BULLETIN OF MARINE SCIENCE. 89(4):1015–1035.

2013
http://dx.doi.org/10.5343/bms.2012.1066

Archaeological Investigations Provide

Late Holocene Baseline Ecological
Data for Bocas del Toro, Panama

Thomas A Wake, Douglas R Doughty, and Michael Kay

Abstract
Recent archaeological investigations at Sitio Drago, Bocas del Toro, Panama, have
provided copious amounts of well-preserved shell and bone. In terms of animal
remains, marine resources dominate the archaeological record in Bocas del Toro.
This is not surprising given the diversity of marine habitats available to the region’s
pre-Columbian inhabitants. The high marine biodiversity of the region provides
a wide variety of potential food sources for humans. Animal remains recovered
from archaeological sites in the region represent subfossil baseline Late Holocene
ecological samples that provide an opportunity to examine past human subsistence
behavior in a coastal environment dominated by bays, coral reefs, and extensive
mangrove stands. We examine marine mollusc and vertebrate remains from the
archaeological locality of Sitio Drago and illustrate evidence of potential pre-
Columbian human impacts on the local environment. The data we present shed
light on past human food preferences, resource focus, and predation effects over
a 700-yr occupation. We highlight the utility of archaeological faunal remains for
providing baseline ecological data for comparison with modern samples.

Archaeologists have been successful at documenting the effects of human hunt-

ing (vertebrates) and collecting (molluscs) on local resources in a variety of coastal
environments, including the Caribbean (Wing 2001, Wing and Wing 2001, Blick
2007, Carder et al. 2007), California (Braje et al. 2007, Erlandson et al. 2008, 2011,
Erlandson and Rick 2010), and Polynesia (Butler 2001, Morrison and Hunt 2007,
Ono and Intoh 2011). Recently there has been considerable debate about establish-
ing baseline data for fisheries and marine vertebrate populations in the Caribbean
(Baisre 2010a,b, Butler 2010, Curet 2010, deFrance 2010, Jones 2010, McClenachan
et al. 2010, Carder and Crock 2012). Baisre (2010a) received criticism from several
archaeologists for his apparent dismissal of archaeological data as useful for estab-
lishing baseline fisheries data for the Caribbean. The establishment of a baseline is
essential if we are to measure the effects of human exploitation on the Caribbean’s
various marine resources.
From a fisheries standpoint, archeological data can provide a great deal of infor-
mation concerning which species were caught from a given time period. What ar-
chaeologists cannot do is provide “unbiased” data prior to the time of human arrival
at archaeological sites and their effects on local resources. Archaeological assem-
blages can be viewed as representing the Holocene fossil record, with one impor-
tant exception—they represent the animals selected for consumption by humans.
Archaeological records can provide insight into human resource preferences, and
once established, the possible effects people have had on these resources.

Bulletin of Marine Science 1015

© 2013 Rosenstiel School of Marine & Atmospheric Science of
the University of Miami
1016 BULLETIN OF MARINE SCIENCE. VOL 89, NO 4. 2013

Archaeology of Bocas del Toro

Panama’s archaeological history is rich, beginning with initial human coloni-

zation during the Terminal Pleistocene (11,100–10,000 BP), moving into an Early
Preceramic phase that saw plant domestication by 8000 BP and to swidden farming
with maize and root crops by 7000 BP (Piperno 2009, 2011a,b). The earliest ceramic
production in Central America (Monagrillo, 4500–3200 BP) occurred in Central
Panama, with subsequent regional diversification (Cooke 2005). Evidence for so-
cial ranking appears ca. 2000 BP and for chiefdoms with structured hierarchies by
1500 BP in some areas of the Pacific watershed (Cooke 2005, Hoopes 2005). The
food remains left behind at archaeological habitation sites can provide a great deal of
information concerning hunting and fishing practices, paleoenvironmental recon-
struction, and baseline ecological data for times soon after humans began to affect
pristine tropical American coasts.
Overshadowed by the archaeology of the central provinces, the archaeology of
Bocas del Toro Province and Isla Colón remained completely unknown until 1949,
when Matthew Stirling led an expedition to the region (Stirling and Stirling 1964).
They recorded and excavated various sites in the Almirante Bay region including one
on the beach at Punta Drago. Gordon (1962, 1982) undertook limited archaeological
investigations in the region and noted a distinct absence of archaeological sites on
Isla Colón.
Linares and Ranere (1980) conducted excavations at four sites on the Aguacate
Peninsula at Cerro Brujo during the 1970s. They estimated the Aguacate Peninsula
population numbered approximately 120 persons living among four dispersed ham-
lets, each consisting of small structures roughly 300 m apart (Linares and Ranere
1980). Five radiocarbon dates illustrate that the Cerro Brujo sites were occupied be-
tween AD 880 and 1250 (Linares 1977, Linares and Ranere 1980). An earlier phase,
dated to approximately 1400 BP based on ceramic association, was also identified.
According to Linares (1977:311), “the archaeological settlements of Bocas prov-
ince appear to represent marginal populations organized on the basis of small family
(?) groups, without status differentiation or political organization of any recogniz-
able kind.” Linares and Ranere (1980:66) suggest that the populace of the Aguacate
Peninsula existed in a cultural “backwater,” especially when compared to the better-
known Panamanian chiefdoms on the Pacific coast (see also Steward 1948, Steward
and Faron 1959, Haberland 1984). While Linares (1977:310) does state that larger
sites may exist in the region, she adds that Cerro Brujo “was fairly typical of settle-
ment and subsistence patterns in Bocas.”
Recent archaeological surveys and excavations in the central Panamanian
Caribbean watersheds of Colón and Veraguas provinces (Griggs 2005), highland
Chiriquí (Holmberg 2009, Palumbo 2009), and greater Bocas del Toro region (Chavez
et al. 1996, Baldi 2001, Wake 2006, Wake et al. 2004, 2012) suggest a more wide-
spread and diverse human presence with extensive settlements (Griggs 2005) and
earlier than anticipated occupations (Drolet 1980, Griggs et al. 2002, Griggs 2005,
Wake 2006). Likewise, Black Creek, in Costa Rica close to the Panamanian border,
was occupied between 1000 and 500 BC (Chavez et al. 1996, Baldi 2001), adding con-
siderable support to the impression that Caribbean coasts of Lower Central America
were occupied considerably earlier than Linares’ 1400 BP estimate for earliest settle-
ment of the Aguacate Peninsula.
 wake et al.: baseline ecological data from sitio drago 1017

Sitio Drago (henceforth SD, Fig. 1), located near the northwest corner of Isla Colón,
is a large (15 ha) site with deep cultural deposits and a diverse artifact assemblage
suggesting a relatively large population. The site also appears to be more internally
complex than the loosely organized farming hamlets seen at Cerro Brujo (hence-
forth CB), with several obvious (1–2 m in height) mounds of domestic refuse that
may represent living structures located among scattered low mounded artifact and
shell concentrations. Ceramic evidence recovered from SD indicates strong links
with the Pacific coastal regions of central Panama (Greater Coclé), western Panama
(Chiriquí), Costa Rica (Diquís), even as far as central and northwestern Costa Rica
(Guanacaste), and suggests SD may have functioned as a regional trading center.
Broken jaguar effigy metates and sculpture fragments from surface contexts, and
high recovery of shell and bone ornaments compared to elsewhere in the region in-
dicate the presence of elite trade goods and the possibility of social ranking (Wake et
al. 2004, 2012, Wake and Mendizabal in press).
Fifteen radiometric age determinations date occupation of Sitio Drago to AD 690–
1410 (Wake et al. 2004, 2012, Wake 2006, Wake and Mendizabal in press), contem-
poraneous with CB. The presence of basin milling stones, carbonized remains of
tree crops, a diverse array of terrestrial forest reptiles and mammals, and a variety of
reef, mangrove, and beachfront molluscs and fish recovered at SD (Wake et al. 2004,
2012, Wake and Mendizabal in press) generally are consistent with Linares’ (1976,
1977, Linares and Ranere 1980) “vegeculture” (reliance on domesticated root crops
and wild tree crops), “garden hunting,” and marine fishing subsistence model for CB
(Wake et al. 2004, Wake 2006, Wake and Mendizabal in press). However, the addi-
tional presence of offshore fish species and flat milling stones at SD may suggest that
maize cultivation and open-ocean fishing were regular activities at SD as opposed to
raising solely root crops, and obtaining local terrestrial vertebrates and fish.

Geological Context

Boca del Drago (9°24´56˝N, 82°19´37˝W) is located on the northwestern corner of

Isla Colón, some 18 km to the northwest of the town of Bocas del Toro (Fig. 1). The
site itself lies on top of a stabilized beach ridge consisting of reworked coralline sands
derived from reef erosion.
Three litho-stratigraphic units are present on Isla Colón (Coates et al. 2005): the
Plio-Pleistocene (3.5–2.0 Ma) Old Bank Formation, the oldest, includes blue-gray
mudstones, fine sandstones, fine volcanic conglomerates, and interwoven beds of
volcanic rock; the Early Pleistocene (2.0–1.77 Ma) La Gruta Formation, covering the
north-central part of the Island, consists of recrystallized reefs and reef rubble; and
the more recent Pleistocene Ground Creek Formation, comprised of newer reef de-
posits inserted and superimposed on the La Gruta Formation.

Archaeological Context

Occupation at SD is divided into two distinct phases based on typological similari-

ties between pottery recovered in the excavations, fifteen radiocarbon dates (Wake et
al. 2012), and radiocarbon-dated pottery samples from other sites in western Panama
and Costa Rica. The earlier phase of occupation, exemplified from the bottom of the
archaeological deposit to 30 cm below the ground surface, dates from approximately
1018 BULLETIN OF MARINE SCIENCE. VOL 89, NO 4. 2013

Figure 1. Bocas del Toro, Isla Colón, Panama, Sitio Drago location.

AD 800 to 1100 and is characterized by a wide variety of ceramic artifacts, dominat-

ed by locally produced utilitarian wares and Bocas Brushed wares (e.g., Kudarauskis
et al. 1980, Linares 1980a). Bocas Brushed wares often have raised figures of marine
organisms such as starfish, octopi, crabs, lobsters, fishes, dolphins, and birds ap-
plied to the shoulders and/or handles of the vessels. A variety of less common high-
quality imported ceramics from the Pacific coast of central Panama from western
Veraguas to Coclé (Conte Polychrome; Cooke 2012), Chiriquí and southwest Costa
Rica (Chocolate Incised), central Costa Rica (Irazú Yellow-on-red), northwest Costa
Rica (Mora Polychrome), and southwest Nicaragua (Papagayo Polychrome) is also
found in the earlier phase deposits.
The second, or later occupational phase dates from approximately AD 1100 to
1400, and is characterized by the presence of high-fired, thin-walled Bisquit Ware
(Linares de Sapir 1968a,b). These ceramics, found only in the upper 30–40 cm of
the deposit, are varying shades of beige with smooth bodies and appliqué decora-
tion, including figurines at the shoulders and geometric designs applied on the neck.
The Bisquit Ware sherds may represent vessels imported from the Pacific slope of
Chiriquí or a locally produced variant.

Archaeological Excavation

A total of 48 1 × 1 m pits have been excavated at SD. The units are numbered
consecutively from those excavated in the first season to the most recent. Each unit
was excavated in 10 cm levels. Diagnostic artifacts, ceramic clusters, large bones,
some shell clusters, and other significant finds were plotted in three dimensions in
 wake et al.: baseline ecological data from sitio drago 1019

Figure 2. Sitio Drago site map surveyed and drawn by H Barnard, August 2006. 100 m grid lines
within Zone 17 P, projected on the WGS84 Geode. 0.2 m contour lines above local sea level.

situ and collected. All excavated soil is wet-screened through 3 mm mesh in the
field. Ceramics, stone, large shells, and other diagnostic artifacts are collected from
the screen and bagged. The remaining material—a mix of shell, bone, carbon, a few
artifacts, and miscellaneous material are bagged, dried, and eventually sorted and
analyzed. Three key areas of the site have received more extensive excavation: a 1 × 2
m pit in Mound 6 (Unit 1), a 3 × 3 m pit in Mound 1 (including Unit 22), and a 2 × 8
m trench (including Unit 14) in the Mortuary Area (Fig. 2).

Animal Remains

Molluscs
Shells are one of the most common objects encountered at SD. Specimens were
identified using field guides (Abbott and Morris 1995) and confirmed in consultation
with comparative collections held at the Smithsonian Tropical Research Institute.
Based on the specimens, three primary mollusc habitats were exploited by the in-
habitants of SD: sandy beaches, shallow water reef flats, and mangroves. In terms of
raw numbers, sandy beach species dominate, with those from the two other habitats
well represented. In general, analysis indicates a logistical mollusc collection strat-
egy targeting specific microhabitats known to contain concentrations of certain spe-
cies. Collection techniques may have included harvesting individual species by hand
(most of the identified Gastropods, see Online Table 1 for mollusc species authori-
ties), forced removal of sessile bivalves (Arca zebra and Anadara spp.), stripping of
mangrove roots (Crassostrea rhizophorae and Isognomon alatus), and perhaps bulk
harvesting with rakes or openwork baskets (Chione cancellata, Donax variabilis,
Anomalocardia cuneimeris).
1020 BULLETIN OF MARINE SCIENCE. VOL 89, NO 4. 2013

Identified Mollusc Species.—Some 86 genera and 76 species of molluscs are iden-

tified at SD (Online Table 1). Of these, 15 taxa represent approximately 92% of the
molluscs encountered (Table 1). The other 8% include large gastropods that are
considered valuable food today (Lobatus gigas, Cassis madagascarensis, Charonia
variegata, Melongena patula, Turbinella angulata, and Vasum muricatum), several
edible bivalves (scallops such as Amusium laurenti, Argopecten gibbus, Nodipecten
nodosus, and other bivalves including Arca imbricata, Barbatia cancellaria, Chama
macerophylla, Chione paphia, Periglypta listeri, and Tivela mactroides, among oth-
ers), and other probable bycatch species commonly associated with the primary prey
in their respective microhabitats. Some of these dietary bycatch species may have
been used for decorative purposes (including Ancilla sp., Cyphoma gibbosum, Conus
spp., Cypraea sp., Oliva reticularis) or for use as dye (Purpura patula). Both L. gigas
and C. variegata were used for shell trumpets.
Discrete concentrations of certain bivalve species (C. cancellata, D. variabilis, A.
cuneimeris, C. rhizophorae, P. imbricata, and I. alatus) were commonly encountered
in various excavation units and levels across the site. These concentrations suggest
that the collectors were quite selective and processed certain species in discrete
events as opposed to opportunistically harvesting and processing whatever mix of
species was encountered.
The gastropods would almost certainly have been collected by hand from their
respective preferred habitats. Few shells of any identified species appear burned, sug-
gesting that most cooking occurred in vessels such as ceramic pots.
Trends through Time in the Molluscs.—To investigate whether human harvest-
ing affected local molluscs at SD, we assessed size through time in two of the most
common species: A. cuneimeris and A. zebra. These two species have different life

Table 1. Fifteen most common mollusc species uncovered at Sitio Drago (number of valves / 2 for
bivalves, Doughty 2011).

Taxon Count Percent

Arca zebra 4,210.5 17.12
Crassostrea rhizophorae 3,367.5 13.68
Anomalocardia cuneimeris 3,075.5 12.50
Chione cancellata 2,823.0 11.48
Donax sp. 2,451.0 9.96
Anadara notabilis 2,384.5 9.69
Pinctada imbricata 812.5 3.31
Strombus pugilis 736.0 2.99
Lobatus raninus 729.0 2.96
Isognomon alatus 714.0 2.91
Codakia orbicularis 484.5 1.97
Chicoreus pomum 438.0 1.78
Fasciolaria tulipa 207.0 0.84
Arcopagia fausta 159.5 0.65
Cittarium pica 131.0 0.53
“Others” 1,877.5 7.63
Total 2,4601.0 100.00
Top 15 22,723.5 92.37
All others 1,877.5 7.63
 wake et al.: baseline ecological data from sitio drago 1021

Figure 3. Relative abundances of select molluscs through occupational phases at units 3, 14, and
22 at Sitio Drago, Bocas del Toro. (A) Crassostrea rhizophorae, (B) Isognomon alatus, (C)
Pinctada imbricata, (D) Arca zebra, (E) Anadara notabilis, (F) Donax sp., (G) Anomalocardia
cuneimeris, (H) Chione cancellata, (I) Codakia orbicularis, (J) Lobatus raninus, and (K)
Strombus pugilis.

histories; A. cuneimeris is a mobile shallow sandy beach inhabitant (Abbot and

Morris 1995). Arca zebra is sessile and attaches to structure on shallow reef flats
(Abbot and Morris 1995).
In total, 14,572 complete representatives of these two species (A. zebra: 8421, A.
cuneimeris: 6151) were selected from 18 excavation units across SD. All whole shells
were measured with digital calipers and differences tested using an M-test. Shell
size remained constant through time in A. zebra (r = −0.062, P < 1.675) and actually
increased slightly in A. cuneimeris (r = −0.043, P < 0.009) (Doughty 2011), suggest-
ing a lack of over exploitation. The slight size increase in A. cuneimeris may indicate
a possible landscape or climate change may have favored these species or, perhaps,
collectors accessed more distant patches (e.g., Broughton 1999).
Relative Frequency of Molluscs through Time.—Within the 15 most common mol-
luscs that are almost always present at SD, several shifts in relative frequency can be
observed through time (Fig. 3; Doughty 2011). While these species are always present,
their numerical representation does change through time. Each of these charts presents
the relative frequency of one of the 15 most common species. Units 3, 14, and 22 rep-
resent two different areas of the SD site (Fig. 2). Units 3 and 22 are at opposite ends of
Mound 1. Unit 14 lies within the mortuary complex, just outside the northern corner
of Tomb 1 (see Wake et al. 2012, Wake and Mendizabal in press).
1022 BULLETIN OF MARINE SCIENCE. VOL 89, NO 4. 2013

Oysters.—Two of the oyster species identified at SD are most commonly found at-
tached to mangrove roots: C. rhizophorae and I. alatus. The other oyster in the top
15, the pearl oyster, Pinctada imbricata, is a deeper water species. All three appear
to have been used as food sources; often encountered in discrete concentrations, and
all share a similar pattern through time: low relative frequencies early and late, with
their highest relative frequencies occurring during the middle of the site’s occupa-
tion in levels 8 and 9 (Fig. 3A–C). The later occupational phase between levels 5 and
3 (30–50 cm below surface) coincides with the appearance of distinct Bisquit Ware
ceramic sherds across the site that date to 1100–1400 AD and may represent over-
exploitation. This and alternative possibilities are discussed in greater detail below.
Sessile Clams.—The two most common sessile clams at SD are the turkey wing, A.
zebra, and the eared ark clam, Anadara notabilis. Relative frequencies of both spe-
cies remain high throughout the stratigraphic sequence at SD (Fig. 3D,E), but both
peak early and late precisely when oysters are less numerous suggesting that these
sessile clam species represent a lower-ranked resource than oysters.
Mobile Clams.—Mobile bivalves in the SD top 15 are Anomalocardia, Chione,
Codakia, and Donax. All are relatively small and labor-intensive to harvest. Donax is
common in the lower levels of Unit 14 and remains present in low numbers through-
out the sequence in the other two units (Fig. 3F). Anomalocardia is present in all
units examined with no clear trend through time (Fig. 3G). Chione increases dramat-
ically in the upper levels of the earlier occupation phase, only to drop during the later,
Bisquit Ware phase (AD 1200–1400, Fig. 3H). Codakia remains steadily represented
throughout the sequence (Fig. 3I).
Snails.—The two gastropod species in the SD top 15 that show patterns through
time are the hawk-wing conch, Lobatus raninus, and the fighting conch, Strombus
pugilis. Lobatus raninus is more common in the early part of the earlier occupational
phase and then steadily declines (Fig. 3J). In contrast, S. pugilis is present in low
numbers only increasing abruptly during the later, Bisquit Ware phase (Fig. 3K). The
shift from L. raninus to S. pugilis may have occurred because of over exploitation of
the local L. raninus population.
Fish
Material from Unit 1 was identified by Kay (2010) using the FLMNH Environmental
Archaeology comparative collection, supervised by KF Emery, in consultation with
Wake. Unit 22 was identified by Wake using the UCLA and LACM comparative col-
lections. Similar methods for determining number of identified specimens (NISP)
and minimum number of individuals (MNI) and assessing general morphology were
applied by both.
We identified 52 genera and 43 species representing 34 families of fish at SD (Table
2). This level of taxonomic richness is expected given SD’s proximity to a diverse
range of marine habitats. Present were taxa from reefs (e.g., certain Lutjanus spp.,
Epinephelus spp., Haemulon spp., Acanthurus spp., and Sparisoma spp.), open wa-
ters [e.g., Caranx spp., Euthynnus aletteratus (see Table 2 for fish species authori-
ties), Scomberomorus spp., and Seriola spp.], estuaries (Megalops atlanticus, Albula
vulpes, Centropomus spp., and some Lutjanus), and soft bottoms (e.g., Micropogonias
 wake et al.: baseline ecological data from sitio drago 1023

Table 2. Fishes from Sitio Drago, Units 1 and 22, and Cerro Brujo. NISP = number of identified specimens,
MNI = minimum number of individuals, empty cells = 0.

Sitio Drago Sitio Drago Cerro Brujo

Unit 1 Unit 22 (total)
Taxon NISP MNI NISP MNI NISP MNI
Carcharhinus cf. limbatus (Müller and Henle, 1839) 2 1
Carcharhinus Blainville, 1816 1 1 1 1
Sphyrnidae 7 2
Dasyatis Rafinesque, 1810 2 2 1 1 5 4
Dasyatidae 4
Aetobatus narinari (Euphrasen, 1790) 2 1 1 1
Rajiformes 2
Megalops atlanticus Valenciennes, 1847 23 3 7 2 9 5
Elops saurus Linnaeus, 1766 2 1
Albula vulpes (Linnaeus, 1758) 13 3 76 3 3 1
Clupeidae 251 2 271
Cathorops Jordan and Gilbert, 1883 106 25
Sciades Müller and Troschel, 1849 9 9 12 6
Ariidae 88
Gymnothorax Bloch, 1795 1 1
Gobiomorus dormitor Lacépède, 1800 2 2
Eleotridae 1 1
Sanopus barbatus (Meek & Hildebrand, 1928) 2 2
Opsanus Rafinesque, 1818 1 1
Batrachoididae 5 2 22 12
Strongylura Van Hasselt, 1824 1 1
Tylosurus crocodilus (Péron and Lesueur, 1821) 4 1 127 1
Belonidae 7 10 7
Scorpaena sp. Linnaeus, 1758 1 1
Epinephelus itajara (Lichtenstein, 1822) 1 1
Epinephelus Bloch, 1793 4 3 6 1 177 25
Mycteroperca bonaci (Poey, 1860) 1 1
Mycteroperca Gill, 1862 1 1 1 1
Serranidae 57 6 80 27
Centropomus Lacépède, 1802 2 1
Centropomidae 403 53
Mugil sp. Linnaeus, 1758 3 1 8 1
Sphyraena barracuda (Edwards, 1771) 6 1
Sphyraena guachancho (Cuvier, 1829) 28 17 3
Sphyraena Artedi, 1793 74 4 66 2 14 6
Rachycentron canadum (Linnaeus, 1766) 1 1
Caranx crysos (Mitchill, 1815) 97 12
Caranx hippos (Linnaeus, 1766) 4 1
Caranx latus Agassiz, 1831 4 1
Caranx Lacépède, 1801 340 161 6 152 19
Chloroscombrus chrysurus (Linnaeus, 1766) 157 2 947 55
Oligoplites Gill,1863 4 1
Selene Lacépède, 1802 7 3 6 2
Seriola Cuvier, 1816 2 1
Trachinotus Lacépède, 1801 2 2 5 3
1024 BULLETIN OF MARINE SCIENCE. VOL 89, NO 4. 2013

Table 2. Continued.

Sitio Drago Sitio Drago Cerro Brujo

Unit 1 Unit 22 (total)
Taxon NISP MNI NISP MNI NISP MNI
Carangidae 661 20 7 2
Euthynnus aleterattus (Rafinesque, 1810) 177 14 119 4
Scomberomorus maculatus (Mitchill, 1815) 2 1
Scomberomorus Lacépède, 1801 11 1 50 1
Scombridae 68 2
Lutjanus analis (Cuvier, 1828) 18 6
Lutjanus apodus (Walbaum, 1792) 7 5
Lutjanus jocu (Bloch and Schneider, 1801) 2 1
Lutjanus synagris (Linnaeus, 1758) 125 47
Lutjanus Bloch, 1790 366 21 112 16 79 38
Ocyurus chrysurus (Bloch, 1791) 6 4
Haemulon flavolineatum (Desmarest, 1823) 17 11
Haemulon parra (Desmarest, 1823) 10 8
Haemulon plumieri (Lacépède, 1801) 27 14
Haemulon sciurus (Shaw, 1803) 7 4
Haemulon Cuvier, 1829 22 9 13 3 17 10
Haemulidae 7 6
Archosargus Gill, 1865 1 1
Calamus Swainson, 1839 1 1 2 2
Sparidae 26 6
Ctenosciaena gracilicirrhus (Metzelaar, 1919) 5 3
Cynoscion jamaicensis (Vaillant and Bocourt, 1883) 1 1
Cynoscion Gill, 1861 1 1
Larimus breviceps Cuvier, 1830 29 20
Micropogonias furnieri (Desmarest, 1823) 5 3 3 2
Stellifer colonensis (Meek and Hildebrand, 1925) 13 6
Sciaenidae 4 1 1 1
Diapterus Ranzani, 1842 1 1
Gerres cinereus (Walbaum, 1792) 3 2
Gerreidae 4 2 2
Kyphosus sectatrix (Linnaeus, 1758) 2 1
Kyphosidae 1 1
Abudefduf Forsskål, 1775 1 1
Bodianus pulchellus (Poey, 1860) 1 1
Bodianus rufus (Linnaeus, 1758) 10 6
Halichoeres Rüppell, 1835 1 1
Lachnolaimus maximus (Walbaum, 1792) 1 1
Scarus Forsskål, 1775 1 8 2
Sparisoma radians (Valenciennes, 1840) 5 2
Sparisoma rubripinne (Valenciennes, 1840) 4 3
Sparisoma viride (Bonnaterre, 1788) 1 1
Sparisoma Swainson, 1839 74 5 7 3
Scaridae 15 4 4
cf. Istiophoridae 1
Acanthurus Forsskål, 1775 35 4 2
Acanthuridae 7 2
 wake et al.: baseline ecological data from sitio drago 1025

Table 2. Continued.

Sitio Drago Sitio Drago Cerro Brujo

Unit 1 Unit 22 (total)
Taxon NISP MNI NISP MNI NISP MNI
Balistes Linnaeus, 1758 20 3 2 1
Balistes vetula Linnaeus, 1758 2 1
Balistidae 1 1
Chilomycterus Brisout de Barneville, 1846 277 8
Diodon Linnaeus, 1758 3 3 1 57 8
Diodontidae 3 3
Teleostei 140
Total 2,588 140 2,727 328 1,334 242

furnieri, Ctenosciaena gracilicirrhus, Larimus breviceps, Stellifer colonensis) (Froese

and Pauly 2012).
While fish remains are ubiquitous at SD, fishing technology remains elusive. No
diagnostic fish hooks made of bone or shell have been recovered. Fishing technology
recovered from SD is limited to a suite of small, notched or grooved net weights.
These weights are relatively light, within the 28–56 g (1–2 oz) range, the kind used
on modern light weight seines or throw nets. However, remains of tuna, sierra,
and tarpon >50 cm in length strongly suggest the use of heavier nets. Exquemelin
(1969[1678]:211) describes a fishing net recovered from an abandoned Indian canoe
on Isla Colón as “four fathoms long and half a fathom broad” (7.3 × 0.9 m, possibly
a beach seine), but does not mention any weights. The use of traps or weirs is also a
possibility, but only inshore, anchored to a substrate on reef flats or in seagrass.
To understand potential human effects on local fisheries, we used fish bone collec-
tions recovered from Unit 1 (Kay 2010) and Unit 22 at SD to compile relative species
abundances and estimate mean trophic levels at earlier and later occupation phases.
Mean trophic level (MTL) was calculated thus:

TL Q MNI V
taxon - n
|
taxon - 1
MTL = TNI
where TL = estimated trophic level for each taxon (FishBase, Froese and Pauly 2012),
MNI = minimum number of individuals for each taxon, and TNI = total number of
individuals for all identified taxa.
Estimated MTL was 3.83 and 3.53 for the earlier occupation period in Units 1 and
22, respectively, and 3.74 and 3.48 for the later period in Units 1 and 22, respectively.
Imported pottery in the two mounds suggests equally high status of the residents
implying that the different MTLs between the mounds were probably caused by dif-
fering harvesting practices rather than differences in diet. MTL in both mounds de-
clines highly significantly over time (Wilcoxon signed-rank test: Unit 1, z = 3.23, P >
| z | = 0.0009, Unit 22, z = −6.189, P > | z | = 0.0000).
Declining MTL across the site suggests a decline in the quality of fishing, cor-
roborating the loss of preferred high trophic level predatory fish (Jackson et al. 2001,
Lotze et al. 2006, Pauly et al. 2009). Similar shifts towards lower-ranking fishes have
been documented elsewhere in the Caribbean (Wing 2001, Wing and Wing 2001,
1026 BULLETIN OF MARINE SCIENCE. VOL 89, NO 4. 2013

Blick 2007, Carder et al. 2007, DeFrance 2010, McChlenachan et al. 2010) and in
Polynesia (Butler 2001, Morrison and Hunt 2007, Ono and Intoh 2011). An alterna-
tive explanation could be a shift toward more communal fishing practices that tend
to extract lower trophic level schooling fish species, but this seems unlikely given the
universal preference for large high trophic level fish if they are available (Lotze et al.
2006).

Marine Reptiles
Crocodiles.—We consider the American crocodile, Crocodylus acutus Cuvier, 1807,
a marine resource because it is commonly found in estuaries and mangroves. Six
American crocodile bone specimens (MNI = 1) were recovered from Unit 14 and
identified by one of us (TAW). Based on the size of a conjoined left maxilla and pre-
maxilla, the crocodile was approximately 5 m long (e.g., Platt et al. 2009). Such an
individual would provide a great deal of meat, teeth for display, and possibly much
prestige to its killer. While the presence of Crocodylus at SD is interesting, it is rare
and does not represent a staple resource.
Sea Turtles.—Sea turtle remains, on the other hand, are ubiquitous at SD and
must have represented an important, predictable source of meat and eggs. One of
us (TAW) identified two species of sea turtle at SD; the hawksbill, Eretmochelys im-
bricata (see Table 3 for higher marine vertebrates species authorities), and the more
common green sea turtle, Chelonia mydas. Loggerhead sea turtle, Caretta caretta,
was found at CB (Wing 1980; Table 3). All of these species, as well as the giant leath-
erback sea turtle, Dermochelys coriacea Vandelli, 1761, nest in the local area today
and almost certainly did so in the past.
The majority of sea turtle specimens are carapace and plastron fragments and pha-
langes; the most common bones in the cheloniid skeleton. The primary limb bones
including shoulder and pelvic girdles and the upper and lower fore- and hind-limb
elements are commonly encountered. Sea turtle represents the most common tetra-
pod in almost all units analyzed so far at SD (Fig. 4), suggesting that the inhabitants
used sea turtles as a primary source of meat and probably other by-products such
as fat, eggs, and shells. The seasonal predictability and large size of mature nesting
individuals represent a pivotal highly ranked resource. It is particularly interesting
that we find no relative decrease in sea turtle representation between the two occu-
pational periods at SD. In fact, Table 4 illustrates, in terms of bone density m−3 (NISP
m−3), that sea turtle bones become much more common, yet no more fragmented,
during the later phase of occupation at SD. This pattern suggests intensification of

Table 3. Number of higher marine vertebrate specimens identified from Sitio Drago and Cerro
Brujo.

Taxon Sitio Drago Cerro Brujo

Crocodylus acutus Cuvier, 1807 6
Crocodilidae 1 5
Chelonia mydas (Linnaeus, 1758) 144 6
Caretta caretta (Linnaeus, 1758) 1
Eretmochelys imbricata (Linnaeus, 1766) 3 5
Cheloniidae 2,300 286
Trichechus manatus Linnaeus, 1758 37 13
 wake et al.: baseline ecological data from sitio drago 1027

Figure 4. Sea turtle distribution at Sitio Drago by excavation unit.

the sea turtle resource between AD 1100 and 1400. Harvesting of sea turtles in Bocas
del Toro continued into and intensified during the early Historic Period in Bocas del
Toro (Exquemelin 1969[1678], Jackson et al 2001, Bjorndal and Jackson 2002).
Marine Mammals
Manatee.—Manatee represented an important source of meat for both the prehis-
toric and early historic occupants of SD and visitors to Bocas del Toro (Exquemelin
1969[1678], Wing 1980). The West-Indian Manatee, Trichecus manatus, is the larg-
est Central American mammal (Reid 1997) and ranges (seasonally) from Virginia,
along the North American eastern seaboard to the Caribbean coast and associated
waterways, the Greater and Lesser Antilles, as far south as central Brazil. Manatees
are still present in the Bocas del Toro area in the San-San Pond Sak Wetland Reserve
(Mou-Sue et al. 1990).
In Central America, manatee was hunted by the Maya (McKillop 1985), the
Miskito (Nietschmann 1973), the Rama (Loveland 1976), at SD and CB (Wing 1980),
and probably anywhere they were available (Bradley 1983). Their meat, fat, hides, and
bones were all important resources for pre-Columbian and Historic Period inhabit-
ants of the Caribbean coast. Manatee bone was an important raw material for pre-
Columbian Panamanian carvers, as testified by the elaborate carved “batons” and
other carved objects recovered from Sitio Conte (Lothrop 1937) and elsewhere in
central Pacific Panama (Ladd 1964, Cooke and Jimenez 2010). These carvings, or at
least the bone raw material, must have traveled from the Caribbean coast.
One of us (TAW) identified 37 manatee bones recovered at SD, representing at
least 9 individuals. Manatee remains are far less common in the upper 30 cm of
the SD archaeological deposits (Table 4). While the overall sample of manatee is

Table 4. Sitio Drago sea turtle bone density m−3 (number of identified specimens m−3).

Unit
Period 1 18 19 22 25
Early 101 55 33 559 120
Late 253 180 117 870 203
1028 BULLETIN OF MARINE SCIENCE. VOL 89, NO 4. 2013

small, this pattern may represent overexploitation of the local manatee population.
Nonetheless, exploitation of manatee at Boca del Drago continued into the Historic
Period. For example, in March 1671 three ships/crews of Henry Morgan’s pirate fleet
that had just sacked and burned Cartagena, Portobelo, and Panama entered Boca del
Drago hungry, with the explicit intent of hunting manatee (Exquemelin 1969[1678]).
The description of manatee by Exquemelin is fairly accurate and illustrates the high
esteem manatee were held in as a food and raw material source.

Discussion and Comparison with Cerro Brujo

The CB site represents the only other professionally excavated archaeological site
in Bocas del Toro. CB was contemporaneous with SD (Linares 1977, Wake et al. 2012)
and produced a substantial amount of information from both shell and bone mate-
rial. While SD and CB share much of their archaeofauna taxa, both the molluscan
and vertebrate assemblages are more numerous and diverse at SD. Table 5 compares
the most common molluscs at CB and SD. Both sites include sessile clams associ-
ated with mangroves (oysters) and reef flats (turkey wings), but smaller, more mobile,
sandy beach and shallow soft bottom taxa are present at SD, but generally absent
from CB, as would be expected by the location of the two sites. CB is surrounded by
mangroves, fringing reefs, and seagrass beds, while the sandy beaches close to SD are
absent on the Aguacate Peninsula where CB is located.
Although Borgogno and Linares (1980) observed no clear trends in the relative fre-
quencies of the most common molluscs in the three shell-bearing strata at CB, con-
cluding that there was little effect of human harvesting on molluscs, our data suggest
substantial effects of human harvesting at SD. The three oyster species at SD reach
their greatest relative frequency during the middle of the occupational sequence fol-
lowed by a dramatic decrease in the later, Bisquit Ware, occupational phase. Dietary
preferences could have changed, but a similar curve in three different species that
inhabit two distinct habitats suggests that intense harvesting pressure reduced their
abundances. Likewise, relative frequency of the conch L. raninus declines in the later
occupational sequence.
In contrast to these declines the bivalve C. cancellata and the conch S. pugilis in-
crease noticeably in the upper 40 cm of Units 3, 14, and 22 at SD, possibly represent-
ing a shift of resources as the three oyster species and the conch L. raninus became
less available.
Sessile (A. zebra and A. notabilis) and mobile (e.g., species of Donax, Anomalocardia,
and Codakia) clams occur consistently abundant through the occupation sequence,
demonstrating their significant contribution to the diet. None of these important
taxa show clear changes in their relative frequency through the occupation sequence
implying no discernible effects of 700 yrs of harvesting at SD.
Fishes
As with the molluscs, fish assemblages from CB and SD reflect each site’s suite of
local habitats. The fish assemblage from CB is dominated by taxa (up to 22 genera)
that prefer low energy reefs, seagrass beds, and mangroves while the fish assemblage
from SD has a greater diversity (53 genera) and representatives from a wider array of
habitat types.
 wake et al.: baseline ecological data from sitio drago 1029

Table 5. Number of individual manatee, Trichecus manatus, remains from Sitio Drago, by
occupation phase.

Unit Early Late

1 6 3
2 1 0
3 1 0
14 16 1
18 1 1
19 1 0
22 4 2
25 1 1
39 1 0
41 2 0
42 4 0
Total 38 8

Wing (1980) noted a slight increase in the amount of serranid fishes (which include
grouper and sea basses) in the more recent occupation phase at CB, but observed no
proportional change in any of the other five important fish families (Centropomidae,
Lutjanidae, Carangidae, Megalopidae, and Haemulidae). Identifications in the pres-
ent study unfortunately are not precise enough to permit trophic level estimates at
CB, but in general no obvious shifts in trophic levels took place.
Linares (1980b) estimated the population of the entire Aguacate peninsula, where
CB is located, at approximately 120 individuals, suggesting a density of only 3–4 per-
sons km−2. In contrast the population of SD was approximately 100 individuals (as-
suming 20 households of five persons) estimating a density of well over 500 persons
km−2 in the SD area. As such, harvesting intensity would have been much greater
than on the Aguacate Peninsula, explaining the significant decline in fish MTL at SD
that was not apparently mirrored at CB.
No indication of any change in fishing methods is seen through time at SD. The
only evidence of fishing technology recovered from the site is a considerable number
of small, notched-pebble net weights found throughout the sequence at SD. The im-
plication is that less desirable generalist and herbivorous fish species were included
in the diet during the later occupational period at SD due to increasing harvest-
ing pressure that reduced the abundance of more highly ranked carnivorous fish
species. As the local population grew, fishing practices may have shifted to a more
communal-based fishing effort that cast a broader net as opposed to hunting of pre-
ferred individuals, resulting in the inclusion of more lower–trophic level species in
the catch. At any rate, we believe that continuous net fishing at SD over time had an
effect of reducing the availability of more highly ranked species in local fish popula-
tions, resulting in the inclusion of greater numbers of lower trophic level species in
the overall harvest.
Sea Turtles
The exploitation of sea turtles by the pre-Columbian inhabitants of Bocas del Toro
represents an important subsistence activity at both CB and SD and sea turtle re-
mains are common at both sites, but particularly at SD. Sea turtle remains become
more common at SD in the most recent occupational period (Table 4) suggesting
1030 BULLETIN OF MARINE SCIENCE. VOL 89, NO 4. 2013

Table 6. Most common mollusc species (as a percentage of the total) at Cerro Brujo (Borgogno and
Linares 1980) and Sitio Drago (Doughty 2011).

Cerro Brujo Sitio Drago

Taxon Percent Taxon Percent
Chama macerophylla 34.5 Arca zebra 17.1
Arca imbricata 24.5 Crassostrea rhizophorae 13.7
Arca zebra 17.5 Anomalocardia cuneimeris 12.5
Crassostrea rhizophorae 11.5 Chione cancellata 11.5
Isognomon alatus 7.0 Donax variabilis 9.9
Other species 5.0 Anadara notabilis 9.7
Other species 11.9

intensification of sea turtle harvesting, a pattern that continued into the Historic
Period as European sailors collected sea turtles for food (Jackson et al. 2001, Bjorndal
and Jackson 2002). Today, sea turtles in Bocas del Toro are relatively rare and appar-
ently declining in abundance (Meylan 1999, Troëng et al. 2004) and the role of inten-
sifying harvesting over >700 yrs must be considered when attempting to explain why.
Manatees
As European sailors learned that manatee were readily available in Bocas del Toro
they came to the region to hunt them, steadily increasing pressure on the existing
population. As with the turtles, however, remains from both SD and CB reveal that
the West-Indian manatee was an important source of food and raw material for the
region’s pre-Columbian inhabitants and the increasing number of manatee remains
in the most recent occupational phase suggest increasing hunting pressure.

Conclusions

Throughout the Caribbean (Wing 2001, Wing and Wing 2001, Blick 2007, Carder
et al. 2007), along the California coast (Braje et al. 2007, Erlandson et al. 2008, 2011,
Erlandson and Rick 2010), and in Polynesia (Butler 2001, Morrison and Hunt 2007,
Ono and Intoh 2011), archaeologists have documented the effects of human hunting
(vertebrates) and collecting (molluscs) on local resources. This is done by using data
from initial occupation periods or lower levels of archaeological sites as “baseline”
data, because there is often no other source of earlier data from a given area.
In the case of SD, human harvesting appears to have driven changes in the avail-
ability of resources amongst the mollusk, fish, and tetropod communities in Bocas.
Desirable conch and clams were substituted by less desirable taxa over time suggest-
ing resource exploitation. The mean trophic level of fish captured over time declined,
which we conclude to represent trophic changes in the local fishery because the net-
based fishing technology used at SD is not very selective and did not change through
time. The hunting of sea turtles and manatees intensified over time. Once Europeans
arrive in the region data concerning molluscs and fish become vague or unavailable,
but historical accounts confirm that the hunting of sea turtles and manatee contin-
ued and intensified.
Fisheries in Bocas are today in a dire state. Conversations with local fishermen
make it apparent that large queen conchs are rare, oyster beds have virtually disap-
peared, and oysters on mangrove roots appear to be smaller than those found in ar-
chaeological shell middens. Similar conversations suggest that lobsters harvested in
 wake et al.: baseline ecological data from sitio drago 1031

Bocas are getting smaller and that red snapper are decreasing in size and are harder
to fish. Sea turtles in the region are rare and imperiled by continued hunting—as
evidenced by the steel sea turtle harpoons carried by many artisanal fishermen in
Bocas del Toro. The few manatee still present in the San San–Pond Sak reserve are
mute testimony to the once denser population documented by historical sources and
archaeological data (Exquemelin 1969[1678], Wing 1980, Wake and Mendizabal in
press).

Acknowledgments

We thank the following persons for their continued support of our archaeological field-
work at Boca del Drago: DA “Bolo” Serracín and the Serracín family, especially A Serracín
de Shaffer, W and J Serracín, B Serracín de Alvarez, and T Serracín de Duke; M and D Wake;
C Campbell and S Wake; M Holmes, H Campbell, J Snowden; associated researchers and
graduate students including PJ Lahanas, C Fitzgerald, T Mendizabal, A Mojica, M Davis, J de
Leon, J Howard, L Martin, D Smith, and all the participants in the 2003–2010 field seasons.
D Doughty identified the majority of the shell specimens reported here (Doughty 2011) in
consultation with T Wake and F Rodriguez. The Franklin Fund of the American Philosophical
Society, The Cotsen Institute of Archaeology at UCLA, The Institute for Tropical Ecology
and Conservation (ITEC), and individual donors provided monetary and material support
for the PASD. We thank A O’Dea and F Rodriguez for their assistance in determining nam-
ing authorities for the molluscan species mentioned here. We referred primarily to WoRMS
(World Register of Marine Species, http://www.marinespecies.org) and ITIS (the Integrated
Taxonomic Information System, http://www.itis.gov) for invertebrate and vertebrate naming
authorities. We thank the UCLA Math Sciences Statistical Consulting Service for their as-
sistance. We thank three anonymous reviewers and the guest editor, A O’Dea, for their thor-
ough, thoughtful, and helpful comments. We thank A O’Dea for his help in drafting Figure
3. We thank STRI and A O’Dea for the invitation to participate in the symposium and our
inclusion in this volume. T Wake is a STRI Research Associate. We thank STRI for its gener-
ous institutional support. Any errors or omissions are strictly our own.

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Date Submitted: 23 August, 2012.

Date Accepted: 14 May, 2013.
Available Online: 10 October, 2013.

Addresses: (TAW) Zooarchaeology Laboratory, The Cotsen Institute of Archaeology at

UCLA, Los Angeles, California 90095-1510. Email: <twake@ucla.edu>. (DRD) 30 Greenfield
Ave., Unit 615, Toronto, Ontario, Canada M2N 6N3. Email: <dougdoughty@sympatico.ca>.
(MK) Greenwood & Associates, Inc., 725 Jacon Way, Pacific Palisades, California 90272.
Email: <michaelmozartkay@gmail.com>.