DOCUMENTA

ARCHAEOBIOLOGIAE 16
Animals and Humans through Time and Space:
Investigating Diverse Relationships
 DOCUMENTA ARCHAEOBIOLOGIAE

Veröffentlichungen der Staatssammlung für Anthropologie München

und der Staatssammlung für Paläoanatomie München

Band 16

Begründet von
Gisela Grupe und Joris Peters

Herausgegeben von
George McGlynn und Joris Peters
 DOCUMENTA
ARCHAEOBIOLOGIAE

Animals and Humans through Time and Space:

Investigating Diverse Relationships
Essays in Honour of Joris Peters

Edited by
Nadja Pöllath, Nora Battermann, Stephanie Emra, Veronika Goebel,
Ptolemaios Paxinos, Martina Schwarzenberger, Simon Trixl,
Michaela Zimmermann

Verlag Marie Leidorf GmbH . Rahden/Westf.

2023
 428 Seiten mit 162 Abbildungen und 61 Tabellen

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Bibliograsche Information der Deutschen Nationalbibliothek

Nadja Pöllath, Nora Battermann, Stephanie Emra, Veronika Goebel,

Ptolemaios Paxinos, Martina Schwarzenberger, Simon Trixl,
Michaela Zimmermann (Eds.):
Animals and Humans through Time and Space: Investigating Diverse
Relationships
Essays in Honour of Joris Peters
Rahden/Westf. : Leidorf, 2023
(Documenta Archaeobiologiae ; Bd. 16)
ISBN 978-3-89646-700-3

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 Inhaltsverzeichnis

Vorwort . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

Grußworte . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

Laudatio . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

Mitgliedschaften, Ehrungen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33

Schriftenverzeichnis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34

Autorenverzeichnis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49

Cattle and conservatism on the central Anatolian plateau:

Bos exploitation at Chalcolithic Köşk Höyük, Niğde, Turkey
Benjamin S. Arbuckle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53

Last hunter-gatherers, rst farmers in Brandenburg – An archaeobiological discourse

Norbert Benecke, Susanne Jahns, Steffen Wolters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69

Iron Age animal gurines made of copper alloy from the sanctuary at Dülük Baba Tepesi
Michael Blömer, Engelbert Winter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85

Molluscs in El Argar culture:

An interpretative proposal of their role in domestic and funerary-symbolic contexts
Cristina Cabrera Taravillo, Ignacio Bize Font, Corina Liesau von Lettow-Vorbeck,
Arturo Morales Muñiz . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93

Stone cold foxes – biology, archaeology, and iconography in Upper Mesopotamia

Stephanie Emra, Nora Battermann, Nadja Pöllath . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107

Da ist der Wurm drin! Streuüberlieferung lateinischer veterinärmedizinischer Rezepte

in frühmittelalterlichen humanmedizinischen Handschriften
Klaus-Dietrich Fischer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125

The archaeozoological remains from Late Chalcolithic Çukuriçi Höyük, on the western Anatolian coast.
Alfred Galik, Stephanie Emra, Christoph Schwall, Barbara Horejs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139

Das Kapaunisieren anhand der Literatur des 19./20. Jahrhunderts und des historischen
Instrumentariums der tiermedizinhistorischen Sammlung der Tierärztlichen Fakultät München
Veronika Goebel, Martina Schwarzenberger, Michaela Pfeuffer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149

Bergen, Sammeln und Bewahren: Menschliche Skelettfunde als Teil des archäologischen Erbes
Michaela Harbeck, Kristin von Heyking, Andrea Grigat, Heiner Schwarzberg,
George McGlynn, Peter Schröter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 163

“May the camels multiply”. More data about camels in the Southern Levant,
the Arabian Peninsula, and Mesopotamia
Martin Heide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 185

Tierknochen in ägyptischen Texten

Friedhelm Hoffmann . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 193
10

Faunal remains from Salat Cami Yanı: An Early Pottery Neolithic site in the Upper Tigris Valley
Hitomi Hongo, Saiji Arai, Can Yumni Gündem, Banu Öksüz, Yutaka Miyake . . . . . . . . . . . . . . . . . . . . . . . . . 201

Der vorptolemäische Gangarm C-D-4 im Ibiotapheion von Tuna el-Gebel

Dieter Kessler . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213

A case of (food) taboo in the Bronze and Iron Ages in south-central Europe?
– An explanation of the lack of freshwater pearls in the archaeological record of prehistoric Europe
Carola Metzner-Nebelsick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 225

Linear Pottery culture and Rössen culture nds from Gudensberg,

North Hesse, and the question of the use of cattle as working animals already in the Rössen culture
Kerstin Pasda, Eberhardt Kettlitz . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 235

The age of rats. Possibilities and limitations of ageing rat bones from
archaeological excavations based on material from the medieval site of Marienhof in Munich, Germany
Ptolemaios Dimitrios Paxinos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 261

Animal bones in the funerary practices at the 1st-3rd century AD necropolis of Potzneusiedl (Austria)
Konstantina Saliari, Erich Pucher, Francesco Boschin, Viola Winkler, Lucia Clara Formato . . . . . . . . . . . . . 277

The evolution of goat husbandry in Tell Halula:

A look back to goat exploitation through biomechanics and stable isotope analysis
Maria Saña, Roger Alcàntara, Carlos Tornero . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 295

Angeln und Fischen in Zeiten der pax Romana im Alpenvorland

Bernd Steidl . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 307

They travelled far - Roman period camelids from Switzerland

Barbara Stopp, Sabine Deschler-Erb, Sven Billo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 325

Zu Rachitis und Osteomalazie bei prähistorischen und historischen Tieren

Wolf-Rüdiger Teegen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 339

Roman frontier regions in the focus of interdisciplinary ancient studies:

A horse skeleton from Heidenheim an der Brenz (Baden-Württemberg) and its context
Simon Trixl, Janette Horvath, Elisabeth Stephan, Andreas Thiel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 363

Tierknochenfunde aus einem Befestigungsgraben der eisenzeitlichen Siedlung Muweilah (Sharjah, VAE)
und ihre Bedeutung für Fragen zur Domestikation und Nutzung des Dromedars
Hans-Peter Uerpmann, Margarethe Uerpmann, Peter Magee . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379

Bronze Age subsistence along the southern coast of Yemen: The example of al-Uriyash
Wim Van Neer, Wim Wouters, Sergey Popov, Vittoria Buffa, Bea De Cupere . . . . . . . . . . . . . . . . . . . . . . . . . . 389

Subsistence intensication: the molluscs from the Chácara I sector at Ligüiqui

(Manabí Province, Ecuador: AD 12th-15th centuries)
Víctor F. Vásquez Sánchez, Teresa E. Rosales Tham, Manuel Castro Priego, Lauro Olmo Enciso,
Juan A. Jijón-Porras, Marcos O. Labrada-Ochoa, Arturo Morales Muñiz . . . . . . . . . . . . . . . . . . . . . . . . . . . 407

Instructions for Authors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 427

 Subsistence intensi cation: the molluscs from the Chácara I sector at
Ligüiqui (Manabí Province, Ecuador: AD 12th-15th centuries)

Víctor F. Vásquez Sánchez1, Teresa E. Rosales Tham2, Manuel Castro Priego3,

Lauro Olmo Enciso3, Juan A. Jijón-Porras4, Marcos O. Labrada-Ochoa5,
Arturo Morales Muñiz6

1
Centro de Investigaciones Arqueobiológicas y Paleoecológicas Andinas, Trujillo, Perú
2
Departamento de Antropología y Arqueología, Universidad Nacional de Trujillo, Perú
3
Departamento de Historia I y Filosofía, Área de Arqueología,
Universidad Alcalá de Henares, Madrid, España
4
Departamento de Ciencias Humanísticas, Universidad Técnica de Manabí, Manabí, Ecuador
5
Departamento de Historia del Arte, Arqueología y Música,
Universidad de Córdoba, Córdoba, España
6
Laboratorio Arqueozoología, Departamento de Biología,
Universidad Autónoma de Madrid, Madrid, España

Abstract / Zusammenfassung

The assemblage of molluscs retrieved in an area lying next to the shing installations in the Ecuadorian coastal
site of Ligüiqui is reported. The small size of the collections, barely 1000 remains, does not preclude the richness
of a sample that includes 55 taxa. The analysis reveals that the molluscs mostly represent consumption refuse from
human meals with scarce evidence of craftwork and only one specimen of the Spiny Oyster, Spondylus princeps
unicolor testifying trade. The faunal remains and the singular shing installations for which this settlement is best
known both indicate that the subsistence of the inhabitants clearly focussed on marine resources. Although the overall
composition of the malacological assemblages appears to remain stable throughout the scarcely two centuries the
occupation lasted, subtle shifts have been detected in the presence, frequency, and size of certain species that hint
at an overexploitation of resources through time coupled with a move towards deeper water whose reasons remain
debatable at this point.

Diese Studie behandelt Funde von Molluskenschalen, die auf einem Areal in der Nähe der prähistorischen Fi-
schereianlagen von Ligüiqui an der ecuadorianischen Küste bei archäologischen Ausgrabungen gefunden wurden.
Trotz dem geringen Umfang des Fundmaterials von knapp 1000 Fragmenten ist es mit 55 Taxa sehr artenreich. Die
Analyse zeigt, dass es sich bei den Mollusken hauptsächlich um Essensreste von menschlichen Mahlzeiten han-
delt, mit wenigen Hinweisen auf handwerkliche Tätigkeiten und nur einem Exemplar der Stachelauster, Spondylus
princeps unicolor, das seinerseits von Handel zeugt. Die Tierreste und die einzigartigen Fischereianlagen, für die
diese Siedlung bekannt ist, zeigen an, dass die Ernährung der Bewohner stark auf marine Ressourcen ausgerichtet
war. Obwohl die Gesamtzusammensetzung der Molluskenfauna während der knapp zwei Jahrhunderte, die die
Besiedlung dauerte, stabil zu bleiben scheint, wurden subtile Verschiebungen im Vorkommen, in der Häu gkeit
und in der Größe bestimmter Arten festgestellt, die auf eine übermäßige Ausbeutung der Ressourcen im Laufe der
Zeit in Verbindung mit einer Verlagerung der Nahrungssuche in tiefere Gewässer hinweisen. Die Gründe dafür
sind derzeit noch nicht geklärt.

Keywords: Ecuador, Manteño culture, molluscs, subsistence, overharvesting

Ecuador, Manteño-Kultur, Mollusken, Subsistenz, Übernutzung
408
1 Introduction
Since the beginning of the 20th century, the study of the
pre-Hispanic societies from Ecuador has been focused
on the coast (Marcos 1986; 2005; Delgado-Espinoza
2010; Valdez 2010). One target period to explore pro-
cesses such as social complexity, production and inten-
si cation, and landscape use is the Manteño-Huancavila
culture that lasted from AD 600-1550 and developed
in the provinces of Guayas, Santa Elena, and Manabí
(Gutiérrez-Usillos 2009; Lunniss 2020). For coastal
communities, one postulated driver of such develop-
ments was the commerce of the spiny oyster (Spondylus)
that presumably fostered the development of chiefdoms
with a de facto state structure in the area. Archaeology
has stressed the religious and ritual connotations of those
shells, and their role as social status indicators, since
adornments (bracelets, collars, pectoral plates) executed
on spiny oyster shell is a common item in the burials of
the pre-Hispanic elites. Although the spiny oyster com-
merce had been acknowledged as a prime mover since
the 3rd millennium BC, in recent years its traditional
relevance in the trade networks that developed between
present-day Peru and Ecuador has been somehow down-
graded (Carter 2011).
On the central Ecuadorian coast, the emergence of this
new social organisation, coupled to that of the local
chiefdoms (i.e. “cacicazgos”), can be dated to AD 600-
900, yet it is not until AD 1050-1200 that settlement
models featuring a complex and hierarchical organisa-
tion develop over wide territories. The urbanism from
this later phase is often associated with monumental con-
structions such as those found at Cerro de Hojas, Agua
Blanca and Salango (McEwan 2004; Lunniss 2020). It
is also at this time that settlements centred on aquatic
resources, as is the case at Ligüiqui, are recorded for
the rst time.
2 The site of Ligüiqui
Located ca. 20 km SW of the town of Manta (province of
Manabí), the site of Ligüiqui (coordinates: 1°1’41.18”S
& 80°52’52.85”W) is a singular Late pre-Hispanic (AD
12th-15th centuries) settlement on the central coast of Ec-
uador (Fig. 1). The 2,500 ha that the site covers spread
over two wide coves, Ligüiqui and Punta Cangrejo, set
between the capes of San Lorenzo and San Mateo, and
feature two different sectors. Urbanised areas are ar-
ranged into independent clusters of up to 15 ha each,
occupying the hilltops of dozens of 20-60 m high knolls
that run parallel to the Paci c Ocean. These knolls are
separated by deep ravines, each one of them harbouring
a seasonal brook. The settlement areas link with an ex-
tensive shing enclave featuring water enclosures that
spread over 4.5 km of the Paci c Ocean shore (Fig. 1C).
Víctor F. Vásquez Sánchez et. al.
The coastal area developed during the Eocene Period
(56 mya) and features a mostly sandstone basement
on top of which sit clays and sand accumulated dur-
ing the Pleistocene. The area lying next to the shore
is mostly constituted by basalts, themselves a result of
the intensive volcanic activity the province of Manabí
witnessed until recent times (Pearsall & Zeidler 1994,
201). Evidence of this activity is also the tetra volcanic
sedimentation recorded on several spots within the site’s
perimeter caused by volcanic eruptions in the Andes
during the last four millennia (Guillaume Gentil 2013).
The pro le of the coast features a gradually sloping and
narrow (ca.100 m) continental shelf whose rift falls deep
into the ocean oor. As a result of that topography and
a gentle surf, snorkelling on the continental platform
is nowadays a popular pastime on these beaches. Tidal
amplitude, depending on the spot, oscillates between
2-5.5 m (https://www.inocar.mil.ec/web/index.php/pro-
ductos/tabla-mareas).
The landscape around the settled area is the typical
dry tropical forest of the Ecuadorian coast, subjected
to intensive erosive episodes magni ed in more recent
times by vegetation losses caused by anthropic activ-
ities. Eastwards, as altitude increases, a humid forest
develops nurtured by the moisture that the coastal fogs,
particularly intense from June to December, bring along.
Although Ligüiqui was rst mentioned as a part of a
“reducción” (i.e. an indigenous community set up under
ecclesiastical or royal authority to facilitate colonisation)
already at the start of 17th century, it has remained “hid-
den” to scienti c enquiry until very recent times (Ponce
Leyva 1994. 35). The work our team started in 2018-
2019 included: (a) a non-invasive survey of the northern
portion of the settled area, (b) an intensive prospection
of the northern shoreline, and (c) open area excavations
and stratigraphic sequencing of the spaces dubbed “Las
Chácaras” and “Los Charcos” on the shing complex
area of the shore. Radiocarbon dates set the urban area
within the Late Manteño culture (AD 11th-15th centuries)
(Beta-538425: 950±30 BP (2σ: 1043-1212 cal. AD);
Beta-613358: 1270± 30 BP (2σ: 1166 - 1413 cal AD);
Hogg et al. 2020). On the shing quarters, deposits from
earlier stages, contextually dated to the 1st millennium
BC, have been also recorded.
The shing installations are structured as a system of
enclosures (local people call them “corrales”), organ-
ised into a series of interconnected semi-circular basalt
slab enclosures occupying a series of platforms, each
one of which plunges deeper and farther away from the
shoreline (Fig. 1C). 3D models of the placement of the
basalt slabs at Punta Cangrejo allowed us to document
the constructive features of the enclosures. The erected
slabs are arranged into arches with maximum diame-
ters of 141 m. Although the sea covers these slabs for
Subsistence intensi cation: the molluscs from the Chácara I sector at Ligüiqui 409

Fig. 1: Ligüiqui: (A) Overview of the area signalling the main settlement areas (yellow rectangles) and shing
structures (white rectangles); (B) Ground plan of the “Chácara I” Sector (C) Partial view of the shing structures
(inset: location of Ligüiqui on the Ecuadorian coast) (photos and maps: (A) World Imagery, copyright ESRI; (B
and C) M. Castro-Priego and L. Olmo-Enciso).

a substantial portion of the day, the top of even those

located farthest away from the shore (ca. 100 m into
the ocean) are still visible at high tide. These complex
structures hint at a dense population of shermen, prob-
ably with different tasks assigned to them. In this way,
on the enclosures nearest to the shore, shell-gathering,
and shing of shallow-water, small-sized, shes were the
main activities. On the outermost, deeper, enclosures that
remained underwater most of the time, shing of deeper
water species undoubtedly implied use of rafts, along
with shing gear such as harpoons and seines.
3 Material and Methods
The mollusc remains reported in this paper derive from
the Chácara I sector, an area associated with the shing
complex where both kitchens and habitation spaces have
been documented (Fig. 2). All sediments were sieved
through 2 mm meshes. The excavation unit here is the
Unidad estratigrá ca (UE hereafter), of which 13 yield-
ed mollusc remains: UE 0, 1, 3, 5, 6, 7, 9, 13, 21, 26, 27,
28 and 32 (Tables 8 and 9).
410 Víctor F. Vásquez Sánchez et. al.

Fig. 2: Location of the main archaeological contexts (units) of the “Chácara I” sector (photo and adaptation by M.
Castro-Priego and L. Olmo-Enciso).

Fig. 3: Relative frequencies, according to biotope, of the molluscs from Ligüiqui (graph by V. F. Vásquez Sánchez,
T. E. Rosales Tham and A. Morales Muñiz).
Subsistence intensi cation: the molluscs from the Chácara I sector at Ligüiqui
Identi cation was made with AMM’s reference collec-
tion housed at the Laboratorio de Arqueozoologia from
Universidad Autónoma de Madrid (LAZ-UAM). Iden-
ti cation of specimens from species not available in this
collection was accomplished with the works of Álamo
& Valdivieso (1987), Breure (1978; 1979), Brito-Vera &
Mora (2016), Dall (1909), Keen (1958, 1971), Marincov-
ich (1973), Olsson (1961), Osorio et al. (1979), Peña
(1970, 1971) and http://www.idscaro.net/sci/01_coll/
indet.htm., http://www.femorale.com/, https://concholo-
gy.be/?t=261. Álamo & Valdivieso (1987), Keen (1958,
1971) and Marincovich (1973) were consulted to check
the biology and spatial plus bathymetric distributions
of the species.
Quanti cation considered the Minimum Number of In-
dividuals (MNI), the Number of Identi ed Specimens
(NR, following the Spanish acronym) and the weight of
the shells, expressed as grams (Reitz & Wing 2008). For
the NR we considered all kinds of specimens, complete
and fragmented. MNI estimations vary depending on
the class of molluscs. For chitons (Polyplacophora), the
highest number of cephalic and/or anal plates was con-
sidered, but also the number and position of the remain-
ing dorsal plates. In addition to complete shells, MNIs
for gastropods took account of the number of apexes and
complete peristomes. In the case of bivalves, the largest
number of right or left valves, but also the size of the
valves, is considered but the grand total also took into
consideration the highest number of right and left umbos
and hinge fragments. Specimens were weighted in a CE
Baxtran FFN digital balance (estimated error: 0.01 g).
Only the maximum length, expressed in millimetres, has
been taken on all complete specimens. Measurements
were made with a Power x® Mitutoyo digital calliper
(estimated error: 0.1 mm). Slightly eroded specimens
and incomplete ones whose lengths could be taken were
systematically measured Biometrical analyses focused
on the specimens from the richest units (UE1 and UE
13). To validate the signi cance of size differences
through a contrast of mean values from paired samples,
the t-Student test was employed (Dawson-Saunders &
Trapp 1993).
Taphonomic group assignment (i.e. food, worked re-
mains, intrusive specimens, background faunas) follows
Gautier (1987). Worked shell protocols follow Guinea
(2006) and Rosales Tham (2016).
4 Results
Table 1 provides an overview of the molluscs assemblag-
es of the 13 UE from the Chácara I sector at Ligüiqui.
A total of 1102 specimens, representing 981 individuals
for a weight slightly over 2.6 kg, constitute a modest
collection despite its richness of 55 taxa. The high (0.89)
411
correspondence between the MNI and the NISP evidenc-
es the completeness of the shells (fragments represent
ca.1% of the retrieved specimens), but also the preva-
lence of marine gastropods that constitute 96% of the
MNI and ca. 90% of the NISP (bivalves ca.10% and
3.6%, respectively). Chitons (Polyplacophora) and land
snails (Pulmonata) represent anecdotical items.
The mollusc remains are not evenly distributed since,
combined, UE 1 and UE13 represent 56% of the grand
MNI total and 54.5% of the grand NISP total (MNI:
40.3% [UE 1] and 15.6% [UE 13]; NISP: 37.4% [UE 1]
and 16.1% [UE 13]), the remaining UE featuring abun-
dance estimators below 10% (Tables 8, 9).
Taxonomically, the 55 recorded taxa include 41 species
(34 gastropods, 7 bivalves), 7 genera (1 gastropod, 4
bivalves, 1 pulmonate), 4 families (3 bivalves, 1 pulmo-
nate) and 3 classes (i.e. Polyplacophora, Gastropoda and
Bivalvia). Another hint of the generally good condition
of the specimens is the fact that only a few fragments
had to remain identi ed at the most general level (i.e.
mollusc indet.).
As is normally the case in mollusc assemblages, in-
cluding natural ones and those accumulated by people,
few species dominate. At Ligüiqui, this dominance
corresponds to the Peruvian tegula, Tegula picta (22%
of the total MNI; ca. 20% of the total NISP), and the
sunburst star turban, Astraea buschii (13% of the total
MNI; 13.5% of the total NISP) which together with the
dove-shell, Columbella columbiformis (12.4% of the
total MNI; 11% of the total NISP), constitute nearly half
of the studied sample (MNI: 47.5%; NISP: 44.3%). The
seven species that follow in abundance are also gastro-
pods, and only the pearl oyster, Pinctada mazatlanica,
on the eleventh position, could be argued to qualify as a
relevant item of the assemblages. The 10 more frequent
species represent 87% of the MNI and 80% of the NISP)
(Fig. 3). Remarkable, given that Ligüiqui had been con-
sidered a spiny oyster “production site”, is that only a
single worked fragment of Spondylus princeps unicolor
(as we will see, a taxon alien to the Ecuadorian coast)
was retrieved in UE3 (Tables 2, 8).
From an ecological standpoint, the bulk of the identi ed
specimens represents species from the mesolittoral and
infralittoral zones of a mostly rocky shore (Fig. 3; Tables
3, 4). Set within such context, it seems remarkable that
not even a single species reaches to the supralittoral
zone and only the periwinkle, Littorina modesta, inhabits
deeper water.
From a biogeographical perspective, 40 out of 41 of the
identi ed species at Ligüiqui occur mainly in the Pan-
amanian province that ranges from 0o-20o N, covering
all the Ecuadorian coast. 21 of these species reach into
412 Víctor F. Vásquez Sánchez et. al.

Taxon Summary
MNI % NR % Weight %
Polyplacophora 1 0.1 1 0.1 0.2 0.0
Fissurella asperella 2 0.2 2 0.2 5.6 0.2
Fissurella sp. 2 0.2 2 0.2 2.7 0.1
Turbo saxosus 61 6.2 64 5.8 223.3 8.5
Tegula picta 217 22.1 217 19.7 379.7 14.4
Astraea buschii 127 12.9 149 13.5 480.1 18.2
Nerita funiculata 1 0.1 1 0.1 0.7 0.0
Littorina modesta 13 1.3 14 1.3 18.2 0.7
Cerithium menkei 1 0.1 1 0.1 6.4 0.2
Cerithium stercusmuscarum 1 0.1 1 0.1 0.8 0.0
Crucibulum scutellatum 1 0.1 1 0.1 11.4 0.4
Crepidula sp. 1 0.1 1 0.1 0.1 0.0
Polinices navidus 1 0.1 1 0.1 5.0 0.2
Trivia solandri 7 0.7 7 0.6 7.3 0.3
Cypraea cervinetta 4 0.4 5 0.5 61.4 2.3
Cypraea arabicula 2 0.2 3 0.3 9.5 0.4
Cassis centiquadrata 1 0.1 1 0.1 18.9 0.7
Bursa caelata 1 0.1 1 0.1 3.9 0.1
Muricopsis zeteki 1 0.1 1 0.1 1.2 0.0
Coralliophila nux 25 2.5 25 2.3 54.7 2.1
Thais triangularis 5 0.5 5 0.5 14.2 0.5
Thais haemastoma 64 6.5 68 6.2 138.4 5.2
Thais delessertiana 81 8.3 81 7.4 139.8 5.3
Thais melones 5 0.5 5 0.5 43.5 1.6
Acanthina brevidentata 22 2.2 22 2.0 42.9 1.6
Neorapana muricata 1 0.1 1 0.1 20.1 0.8
Cantharus elegans 3 0.3 3 0.3 17.1 0.6
Cantharus gemmatus 93 9.5 93 8.4 178.4 6.8
Cantharus vibex 13 1.3 13 1.2 19.3 0.7
Columbella labiosa 41 4.2 41 3.7 59.7 2.3
Columbella strombiformis 122 12.4 122 11.1 151.6 5.7
Anachis costellata 2 0.2 2 0.2 0.7 0.0
Mazatlania fulgurata 1 0.1 1 0.1 0.2 0.0
Oliva incrassata 4 0.4 4 0.4 29.7 1.1
Olivella columellaris 3 0.3 3 0.3 0.7 0.0
Conus gladiator 1 0.1 1 0.1 3.1 0.1
Conus patricius 1 0.1 1 0.1 12.3 0.5
Bulimulidae 2 0.2 8 0.7 2.3 0.1
Drymaeus sp. 2 0.2 2 0.2 0.3 0.0
Marine gastrop indet 8 0.8 13 1.2 6.2 0.2
Arcidae 1 0.1 3 0.3 2.7 0.1
Barbatia sp. 1 0.1 1 0.1 1.2 0.0
Anadara formosa 1 0.1 1 0.1 33.9 1.3
Mytilidae 1 0.1 8 0.7 9.9 0.4
Pteriidae 5 0.5 11 1.0 11.2 0.4
Pteria sterna 3 0.3 26 2.4 67.7 2.6
Pinctada mazatlanica 15 1.5 51 4.6 151.4 5.7
Ostrea sp. 1 0.1 1 0.1 37.4 1.4
Lyropecten subnodosus 1 0.1 1 0.1 54.7 2.1
Plicatula sp. 1 0.1 1 0.1 2.1 0.1
Spondylus prínceps unicolor 1 0.1 1 0.1 46.5 1.8
Pitar sp. 1 0.1 1 0.1 1.7 0.1
Protothaca zorritensis 1 0.1 2 0.2 18.2 0.7
Protothaca columbiensis 1 0.1 1 0.1 19.1 0.7
Marine bivalve indet 2 0.2 5 0.5 8.1 0.3
Marine mollusc indet 1 0.1 1 0.1 0.1 0.0
Total 981 1102 2637.5
Table 1: Abundance, expressed as MNI, NISP (NR) and weight, in grams, of the mollusc assemblages retrieved in
the Chácara I Sector from Ligüiqui (taxa in bold signal highly esteemed food items).
 TAXON Californian Province Panamian Province Peruvian Province Magallanic Province
40º N 30º N 20º N 10º N 0º N 10º S 20ºS 30ºS 40ºS 50ºS
Fissurella asperella
Solariella triplostophanus
Tegula picta
Astraea buschii
Nerita funiculata
Littorina modesta
Cerithium menkei
Cerithium stercusmuscar
Crucibulum scutellatum
Polinices navidus
Trivia solandri
Cypraea cervinetta
Cypraea arabicula
Cassis centiquadrata
Bursa caelata
Muricopsis zeteki
Coralliophila nux
Thais triangularis
Thais haemastoma
Thais delessertiana
Thais melones
Acanthina brevidentata
Neorapana muricata
Cantharus elegans
Cantharus gemmatus
Cantharus vibex
Columbella labiosa
Columbella strombiformis
Subsistence intensi cation: the molluscs from the Chácara I sector at Ligüiqui

Anachis costellata
Mazatlania fulgurata
Oliva incrassata
Olivella columellaris
Conus gladiator
Conus patricius
Anadara formosa
Pteria sterna
Pinctada mazatlanica
Lyropecten subnodosus
Spondylus prínceps unic
Protothaca zorritensis
Protothaca columbiensis

Widespread Distribution Tropical

413

Table 2: Biogeographical range of selected mollusc species from Ligüiqui.

414 Víctor F. Vásquez Sánchez et. al.

Taxon Supralittoral Mesolittoral Infralittoral Archibenthonic

Fissurella asperella
Turbo saxosus
Tegula picta
Astraea buschii
Nerita funiculata
Littorina modesta
Cerithium menkei
Crucibulum scutellatum
Crepidula sp.
Triva solandri
Bursa caelata
Muricopsis zeteki
Coralliophila nux
Thais triangularis
Thais haemastoma
Thais delessertiana
Thais melones
Acanthina brevidentata
Neorapana muricata
Cantharus elegans
Cantharus gemmatus
Cantharus vibex
Columbella labiosa
Columbella strombiformis
Anachis costellata
Conus gladiator
Conus patricius
Barbatia sp.
Ostrea sp.
Plicatula sp.
Spondylus princeps unicolor

Table 3: Zonal distribution of selected rocky substrate molluscs from Ligüiqui.

the northern Californian province (30o-40o N) and a fur-
ther 24 into the southern Peruvian province (10o-40o S)
(Keen 1971; Marinkovich 1973; Table 2). Only one clam
species, Protothaca zorritensis, is nowadays restricted
to the later province although it reaches to the southern
border of the Panamanian province.
5 Discussion
The features of the mollusc assemblages from the Chá-
cara I Sector re ect taphonomic, ecological, and util-
itarian (ethnographic) reasons, intermingled to such a
degree that it is often dif cult to ascribe the abundance
of any given taxon to a particular cause or a precise
combination of them.
From a utilitarian standpoint, the collection features only
six worked items. The most remarkable of these are a
rectangular fragment of a Spiny Oyster dorsal valve of
unknown functionality in UE 3 (Fig. 5A) and a perfo-
rated portion of a Pearl Oyster dorsal valve (worked as a
pendant?) from UE 32 (Fig. 5B). Pearl oysters worked as
pendants also appear on UE 13 and UE 3, with a possible
fourth pendant, too fragmented to allow identi cation,
retrieved in UE 0. The latter unit also yielded a shell bead
that was impossible to identify taxonomically. Absence
of fragments that signal the stages of the chaîne opéra-
Subsistence intensi cation: the molluscs from the Chácara I sector at Ligüiqui 415

Taxon Supralittoral Mesolittoral Infralittoral

Cerithium stercusmuscaron
Polinices ravidus
Cypraea cervinetta
Cypraea arabicula
Cassis centiquadrata
Mazatlania fulgurata
Oliva incrassata
Olivella columellaris
Anadara formosa
Pteria sterna
Pinctada maztlanica
Lyropecten subnodosus
Pitar sp.
Protothaca zorritensis
Protothaca columbiensis

Table 4: Zonal distribution of selected sandy substrate molluscs from Ligüiqui.

UE 1
TAXON
MNI % NR % Weight %
Turbo saxosus 39 9.8 39 9.5 115.0 13.0
Tegula picta 79 19.9 79 19.2 150.0 17.0
Astraea buschii 61 15.4 64 15.5 224.1 25.4
Nerita funiculata 1 0.3 1 0.2 0.7 0.1
Littorina modesta 1 0.3 1 0.2 1.3 0.1
Polinices navidus 1 0.3 1 0.2 5.0 0.6
Trivia solandri 3 0.8 3 0.7 2.8 0.3
Cypraea cervinetta 1 0.3 2 0.5 15.0 1.7
Bursa caelata 1 0.3 1 0.2 3.9 0.4
Muricopsis zeteki 1 0.3 1 0.2 1.2 0.1
Thais triangularis 1 0.3 1 0.2 1.3 0.1
Thais delessertiana 79 19.9 79 19.2 137.5 15.6
Acanthina brevidentata 1 0.3 1 0.2 1.4 0.2
Cantharus elegans 2 0.5 2 0.5 12.6 1.4
Cantharus gemmatus 38 9.6 38 9.2 75.0 8.5
Cantharus vibex 3 0.8 3 0.7 8.4 1.0
Columbella labiosa 2 0.5 2 0.5 4.2 0.5
Columbella strombiformis 76 19.2 76 18.4 106.4 12.0
Mazatlania fulgurata 1 0.3 1 0.2 0.2 0.0
Oliva incrassata 1 0.3 1 0.2 4.2 0.5
Olivella columellaris 1 0.3 1 0.2 0.2 0.0
Bulimulidae 1 0.3 4 1.0 0.2 0.0
Arcidae 1 0.3 3 0.7 2.7 0.3
Mytilidae 1 0.3 8 1.9 9.9 1.1
Total 396 412 883.2

Table 5: Abundance, expressed as MNI, NISP (NR) and weight, in grams, of the mollusc assemblage from UE 1.
416 Víctor F. Vásquez Sánchez et. al.

Fig. 4: The dominant molluscs at Ligüiqui: (A) Cantharus gemmatus (B)Tegula picta (C) Astraea buschii (photos
by V. F. Vásquez Sánchez, T. E. Rosales Tham and M. Castro Priego).

toire suggest that the scarce worked specimens must
have been produced elsewhere (Çakirlar 2009).
The 2mm mesh through which sediments were sieved is
large enough to allow a substantial number of items to
be lost in the retrieval process. This fraction would pref-
erentially affect small-sized taxa, but also shell splinters
from larger-sized taxa. We suspect that it this taphonomic
loss what could explain the scarce presence of the high-
ly esteemed local food item chitons (Polyplacophora),
whose shell is composed of small articulated plates. In
general, one may surmise that since shell splinters would

UE13
TAXON
MNI % NISP % Weight NISP
Polyplacophora 1 0.7 1 0.6 0.2 0.0
Turbo saxosus 2 1.3 2 1.1 26.2 6.0
Tegula picta 28 18.3 28 15.7 47.8 10.9
Astraea buschii 15 9.8 17 9.6 64.2 14.6
Littorina modesta 2 1.3 2 1.1 4.0 0.9
Cerithium menkei 1 0.7 1 0.6 6.4 1.5
Trivia solandri 1 0.7 1 0.6 0.7 0.2
Cypraea cervinetta 2 1.3 2 1.1 19.1 4.3
Cypraea arabicula 2 1.3 3 1.7 9.5 2.2
Cassis centiquadrata 1 0.7 1 0.6 18.9 4.3
Coralliophila nux 11 7.2 11 6.2 23.6 5.4
Thais triangularis 2 1.3 2 1.1 8.3 1.9
Thais haemastoma 25 16.3 26 14.6 52.2 11.9
Acanthina brevidentata 16 10.5 16 9.0 31.9 7.3
Cantharus gemmatus 15 9.8 15 8.4 29.3 6.7
Cantharus vibex 5 3.3 5 2.8 5.0 1.1
Columbella labiosa 10 6.5 10 5.6 12.8 2.9
Columbella strombiformis 3 2.0 3 1.7 2.2 0.5
Oliva incrassata 1 0.7 1 0.6 5.9 1.3
Conus gladiator 1 0.7 1 0.6 3.1 0.7
Barbatia sp. 1 0.7 1 0.6 1.2 0.3
Pteriidae 1 0.7 1 0.6 1.0 0.2
Pinctada mazatlanica 7 4.6 28 15.7 65.7 15.0
Total 153 178 439.2

Table 6: Abundance, expressed as MNI, NISP (NR) and weight, in grams, of the mollusc assemblage from UE 13.
Subsistence intensi cation: the molluscs from the Chácara I sector at Ligüiqui
Fig. 5: Worked shells from Ligüiqui: (A) Spiny Oyster (Spondylus princeps) (B) Pearl Oyster (Pinctada mazatlan-
ica) (photos by V. F. Vásquez Sánchez, T. E. Rosales Tham and M. Castro Priego).
be, for the most part, non-informative and the smallest
molluscs never consumed, this presumed loss of re-
mains most likely affected specimens used as ornaments
(beads) and small molluscs accumulated through natural
processes. The lack of background fauna precludes us
from making a reliable reconstruction of the original
mollusc community that existed in this area at the time
of the occupation.
To assess to what extent larger, thin-shelled, molluscs
may be also underrepresented, ecological data may
prove of help. As reported, our assemblages re ect a
rocky shore, as is today the case in the area, yet mus-
sels -a conspicuous and readily accessible resource on
rocky shores- are extremely scarce (Table 1). Given the
fragility of mussel shells, one may think that such low
abundance also re ects a huge taphonomic loss, but in
this case, ecology tells us otherwise. With exceptions, the
Ecuadorian coast does not have the cold and productive
waters of Perú, where mussels abound (Wilson 1981).
The low frequencies of mussels at Ligüiqui could thus
be most parsimoniously taken to re ect this low produc-
tivity of tropical waters, and render availability, rather
than taphonomic loss, the reason explaining that scarcity.
Ecological constraints could likewise help us assess the
scarcity of other thin-shelled taxa such as the terrestrial
snails (Bulimidae, Drymaeus sp.). Pulmonates, however,
are always scarce on the dry Ecuadorian and Peruvian
littoral and only become frequent in the more humid hab-
itats lying next to it (Gálvez 2010). Thin-shelled marine
molluscs -bivalves for the most part- in turn require loose
substrates to bury themselves and these are not readily
available in this sector of the coast (Alexander et al.
1993; Table 1). In other words, whereas taphonomic loss
is certainly a factor to reckon with, and, as we will see,
the assemblages does seem to re ect human activity for
the most part, ecological features suf ce to explain the
scarcity of taxa most likely to be missed by an inadequate
retrieval of remains.
417
In contrast, choice emerges as a powerful force dictat-
ing presence and abundance of most taxa at Ligüiqui.
Fully 21 species, representing more than 40% of the 49
marine taxa, are highly esteemed as food items in the
area today (Table 1, taxa in bold). Quantitatively, these
species represent some 70% of the items at the Chácara
I Sector, with independence of the estimator consid-
ered (i.e. MNI: 73.5%; NISP: 69.3%; weight: 71.6%).
These species evidence that the retrieved remains mostly
represent consumption refuse in all units (Tables 8, 9).
Availability may still be the reason explaining the low
frequencies of the most productive taxa (e.g. the bur-
ying venerid clams, Protothaca columbiensis, and P.
zorritensis), and of molluscs featuring the best quality
meat (e.g. the murex shell, Muricopsis zeteki, the ark
shell, Anadara formosa and, above all, the spiny oyster,
Spondylus princeps unicolor), since the latter three are
denizens of the less accessible infralittoral zone (Tables
3, 4). The case of the spiny oyster, one of the most highly
appreciated mollusc meat in the world, is noteworthy
in that the only specimen retrieved is not only worked
but also represents an exotic species that only reaches
to the upper fringes of the Panamanian province (i.e.
20o N; Table 2). Be it as it may, the scarcity/absence of
these food items suggests that the diet exempli ed by
the remaining edible species re ect that of a low social
level group as presumed for the workers that inhabited
the “Chácara I” Sector (M. Castro, unpublished).
The completeness of most shells and the absence of trac-
es of re hint at a non-destructive method for extracting
the meat. In the case of gastropods, this most probably
involved spikes or hooks to poke the soft tissue out. The
absence of any artefacts of this kind in the excavation
hints at organic instruments of which the local vegeta-
tion features two frequent spine-bearing species (Fig.
6). Interesting here is that this presumed non-destructive
meat-extraction technique is not the method locals today
employ, since shells are systematically smashed with
418
Fig. 6: Frequent spiny bushes around Ligüiqui today: (A) Prosopis pallida (“Algarrobo”) (B) Pilosocereus
tweedyanus (Cactus), with a close-up of its spine (photos by L. Alonzo and M. Castro Priego
stones. Perhaps the fact that the Ligüiqui community is a
newcomer into this area (they settled in the last century
from nearby areas) may explain the “clumsy” way in
which they access the meat of snails.
Although the archaeological sequence at Ligüiqui covers
a lapse of less than 200 years (see above), a comparison
of the richest units in terms of NISP (NR), that also hap-
pen to coincide with the earliest (UE1: Tola phase) and
latest (UE13: kitchen oor) moments of the occupation,
reveals interesting trends (Tables 5, 6; Figs. 5, 6).
One rst difference relates to the increase of bivalves
with time. At UE1, these only constitute 0.5% of the
total MNI (2.6% of the NISP and 1.4% of the weight),
whereas in UE 13, bivalves rise to ca. 6% of the MNI,
representing 17% of the NISP and 15.5% of the weight
(Tables 5, 6). In other words, a ca. tenfold increase in
bivalves from the start to the end of the occupation.
That increase mimics the more than tenfold increase
of deeper-water (infralittoral) taxa from UE1 to UE13.
Indeed, infralittoral molluscs represent 1.2% of the MNI
in UE1 but ca. 10% at UE 13. Other estimators follow
Víctor F. Vásquez Sánchez et. al.
this same trend (NISP: 1.1% (UE1) vs. 20.8% (UE13);
Weight: 2.4% (UE1) vs. 24% (UE13)). The taxa most-
ly responsible for this increment are the pearl oysters,
Pinctada mazatlanica and Pteriidae, absent at UE1 and
hinting at an expansion of the shell shing activity into
deeper waters (Tables 5, 6). Since pearl oysters remain
always below water, the move towards deeper water
may concomitantly imply an implementation of diving
as part of the shell shing activity, re ecting an interest
in pearls for which no evidence exists at the time the
occupation started.
Other taxonomic differences between units UE1 and
UE13 are more dif cult to account for. The presence of
mussels at UE1 and their absence at UE13, for example,
could be read along the line of the move towards deeper
water, yet may also re ect a “stochastic construct” of
sorts caused by the negligible NISP of mussels. In terms
of molluscs considered esteemed foodstuffs, some differ-
ences may also be worth commenting. The most striking
ones are the absences in UE1 of certain species that
constitute signi cant items at UE13 (e.g. Coralliophyla
nux and Thais haemastoma) and viceversa (e.g. Thais
Subsistence intensi cation: the molluscs from the Chácara I sector at Ligüiqui
UE1
Species
N Min Max Y S
Astraea buschii 30 16.5 38.9 22.4
Tegula picta 30 13.8 21.6 16.8
Cantharus gemmatus 30 10.08 30.4 24.3
Table 7: Descriptive statistics of the total lengths of the three most common mollusc species at Ligüiqui found in
the oldest (UE1) and one of the young (UE13) archaeological units.
desertiana only present at UE1). Essentially the same
would apply to the tenfold increase of certain species
(e.g. Solariella triplostophanus: 10% (UE1) vs. 1.3%
(UE13); Acanthina brevidentata: 0.3% (UE1) vs. 10.5%
(UE13); Columbella strombiformis: 19.2% (UE1) vs.
2% (UE13) (percentages refer to MNI). Could these
differences indicate environmental changes, or, alterna-
tively, overexploitation events or merely re ect a shift-
ing desirability of species through time? Although it is
dif cult to provide de nitive answers to these questions
with the data at hand, we need to note that the marine
environment around Ligüiqui remained essentially stable
throughout the time the occupation lasted, as also did
the frequencies of the dominant taxa, T. picta and A.
buschii (Tables 5, 6; Figs. 5, 6). In principle, this leaves
little room to address the reasons behind the taxonomic
“shifts” recorded between UE1 and UE13. The question
of a putative overexploitation of resources leading to
local extinctions of the less resilient species, on the other
hand, can be assessed through a biometrical analysis of
the collections.
Table 7 summarises the descriptive statistics of the TL
(total length) values from three snail species found in
UE1 and UE13, including the two most common mol-
luscs at Ligüiqui, Tegula picta and Astraea buschii. Al-
though the samples from UE13 do not meet the N≥30
criterion when a t-Student test is used to compare the
means from two paired samples, the results are never-
theless revealing. As can be seen, the means and ranges
of TL do not differ noticeably between both units in the
case of Cantharus gemmatus, an impression that gets sta-
tistical con rmation through a t-Student test that con rm
the differences not being signi cant (p1= 0.092 ≥0.05;
p2= 0.184 ≥0.05). On the other hand, both the minimum,
maximum and mean TL values of T. picta and A. buschii
are systematically higher at UE1 and lower at UE13,
again con rmed by the t-Student tests. In this way, for
T. picta p-values are p1= 0.01 ≤ 0.05 and p2= 0.02 ≤
0.05, whereas for A. buschii p-values are p1= 0.0005 ≤
0.05 and p2= 0.001 ≤ 0.05 (i.e. no signi cant differences
exist). In other words, what the statistical tests tell us is
that the two dominant species did get smaller in a mere
150 years of occupation, a feature taken as a hallmark
of processes of intensi ed harvesting as documented
since the MSA (Middle Stone Age) in Africa (Steele &
419
UE13
CV N Min Max Y S CV
6.14 27.41 12 12.91 27.7 20.9 4.85 23.21
1.74 10.36 18 8.77 20 14.6 3.11 21.29
3.9 16.05 11 13.72 32.49 23 5.8 25.22
Klein 2008; Mannino et al. 2014). According to these
authors, overexploitation does not necessarily require
large human populations nor intensi ed production to
take place provided a population remains in an area for
a prolonged period. In other words, sedentism fosters
overexploitation, and eventually local extinctions, even
in low-density demographic conditions with the lowest
levels of technological development. For these reasons,
one has here ground to suspect that the ups and downs
seen in the abundance of species from UE1 and UE13, or
even their disappearance in the youngest unit, may re ect
this phenomenon. Overexploitation could be thus taken
as a proxy of the sedentism of the population stationed at
Ligüiqui. When one combines these shifting abundances
and diminishing sizes with the presumed move offshore,
and the inferred development of diving activity to collect
pearl shells, the conclusion one reaches is that during this
presumably restricted time interval an intensi cation of
shell shing activities did occur in this spot of the Ec-
uadorian coastline, an issue that other records, faunal or
not, will help us con rm in a near future.
Though Ligüiqui represents a singular site within the
context of the Manteño culture due to its complex sys-
tem of shing installations, an intense marine exploita-
tion has been archaeozoologically documented on the
coast of Manabí during the Chorrera (1050-50 BC) and
Guangala (50 BC-950 AD) cultural stages at the site of
Salango (Béarez et al. 2012; Damp 2014, 17). As such,
it is to Salango that one will need to look to frame the
Ligüiqui data presented in this paper.
6 Conclusions
Considering the intense production activity centred on
Spondylus that had been hitherto postulated for Ligüiqui,
the most remarkable result of this study is that molluscs
provide scarce evidence suggesting production, includ-
ing craftwork, and/or trade at any signi cant level be-
yond subsistence. For this reason, that hypothesis, brie y
entertained in the introductory section, must be provi-
sionally rejected. The data, on the other hand, suggest
that these molluscs represent the food refuse of a group
that the archaeological evidence hints as the operators
of the spectacular shing installations. The homogeneity
of the mollusc assemblages through time additionally
 UE 0: Road UE 1 UE 3 UE5 UE 6 UE 7 UE 9
420

TAXON
MNI NR Weight MNI NR Weight MNI NR Weight MNI NR Weight MNI NR Weight MNI NR Weight MNI NR Weight
Polyplacophora
Fissurella asperella 2 2 5.6
Fissurella sp. 1 1 0.2
Turbo saxosus 39 39 115.0 2 2 14.0 4 5 18.3 3 5 16.4 1 1 4.2
Tegula picta 79 79 150.0 3 3 4.6 29 29 41.1 4 4 7.1 1 1 0.7 20 20 29.6
Astraea buschii 61 64 224.1 7 18 16.0 7 9 20.2 2 4 7.8
Nerita funiculata 1 1 0.7
Littorina modesta 1 1 1.3 1 1 1.2 3 3 4.7 2 2 1.6 2 3 2.6
Cerithium menkei
Cerithium stercusmuscarum 1 1 0.8
Crucibulum scutellatum 1 1 11.4
Crepidula sp. 1 1 0.1
Polinices navidus 1 1 5.0
Trivia solandri 3 3 2.8
Cypraea cervinetta 1 2 15.0 1 1 27.3
Cypraea arabicula
Cassis centiquadrata
Bursa caelata 1 1 3.9
Muricopsis zeteki 1 1 1.2
Coralliophila nux
Thais triangularis 1 1 1.3 1 1 2.0
Thais haemastoma 7 8 14.6 6 7 10.7 3 3 4.1 5 5 9.4
Thais delessertiana 79 79 137.5 2 2 2.3
Thais melones 2 2 8.8
Acanthina brevidentata 1 1 1.4 1 1 3.2
Neorapana muricata
Cantharus elegans 2 2 12.6
Cantharus gemmatus 38 38 75.0 1 1 2.4 1 1 1.4 8 8 11.8 2 2 4.4
Cantharus vibex 3 3 8.4 1 1 1.6 1 1 1.3
Columbella labiosa 2 2 4.2 5 5 4.4 3 3 3.0 2 2 3.5 5 5 6.1
Columbella strombiformis 76 76 106.4 4 4 6.2 10 10 6.3 7 7 7.1 1 1 1.3
Víctor F. Vásquez Sánchez et. al.
 UE 0: Road UE 1 UE 3 UE5 UE 6 UE 7 UE 9
TAXON
MNI NR Weight MNI NR Weight MNI NR Weight MNI NR Weight MNI NR Weight MNI NR Weight MNI NR Weight
Anachis costellata
Mazatlania fulgurata 1 1 0.2

Oliva incrassata 1 1 4.2 2 2 19.6

Olivella columellaris 1 1 0.2 1 1 0.3

Conus gladiator
Conus patricius 1 1 12.3

Bulimulidae 1 4 0.2 1 4 2.1

Drymaeus sp.
Marine gastrop indet 5 10 4.0 3 3 2.2

Arcidae 1 3 2.7

Barbatia sp.
Anadara formosa
Mytilidae 1 8 9.9

Pteriidae 2 3 3.6 1 4 2.2

Pteria sterna 2 23 66.0 1 3 1.7

Pinctada mazatlanica 1 5 21.0

Ostrea sp.
Lyropecten subnodosus
Subsistence intensi cation: the molluscs from the Chácara I sector at Ligüiqui

Plicatula sp.
Spondylus prínceps unicolor 1 1 46.5

Pitar sp. 1 1 1.7

Protothaca zorritensis 1 2 18.2

Protothaca columbiensis
Marine bivalve indet 1 1 0.2 1 4 7.9

Marine mollusc indet 1 1 0.1

Total 3 3 0.5 396 412 883.2 24 26 114.2 86 132 218.7 41 48 104.5 5 7 7.2 43 50 115.9
421

Table 8: Ligüiqui: Molluscs from units 0, 1, 3, 5, 6, 7 & 9.

 UE 13 UE 21 UE 26 UE 27 UE 28 UE 32
422

TAXON
MNI NR Weight MNI NR Weight MNI NR Weight MNI NR Weight MNI NR Weight MNI NR Weight
Polyplacophora 1 1 0.2
Fissurella asperella
Fissurella sp. 1 1 2.5
Turbo saxosus 2 2 26.2 9 9 25.1 1 1 4.1
Tegula picta 28 28 47.8 5 5 9.1 21 21 43.3 16 16 26.6 11 11 19.8
Astraea buschii 15 17 64.2 1 1 8.6 3 3 21.6 23 23 86.5 7 9 26.3 1 1 4.8
Nerita funiculata
Littorina modesta 2 2 4.0 2 2 2.8
Cerithium menkei 1 1 6.4
Cerithium stercusmuscarum
Crucibulum scutellatum
Crepidula sp.
Polinices navidus
Trivia solandri 1 1 0.7 1 1 1.4 1 1 1.8 1 1 0.6
Cypraea cervinetta 2 2 19.1
Cypraea arabicula 2 3 9.5
Cassis centiquadrata 1 1 18.9
Bursa caelata
Muricopsis zeteki
Coralliophila nux 11 11 23.6 8 8 16.1 6 6 15.0
Thais triangularis 2 2 8.3 1 1 2.6
Thais haemastoma 25 26 52.2 2 3 2.5 3 3 5.4 11 11 23.5 2 2 16.0
Thais delessertiana
Thais melones 1 1 20.5 1 1 4.6 1 1 9.6
Acanthina brevidentata 16 16 31.9 2 2 2.7 2 2 3.7
Neorapana muricata 1 1 20.1
Cantharus elegans 1 1 4.5
Cantharus gemmatus 15 15 29.3 3 3 6.9 9 9 15.1 12 12 21.0 4 4 11.1
Cantharus vibex 5 5 5.0 3 3 3.0
Columbella labiosa 10 10 12.8 2 2 2.7 11 11 20.3 1 1 2.7
Víctor F. Vásquez Sánchez et. al.
 UE 13 UE 21 UE 26 UE 27 UE 28 UE 32
TAXON
MNI NR Weight MNI NR Weight MNI NR Weight MNI NR Weight MNI NR Weight MNI NR Weight
Columbella strombiformis 3 3 2.2 4 4 3.6 4 4 4.5 13 13 14.0
Anachis costellata 1 1 0.3 1 1 0.4
Mazatlania fulgurata
Oliva incrassata 1 1 5.9
Olivella columellaris 1 1 0.2
Conus gladiator 1 1 3.1
Conus patricius
Bulimulidae
Drymaeus sp. 1 1 0.2 1 1 0.1
Marine gastrop indet
Arcidae
Barbatia sp. 1 1 1.2
Anadara formosa 1 1 33.9
Mytilidae
Pteriidae 1 1 1.0 1 3 4.4
Pteria sterna
Pinctada mazatlanica 7 28 65.7 2 4 13.6 3 8 40.1 2 6 11.0
Ostrea sp. 1 1 37.4
Lyropecten subnodosus 1 1 54.7
Subsistence intensi cation: the molluscs from the Chácara I sector at Ligüiqui

Plicatula sp 1 1 2.1
Spondylus prínceps unicolor
Pitar sp.
Protothaca zorritensis
Protothaca columbiensis 1 1 19.1
Marine bivalve indet
Marine mollusc indet
Total 153 178 439.2 2 2 29.1 30 33 74.1 96 98 331.5 76 83 177.9 26 30 141.5
423

Table 9: Ligüiqui: Molluscs from units 13, 21, 26, 27, 28 & 32.
424
hints at a culinary tradition that did not change through-
out the study period. Such culinary tradition does not
imply stasis, since both subtle shifts in the frequency
of certain species, along with the disappearance and the
appearance of some others, appear when the faunas from
the oldest and youngest deposits are confronted. These
trends, in combination with the deeper bathymetrical
ranges of the most recent taxa and the size reduction of
the most common ones, hint at signi cant developments
of the exploitation strategies through time. Whether the
diminishing frequencies of the common molluscs and
the presumed displacement of shell- shing into deeper
waters re ect a depletion of resources, a preferential
cropping of pearl oysters for reasons disconnected with
subsistence, or simply a construct due to an inadequate
retrieval, are issues that data retrieved in a more reliable
way should allow us to eventually con rm.
Acknowledgements
Support was received by the Spanish Ministry of Econ-
omy and Competitiveness projects HAR 2014-55722-P
(Ictioarqueologia de la Prehistoria cantábrica: Modelos
para la caracterización de las primeras pesquerías eu-
ropeas), HAR 2017-88325-P (Historical archaeobiology
of the European hake (Merluccius merluccius, L.1758)
in the NE Atlantic: The Iberian evidence (AD 10th–18th)
and the PID2020-118662GB-100 (FISHCIIS - Fishing
Isotopes) project from the Ministry of Science and In-
novation.
The Ligüiqui Research Project is a part of Perduraciones,
continuidad y ruptura. Nuevas realidades de desigual-
dad en la costa ecuatoriana central (ss. XVI-XVII):
indicadores arqueológicos y transformaciones medio-
ambientales, funded by the Spanish Ministry of Culture
and Sports (Spanish abroad archaeological Programme),
during 2018 and 2019 (Ref: 43- T002018N0000042859
and T002019N0000038722), Palarq Foundation (2018-
2020), together with the Research Programme of the
National Institute of Cultural Heritage of Ecuador. This
publication is a part of the research group Archaeolo-
gy: Landscapes, Colonialism and Cultural Heritage of
Alcalá University.
We are grateful for the information provided by the
Ligüiqui community, especially by Leonardo Alonzo.
Víctor F. Vásquez Sánchez et. al.
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