (lagoonal) in origin (Fig. 3). exclusively of bluish to yellow, intensely bioturbated
Fig. 2. Composite stratigraphic section for the Lower Pleistocene sedimentary succession exposed between Punta Cabuya and Punta Alcatraz
(Punta Ballena Member). The eight depositional sequences identified are subdivided into component systems tracts and tentatively correlated
with the astronomically tuned oxygen isotope time scale of Shackleton et al. (1990). Since a single sequence and associated lower boundary
should represent a glacial–interglacial stage couplet, they seem to correspond to the record of oxygen isotope stages 35–49 defined in deep sea
deposits. Facies association codes refer to Table 1.
lower shoreface sandstone containing fossil remains vertical burrows 4–8 mm in diameter assigned to the
scattered or concentrated along the base. The ichno- ichnogenus Skolithos, and horizontal Thalassinoides
logical suite, which may be regarded as a Skolithos networks (Pemberton et al., 1992). Between Punta
assemblage, includes vertical shafts of fully lined, Cabuya and La Cabuya, the marine deposits of
knobby walled burrows up to 4 cm in diameter of the sequence PB3 are underlain by trough-cross bedded
ichnofossil Ophiomorpha nodosa, rare unbranched non-marine pebbly coarse sandstones that are strik-
ingly similar to those composing the fluvial facies sequence was probably emplaced during interglacial
assemblage of the El Matal Mbr (Fig. 3). oxygen isotope stage 35 and the Punta Ballena Mbr
The remaining four sequences of the Punta Ballena started to accumulate about 1.5 Ma BP, during
Mbr (namely, sequences PB5, PB6, PB7, PB8) are interglacial oxygen isotope stage 49.
composed of fining-upward facies successions with
fossiliferous and strongly cemented trough cross-
bedded sandstones at the base, followed by perva- 5. Shell concentrations
sively bioturbated bluish siltstone with sparse frag-
ments of carbonized wood and pristine molluscan Shell concentrations have been defined by Kidwell
remains scattered or concentrated in conspicuously (1991a) as concentrations of biomineralized remains
dense patches. The RS at the base of sequence PB5 is more than 2 mm in size from any invertebrate animal.
underlain by laterally discontinuous sets of tidally Those in the Punta Ballena Mbr recur systematically
influenced estuarine-fluvial strata (incised-valley fill), in a well-defined basal and mid-cycle stratigraphic
consisting of well-developed inclined heterolithic position within depositional sequences. Based on
units that are incised into the subjacent depositional taxonomic composition and taphonomic attributes of
sequence. their individual components (i.e. articulation versus
disarticulation, abrasion, bioerosion, orientation,
4.1.2. Controls on sequence development encrustation, degree of fragmentation and sorting by
Shallow-marine and coastal strata of the Punta size and shape), these shell beds are interpreted to
Ballena Mbr are arranged in a high-frequency cyclic represent two types of concentrations, each showing a
stacking pattern and, using the dated tephra for markedly different mode of formation: current/wave-
guidance, are likely to be Early Pleistocene in age. winnowed shell beds and community shell concen-
The origin of sea-level oscillations that generate trations (Meldahl, 1993) (Table 3).
depositional sequences is difficult to assess with the The skeletal remains contained within the marine
limited data at hand. However, because of the deposits of the Jama Formation are predominantly
extremely fast rates of eustatic sea-level changes molluscan but also include abundant fish otoliths,
during the Plio-Pleistocene, which would have had selachian teeth, bryozoans, barnacle plates and frag-
the effects of swamping the long-term rates of vertical ments of echinoid tests (Cantalamessa et al., 2001,
tectonic movements, it is safe to assume that sequence 2002). Although of little use for age determination,
development was controlled by eustasy, and that this many species of the molluscan fauna recovered within
was related to the orbital time series of obliquity, the this formation are still extant, and, therefore, represent
dominant periodicity during late Pliocene and Early a valuable source of paleoecological information.
Pleistocene (Shackleton et al., 1990, 1995).
Figure 2 is our attempt to correlate these small- 5.1. Current/wave-winnowed shell beds
scale depositional sequences directly with the oxygen
isotope sea-level index of Early Pleistocene. Although Shell beds categorized as current/wave-winnowed
tentative, this attribution is consistent with the always occur at the base of the transgressive shelf
numeric age obtained from the dated volcanic ash wedge (BSW) and each can be traced for hundreds to
layer (1.16F0.06 Ma), which implies that the PB8 thousand of meters laterally along the outcrop. These
Fig. 3. Detail of (a) the trough cross-stratified conglomerate at the base of sequence PB1 and (b) of the angular unconformity between the Punta
Pasa Borracho Mbr and the Punta Ballena Mbr. (c) Mosaic of outcrop at Punta Cabuya. SB/TS1 is the sequence boundary coincident with the
transgressive surface at the base of sequence PB1. The back-barrier wedge (BBW), resting on the SB/TS1 and topped by RS1, fines upward and
is a composition (from bottom to top) of conglomerate, pebble-rich sanstone, cross-bedded sandstone with variable amounts of bioturbation
(Thalassinoides) (facies association Nm1), and mudstone (facies association Mm1) interpreted as fluvial, tidally influenced fluvial and lagoon/
marsh deposits erosionally overlain by wave-dominated shoreline strata (facies association Om3) (BSW, backstepping shelf wedge). (d)
Panoramic view of the lower portion of the Punta Ballena Mbr. Note the planar nature of sequence stratigraphic key surfaces. (e) Close-up of the
5-m-thick transgressive incised-valley fill at the base of sequence JF3. The fill comprises fluvial, trough cross-stratified grayish sandstone with
sparse pebbles (facies association Nm2) and is sharply overlain by the facies association Om3.
12
Table 3
Summary of the characteristic features of mollusk shell concentrations from the Punta Ballena Mbr of the Jama Formation
Depositional Shell bed Associated Genetic Thickness Biofabric Shell Trophic (b) Upper contact;
sequence type stratal classificationa (m) condition characteristics (a) Lower contact
attenuation
PB2 Current/ Onlap Sedimentologic 0.8 Closely packed to dispersed. Fresh to variably Highly diverse (b) Gradational;
G. Cantalamessa et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 216 (2005) 1–25

wave-winnowed Shells are mostly preserved broken, abraded concentration (a) Sharp, erosion soled
(dispersed) aligned with the bedding and and rounded. (104 taxa) dominated
oriented both randomly and Low to high by infaunal
convex-up fragmentation, suspension feeders
minor encrustations
and common bioerosion
PB4 Current/wave- Onlap Sedimentologic 0.05 Closely packed and randomly Mainly finely N/A (b) Gradational;
winnowed oriented comminuted shell (a) Sharp, erosion soled
(simple) debris. Broken and
abraded shells are also
present
PB5 Current/wave- Onlap Sedimentologic 3 Well-sorted, trough cross- Ditto N/A (b) Rapidly gradational;
winnowed stratified, coarse to finely (a) Sharp, erosion soled
(simple) comminuted shell debris
with a discontinuous,
parautochthonous oyster
bank at the top
PB6 Current/wave- Onlap Sedimentologic 5 Ditto Ditto N/A (b) Rapidly gradational;
winnowed (a) Sharp, erosion soled
(simple)
PB7 Current/wave- Onlap Sedimentologic 4 Well-sorted, trough cross- Ditto N/A (b) Rapidly gradational;
winnowed stratified, coarse to finely (a) Sharp, erosion soled
(simple) comminuted shell debris
PB8 Current/wave- Onlap Sedimentologic 3 Ditto Ditto N/A (b) Rapidly gradational;
winnowed (a) Sharp, erosion soled
(simple)
PB5 Community Backlap Biogenic ~0.40 High percentage of conjoined Mainly freshly Diverse concentration (b) Rapidly gradational;
specimens either in life preserved whole shells (82 taxa) dominated (a) Variably Thalassinoides
position or with butterflied by infaunal burrowed, rapidly gradational
valves suspension feeders.
Deposit-feeders and
carnivores are
common
PB6 Community Backlap Biogenic ~0.40 Ditto Ditto Ditto (b) Rapidly gradational;
(a) Variably Thalassinoides
burrowed, rapidly
gradational
a
From Kidwell et al. (1986).
shell beds have erosional bases and overlie sediments aligned with bedding. Occasionally, pebbles and
of a distinctly different facies, and therefore are large bone fragments are found in the skeletal
associated with significant erosion. In addition, their assemblage (Fig. 4).
upper contacts are rapidly gradational. On the basis of At the top of the shell bed located at the base of the
grain size, bioclast composition, and internal archi- PB5 sequence is a discontinuous oyster bank up to
tecture, they are classified into two subtypes: (1) 60–70 cm thick and a few meters in length (Fig. 4c).
simple; and (2) dispersed. These fossils probably represent a new species of the
genus Crassostrea (Ragaini et al., 2004). Specimens
5.1.1. Taphonomic features of simple current/wave- constitute a bioclast-supported monospecific assem-
winnowed shell beds blage where individuals of large size (up to 40 cm in
Thickness of the basal coquinas occurring within height) are densely packed in an unconsolidated sandy
sequences PB4, PB5, PB6, PB7, and PB8 changes matrix without any distinguishable attachment area. A
significantly from one cycle to another. They may few among the conjoined oysters held the upright
be up to 5 m thick and characterized by trough growth position, but most are tilted or dislodged to
cross-stratification, or instead be only a few oblique or nearly horizontal position (Fig. 4d).
centimeters thick and internally structureless. As a According to Jimenez et al. (1991), individuals that
rule, these concentrations are composed of well- tip over after a period of upright growth and revert to
ordered and -rounded convex-up valves, silt-filled upright growth result in particular morphology (i.e.
internal moulds, large heavy-shelled oysters, com- cup-shaped). Taking into account the taphonomic
minuted shell debris and well-sorted, bioclast- evidence previously mentioned and the spatulate shell
supported disarticulated and abraded shells that are shape of our individuals, we infer that the oblique/
Fig. 4. Details of simple current/wave winnowed shell bed (facies association Om1s). (a) Well-sorted, trough cross-stratified and medium to very
coarse grained shell-rich sands containing (b) sporadic vertebrate remains (bedding view). (c) Discontinuous concentration of large specimens of
Crassostrea at the top of facies association Om1s. Sequence PB5, Punta Ballena. Knife (in brackets) 12 cm long. (d) Tilted or dislodged
individuals of Crassostrea sp. nov. from vertical to oblique or nearly horizontal position.
horizontal positions reflect a post-mortem dislodging

due to wave/current sweeping action. However, the
presence of both articulated and disarticulated speci-
mens displaying broken but nearly whole valves (i.e.
each piece of the shell is juxtaposed with the
surrounding ones) together with the general good
preservation of the shells allow us to infer minimal to
no post-mortem transport.
Simple current/wave-winnowed shell beds are
invariably strongly cemented and therefore they are
able to form small headlands where they intersect the
beach (e.g. Punta Ballena, Punta La Cereza, Punta
Alcatraz). For the same reason, molluscan remains are
poorly preserved, and impossible to extract in a
suitable manner for detailed taxonomic determination.
5.1.2. Taphonomic features of dispersed current/

wave-winnowed shell beds
This subtype of current/wave-winnowed shell bed
is present only at the base of sequence PB2, which
is exposed along the cliff between Punta Cabuya
and La Cabuya (Fig. 5). It occurs as multiple, thin
shell beds (from one-valve-thick to 35 cm) within a Fig. 5. Details of dispersed current/wave-winnowed shell bed
thicker, weakly lithified, massive and extensively (facies association Om1d). (a) Cross-section of the multi-event
bioturbated silty sand interval that is about 80 cm shell concentration at the base of sequence PB2 (La Cabuya). Note
the dispersing-upward biofabric trend. Pen is 15 cm long. The wavy
thick. Its upper contact is gradational, whereas the line indicates the sequence boundary. (b) Bedding plane view of the
lower contact is sharply erosional on the fine- same shell bed. Note the mostly concave-down orientation of shells.
grained sediments of the underlying sequence.
Individual shelly intervals thin upward and are both
shell- and matrix-supported. Bivalve shells, domi- 5.1.3. Dominant faunal elements of dispersed current/
nantly disarticulated, are preserved aligned with the wave-winnowed shell beds (Macoma Association)
bedding and oriented both randomly and convex-up; The mollusk assemblage is highly diverse (104
stacked arrangements with concave-down valves taxa) and composed of 62 bivalve and 42 gastropod
and nesting of shells are common in the lowermost taxa. Some characterizing faunal elements at the
and thickest interval, where shell density is higher. outcrop level were not present in the bulk samples
The biogenic hardparts of this concentration show a (e.g. Macoma lamproleuca, Sanguinolaria bertini,
variable state of preservation from pristine whole Northia northiae) or are represented only by scarce
shells to shell debris. The specimens are not usually fragments (e.g. Temnoconcha cognata). Based on the
found in life position and display, in general terms, pooled quantitative bulk sample, bivalves (54 taxa)
heavy signs of abrasion, low to high fragmentation, constitute 60.84% of individuals, whereas gastropods
rare encrusters (barnacles, bryozoans, calcareous (32 taxa) 39.13%. The rest of the fossil fauna consists
algae, etc.) and common bioerosion traces (clionid of crab pincers, barnacle plates, bryozoans, test
sponges, spionid polychaetes, etc.). Abrasion of the fragments of regular and irregular echinoids, and
openings of most endolith borings suggests the abundant fish otoliths; scaphopods are absent. Almost
borings were acquired before the shells were all taxa are represented by juvenile forms, and only
abraded. The rare specimens with conjoined valves some species of corbulids, mactrids, nuculids and
(Harvella elegans, for example) are dislodged from nuculanids include articulated specimens. The assem-
life position. blage is never dominated overwhelmingly by a single
species; among the most dominant taxa were Olivella noides burrowed interval, whereas their upper
spp. (14.1%), Corbula porcella (12.37%), Trigono- contacts are rapidly gradational from relatively
cardia obovalis (6.77%), Crepidula sp. (5.1%), and shell-rich to shell-poor, fine-grained lithofacies
Nucula exigua (4.54%). (Fig. 6).
Regarding the trophic structure, the largest group
corresponds to suspension feeders (62.62%), domi- 5.2.1. Taphonomic features of community shell
nated by infaunal bivalves inhabiting soft/firm sub- concentrations
strates (39.79%). Among the epifaunal component The skeletal elements of these shell concentra-
(14.95%) the most prominent species is Trigonocardia tions consist primarily of mud-supported, well-
obovalis (6.77%) followed by Crepidula sp. (5.1%) preserved mollusks most of which are in pristine
and Pacipecten tumbezensis (2.87%). Deposit feeders condition and retain original shell sheen and/or fine
account for 4.78% and mainly consist of the remain- details of ornamentation. Bioerosion and encrusta-
ing families of bivalves: nuculoids, nuculanids, tion are scarce, and signs of fragmentation and
lucinids, and tellinids. However, some tellinid species abrasion inconspicuous. Bivalves display a very
among surface deposit feeders may occasionally high percentage of conjoined specimens; most of
suspension feed and lucinids are chemiosymbiontic these are either in life position or are concave-up
deposit feeders. Carnivorous gastropods, along with and concave-down butterflied valves (i.e. valves
the bivalve Cardiomya ecuadoriana, constitute still articulated but sprung open upon death) (Fig.
27.61% of individuals, whereas herbivorous and 6); disarticulated individuals show a balanced
parasitic gastropods bear very low percentages proportion of right and left valves. The concave-
(3.18% and 1.78%, respectively). up butterflied mode of bivalve occurrence is a
Although some bivalves (Anadara grandis, Corb- powerful paleoenvironmental indicator (Allmon,
ula tenuis, Sanguinolaria bertini, Strigilla interrupta, 1985) and requires a well-defined array of bio-
Tellina laplata, etc.) are regarded as intertidal to inner stratinomic processes. In temporal order, this suite
subtidal (shoreface), a certain number of taxa indicate of processes consists of a phase of exhumation of
deeper or broader bathymetric ranges, including live infaunal bivalves, a subsequent short period of
Corbula porcella and Trigonocardia obovalis, which exposure on the sea floor during which decom-
are the most prominent species of the assemblage. The position of adductor muscle and shell gaping take
former is known today from depths of 60 to 100 m, place and finally, before the ligamental decay,
whereas the latter lives from the intertidal zone to 50 burial of shells beneath muddy blankets of sediment
m. In this case the presence of consistently disarticu- settling out from suspension. If energy conditions
lated specimens of C. porcella may be explained by are relatively high during this last phase, currents
reworking from older fossil assemblages exhumed may overturn the butterflied shells into the more
and erosionally recycled during shoreface retreat by hydrodynamically stable convex-up position, and so
the erosion of the subjacent fine-grained, deeper-water preservation concave-up implies low energy after
sediments of the underlying depositional sequence. the exhumation event, as well as relatively rapid
subsequent burial, disallowing the disarticulation
5.2. Community shell concentrations (Raeta and bioerosion/encrustation of valves. Further evi-
Association) dence indicating relatively rapid burial is the
abundance of disarticulated but unbroken epifaunal
In addition to the current/wave-winnowed beds, shells that, in spite of being out of life position,
the record of the Punta Ballena Mbr includes two apparently suffered negligible transport and are
well-defined community shell concentrations, con- preserved in a pristine, nearly fresh state. Such
taining relatively deeper water molluscan species. taphonomic attributes are consistent with autoch-
These 40-cm-thick intervals occur only within the thonous/parautochthonous assemblages (sensu Kid-
sequences PB5 and PB6, where they are hosted in well et al., 1986) and indicate minimal post-mortem
a mid-cycle position. These mollusk concentrations disturbances being related to very little reworking
overlie, and are partially encased in, a Thalassi- and scarce transportation.
Fig. 6. (a) Thalassinoides burrow system exposed just below the community shell concentration of sequence PB5 (Punta Ballena). Bedding
plane view of (b) conjoined specimens of Raeta undulata preserving life position (community shell concentration), of (c) a concave-down
butterflied specimen of Macoma lamproleuca with slightly disarticulated valves (knife is 15 cm long), and (d) a concave-up butterflied bivalve
in the same shell concentration.
5.2.2. Dominant faunal elements of community shell shellQ substrates (Hertlein and Strong, 1950). Because
concentrations more detailed information on the mode of life of R.
This is a diverse mollusk assemblage in which undulata is not available, some ecological insights
bivalves and gastropods are equivalent in number of may be deduced from the closely related western
taxa (41 each), whereas scaphopods are represented Atlantic Raeta plicatella (Lamarck) (Harry, 1969).
only by three species. The fossil remains also This taxon retains a vertical life position with an
include crab pincers, barnacle plates, bryozoans, test upward orientation of the posterior gape. The foot is
fragments of irregular echinoids and abundant atrophied, permitting only scanty movements beyond
otoliths. the shell margin. Among mactrids this feature seems
The mactrid bivalve Raeta undulata (Gould), after to be typical of species inhabiting muddier substrates,
which the association is named, is the characterizing whereas species occupying coarser bottoms have a
faunal element at the outcrop level, forming clumps better developed foot and actively burrowing lifestyle
composed of several specimens. Many individuals of (Allen, 1985). According to Harry (1969), the siphon
this species are preserved in a vertical, posterior-up length (2/3 as long as the shell) and the atrophied foot
orientation (life position) whilst others show a weak of R. plicatella suggest a deep-burrowing life habit
oblique orientation (Fig. 6). Raeta undulata is an with a scarce ability to move once life position is
infaunal suspension-feeder known today from very attained.
shallow waters (no more than 20–30 m in depth) and Although no information is available on the depth
mostly sandy substrates (Coan et al., 2000), although of burial for Raeta undulata, taking into account the
it has also been recovered from muddy and bcrushed- morphological features of Raeta plicatella and the
maximum size of our individuals of R. undulata (8 to Tellina cf. hertleini, Tellina laplata and Temnoconcha
9 cm in length), this species is probably also a deep- cognata.
burrower (sensu Stanley, 1970) or, more probably, a Arcids are epifaunal, semi-infaunal or shallow
moderate deep-burrower with a slow burrowing rate. infaunal suspension-feeders that usually live with
Consequently, if individuals of this species are the sagittal plane in vertical position; however, shell
exhumed while alive due to some substrate distur- shape (length/height ratio sensu Stanley, 1970;
bances (current scour, wave action, etc.), they are Alexander, 1993) of above mentioned arcid species
unlikely to be able to quickly reburrow themselves. In is consistent with an infaunal or semi-infaunal mode
this case some individuals may die and subsequently of life. Our specimens are frequently disarticulated,
be preserved conjoined in a horizontal (or slightly with nearly equal numbers of right and left valves, and
beak-up) and hydrodynamically stable orientation or oriented both concave-up and -down, whereas many
with disarticulated valves. Similarly, given this low conjoined individuals exhibit only minor reorientation
activity, they are probably also unable to escape burial from the life position. Many shells are well preserved
by sudden increments of sediments during life (e.g. and lack encrusters and bioerosion signs; boreholes
Kranz, 1974), leading to their preservation in life- from predatory gastropods occur only on a few
position. The preservation of most specimens of R. specimens and evidence of moderate abrasion is
undulata in life orientation suggests a low-energy mainly restricted to the smallest specimens.
environment with rare events of physical disturbances Tellinids are deep-infaunal detritus-feeders most of
having sufficient energy to exhume infaunal forms. which show a horizontal life orientation. Most of our
None of the life-positioned R. undulata show an specimens are articulated and in life position showing
inverted orientation, i.e. a vertical anterior-up posi- good preservation in general terms, i.e. unbroken
tion, usually interpreted as escape responses of valves with minimal damage by bioerosion and
infaunal bivalves to rapid sedimentation (Kondo, abrasion. Several individuals of Macoma lamproleuca
1997). If R. undulata were a rapid, high-energy display butterflied occurrences with prevailing con-
burrower, the absence of such inverted positions cave-up orientations. These attributes of arcids and
would argue against high rates of sedimentation. tellinids are consistent with a relatively stable seafloor
However, since this species was probably a slow with rare events of exhumation/erosion and mud
burrower based on other observations, the taphonomic deposition.
evidence in favor of mortality via smothering is The remaining bivalve taxa, each represented by
relatively high. The overall setting implied by the very few specimens, consist of infaunal suspension-
fauna is a seafloor affected by rare high-energy events feeders (13.61%) and very shallow infaunal detritus-
(capable of exhuming infauna and leading to butterf- feeders such as nuculids (0.96%) and nuculanids
lied specimens) and also by more common events of (1.35%). Epifaunal bivalves Pacipecten tumbezensis
mud deposition that were capable of trapping slow- and Trigonocardia obovalis are very scarce (1.14%),
burrowing infauna in their burrows. These killer as might be expected given other evidence for a soft
depositional events were still clearly infrequent, given substratum.
that the preserved fauna consists of large-bodied Gastropods show the same number of taxa as
adults: only widely spaced events in time would bivalves (41), but each gastropod taxon is represented
permit the fauna to obtain adulthood between succes- by few individuals. Among the most common, are the
sive catastrophic burial (or exhumation) events large Oliva incrassata (2.98%) and olivid species
(Johnson, 1989). belonging to Olivella (1.73%). The state of preserva-
Arcids (17.73%) and tellinids (20.22%) are another tion of gastropod shells is generally good even though
significant component of the paleocommunity. The fragmentation and abrasion appear more marked and
former are represented by Anadara aequatorialis, frequent in comparison with bivalves. This fact is
Anadaranux, Anadara obesa, Grandiarca grandis, probably related to differences among the prevailing
and Lunarca brevifrons in addition to Arcidae indet.; life habits of bivalves and gastropods; the latter, in
the tellinids are represented mainly by Macoma fact, are mostly epifaunal/semi-infaunal forms and
grandis, Macoma lamproleuca, Tellina aequizonata, therefore they may be more susceptible to (moderate)
post-mortem reworking and transport even in settings during times of low rates of net siliciclastic accumu-
of weak environmental energy. lation (i.e. they represent depositional hiatuses). The
From a trophic standpoint, suspension-feeders coarse-grained nature of the sediments, and trough
(most of the bivalve taxa) are the prominent elements cross-stratification reflecting the lateral migration of
(56.36%), but deposit-feeders (nuculids, nuculanids, 3-D bedforms, indicate that high flow velocities were
tellinids and scaphopods) and carnivores (gastropods) responsible for their deposition. This shell-rich facies
are common. The high species diversity of predatory is interpreted as skeletal banks that were winnowed,
gastropods (20 taxa at least) in comparison with the extensively reworked and sometimes transported by
scarcity of the related signs of attack on the mollusk strong storm-driven waves and currents.
shells suggests the presence of potential prey among In conclusion, both the taphonomic signatures of
some soft-bodied taxa (e.g. ascidians). The relative fossils (highly variable state of preservation, rare
insignificance of herbivores and browsers (only two articulated bivalves, high degree of breakage and
taxa accounting for 0.3%) is noteworthy, and suggests fragmentation, concave-down arrangement of whole
a lack of seafloor vegetation. valves, etc.) and the co-occurrence of ecologically
The bathymetric ranges of extant species in this unrelated groups of species (different substrates,
assemblage lead us to infer an inner subtidal setting bathymetry, etc.) suggest that this shell bed can be
no more than 20–30 m in depth. This inference is regarded as a mixture of mollusks originally inhabit-
also supported by the presence of Raeta undulata, ing the surf and the shallow subtidal zones. Shells
which today lives in the foreshore and the inner part were reworked under conditions of medium to high
of the subtidal zone. Keen (1971) reported that wave and current energy and may have suffered
bbeach valves are fairly common, but entire speci- transport both within- and out-of-habitat.
mens are hard to find, even by dredgingQ, and so the Shell-rich facies underlain by sequence-bounding
occurrence of most fossil individuals as either ravinement surfaces and similar to the current/wave
conjoined or in (nearly) life position suggests that winnowed shell beds described here, have also been
the fossil community inhabited the deeper environ- described, among other places, at the base of late
ments of the bathymetric range of this species. Only Pliocene–Pleistocene cyclothems of New Zealand
Grandiarca grandis (0.76%) and Melongena sp. (Abbott and Carter, 1994; Naish and Kamp, 1997;
(0.1%) indicate shallower settings than those men- Abbott, 1998), Pleistocene of California (Carter et
tioned previously: the former inhabits the foreshore/ al., 2002) and Japan (Kitamura et al., 2000), in Plio-
shoreface zone to 5 m in depth, and the latter is Pleistocene shallow-marine deposits of the Gulf of
common on intertidal mud flats, and both species are California (Meldahl, 1993) and Ecuador (Di Celma,
linked to marine/estuarine environments with fluctu- 2002), in Miocene sediments of the eastern United
ating salinity values. Their presence in the paleo- States (Kidwell, 1989) and Argentina (Del Rı́o et al.,
community may be related to post-mortem 2001), and in Upper Jurassic–Lower Cretaceous
displacement to subtidal settings of accumulation. successions of India (Fürsich and Pandey, 2003). In
particular, the well-preserved, infauna-dominated,
parautochthonous assemblages of dispersed current/
6. Discussion wave-winnowed shell beds are very similar for
paleontological and depositional features to those at
6.1. Origin of current/wave winnowed shell beds the base of the Early Pleistocene depositional
sequences of the Omma Formation, in central Japan
When combined, all the biostratinomic, sedimen- (Kitamura et al., 1994). These latter contrast only for
tological and paleoecological features described above faunal diversity that is significantly higher in the
reveal that skeletal components of the simple current/ basal shell beds of Punta Ballena Mbr (Jama
wave winnowed shell beds persisted on the seafloor Formation).
for a relatively long time, suffering post-mortem Many of the contrasting features existing between
modification before final burial. Specifically they condensed deposits at the base of 3rd-order Miocene
were exposed to repeated and extensive reworking sequences and 5th to 7th order Plio-Pleistocene
sequences, such as the more complex taphonomic 6.2. Origin of community shell concentrations
processes and internal stratigraphy recorded in the
former, may be explained by the consistent difference Dense layers of infaunal bivalves occurring above
in time involved in their formation (Kondo et al., 1998). intensely burrowed intervals, clearly reflect periods of
According to Norris (1986), current/wave depos- reduced net terrigenous sedimentation when, owing to
ited shell concentrations are largely restricted to the the presence of a slowly- or non-aggrading sediment
zone regularly swept by storms. Within this environ- surface, bivalves burrowing almost at the same depth
ment, skeletons included in these bedforms may be below the sea floor may be clustered together beyond
derived from reworking of indigenous living mollusks likely living densities. In addition, concentrations of
(parautochthonous assemblages sensu Kidwell et al., fish otoliths extremely higher than those of adjacent
1986), from older fossil assemblages exhumed and strata further corroborate the hypothesis that back-
erosionally recycled during shoreface retreat by the ground processes included long times of low to zero
erosion of the subjacent depositional sequences sedimentation. In this case, as concentrations occurred
(remanié of Craig and Hallam, 1963), or from shells under conditions of relative environmental stability
transported from immediately adjacent, roughly and only specimens from a single temporally persis-
coeval, environments (allochthonous assemblages tent community accumulate, shell concentrations are
sensu Kidwell et al., 1986). According to Kidwell within-habitat time-averaged assemblages (Kidwell
(1991a) assemblages composed in large part of and Bosence, 1991) characterized by a faunal diver-
skeletons reworked from the underlying strata and sity relatively lower than that recorded for dispersed
thus, pre-dating local transgressive erosion, are current/wave-winnowed shell beds.
referred as transgressive lags, whereas assemblages Community shell concentrations are interpreted as
created by colonization and passive accumulation of biogenic or mixed biogenic–sedimentologic (sensu
autochthonous/parautochthonous specimens post-date Kidwell et al., 1986), mainly autochthonous assemb-
transgressive erosion and are referred as condensed lages formed by in-place infauna accumulated on
onlap assemblages; many skeletal concentrations that inner shelf settings during periods of reduced terri-
mantle disconformities will be mixtures of these two genous net sedimentation due to the concomitant
sources. As a consequence of the different origin of action of reduced sediment supply and gentle win-
component skeletons, the species diversity of these nowing. This means that these beds are probably
assemblages is usually high. However, as corrobo- stratigraphically condensed (i.e. significantly thinner
rated by several studies in modern environments (see than coeval strata laid down more landward) and,
literature cited in Tables from II to IV in Kidwell and hence, may be typical examples of hiatal shell
Bosence, 1991; Russell, 1991; Kidwell and Flessa, concentrations (Kidwell, 1991a).
1996), shell transport in level-bottom settings appears The Raeta Association shares a few taphonomic,
to be largely a within-habitat process. Even when paleoecologic, and sedimentologic attributes, such as
shells derived from adjacent habitats account for a the nearly pristine condition of shells, the dominance
large part of the total species present in the final death of infaunal species and the low-density biofabric, with
assemblages, it usually seems to involve only small- the Serratina Association described by Abbott (1997)
shelled individuals (Cadeé, 2002) and, therefore, a in the mid-Pleistocene sequences of the Wanganui
minority of all specimens in the final assemblage. The Basin. However, the Raeta Association is distin-
high species diversity, the grain-size and lithologic guished by the dominance of mud-tolerant suspen-
nature of the internal moulds that differ from those of sion-feeders, and by the abundance of bivalves in a
the surrounding sediment, the wide range of preser- normal life orientation. Because suspension-feeders
vational state, and the co-occurrence of ecologically generally are better adapted to reduced sediment input
unrelated species, are all clear evidence implying that and low turbidity conditions, the prevalence of this
the assemblages occurring within these beds preserve trophic group and mode of bivalves occurrence in the
a mixture of non-contemporaneous remains and, Raeta Association is likely to be sponsored by low
therefore, are significantly environmentally time- sedimentation rates and relative stability of the
averaged (Fürsich, 1995). substrate (Fürsich, 1995; Brett, 1998).
6.3. Sequence stratigraphic interpretation temporary, transgressive, shore-connected clastic

wedge, where conditions of reduced terrigenous net
In the Punta Ballena Mbr, conventional sedimento- sediment-accumulation rate pertain due to the land-
logic analysis indicates that unconformities interposed ward translation of coastal depocenters. Since these
between successive cyclothems represent subaerial skeletal accumulations occupy a mid-cycle position
erosion during sea-level lowstands, marine ravine- within depositional sequences, Kidwell (1991b) and
ment during successive transgressions, or both. The Abbott (1997) referred to them using the more
erosion surfaces underneath non-marine or marginal- appropriate sequence stratigraphic terms of backlap
marine deposits (sequence boundaries), are interpreted shell beds (i.e. tracking the backstepping distal edge
as broad incised valleys developed during relative sea- of the TST) and mid-cycle condensed shellbeds
level lowering and filled during the following phase of (MCS), respectively.
sea-level rise (incised valley fill, IVF). Where IVF According to Abbott (1997) and Abbott and
deposits are absent, the vertical contact between the Carter (1997), the formation of a variably eroded,
subsequent depositional sequences occurs by means bioturbated and/or burrowed transgressive surface
of typical low-relief erosional scours that are usually (local flooding surface, LFS) that rests directly below
attributed to current- and wave-action associated with the backlap shell bed, reflects time intervals charac-
processes of shoreface retreat during each trans- terized by relatively low rates of terrigenous supply
gression phase (Nummedal and Swift, 1987). Gen- at the distal margins of TST, while seaward of this
erally these surfaces are carved directly into the surface sediment-starved conditions foster the con-
truncated offshore massive siltstone of the subjacent centration of well-preserved, parautochthonous and/
sequence and are typically crossed by Thalassinoides. or autochthonous fossil assemblages (community
These burrows, belonging to the Glossifungites shell beds). Shell concentration in the community
ichnofacies, have a prominent vertical component beds terminates when conditions of reduced silici-
and penetrate into the semi-consolidated substrate to clastic sediment supply on the inner shelf are
depths of as much as 1.5 m below the RS. The fact replaced by conditions of normal terrigenous sed-
that these burrows are emplaced over firm, generally imentation because of the progradation of shore-
fine-grained sediments of the subjacent depositional connected siliciclastic wedges (Kidwell, 1989,
sequence and are filled with sediments originating 1991b) or the lateral growth of shelf mud-blankets
from the subsequent one, indicates that periods of (Abbott, 2000). In this case shell-poor, fine-grained
omission occurred after generation of the RSs via sediments can sit disconformably on the backlap
erosional exhumation of overcompacted substrates, shell beds. On the basis of the degree of taphonomic
and before the onset of sedimentation of the back- and taxonomic changes across the disconformity
stepping shelf wedge. The current/wave-winnowed surfaces bounding backlap shell beds (local flooding
beds always sit directly above the RSs and, since they surface and downlap surface), Abbott and Carter
commonly mark the onset of transgressive conditions (1997) also stated that they represent diastems
in shallow marine settings, skeletal concentrations of associated with little or no erosion and are charac-
this type have been called onlap shell beds by Kidwell terized by breaks in sedimentation that are compa-
(1991b). Recently Naish and Kamp (1997) and Kondo ratively lower in magnitude than those associated
et al. (1998) developed Kidwell’s scheme for with sequence-bounding surfaces.
sequence stratigraphic classification of shell beds in Although, on the whole, both these kinds of shell
the Plio-Pleistocene. concentration of the Jama Formation show features
In contrast, community shell concentrations from consistent with a genuine condensation due to reduced
depositional sequences of the Punta Ballena Mbr are siliciclastic sediment supply, their hiatal nature can
relatively shell-poor and loosely packed hiatal con- only be inferred rather than demonstrated unequiv-
centrations within which macrofossils are preserved ocally within the available outcrop belt, owing to the
in-situ or nearly so within inner shelf fine-sandy absence of exposures demonstrating that they are the
siltstone. They are regarded as forming in low energy product of landward and seaward thinning of thicker
settings at the basinward-tapering edge of the con- coeval strata.
6.4. Patterns of sedimentary regimes during trans- of mainly infaunal bivalves, resulting in the concen-
gressions and genetic models of shell concentrations tration of matrix-rich shelly material (community shell
beds), whose biofabric has been only partially
The existence in these depositional sequences of modified by burrowing organisms (backlap shell
two different onshore–offshore patterns of sedimen- bed). Shell concentration ceased with the onset of
tary regimes during transgressions is indicated by the downlap at the distal edge of HST when, owing to the
occurrence of two distinct TST motifs. Assuming seaward shift of the coastline, areas that were
Walther’s principle, the vertical changes in lithology sediment starved during transgression started to
and shell concentration features within transgressive receive increasing amounts of siliciclastics.
layers reflect the onshore–offshore progressive reduc-
tion of environmental energy and the pattern of net
siliciclastic sediment supply down depositional dip. 7. Conclusions
Therefore, across an idealized transect of contempo-
raneous shoreface to offshore environments, current/ (1) The sedimentary succession exposed along the
wave-winnowed shell beds accumulate directly on the sea-cliff west of Jama, northern Ecuador, is a north-
RSs and represent prominent breaks in sedimentation eastward-dipping Early Pleistocene clastic wedge. A
within shallow-marine settings regularly swept by detailed sequence stratigraphic study, integrating
storms and currents. This kind of condensation has sedimentologic, stratigraphic, and paleontologic
been called scour condensation by Storms and Swift observations, reveals a cyclic sedimentation pattern
(2003). Given the scarcity of siliciclastic, sand-grade consisting of both deepening-upward and deepening-
deposits within the TSTs of depositional sequences shallowing facies successions, which may be inter-
PB5, PB6, PB7, and PB8, these shallow settings were preted as repeated sea-level oscillations (i.e. they are
probably starved of sand during transgressions. depositional sequences).
Because of the high environmental energy, these (2) Within sequences, shell concentration may
settings were also characterized by the total passing occurs at the base and at the top of the backstepping
of fines which, in turn, were delivered as suspended shelf wedge that constitutes the marine portion of the
load in relatively deeper water and laid down below transgressive systems tract. Two fundamental types of
storm wave-base. In vertical section this area of hiatal shell concentrations, differing from each other
deposition is represented by the monotonous fine- both in their stratigraphic position and their tapho-
sandy siltstone interposed between the onlap and nomic and ecological attributes, can be distinguished:
backlap shell beds (facies association Om4 in Table current/wave-winnowed and community shell con-
2). Within sequences PB1, PB3, and PB4, current/ centrations. The current/wave-winnowed shell beds
wave-winnowed (onlap) shell beds are absent or lie directly upon ravinement surfaces (onlap shell
poorly developed and the TST is composed of sand beds) and exhibit high-diversity faunas. Sediment
sized sediment. This is interpreted as resulting from facies as well as taphonomic and paleoecologic
high siliciclastic sediment supply during transgressive features are all suggestive of a shallow-marine, high-
phases associated with development of sediment energy shoreface environment characterized by fre-
trapping nearshore wedges, with conditions of scour quent transport and reworking and where bioclasts are
condensation being more spatially restricted and representative of a mixing of ecologically unrelated
difficult to attain. species. In contrast, community shell concentrations
Seaward of the onshore area of scour and/or sand occur in a roughly mid-cycle position and consist of
accumulation, the inner shelf received decreasing autochthonous/parautochthonous assemblages with a
amounts of suspended terrigenous deposits, and at relatively lower diversity. Visually, they are domi-
some offshore point became nearly starved (starvation nated by articulated individuals of the infaunal bivalve
condensation of Storms and Swift, 2003). Within Raeta undulata that are preserved in life orientation,
these low-energy, nearly sediment-starved offshore or have undergone only minor reorientation, appa-
areas, a fine-grained and slowly aggrading seafloor rently through bioturbation. Based on their tapho-
was colonized by a number of successive generations nomic and paleoecologic attributes, which are
suggestive of accumulation in low-energy environ- Abbott and Carlton E. Brett are also acknowledged
ments characterized by reduced siliciclastic supply for their useful comments as journal reviewers.
and sporadic removal of finer material, these concen-
trations are inferred to be formed in inner-shelf
settings, at the distal feather-edge of the transgressive References
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Palaeogeography, Palaeoclimatology, Palaeoecology 216 (2005) 1 – 25

Sequence stratigraphy of the Punta Ballena Member of the Jama

1. Introduction Tschopp (1948) the Jama Formation comprises upper

Table 1 Table 1 (continued)

Table 1 (continued) Di Geronimo and Robba (1976). Faunal compositions

TOTAL 2823 99.97 1042 99.94

G. Cantalamessa et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 216 (2005) 1–25

horizontal positions reflect a post-mortem dislodging

5.1.2. Taphonomic features of dispersed current/

6.3. Sequence stratigraphic interpretation temporary, transgressive, shore-connected clastic

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