Received: 15 August 2018 Revised: 3 February 2019 Accepted: 24 May 2019

DOI: 10.1002/oa.2790

SPECIAL ISSUE PAPER

Pre‐Hispanic Maya fisheries and coastal adaptations in the

northern lowlands from the Classic (500–900 AD) to
Postclassic (900–1400 AD) periods

Nayeli G. Jiménez Cano

Facultad de Ciencias Antropológicas,

Universidad Autónoma de Yucatán, Mérida, Abstract
Mexico Recent studies of fish remains at Mayan settlements from the Classic (500–900 AD)

Correspondence
Nayeli G. Jiménez Cano, Facultad de Ciencias
Antropológicas, Universidad Autónoma de
Yucatán, Km. 1 Carretera Mérida‐Tizimín,
Cholul, Mérida 97305, Mexico.
Email: nayeli.jimenez@correo.uady.mx
Funding information
Mexican Consejo Nacional de Ciencia y
Tecnología, Grant/Award Number: 215047
and Postclassic (900–1400 AD) periods are examined. These analyses deepen our
understanding of ancient Maya fishing practices and coastal ecosystems and call into
question traditional paradigms of environmental stability in the northern Maya low-
lands. The effects of droughts recorded during the Maya collapse (800–1000 AD)
led to an overall reduction in freshwater inflow into the estuaries combined with epi-
sodic increases of sedimentary output. The latter fostered the development of muddy
bottoms along the coast, in particular around estuaries formed by underwater river
discharges. Ecological characterization and trophic level shifts of fish assemblages
from the Gulf of Mexico and Caribbean shores help elucidate environmental stress
episodes. These findings set a preliminary baseline for pre‐Hispanic Maya fisheries,
which can further our understanding of present day fisheries along the Maya coast.

K E Y W OR D S

coastal adaptations, Maya, Maya collapse, paleoecology, pre‐Hispanic fisheries

1 | I N T RO D U CT I O N Previously, it was postulated that environmental pressures at the

The Maya region is exceptionally well positioned for procuring fish
because its coasts serve as home and nursery grounds for a high diver-
sity of fish species. Pre‐Hispanic Maya fisheries were characterized by
the preferential exploitation of estuarine environments and reached
their maximum productivity during the Classic (250–1000 AD) and
Postclassic (1000–1500 AD) periods (Jiménez Cano, 2017).
The transition between these periods, also known as the Terminal
Classic (800–1000 AD), represents a crucial episode in Maya history.
This period corresponds to the so‐called Maya “collapse” in the
southern Maya lowlands coincident with a moment of social unrest
and environmental crisis. This episode was presumably fostered by
demographic pressure and recurrent droughts that led to an unsus-
tainable use of natural resources causing deforestation and loss of
soil fertility (Abrams & Rue, 1988; Culbert, 1988; Curtis et al.,
1998; Hodell, Brenner, Curtis, & Guilderson, 2001; Hodell, Curtis, &
Brenner, 1995).
end of the Classic period might have led to intensive hunting of
already stressed animal populations (Pohl, 1990). However,
zooarchaeological studies on the isotopic composition of mammal
remains from the Guatemalan Petexbatún region failed to confirm
the ecological collapse model (Emery, 2004, 2010).
In the northern Maya lowlands (NML), the collapse apparently did
not have such environmental and social devastating effects. Instead,
settlements continued to be inhabited, and the Puuc region even
flourished at the end of the Classic period. In addition, large cities such
as Chichén Itzá and Mayapán managed to extend their commercial
networks after the Terminal Classic (Chase & Chase, 1992; Freidel,
1986; McKillop, 2004).
Notwithstanding the idea of an apparent social stability, this situa-
tion has been recently questioned by paleoenvironmental records of
stable δ18O isotopes from several localities in northern Yucatan.
Between 800 and 1000 AD, a series of droughts have been recorded
through isotopic values of ostracods and stalagmites from Yucatan

Int J Osteoarchaeol. 2019;29:469–476. wileyonlinelibrary.com/journal/oa © 2019 John Wiley & Sons, Ltd. 469
470
(Brenner et al., 2001; Curtis, Hodell, & Brenner, 1996; Hodell et al.,
1995; Medina Elizalde et al., 2010). The data suggest that this period
was the driest over the past 35,000 years (Medina Elizalde et al.,
2010; Medina Elizalde & Rohling, 2012), which affected inland ecosys-
tems (Dunning, Beach, & Luzzadder‐Beach, 2012). As of this writing,
this period of presumed environmental stress remains scarcely
explored in coastal scenarios, and additionally, it has been postulated
that coasts remained ecologically stable over the past 3,000 years
(Eaton, 1978, p. 9). In fact, such ecological stability might not be prob-
able because droughts would have affected NML coasts because estu-
aries are conformed mainly through freshwater input from underwater
rivers, themselves nourished by pluvial activity. One should thus
expect changes of the seascape involving at least sedimentation and
changes in salinity. Archaeological fish remains can serve as proxies
to track changes in ancient coastal environments because fish physiol-
ogy responds to changes in the temperature and salinity of the water
affected by pluviometry. From such a perspective, it seems inescap-
able that droughts fostering seascape changes must have affected fish
and fishing practices.
This paper aims at assessing putative changes taking place along
the coasts of the NML through shifts in fish procurement strategies
resulting from environmental stress episodes during the transition
from the Classic to the Postclassic periods. This paper aims to infer
ecological changes during the droughts reported at the Terminal Clas-
sic along the NML coasts in light of robust ichthyoarchaeological
JIMÉNEZ CANO
evidence in combination with ecological measurements to provide a
baseline for future studies of pre‐Hispanic Maya fishery ecodynamics.
2 | MATERIALS AND METHODS
To assess whether and how fish procurement strategies were affected
due to environmental changes on the NML, the ichthyoarchaeological
data of nine settlements are compared. These ichthyoarchaeological
assemblages date to the Classic and Postclassic periods and come
from two coastal zones: the Gulf of Mexico and the Caribbean
(Figure 1).
Of these settlements, Mayapán and Xcambó were analysed by the
author using the zooarchaeological reference collections housed at
the Laboratorio de Arqueozoología, Universidad Autónoma de Madrid,
the Laboratorio de Zooarqueología, Universidad Autónoma de
Yucatán, and the Zooarchaeology Collection of the Florida Museum
of Natural History. Fish remains from Caracol, Champotón, Cozumel,
Marco González, Northern River Lagoon, Isla Cerritos, and Vista
Alegre were studied by other authors (García Paz, 2016; Götz,
2008; Hamblin, 1984; Masson, 2004; Seymur, 1991). All selected
ichthyoarchaeological assemblages share similar cultural characteris-
tics and recovery methods (see Table 1).
Ichthyoarchaeological data are presented as number of identified
specimens (NISP) for both bony (Teleost) and cartilaginous
FIGURE 1 Map of the Maya area showing
the location of the archaeological settlements
referred to in this paper
JIMÉNEZ CANO 471

TABLE 1 Information on selected archaeological sites and materials 2.1.2 | TL analyses

mentioned in this work

Site Area Period NISP Recovery (mm) Recent fisheries studies often incorporate TL analyses as tools to cali-
brate environmental and human pressure on fish populations (Pauly,
Vista Alegre GM Classic 2,203 0.36
Christensen, Dalsgaard, Froese, & Torrres, 1998). The TL values
Xcambó GM Classic 2,025 0.36
inferred according to the database compiled by Froese and Pauly
Isla Cerritos GM Classic 459 0.5 (2017) refer to the position occupied by an organism along the trophic
Caracol C Classic 29 0.36 chain, from producers (TL = 1) to highest‐level predators (TL = 5).
Northern River Lagoon C Classic 1,497 0.36 Shifts in TL values have proved crucial to explore fishing patterns
Mayapán GM Postclassic 642 0.36 over recent decades and have occasionally been applied to
Champotón GM Postclassic 197 0.5 ichthyoarchaeological assemblages (Morales & Roselló, 2004; Reitz,
2004). In this paper, the equation of Morales and Roselló (2004),
Cozumel C Postclassic 80 0.36
adapted from Reitz (2004), was used.
Marco González C Postclassic 526 0.36

Abbreviations: GM, Gulf of Mexico; C, Caribbean.

TLi ¼ Σ ðTLijÞ ðNISPijÞ= ΣNISPi;

(Chondricthian) fishes. The NISP values were used as the basis for
calculating ecological indexes and trophic level (TL) values because
alternative abundance estimators, including minimum number of indi-
viduals, systematically underestimate the frequencies of fish taxa, car-
tilaginous fishes in particular, deflating already small assemblages.
where TLi (means TL for a determined period) is the sum of the TL for
each taxon (j) for a given period of time (i) multiplied by the NISP of
that taxon (j) for that period (j). Then, it is divided by the sum of the
NISP of that period. This will represent the mean TL value of each site.

2.1 | Ecological measurements

3 | RESULTS AND DISCUSSION
Ecological indexes and mean TLs were estimated in all nine
The taxonomic composition of each site is provided in Supporting
assemblages. These were obtained by using low taxonomic levels such
Information. Values of richness, diversity, and equitability, as well as
as genus and species and using the statistical package PAST 3.x
the mean TL for each assemblage, are presented in Table 2. In
(Hammer, Harper, & Ryan, 2001).
Table 3, these values are grouped according to their regional
(Caribbean or Gulf of Mexico) and chronological (Classic or Postclassic)
2.1.1 | Ecological indexes
settings.

The estimated ecological indexes were species richness, equitability,

and diversity. Species richness (S') refers to the number of taxa in a
region (Reitz & Wing, 2008), and in this paper, it refers to the number
of taxonomical categories, genus and species, presented in each
collection.
To quantify species diversity, the Shannon–Weiner index (H') was
calculated. This index combines taxa and abundance within each taxo-
nomical category in order to express heterogeneity of the sample
(Shannon & Weaver, 1949), as expressed by the formula

parameters were higher during the Classic period and slightly lower
H0 ¼ Σpi Log10pi ;
where pi is the relative abundance of the taxon (NISP) and Log10pi is
the logarithm of pi (Lloyd & Ghelardi, 1964). In this paper, the 10 base
of the logarithm for this index was applied.
Equitability (V') index refers to the relative evenness of the numer-
ical importance of a species in a sample (Colinvaux, 1986, p. 650), and
it is expressed by the formula
0 0
V ¼ H = loge S;
where H' is the diversity index and logeS is the natural logarithm of the
observed species.
3.1 | Ecological indexes
In terms of sample size, Classic ichthyoarchaeological assemblages of
the Gulf of Mexico featured the greatest number of remains, whereas
Caribbean Postclassic sites were the smallest. In accordance with this,
species richness (S') was also higher in the Gulf of Mexico than in the
Caribbean.
Diversity (H') and equitability (V') indexes indicated that both
during the Postclassic in the Gulf of Mexico, similar to Postclassic
Caribbean fish composition (Figure 2).
This aggregation reflects a more diverse ichthyofauna, with no par-
ticular species dominating the samples. This is also in concordance
with the high species richness recorded in the region. On the other
hand, the Classic Caribbean assemblages reported lower diversity
and equitability indexes. This may reflect the dominance of catfishes
(Ariidae) in the samples, especially at Northern River Lagoon. At first
impression, this suggests a selective exploitation of catfishes in the
Classic Caribbean. In the following section, I will explore whether this
is a response to cultural selection or if this pattern possibly responds
to environmental circumstances.
472 JIMÉNEZ CANO

TABLE 2 Ecological indexes and trophic level values of selected sites from the Maya lowlands

Site VAL XCM ICE CAR NRL MAY CHA COZ MGO

NISP 1,203 2,025 439 29 1,497 642 197 80 526

S' 40 50 36 6 12 36 7 9 17
H' 2.81 2.34 2.70 1.11 0.73 2.54 1.37 1.71 2.10
V' 0.76 0.57 0.75 0.62 0,29 0.71 0.71 0.78 0.74
TL 3.92 4.12 3.91 3.75 3.41 3.67 3.79 4.11 3.73

Abbreviations: S', species richness; H', diversity; V', equitability; NISP, number of identified specimens; TL, trophic level; VAL, Vista Alegre; XCM, Xcambó;
ICE, Isla Cerritos; CAR, Caracol; NRL, Northern River Lagoon; MAY, Mayapán; CHA, Champotón; COZ, Cozumel; MGO, Marco González.

TABLE 3 Pooled ecological and TL global values by cultural periods sites of the Gulf of Mexico and are similar to the global TL values of
and zones the Postclassic period of the Caribbean.
This pattern is best understood when TL values are evaluated in
Classic Postclassic
Period/ terms of taxa relative frequency. From this perspective, all assem-
zone GM C GM C
blages incorporate four TL categories: (a) species with low TL values
S' 42 9 22 13 (TL = 2–3.2): these include detritivores and herbivores fishes such as
H' 2.54 0.92 2.02 1.90 catfishes (Ariidae) and mullets (Mugilidae); (b) species with medium
V' 0.70 0.46 0.71 0.76 TL values (TL = 3.3–3.5): included here are omnivorous fishes such
TL 3.99 3.58 3.73 3.92 asporgies, seabreams (Sparidae), and croakers (Sciaenidae) that feed
on invertebrates and small fishes; (c) species with medium‐high TL
Abbreviations: S', species richness; H', diversity; V', equitability; GM, Gulf of
Mexico; C, Caribbean; TL, trophic level.
values (TL = 3.6–4): secondary carnivores, piscivorous fishes such as
small and medium groupers (Serranidae), snappers (Lutjnidae), and
jacks (Carangidae); and (d) species with high TL values (TL = 4–4.5) that
include top predators such as requiem sharks (Carcharhinus sp.),
hammerhead sharks (Sphyrna sp.), large serranids (Epinephelus itajara),
and barracudas (Sphyraena barracuda).
The regional analyses of these frequencies will aid in understand-
ing fish composition due to environmental stress episodes or cultural
exploitation.

3.2.1 | Gulf of Mexico

Fish acquisition during the Classic period is characterized by the dom-

FIGURE 2 Distribution of diversity and equitability global values
from Classic and Postclassic. GM, Gulf of Mexico; C, Caribbean
3.2 | Trophic levels
Fish assemblages produced interesting TL global values, which corre-
spond to an “inverted” model trend between the Classic and Postclas-
sic periods when the Gulf of Mexico and Caribbean are compared
(Figure 3). There is a temporal trend with higher TL values of the
Caribbean during the Postclassic period and lower TL values of the
Gulf of Mexico at this time. The highest values are recorded in Classic
inance of top predators followed by secondary predators. Detritivores,
composed mainly of catfish, followed in frequency and omnivorous
fishes were less frequent. Interestingly, during the Postclassic, global
TL values inverted the pattern of the Classic (Figure 4). The most sig-
nificant difference is that the frequency of detritivores increased to
~40% of the NISP, equalling that of the top predators that slightly
decreased during this period. The fishes with intermediate TLs main-
tained similar frequencies to those of the Classic period.
These values reflect a healthy marine ecosystem along the Gulf of
Mexico coast during the Classic period given the abundance of sharks,
traditionally considered bioindicators of nondisturbed or slightly
disturbed trophic chains (Bennet, 2005; Ferretti, Worm, Britten,
Heithaus, & Lotze, 2010). Fishing practices took advantage of
favourable marine conditions that allowed an intense exploitation of
sharks during the Classic period.
The shift to detritivores fishes during the Postclassic is likely in line
with environmental changes in estuaries due to rainfall reduction. The
abundance of catfishes suggests a coastal scenario with muddy
JIMÉNEZ CANO
FIGURE 3 Trophic level fluctuations at the Gulf of Mexico and the
Caribbean. TL, trophic level
bottoms probably because of limited freshwater input to the estuaries.
It is likely that these changes were caused by a combination of envi-
ronmental stress episodes rather than purely cultural selection. In this
sense, frequency of fishes with medium and high‐medium TL values
remained stable through time, whereas extremes of the trophic
chained shift. In this case, we are not observing a cascade effect
caused by fishing down the food web trough anthropogenic pressure
(Ferretti et al., 2010; Pauly, 1995; Pauly et al., 1998).
In this sense, differential fish acquisition during these two episodes
of Maya history appears to constitute a signal of coastal geomorpho-
logical shifts conforming with the droughts reported from 800–1000
AD that played an important role as modulators of the shore ecosys-
tems. Specifically, restricted freshwater availability coupled with a
more torrential marine water regime must have raised sedimentation
rates and the turbidity of the waters on the estuaries.
Marine regressions are a phenomenon associated with this climatic
shift. Such changes have been reported along the NML coast in Vista
Alegre where sedimentological studies have revealed superimposed
sequences and transgression episodes at the end of the Classic period
(Beddows & Carter, 2013) as well as changes in sea levels during the
last 3,000 years (Jaijel et al., 2018). The discharge reduction of fresh-
water as the result of diminishing rains resulted in a gradual elevation
FIGURE 4 Frequency of TL categories from the Gulf of Mexico during the Classic and Postclassic. NISP, number of identified specimens; TL,
trophic level
473
of the sea level that flooded the Yucatán coastline (Beddows & Carter,
2013; Goodman & Rohi, 2013). The sequence reported at Vista Alegre
correlates with phases of occupation and abandonment of the site. At
the end of the Classic period, these episodes probably forced the
inhabitants of the site to build elevated platforms to protect them-
selves against sea level rise (Beddows & Carter, 2013; Goodman &
Rohi, 2013). The raised platforms at Vista Alegre resemble those
found at Xcambó, which served to protect the population against
floods (Sierra Sosa, 1996, 2004a, 2004b). Although sedimentological
investigations are still pending at Xcambó, it would be interesting to
question if the abandonment of Xcambó was induced by a restriction
to freshwater access during the end of the Classic period.
Furthermore, our ichthyoarchaeological evidences coincide with an
overall pattern of global climate anomalies called the Medieval Warm
Period that took place between the 10th and 15th century AD, affect-
ing the North Atlantic and defining a period of increased temperatures
and droughts (Bradley, Hughes, & Díaz, 2003; Crowley & Lowery,
2000; Haug et al., 2003). Far away from these northern regions, it is
evident that those climatic alterations influenced fishing patterns of
the coastal Maya population during Classic and Postclassic times.
However, more sedimentological and geomorphological investigations
are needed in order to contrast the recent ichthyoarchaeological
evidence.
Thus, the gathered data confirm an incipient model of Maya sea-
scapes of the Gulf of Mexico coasts where estuarine fishing grounds
did not remain stable during the Classic–Postclassic transition. As a
consequence, coastal transformations modelled fishing practices in
the northern coast of the Yucatan peninsula.
3.2.2 | Caribbean
The Caribbean revealed an inverted pattern in the evolution of the
global TL values to those recorded for the Gulf of Mexico. Classic
period samples were dominated by detritivores, mainly catfish,
followed by large predators, mostly barracudas, whereas species with
medium TL values were poorly represented. During the Postclassic,
there was a sharp increase of top predators, including both sharks
and barracudas, and a decrease of detritivores fishes. Interestingly,
omnivorous fishes increased in frequency surpassing detritivores
474
FIGURE 5 Frequency of TL categories from the Caribbean during the Classic and Postclassic. NISP, number of identified specimens; TL, trophic
level
fishes, wherease secondary carnivores equalled lower TL values for
fishes (Figure 5).
The TL values of the Caribbean ichthyoarchaeological assemblages
reflect an interesting fish acquisition pattern. The dominance of
catfishes in the Classic likely indicates signs of stress due to a virtually
null presence of large predators, such as sharks and groupers who tra-
ditionally thrive in Caribbean waters (Nelson, 2008). However, it is
important to understand the context of provenance because most of
the samples came from Northern River Lagoon, a settlement located
by mangroves where catfishes are common inhabitants. Notwith-
standing this apparent cultural selection driven by the exploitation of
a specific environment, paleoclimatological records on the southern
lowlands have revealed intense soil erosion caused by droughts that
led to a substantial sedimentation of the floodplains in the Caribbean
coast since the Preclassic (Dunning et al., 2002; Dunning & Beach,
2000; Dunning, Scarborough, Valdez, Luzzadder‐ Beach, & Jones,
1999; Jacob, 1995). In this sense, it may be possible that the intense
exploitation of catfishes in the Classic Caribbean was also a conse-
quence of geomorphological changes on the coast.
On the other hand, TL values increased in the Postclassic due to
an increment of large predators, especially sharks. One could argue
that a recovery of the Caribbean marine ecosystem had taken place
from the former period. However, this pattern must be viewed with
caution because the ichthyoarchaeological data were derived from
Postclassical settlements located in the northern part of the Carib-
bean coast, not on the south, near to the southern lowlands, as
was the case for Classic settlements. In fact, it would be interesting
to confirm whether the “environmental recovery” documented on
the southern lowlands after the collapse period also occurred on
the southern coast of the Caribbean, as suggested by Curtis et al.
(1998) and Rosenmeier, Hodell, Brennrer, Curtis, and Guilderson
(2002). There exist few Postclassic settlements in the southern region
of the Yucatan peninsula as the population apparently migrated to
the north when the crisis on the southern lowlands triggered the Ter-
minal Classic (McKillop, 2004). It might have been possible that this
allegedly migration was in fact the result of environmental pressure.
If that was the case, this would explain the apparently “healthy state”
of the coastal fish populations in the northern Caribbean during the
Postclassic when the levels of anthropic pressure were less
developed.
JIMÉNEZ CANO
The Mayan coast of the Caribbean constitutes a region
that needs to be explored further, both archaeologically and
paleoenvironmentally, and for this reason, the ichthyoarchaeological
data will need to be contrasted with additional studies before reliable
baselines can be established in this region.
4 | CO NC LUSIO NS
The ichthyoarchaeological evidence from the Maya lowlands proved
to be an excellent tool for bioindication. The data gathered indicate
a fluctuating seascape model where droughts reported at the Terminal
Classic modified the coastal scenario and, consequently, affected fish
acquisition in the Yucatan peninsula. The evidence gathered so
far suggests that the coasts of the NML did not remain ecologically
stable through time. Notwithstanding the potential of the
ichthyoarchaeological evidence as a powerful paleoecological tool,
more sedimentological and geomorphological investigations need to
be done to confirm a more robust characterization of the paleo‐coast
along the NML.
Nevertheless, the results presented in this paper constitute an
emerging frame of reference to understand the implications of pre‐
Hispanic Mayan fishing strategies, thereby providing an important
contribution to our knowledge of historical changes in Maya fisheries.
ACKNOWLEDGEMENTS
This research was benefited from Grant 215047 from the Mexican
Consejo Nacional de Ciencia y Tecnología (CONACYT). Permission
to analyse the Xcambó and Mayapán material was kindly provided
by Thelma Sierra Sosa (Centro INAH Yucatán), Marilyn Masson (State
University of New York at Albany), and Carlos Peraza Lope (Centro
INAH Yucatán). Carlos García Paz is also thanked for granting permis-
sion to include the data of his Masters Dissertation. Kitty Emery, Irv
Quitmyer, and Christopher Götz are thanked for permission to access
the reference collections in their respective institutions. Arturo
Morales and Eufrasia Roselló revised earlier versions of this paper
and contributed with thoughtful comments. Three anonymous
reviewers are also thanked for their insightful comments that
improved the quality of the paper.
JIMÉNEZ CANO
ORCID
Nayeli G. Jiménez Cano https://orcid.org/0000-0001-9973-2452
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SUPPORTING INFORMATION
Additional supporting information may be found online in the
Supporting Information section at the end of the article.
How to cite this article: Jiménez Cano NG. Pre‐Hispanic
Maya fisheries and coastal adaptations in the northern low-
lands from the Classic (500–900 AD) to Postclassic (900–
1400 AD) periods. Int J Osteoarchaeol. 2019;29:469–476.
https://doi.org/10.1002/oa.2790