Depletion of a resource?

The impact
of prehistoric human foraging on
intertidal mollusc communities and
its signi cance for human settlement,
mobility and dispersal

Marcello A. Mannino and Kenneth D. Thomas

Abstract

Ethnoecological studies have demonstrated the impacts that even relatively small-scale human
foraging has on targeted species of shellsh and the structure of biological communities in intertidal
zones. There is compelling archaeological evidence that people in various parts of the world often
had a depleting effect on shellsh populations. Shellsh and other marine resources have sometimes
been perceived as lowly ranked foods and coastal archaeological sites have often been interpreted
as temporary (possibly seasonal) sites for the exploitation of these inferior food resources. This
model has been challenged by studies of mid-Holocene Mesolithic hunter-gatherer sites in Atlan-
tic Europe, which have shown that marine foods were the main component of the total diet and that
human foraging can deplete shellsh resources. Although subsistence systems based on coastal
resources might have been both viable and acceptable in dietary terms, regular mobility would have
been necessary for them to be sustainable. On longer time scales, such coastal mobility might result
in population dispersal. Sites associated with early anatomically modern humans show the antiquity
of coastal adaptations, including the consumption of shellsh, and the dispersal of early modern
humans out of Africa into south-east Asia and Greater Australia could have been through coastal
environments. This coastal dispersal could have been driven, at least in part, by the impact of early
human foragers on intertidal food resources. Resource depletion in coastal zones was probably
among the rst signicant, but small-scale, ecological impacts of human beings.

Keywords

Coastal resources; colonization; early modern humans; hunter-gatherer mobility; over-exploitation;

resource depletion; shellsh; shell middens.

World Archaeology Vol. 33(3): 452474 Ancient Ecodisasters

2002 Taylor & Francis Ltd ISSN 0043-8243 print/1470-1375 online
DOI: 10.1080/0043824012010747 7
 Depletion of a resource 453

Introduction

Archaeologists and Quaternary scientists generally accept that human populations in the
past sometimes brought about signicant changes to their environments. Agricultural
practices, in particular, have been invoked as signicant movers behind environmental
change. The effect that hunters, gatherers or foragers might have had on their environ-
ments is, however, less certain. On landscape scales, re would probably have been the
most likely tool for effecting, either deliberately or unintentionally, changes in the
environment, while skilled and organized hunting has been cited as a factor in the extinc-
tion of species of large mammals and birds in various parts of the world (including overkill
of Pleistocene megafaunas). The potential impacts of gathering and foraging practices
have generally been assumed to be insignicant on landscape scales (unless re was used)
and of only minor, probably temporary, signicance on local spatial scales. In this paper
we explore the impacts of human foraging on intertidal marine ecosystems, in particular
the evidence for the impact of human exploitation on marine molluscs and how over-
exploitation might be detected in the archaeological record (with a small case study
presenting aspects of our own site-based work), followed by a wider discussion of prehis-
toric human exploitation of intertidal resources and how this might affect patterns of
human mobility and longer-distance dispersal.
Intertidal environments, with a range of resources that are continuously available and
easily accessed, are potential theatres for the enactment of the tragedy of the commons
(Hardin 1968). Hardin reasoned that free access to exploit a common resource would ulti-
mately bring ruin upon all who exploit it. Each exploiter would seek to maximize their
share of the resource, because others also seek to do so. From the perspective of an indi-
vidual exploiter, the negative consequences of acting selshly do not fall upon that indi-
vidual alone but are shared between all exploiters. Therefore, the advantages of acting
selshly outweigh those of not and, because it is in the individual interests of all exploiters
to act in this way, the resource is ultimately destroyed. Whether or not intertidal resources
would ever, in practice, become destroyed is a matter of debate; it is equally likely that
people would move away to exploit less depleted areas of coastline (if such were avail-
able), or to exploit resources in alternative ecosystems, allowing the depleted resources
to recover. Yesner (1987: 293) suggested that human groups exploiting shellsh resources
might evolve strategies to allow fallowing of depleted locations by switching exploitation
between alternative coastal localities. Such indirect resource management could, there-
fore, ensure the long-term viability of life styles based on coastal foraging.
Claassen (1998) has recently challenged the view that people might have a depleting
impact on shellsh resources. She outlines various sources of environmental and biotic
pressures that species of shellsh encounter in order to counter, as she puts it, the fallacy
in assuming that all of Gods dangers for molluscs involve humans (Claassen 1998: 49).
That animal species, including shellsh, are subject to a diverse range of factors that cause
signicant mortality is, of course, well understood by ecologists and many archaeologists.
For example, Hockey (1994: 1925) pointed out that shellsh species have evolved life-
cycle strategies, fecundity rates, behavioural mechanisms and other biological character-
istics to enable them to survive vicissitudes in their physical and biotic environment. The
ecological relationships between prey species and their predators are complex and
454 Marcello A. Mannino and Kenneth D. Thomas

co-evolutionary (Hockey 1994: 20), but human predation on shellsh is, by contrast, a
relatively recent phenomenon in evolutionary terms. Human predators are dextrous tool-
users with good visual acuity, able quickly to harvest large quantities of shellsh (especi-
ally of species which live in dense colonies). Their activities are carried out at low tide
when shellsh are especially susceptible to human predators (although predatory birds
might also exact a signicant toll). Predation during low tides is therefore additional to
the predation loads incurred by many species of shellsh during periods of submersion.
Contra Claassen, we believe there is a potential for this unique form of predation to cause
signicant disequilibrium in shellsh populations, but is there any evidence that it actu-
ally happens?

Intertidal ecology and the impact of human foraging on modern shore communities

Recent ecological studies have demonstrated that even relatively small-scale human
foraging might have signicant effects both on targeted species of shellsh and on the
structure of biological communities in intertidal zones. Much of this research has been
undertaken on rocky shores (e.g. Siegfried 1994), but studies of sandy and muddy shores
(e.g. Okera 1976; McLachlan et al. 1996) show similar effects.
Predation is often an important factor in structuring intertidal communities (Little and
Kitching 1996: 172). Pioneering experimental work by Paine (1974, 1994) demonstrated
that signicant changes in the dominance relations of various intertidal species could be
brought about by selective removal of certain keystone species from areas of the shore.
A view is emerging among ecologists that alternative stable communities can exist on
rocky shores when even fairly small perturbations cross certain thresholds (Petraitis and
Dudgeon 1999). It has been shown that humans can either directly or indirectly affect
the biotic composition of shores. A study of rocky shores in southern and central Cali-
fornia by Lindberg et al. (1998) demonstrated, both experimentally and by observation
of shores exploited by humans, that reduced abundance of large-shelled species of
limpets led to an increase in both large brown algae and in various species of small-
shelled limpets. The intensity of disturbance (by trampling or removal for food, bait or
aquarium specimens) by humans of rocky shore communities in California bears a direct
relationship to reduced population density and diversity of intertidal organisms (Addessi
1994).
Other ecological studies, notably in Chile and southern Africa, as well as in some other
parts of the world (Siegfried 1994; Castilla 1999), have focused on the impact of recent
subsistence foraging on particular species. Human foragers were excluded for nearly two
years from an area of rocky shore in Chile (Castilla and Durn 1985), preventing them
from exploiting the economically important gastropod Concholepas concholepas (a
predator on mussels). C. concholepas increased in abundance while the previously domi-
nant mussel species showed a dramatic decline, paralleled by an increase in barnacles
which had previously been out-competed by the mussels. In adjacent uncontrolled areas
mussels continued to be the dominant species. The longer-term effects of human exclu-
sion are shown in marine reserves on Vancouver Island, British Columbia (Canada),
where the otherwise severely exploited northern abalone Haliotis kamtschatkana has
 Depletion of a resource 455

highest population densities in the oldest reserve (established forty years ago), with the
animals reaching much larger sizes and older ages (Wallace 1999). Keough et al. (1993)
compared mollusc populations on a rocky shore in Victoria (Australia) that had been
protected from human foragers for seventy years with those on unprotected shores. They
showed that species exploited for food or bait were less abundant and had smaller size
distributions on exploited shores, while species that were rarely if ever collected showed
no signicant differences between the shores.
Hockey and Bosman (1986) showed that shell size of populations of exploited species
(mainly various limpets) was reduced compared to those on non-exploited shores in the
Transkei, southern Africa. Species diversity was greater on the exploited shores, suggest-
ing that disturbance through selective predation by humans allowed a wider diversity of
species to co-exist. Swadlings (1976) study of exploited reef-platform populations of
Strombus luhuanus in Papua New Guinea showed they were signicantly smaller and had
higher frequencies of individuals with juvenile shell traits than lightly exploited popu-
lations. Human foragers in the Transkei select larger shells (Hockey 1994), leaving the
exploited shore populations with smaller shells. Branch (1975) found that human preda-
tion on the limpet Patella concolor on the east coast of South Africa led to a change in
population structure, with reduced maximum shell size, biomass, gonad size and (poten-
tially) gonad output. Recently accumulating shell middens have also been analysed and a
reduction of shell size through them has been demonstrated (Hockey and Bosman 1986;
Hockey et al. 1988).
Lasiak (1991a, 1991b, 1992, 1993) has, however, highlighted the problems of consider-
ing a decrease in either population density or shell size as being due only to exploitation
by humans. Her study of the limpet Cellana capensis on exploited and non-exploited
shores showed no evidence for a decline in density on the exploited shore (Lasiak 1993);
rather there was a decline in density on the non-exploited shore resulting from poor
recruitment and high mortality of older limpets. Shifts over time in the size of limpets on
the two shores, as well as inter-annual differences in recruitment, could not be related to
human exploitation. Short-term changes in factors such as recruitment could, therefore,
offset or mask any impacts that human foraging might have had.
Population and community dynamics in littoral zones are complex and susceptible to
marked scalar effects over both time and space. The population dynamics of intertidal
organisms is strongly inuenced by the rate of supply of new recruits, for which the term
supply-side ecology has been coined (Roughgarden et al. 1987). Where settlement rates
are high, inter-annual variation in population density is low but density varies through
each year as a consequence of high juvenile mortality (Gaines and Roughgarden 1985).
Where recruitment varies from year to year, there is marked inter-annual variation in
population density but density varies little within each year because of reduced biotic
interactions. Clearly, species with different life-cycle strategies will show differing patterns
of abundance according to the temporal and spatial scales of observation. Equally, for any
given species, the local properties of each shore type, or habitat, will affect recruitment
and survival and, in turn, population densities. In addition, the trophic level occupied by
species or groups of species (guilds) might have a signicant effect on both biomass and
population densities (Barnes and Hughes 1988). In general, primary consumers such as
grazers and lter-feeders will have higher total biomass and denser populations than
456 Marcello A. Mannino and Kenneth D. Thomas

predatory species, with detritivores (including scavengers) probably being somewhere in

between.
Catterall and Poiner (1987) assessed the potential for depletion of shellsh species by
human predators by taking into account both habitat and life history attributes and Lasiak
(1991b, 1992) adopted a similar approach in a study of intertidal rocky shore gastropods.
The main factors inuencing the susceptibility of species to stock depletion are:
geographic distribution, habitat preferences (cryptic behaviour), presence of adjacent
refuge populations, mobility of benthic stages in the life cycle, duration of a viable larval
phase and size at sexual maturity. Each of these factors can bring advantages or disadvan-
tages to a species. For example, a species that is sessile and gregarious is very vulnerable
to depletion. A species that has adjacent refuge populations, either sub-tidally or on inac-
cessible shores, can repopulate more easily and counteract the depleting effects of exploi-
tation. The likelihood of re-population is, however, dependent on other factors such as
the longevity of planktonic larval stages, the swimming ability of the larvae and the pres-
ence of currents to disperse the larvae to the exploited shores. Size at sexual maturity is
also very important: if the size at sexual maturity is less than the limital size at which a
human predator would consider collecting it, the exploited shore will always have repro-
ducing individuals (Lasiak 1991b).
Different types of shores, and organisms with different biological and ecological prop-
erties, might be varyingly susceptible to over-exploitation, but patterns of human forag-
ing, often related to social and cultural factors, also play an important role. For example,
Kyle et al. (1997a, 1997b) analysed over a seven-year period yields of coastal harvests by
subsistence foragers in Maputaland Marine Reserve, northern Kwazulu-Natal (South
Africa). Yields of rocky shore organisms (mainly mussels, oysters, limpets and tunicates)
showed no tendency to decline, partly due to a reduction of effort by foragers during the
period (Kyle et al. 1997a). A parallel study of foraging on sandy shores, mainly for burrow-
ing crabs, produced similar results (Kyle et al. 1997b).
These changes in foraging intensity resulted from a number of social and cultural
factors: a growing perception that coastal foraging was a poor persons occupation; an
increase in the development of a cash economy; linked to this, an improvement in local
employment prospects; and a fall in the number of harvesters because refugees from the
civil war returned home to Mozambique (Kyle et al. 1997b: 180).

Role of human foraging in the extinction of marine species

Although there is compelling evidence that human foragers can over-exploit (i.e. deplete)
shellsh stocks, it is less certain whether they are ever likely to cause extinction of species
(Hockey 1994). While some species might be at risk, especially those near the limits of
their geographical range, such as some species of giant clam in southern Africa (Herbert
1998), it has generally been assumed that marine resources are abundant and self-
renewing. This view is now changing and ecologists (e.g. Roberts and Hawkins 1999) now
consider human foraging to be a potential contributory factor in extinctions, although
most of the contemporary threats to intertidal and other marine communities come from
 Depletion of a resource 457

domestic or industrial pollution, including accidental or deliberate oil spillages. These

often also lead to shellsh becoming unt for human consumption (IUCN 1983).
The scale or intensity of exploitation is an important factor, with mechanized extrac-
tion by dredging of species living on seabed deposits having a signicant impact (e.g.
Briggs 2000). Diving can also have an impact, even on species which are difcult to gather
such as rock-boring bivalves (Lithophaga), highly prized as food in southern Italy, but
which are increasingly threatened by over-exploitation (Fanelli et al. 1994). Organisms
inhabiting the intertidal zone of rocky shores are probably the most susceptible to over-
exploitation and the possibility of extinction. The white abalone (Haliotis sorenseni) is by
now almost certainly extinct as a result of over-collection, despite it once having an exten-
sive range along the west coast of North America (Tegner et al. 1996).
Wolff (2000), using archaeological, historical and biological sources, documented the
extirpation of various species (10 algae, 10 invertebrates, 13 sh, 5 birds and 4 marine
mammals) in the Wadden Sea (The Netherlands) over the past 2000 years. While habitat
change could account for many of the local extinctions, at least seventeen of the cases could
be ascribed to over-exploitation. For a recent general review of historical extinctions of
both vertebrate and invertebrate species in marine ecosystems, see Carlton et al. (1999).
That extinction through over-exploitation has not occurred more often is probably
related to two main factors: the existence of inaccessible populations providing new
recruits to exploited areas and adaptive human foraging behaviour. For many species the
rejection size by human foragers is larger than size at rst reproduction, although this is
not the case for late maturing taxa such as Haliotis. Forager behaviour also plays against
over-exploitation through prey switching or shifting to other locations where returns are
more productive (i.e. optimal foraging strategies).

Archaeological evidence of the impact of shellsh gathering

Although molluscs might be represented (by their shells) in archaeological deposits

because they were exploited for a range of different reasons (e.g. as discussed by Thomas
and Mannino 2001), the presence of many species, often in large numbers, is most
frequently attributable to their value as a food resource. Over-exploitation has been used
to explain changes in the abundance of species within shell-containing archaeological
deposits, especially when coupled with reductions in shell size (e.g. Botkin 1980; Kent
1988; Spennemann 1989). In addition to these, Botkin noted a shift from more easily gath-
ered and processed species to those that were less easy to collect and process, while Yesner
(1984, 1987) observed species switching over time, possibly to spread the exploitation
load and to allow populations to recover.
Raab (1992) used optimal foraging models to explain shifts in exploitation of shellsh
by the prehistoric inhabitants of San Clemente Island (California). Abalone (easily
processed and high meat-yielding) was initially the dominant shellsh but Tegula (small,
hard to process and low yielding) increased over time while abalone declined, leading Raab
(1992) to suggest that the latter had been over-exploited. Anderson (1981) used a similar
approach for modelling prehistoric rocky shore collecting in Palliser Bay (New Zealand),
458 Marcello A. Mannino and Kenneth D. Thomas

where shell middens showed a shift from a limited range of larger species to a wider spec-
trum of smaller ones. This could not reasonably be explained by changes in choice against
the larger species and was thought to reect over-exploitation of the larger species.
There are problems in using optimal foraging models as the primary basis for inferring
over-exploitation. Meehans (1982) classic studies showed that human foraging strategies
and processing decisions can have a marked impact on what is incorporated into a
developing midden (presumably representing what was taken back to a camp for
consumption), as opposed to what is snacked during foraging and therefore lost to the
archaeological record. Bird and Bliege Bird (1997) demonstrated similar effects in their
study of shellsh gathering strategies among the Meriam of Torres Strait (Australia). The
Meriam collected 90 per cent of their shellsh from reef ats and only a small part from
rocky shores. Many species from reef ats (especially Hippopus and Tridacna) had rela-
tively low processing costs and were mainly processed in the eld. Shellsh species from
the rocky shore (mainly Asaphis and Nerita) required longer processing and as a result
were more frequently transported back to the base camp. In consequence, shells of higher-
ranked shellsh were under-represented at the home base while lower-ranked ones were
over-represented. This model could account for the species composition of the later
midden deposits studied by both Anderson (1981) and Raab (1992), although it cannot
explain why higher-ranked species dominated the earlier phases. In a further analysis of
Meriam shellshing strategies, Bird and Bliege Bird (2000) demonstrated that, where chil-
dren were also active foragers, their contribution to the array of species collected tended
to broaden the spectrum represented by including lower-ranked species that were rarely
collected by adults. If such behaviour was represented in discrete assemblages of shell-
sh, these might erroneously be interpreted either as an intensication of coastal exploi-
tation or as a shift to a broader range of species following possible over-exploitation of
certain preferred species.
Given such problems of interpretation, what criteria recognizable in the archaeological
record might validly indicate over-exploitation of shellsh, as opposed to changes caused
by other factors such as environmental change or changes in human foraging and trans-
portation strategies?

Criteria for inferring over-exploitation from shell midden samples

Reduction in abundance and size of shells through a midden has often, as noted above,
been interpreted as evidence of over-exploitation, but such changes might also have
environmental causes. The main criteria that have been used to suggest over-exploitation
(based on Claassen 1986, 1998) are:
absolute abundance of preferred species will decrease through a midden deposit
mean shell size will decrease through samples taken from the bottom of a midden to
the top
mean or modal shell size of the archaeological samples of a species will be signicantly
smaller than in a non-exploited population.
less easily procured species will increase in number up through a midden deposit
less easily processed species will increase in number.
 Depletion of a resource 459

Each, or all, of these criteria could also be construed as evidence for environmental
change and therefore Claassen (1986: 130) proposed an additional criterion: there should
be a reduction in the mean age of the shells from bottom to top of a midden with no atten-
dant difference in mean shell size within each of the age groups through the midden.
Some studies have explored the impact of human predation by means of the age struc-
tures of shellsh assemblages. Koike (1986), working on shellsh remains from the Jomon
Period site of Kidosaku (Japan), showed that during the Late Jomon period the age struc-
tures of Meretrix lusoria were dominated by three-year-olds, as was a heavily exploited
present-day population. Later (Kofun period) assemblages were dominated by ve-
year-old shells, suggesting a lesser impact at that time.
Heavy harvesting of shellsh was inferred by Lightfoot and Cerrato (1988) from their
analysis of growth lines in shells of Mercenaria mercenaria from the Sungic midden site
on Shelter Island (New York). This species can live for fteen years, exceptionally up to
twenty years. The age distributions of the archaeological samples resembled those of an
intensively harvested population on Long Island, with very few shells in the eight- to
fteen-year age cohorts. Swadling (1976) interpreted changes in shell size and age (from
growth lines) in prehistoric assemblages of the bivalve Chione stuchburyi in northern New
Zealand as indicating over-exploitation. In the same article, Swadling noted that archaeo-
logical samples of Strombus luhuanus from Papua New Guinea were very similar, in both
shell size and high frequency of individuals with juvenile shell traits, to modern heavily
exploited populations.
In all cases where over-exploitation is inferred it is important that all other likely
hypotheses, especially the possible effects of environmental change, have also been evalu-
ated. This is especially true for species for which ageing is not possible. Jerardino (1997)
examined changing species composition and the frequencies and mean shell sizes of the
black mussel Choromytilus meridionalis through a shell midden in southern Africa. Stable
isotope studies showed marked shifts in sea temperatures during the period of the midden,
but these were not correlated with changes either in species composition or in mean size
of mussels. Patterns in abundance and mean size of the latter could in part be correlated
with changes in water turbidity, but variations around 3000 BP were attributed to more
intensive human predation.

A case study: the Mesolithic shell midden at Culverwell, Isle of Portland

Our own studies, reported in a preliminary form (Thomas and Mannino 1999) and more
denitively (Mannino and Thomas 2001), have demonstrated over-exploitation of shell-
sh resources at the Culverwell Mesolithic site (Portland, southern Britain), with a fall-
off in the absolute abundance of the main species through the midden sequence. The
shells of one of these species, the toothed top shell Monodonta lineata, can be aged by
counting the growth lines (varices), which have been shown to be annual (Williamson and
Kendall 1981), although only intact shells can be aged by this method.
It is likely that human foragers would have collected shellsh on the basis of their size
rather than age, but size in M. lineata is not independent of age. Younger age classes (up
to three or four varices), especially, show signicant inter-age size differences that become
460 Marcello A. Mannino and Kenneth D. Thomas

less between older age groups. Human gatherers would have selected larger, higher meat-
yielding shells, and therefore larger and older shells would be expected to dominate the
midden samples. The age structures of the assemblages from the main layers of the
Culverwell midden are shown in Figure 1, for three independent columns of samples
taken through the midden: trench 4 column 1 (MC1), trench 4 column 2 (MC2) and trench
41. The mean sizes and mean ages of these assemblages are shown in Table 1a. The assem-
blages of intact shells from MC1 and MC2 were sufciently large to analyse separately
the sub-samples within layer 8 (the most intensively occupied phase of the midden). The
age-frequency histograms for these are in Figure 2 and mean sizes and ages in Table 1b.
Taking these data sets together, the following results are apparent:
1 As expected, all of the assemblages are biased away from youngest age classes (i.e.
those too small either to nd or be worth collecting).
2 The lowest assemblages in MC1 and MC2 have irregular age distributions, suggesting
either bias in collectors behaviour or uctuations in recruitment to the ancient shore.
The assemblages from layers 8 and 9 (taking into account biases against the younger
age cohorts) have age structures more similar to those of age-stable populations. This
suggests that gathering of shellsh had become more intensive, resulting in assemblages
that, for the older collectable ages, probably more closely reect the age structures of
the shore populations from which they were derived.
3 In each case there is an apparent reduction in mean shell size through the depositional
sequence, although this is formally signicant only between samples 7 and 6 of MC1 (p
= 0.0325, Mann-Whitney U test).
4 In each case there is a reduction in mean age, with those between layers 9 and 8 and
layers 12 and 8 of trench 41 being statistically signicant (p = 0.0130 and p = 0.0004,
respectively, Mann-Whitney U test).
5 The age histograms of M. lineata through layer 8, MC1 (Fig. 2), show a progressive shift
in the dominant age classes towards the younger age ranges. There is a marked reduc-
tion in the 5, 6, 7 and 8 varix classes. For MC2 (Fig. 2) the pattern is less clear-cut, but
there are broadly similar trends. The reduction in varix classes 810, inclusive, can also
be seen in the assemblages from layer 8 in trench 41 (Fig. 1).
Environmental change cannot explain these results, as discussed in detail by Mannino and
Thomas (2001). Ecological studies of M. lineata have shown that mean shell size and popu-
lation density are negatively correlated, while studies of 24 populations across the modern
range of the species showed that age classes of 7, 8 and more varices were consistently
represented (Mannino 2000; Mannino and Thomas in preparation). Populations in less
favourable environments had relatively higher proportions of older age categories,
suggesting that mortality was exacting a heavier toll on planktonic and newly settled indi-
viduals than on well-established older ones. Had the fall-off in abundance through the
midden been the result of environmental change on the local shores, both mean shell size
and age should have increased rather than decreased. Environmental factors are there-
fore unlikely to lie behind the trends in layer 8 of the midden.
The changes in size in M. lineata would have been difcult to interpret without exten-
sive modern ecological surveys of the species. These have shown (Mannino and Thomas
in preparation) that the mean size to which animals grow is dependent on a number of
 T r e n c h 4 C o lu m n 1 T r e n c h 4 C o lu m n 2 T r e n c h 4 1

Layer
L a y e r8 (n = 1 3 8 )
8 (n=138) L a y e r 8 (n = 8 5 ) L a y e r 8= 5 9 )
Layer 8 (n=85) Layer 8( n(n=59)
35 35 35
30 30 30
25 25 25
20 20 20
% % %
15 15 15
10 10 10
5 5 5
0 0 0
1 2 3 4 5 6 7 8 9 10 1 2 3 4 5 6 7 8 9 10 1 2 3 4 5 6 7 8 9 10
varices varices varices

n = 1 4 8 ) n = 3 5 ) L a y e r 9 = 5 9 )
Layer
L a y e r 9
9 ((n=148)
Layer
L a y e r 9
9 ((n=35)
Layer 9( n (n=59)
35 35 35
30 30 30
25 25 25
20 20 20
% % %
15 15 15
10 10 10
5 5 5
0 0 0
1 2 3 4 5 6 7 8 9 10 1 2 3 4 5 6 7 8 9 10 1 2 3 4 5 6 7 8 9 10
varices varices varices
Layer
L a y e r 9-12 (n = 3 2 )
9 - 1 2 (n=32) Layer
L a y e r 12 (n = 1 1 )
1 2 (n=11) Layer
L a y e r 12 n = 1 1 9 )
1 2 ((n=119)
35 35 35
30 30 30
25 25 25
20 20 20
% % %
15 15 15
10 10 10
5 5 5
0 0 0
1 2 3 4 5 6 7 8 9 10 1 2 3 4 5 6 7 8 9 10 1 2 3 4 5 6 7 8 9 10
varices varices varices

Figure 1 Culverwell shell midden. Age proles, based on shell varix counts, of Monodonta lineata in three columns of samples taken through the
midden (Layer 8 is the uppermost, youngest, layer) (n = number of countable shells in each sample).
462 Marcello A. Mannino and Kenneth D. Thomas

Figure 2 Culverwell shell midden. Age proles of Monodonta lineata in vertically ordered sub-
samples within Layer 8 (see Fig. 1).

biological and ecological factors, including temperature, population density and substra-
tum. Such observations are in general accord with those of Lasiak (1993), who showed
that differences in mean shell size varied considerably between different shores whether
they were being exploited or not. Our work has suggested that age is less directly inu-
enced by environmental factors than is size and, compared with size, is less likely to be
the immediate criterion inuencing human choice. Arguments for over-exploitation based
on age distributions are more likely to be convincing than those based solely on size, but
only if sufcient knowledge is available on the age structures and dynamics of modern
 Depletion of a resource 463

Table 1 (A) Mean shell sizes (diameter, in mm) and mean ages (based on varix counts) of M. lineata
from the three sample columns (see Fig. 1). Figures in brackets are sample sizes; sample sizes for
mean shell size are smaller because size is measured at the point of maximum curvature of the shell
aperture in undamaged specimens. (B): As for (A) but for the sub-samples within Layer 8 (see Fig.
2).
A

Trench 4 Column 1 Trench 4 Column 2 Trench 41

Layer Mean Mean Layer Mean Mean Layer Mean Mean
size varices size varices size varices

L8 17.70 (56) 4.0 (138) L8 17.90 (37) 4.0 (85) 8 19.00 (12) 4.4 (59)
L9 18.41 (82) 4.3 (148) L9 18.70 (11) 3.7 (35) 9 18.91 (21) 5.1 (59)
L912 17.67 (20) 4.3 (32) L12 19.16 (7) 4.6 (11) 12 19.82 (38) 5.4 (120)

Trench 4 Column 1 Trench 4 Column 2

Sample/ Mean Mean Sample/ Mean Mean
layer size varices layer size varices

S5/L8 16.73 (4) 3.6 (15) S8/L8 17.67 (3) 3.8 (15)
S6/L8 17.01 (19) 4.0 (45) S9/L8 17.60 (12) 4.0 (30)
S7/L8 18.15 (33) 4.2 (78) S10/L8 18.08 (22) 4.2 (40)

populations of the species. Age has been used to demonstrate over-exploitation in a few
archaeological studies (e.g. Swadling 1976; Koike 1986; Lightfoot and Cerrato 1988), but
in these comparisons were made only between the archaeological assemblages and
modern populations from localities close to the archaeological sites, which were not
necessarily the most appropriate analogues.
Claassens (1986) test for over-exploitation proposed that there should be a reduction
in mean age with no difference in size within the same age cohort. Although in theory
such a test would be useful, in practice it is rarely possible to have sufciently large
samples of intact and measurable shells for each age class. This was the case in our Culver-
well study, but detailed knowledge of the ecology of the species involved provided a valu-
able framework for interpreting the archaeological assemblages. The results suggest,
therefore, that changes in abundance of the main species during the occupation of the
Culverwell midden reect the impact of increased levels of human predation on nearby
shores. The failure, over time, for individuals of M. lineata to be recruited (by shell
growth) into the older age classes suggests that human predation must have been both
intense and frequent (at least annual).

Over-exploitation in retrospect

Our review of the ethnoecological and archaeological evidence for over-exploitation of

intertidal marine molluscs has demonstrated, for various parts of the world, that there is
compelling evidence that foragers have had a signicant impact on mollusc populations,
464 Marcello A. Mannino and Kenneth D. Thomas

often leading to over-exploitation through depletion. Over-exploitation is a rather

general term that, as used here, represents a range of possibilities:
reduction in the average size and age of the exploited species, below those which would
be found in natural populations not exposed to human predation
reduction in the age structure of populations which might reduce breeding capacity and
recruitment, and hence population levels of molluscs on shores
reduction in the abundance (density) of molluscs on shores to levels at which they are
not worth exploiting by human foragers
reduction in breeding capacity which, in the lack of external recruitment, might lead to
local, regional or global extinction
These four possibilities are not, of course, discrete alternatives. Rather they are part of a
continuum of possibilities, each being related to the adjacent ones in functional or
response terms. In archaeological terms, only the rst category might be demonstrable,
with the second being inferred as a likely consequence. The third category might be
inferred from the archaeological record, if data on absolute abundance of species are
available and if alternative (including taphonomic) explanations can be discounted (e.g.
Mannino and Thomas 2001). The fourth might be inferred by the total disappearance of
a species in a depositional sequence (and if it is absent today in the area, or more widely),
again provided that alternative explanations could be effectively discounted. Even if only
the rst can be demonstrated, the result is still very signicant in human terms because it
demonstrates that human exploitation of shellsh in the past was sufciently intense
and/or frequent to alter the age structure of exploited populations signicantly. This, in
turn, has important implications for understanding the role of shellsh, and possibly other
coastal resources, in past human diets.

Coastal resources and shellsh in past human diets

The potential for human predation to have an impact on resources in intertidal environ-
ments depends on the relative importance of those resources in past human diets, and
especially on factors such as the quantities harvested and the frequency of harvesting at
particular localities. Environmental impacts might be fairly minimal if coastal environments
were inhabited for only brief periods, perhaps only to exploit second rate food resources
(Osborn 1977: 171) at times (or seasons) when preferred terrestrial foods were in short
supply. The role of shellsh in human diets has often been assumed to be minor (e.g. Bailey
1975) and possibly only of short-term seasonal signicance. It has also been suggested that
human occupation of coastal habitats and exploitation of inferior marine food resources
began only when population pressure forced people to do so (Cohen 1977; Osborn 1977),
and is therefore a relatively late human adaptation. The archaeological record, showing a
marked increase of early coastal occupation sites in the mid-Holocene, appears to conrm
this (Yesner 1984, 1987), although this might merely reect site preservation. By the mid-
Holocene rising sea levels had stabilized, having drowned Late Pleistocene and early
Holocene coastal landscapes along with any evidence of earlier coastal settlement and
marine resource exploitation (Jones 1991: 422).
 Depletion of a resource 465

The signicance of marine foods, especially shellsh, in prehistoric human diets has
been re-assessed by Perlman (1980), Yesner (1987) and Erlandson (1988, 1994), who
suggest that they might have been protein staples during periods when other meat was
not available. Many seafoods are potentially storable (Yesner 1987: 294; Henshilwood et
al. 1994), and seasonal species such as salmon can be smoked and dried for later consump-
tion. Shellsh are rich in protein and many essential minerals, but low in fats and carbo-
hydrates (Paul and Southgate 1978: 1517), and shellsh protein could be an essential
supplement to predominantly carbohydrate diets that might otherwise cause protein-
deciency conditions such as kwashiorkor (Buchanan 1988: 64). Proteins can be metabo-
lized to yield energy, but a diet high in protein and low in fat and carbohydrate could have
serious health consequences because of the need to excrete large quantities of toxic
nitrogenous waste products. Noli and Avery (1988) have, therefore, questioned if shell-
sh could ever have been a dietary staple. Buchanan (1988: 604) surveyed a range of
recent hunter-forager groups with high protein diets and found no evidence of any adverse
health effects caused by so-called protein poisoning.
Extreme diets based solely on protein are improbable in most coastal environments in
which there is access to a wide diversity of marine intertidal resources, including shellsh,
sh (oily sh are rich in essential fats), large marine algae, large marine mammals and a
range of terrestrial animal and plant foods. Many of these resources are available all year
round, but some may be seasonal. For example, Fitzhugh (1995), in a study of prehistoric
human subsistence on Kodiak Island (Alaska), showed that clams were an important
resource exploited through autumn, winter and spring, but not during the summer when
migratory salmon became the major resource. Fitzhugh (1995) also suggested there was
deliberate avoidance of shellsh during the summer to avoid poisoning by toxic blooms
(red tides) of algae, especially dinoagellates. Other scholars (e.g Waselkov 1987: 111;
Yesner 1987: 300) have also pointed to this hazard that might make all year round exploi-
tation of shellsh unlikely. This is a possibility but, as Claassen (1998: 334) has persua-
sively argued, it is likely that red tides and other algal blooms associated with shellsh
toxicity are relatively recent phenomena resulting from eutrophication of coastal waters
by efuents, including organic waste and agro-chemicals in run-off and eroded soil
material. It might not have been a problem in the past.
A major problem with the debates that have surrounded the significance of marine
foods, especially shellfish, in past human diets is that they have focused only on the
marine foods themselves (e.g. Yesner 1987) and not on the broader range of foods that
are almost always available in coastal environments. Osborn (1977: 195) contended that
primary production in marine ecosystems is very low, rendering them unsuited to
sustained human exploitation. He cited only productivity levels for the open seas, which
are well known to be far lower than for in-shore or coastal environments such as
lagoons, estuaries and reefs (Whittaker 1975: 224). Often these sorts of coastal environ-
ment are more productive than many fully terrestrial ones, with a high proportion of
plant and animal biomass directly available to humans. Recent work has shown that
terrestrial environments in coastal areas can be subsidized by significant energy and
nutrient inputs (mainly detritus) from the sea (Polis and Hurd 1996). Stable carbon and
nitrogen isotopes in terrestrial consumers in coastal areas have shown a strong marine
isotopic signal, even in species that do not feed directly on marine foods (Anderson and
466 Marcello A. Mannino and Kenneth D. Thomas

Polis 1998). A comparative study of coastal and inland coyotes (Canis latrans) in Baja
California (Rose and Polis 1998) showed the coastal populations had higher densities.
The contents of faecal scats revealed that coastal coyotes had a greater breadth of
dietary items, with approximately equal proportions of terrestrial and marine foods, and
higher absolute levels of food intake. In short, these coastal carnivores had access to
more food and to a greater diversity of foods, which probably accounted for their
greater biological success (higher population densities) compared with their purely
terrestrial counterparts. In general, the beneficial consequences of marine subsidies are
more likely to be manifest in coastal regions adjacent to low-productivity arid and semi-
arid terrestrial environments (Polis and Hurd 1996; Anderson and Polis 1998), which
might have consequences for past human exploitation of such environments, as we
discuss later in this article.
The importance of coastal (including marine) resources in Mesolithic subsistence
strategies in north-western Europe has been the subject of much recent debate. A case
could be made for late Mesolithic coastal sedentism on the small island of Oronsay
(southern Hebrides). Evidence of seasonality from shell middens on the island suggests
year-round exploitation of resources on the island (Mellars and Wilkinson 1980) and
the stable isotope composition of human bone collagen from the sites suggests that
marine foods were a major part of the diet (Richards and Mellars 1998). The results
from our own studies at Culverwell (Thomas and Mannino 1999; Mannino and Thomas
2001), summarized above, suggest that Mesolithic exploitation of intertidal marine
molluscs was probably both frequent and intense. On a broader geographical scale,
stable isotope evidence from late Mesolithic human remains from a range of sites along
the Atlantic coast of Europe suggests that marine foods were a significant part of the
diet of those individuals studied (Richards and Hedges 1999a), in marked contrast to
Neolithic individuals who had fully terrestrial diets (Richards and Hedges 1999b).
Richards and Hedges (1999a) suggested that the late Mesolithic individuals analysed
were probably eating mostly fish, or other secondary or tertiary consumers in marine
food chains, with primary consumers such as most marine molluscs constituting a rela-
tively minor part of the food intake.
Larsson (1999) has shown that the scarcity of evidence of coastal settlement in the early
Mesolithic of southern Scandinavia is partly due to rapid and signicant rises of sea level
during the Late Boreal and Atlantic periods, leaving only the later Mesolithic sites
represented. Another factor is that the sea (and its resources) only arrived in the later
Mesolithic of Denmark (Rowley-Conwy 1999: 137), as a result of rising sea levels ood-
ing an erstwhile freshwater lake, creating a diverse coastline with numerous islands
(Pedersen et al. 1997: 23, g. 2). This was the resource-rich ecological context within which
complex and possibly more sedentary coastal societies developed (Rowley-Conwy 1999:
137; Andersen 2000). The broader signicance of all this work is in showing that people
responded readily to the opportunities of this new and productive marine coastal environ-
ment, rather than being forced to do so as a consequence of population pressure. Indeed,
it has been argued (e.g. by Yesner 1987) that coastal subsistence strategies can be a causal
factor behind population growth, rather than being a mere response to population growth
that occurred elsewhere.
 Depletion of a resource 467

Consequences: ecodisaster or a little local difculty?

From the foregoing discussion it appears that in many regions of the world, human settle-
ment and subsistence in coastal environments would have been viable and in some cases
preferable to more fully terrestrial environments. Rather than being second rate foods,
marine foods would have formed an important part of a diverse marine and terrestrial
diet available in coastal areas. The potential over-exploitation of intertidal resources
could have been a problem, which might have necessitated regular mobility for these
resources to be sustainable. Such mobility might have involved seasonal or other episodic
patterns of movement within broad coastal territories, or into non-coastal areas. For
coastal-adapted foragers, the former option would probably have been preferred. Instead
of interpreting the archaeological record as indicating seasonal movements to coastal
environments to exploit marine resources when preferred terrestrial ones were scarce, it
is possible to argue (e.g. Yesner 1987) that the record indicates seasonal movements within
coastal areas, which was driven by local coastal resource depletion. Depending on popu-
lation levels and on forms of social organization, in certain circumstances these move-
ments might have been constrained to local territorial areas, but in the absence of such
social and territorial constraints we suggest that the longer-term consequences might have
been population dispersal to new coastal territories. On a local scale, coastal environ-
ments are usually anked by other, often broadly similar, coastal environments. People
can therefore move between them and exploit them with little new technological and
social adaptation being required.

The world was their oyster

Coastal environments might be seen, in a biogeographical sense (e.g. Spellerberg and

Sawyer 1999: 1815), as linear corridors along which dispersal might be channelled; they
might therefore have been an effective route by which early human groups colonized
previously unoccupied areas. Sauer (1962) was an early champion of coastal environments
as a productive habitat for early humans, offering a diverse range of edible and material
resources as well as being a relatively barrier-free zone for human dispersal and coloni-
sation of new regions. In the following, admittedly speculative, discussion, we consider
the possible relationship between coastal resource depletion, human dispersal and
colonization of new regions. Implicit in this is the assumption that early human dispersal
over long distances and long time scales is the cumulative outcome of small-scale disper-
sals over short time scales. This assumption could be wrong (as we acknowledge for the
special case of Greater Australia) and dispersals over long distances and time scales could
have involved completely different processes to those operating on smaller scales
although just what these special processes might have been is not easily dened.
There is an increasing body of evidence to show that both Neanderthals and early
modern humans exploited intertidal resources, particularly shellsh. The Gibraltar Nean-
derthals exploited a range of marine foods, including mussels, limpets, cockles and barna-
cles, many of which showed signs of burning or heating (Barton et al. 1999: 21; Barton
2000). Stiners (1993, 1994) detailed taphonomic studies of small animals from the
468 Marcello A. Mannino and Kenneth D. Thomas

Neanderthal site of Moscerini Cave (Italy) have also shown the early signicance of inter-
tidal resources, especially shellsh.
The early human occupants (Middle Stone Age) of sites in southern Africa exploited
intertidal shellsh from around 120,000 years ago (Volman 1978; Klein 1999), although the
exact antiquity of some Middle Stone Age mollusc exploitation, such as at Blombos Cave
(Henshilwood et al. 2001), has yet to be claried. Recently, Walter et al. (2000) reported
on a Middle Stone Age site in Eritrea, on the African coast of the Red Sea, dated to about
125,000 years ago and with evidence of exploitation of a range of near-shore gastropods,
bivalves, crustaceans and echinoids. Walter et al. (2000: 69) argue that highly arid
conditions in the region at the peak of the last interglacial, coupled with relatively reduced
levels of competition in coastal zones from other terrestrial mammals, might explain this
apparently sudden and widespread coastal marine adaptation by early humans by the last
interglacial. (To this we might add, as noted above, the benecial effects of marine subsi-
dies to productivity in coastal ecosystems in arid and semiarid environments.) If early
humans were adapted to coastal environments, then coasting out of Africa (Stringer 2000)
becomes a plausible model for early human dispersal from Africa through to south-east
Asia along coastal routes. Stringer (2000) suggests that a possible test for this hypothesis
would be to seek early human sites in association with fossil beaches in areas such as Arabia
and India, which sets up a challenge for future eldwork.
Colonization by coastally adapted foragers has been proposed for both Australia and
the Americas. Bowdler (1977) argued persuasively for colonization of Australia by coastal
groups, but Beaton (1985) pointed out the lack of early sites associated with intertidal
foragers, although any such sites must now lie below the sea. It is currently believed that
Australia was colonized by at least 5060,000 BP (Stringer 1999) and because no land
bridge with south-east Asia ever existed, this must have been achieved using some sort of
sea-borne craft. This, itself, would strongly support the idea of colonization by people who
were adapted to a coastal way of life, being completely familiar with sea-going craft, and
accustomed to undertaking voyages out to sea.
The case for a coastal route of initial colonization of the Americas is less clear-cut,
because both coastal and land routes were possible at times of low sea levels, although
colonization by boat would also have been possible at higher sea-level stands. Fladmark
(1978, 1979) and Gruhn (1988) consider the evidence in favour of a coastal route of entry
across the southern Beringian littoral and down the western coastal strip of North
America. The archaeological evidence is not strong, and any that might have existed must
now be under a considerable depth of sea and sediments. Jones (1991) reviewed the
evidence for early coastal settlement in California and concluded (ibid.: 429) that early
colonists preferentially exploited estuarine habitats, with shellsh being exploited in
signicant amounts (ibid.: 436). Early evidence of coastal subsistence, with shellsh
consumption, has also been found in Brazil (Fairbridge 1976). Recent modelling of
palaeoindian colonization of the Americas (Anderson and Gillam 2000) supports the idea
of a rapid spread of population and suggests that initial colonization and dispersal would
have been most likely through coastal and riverine routes.
Despite the uncertainties and ambiguities about the colonization of the Americas, there
is evidence to support the hypothesis that the primary routes of dispersal of early modern
humans out of Africa into south-east Asia and Australia were through coastal environ-
ments. Such migrations were undertaken by people adapted to coastal foraging, including
 Depletion of a resource 469

the exploitation of shellsh. As we have seen, coastlines are attractive environments for
human foragers and the exploitation of new coastal areas would require minimal adap-
tation. Coastlines, therefore, offer corridors for human population movement into new
areas, and passive diffusion of growing populations along such corridors might, on its own,
account for much early human dispersal. We have also, however, considered how suscep-
tible various categories of intertidal shellsh are to over-exploitation, and how this might
affect human movement strategies in coastal areas. Once the resources in a particular local-
ity had become depleted, early human foragers would have been freely able to move on
to exploit completely new shores. On longer time scales, the additive effect of such small-
scale local movements might have been longer distance dispersal, although in the case of
Greater Australia it is clear that a model based on cumulative random dispersal is not
sufcient on its own. It is also necessary to infer a rapid phase of jump dispersal, prob-
ably involving the use of sea-going craft. Nonetheless, it is possible that the localized impact
of human foraging on intertidal food resources was a contributing factor in a relatively
rapid coastal dispersal of people out of Africa and into some other parts of the world. If
so, resource depletion in coastal zones might have been among the rst signicant ecolog-
ical impacts of anatomically modern human beings.

Acknowledgements

We thank Susann Palmer, Myfanwy Stewart, Daniel Clark and Chris Stevens for advice
and support in the eld and the laboratory. Geoff Bailey, David Harris, Richard Preece
and Chris Stringer provided valuable comments and specic information. Terry OCon-
nor and an anonymous referee made useful critical comments on the initial draft of this
paper, helping to clarify our thoughts and arguments as well as our presentation of them.
Our eld and laboratory work was supported by grants from the Institute of Archaeology
(University College London), the Graduate School of University College London and the
Central Research Fund, University of London. We are grateful for the support (through
Radiocarbon Dating Allocation No. 677/1296) of the Natural Environment Research
Council. MAM acknowledges a fees-only research studentship from the Natural Environ-
ment Research Council.

Marcello A. Mannino and Kenneth D. Thomas (author for correspondence), Institute of

Archaeology, University College London, 314 Gordon Square, London WC1H OPY, UK,
email: <marcello_mannino@hotmail.com; k.thomas@ucl.ac.uk>

Note added in proofs

A recently published article by Jackson et al. (2001) on historical overshing, and relat-
ing this to the collapse of coastal ecosystems, echoes many of the points made in this
article about the potential for humans to deplete resources in coastal ecosystems.
470 Marcello A. Mannino and Kenneth D. Thomas

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