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Quaternary Science Reviews 81 (2013) 1e11

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Quaternary Science Reviews

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Human response to Holocene warming on the Cantabrian Coast(northern Spain): an unexpected outcome

Ana B. Marín-ArroyoInstituto Internacional de Investigaciones Prehistóricas, Universidad de Cantabria, Avd. Los Castros, s/n, E-39005 Santander, Spain

a r t i c l e i n f o

Article history:Received 25 June 2013Received in revised form5 September 2013Accepted 7 September 2013Available online 10 October 2013

Keywords:Pleistocene/Holocene transitionMesolithicCantabrian CoastUngulateShellfishResource productivityOverhuntingClimate change

E-mail address: anabelen.marin@unican.es.

0277-3791/$ e see front matter � 2013 Elsevier Ltd.http://dx.doi.org/10.1016/j.quascirev.2013.09.006

a b s t r a c t

Subsistence was characterized during the Pleistocene to Holocene transition on the Cantabrian Coast(northern Spain) by a progressive diet widening, with a greater exploitation of marine environmentsand a more intense consumption of low-ranked species. This trend was also accompanied by a generaland noticeable decrease in the amount of ungulates that were recovered from a set of archaeologicalsites clearly dominated by shells. The causes behind this change in the economic practice of the lasthunteregatherer groups are still being debated. There are currently two opposing views on the matter,with some scholars defending the role of demographic pressure as the main driving force, while otherresearchers invoke the importance of the environment in the food procurement preferences that wereadopted. Due to their overwhelming abundance, the debate has been mainly focused on marine re-sources, whereas the comparatively less-represented macromammal assemblages have been poorlyinterpreted. However, it is precisely this scarcity that makes them so remarkable. Here, a new inter-pretation of the available data is presented, with a special focus on the identification of overhuntingevidence and on the comparative productivity of each type of resource. Altogether, the demographichypothesis seems to be more coherent with the existing facts.

� 2013 Elsevier Ltd. All rights reserved.

1. Introduction

The Mesolithic witnessed a progressive increase in socio-economic complexity around the world towards the final appear-ance of agriculture. Not surprisingly, the diversity of environments,cultures and economies was so great (Spikins, 2007) that theestablishment of a general shared pattern to define this period isunrealistic (Bailey, 2008). The reasons behind those changes arestill being debated, with two main driving forces commonlyinvoked as triggering factors. On the one hand, the developmentaltheory argues that the intensification of resource procurement andthe related territorial organizationwas intrinsic to human societies,caused either by demographic pressure pushing populationsbeyond the carrying capacity of the ecosystem (Cohen, 1977) or bythe desire to increase social inter-relationships (Bender, 1979). Onthe other hand, the adaptationist theory claims that certain be-haviours would be more productive and thus more suited withincertain ecological conditions (Rowley-Conwy, 2004).

The Cantabrian Coast, located in the north of the IberianPeninsula, has long been a focus area of such debate. At thebeginning of the Holocene the region underwent dramatic changes

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in the economic, social and territorial organization of hunteregatherers societies. Thus, there was a clear simplification of lithictechnology (Fernández-Tresguerres, 1983; Clark, 1995), a neardisappearance of artistic manifestations (Straus, 1992) and areduction of mobility with an intensification of coastal occupation(Fano, 1996; González Morales, 1999; Straus and González Morales,2003). In terms of economy, the Mesolithic diet broadened toinclude low-ranked prey, such as fast moving or dangerous animals,but in particular marine resources, the exploitation of whichincreased noticeably. In fact, the typical site of that period was theso-called “conchero” or shell midden. They were medium-sizedcaves located no more than 3 km from the present sea-shore con-taining great numbers of limpets and top shells, some animalbones, many quartzite picks and choppers and a limited number ofsmall retouched tools (Vega del Sella, 1923).

Although the consumption of secondary resources had beenincreasing during the Upper Palaeolithic (Álvarez-Fernández, 2011),the Cantabrian Mesolithic showed for the first time a clear strategyof coastal exploitation, including marine molluscs, ocean fish, crabsand sea urchins (Gutierrez-Zugasti et al., 2011). This trend wasaccompanied by a significant decrease in the number of terrestrialmammal remains recovered from the available sites, in particularungulates, which contrasts with the typical strategy of subsistenceduring the Upper Palaeolithic (Fano, 2004; González Morales et al.,

Table 1MNI values of mollusc species in the Cantabrian Pleistocene/Holocene transition.

Site Level MNI marine mollusc

Late MagdalenianOscura de Ania IIIA 2Las Caldas III-0 3Tito Bustillo 1Ae1B 4451Los Canes 2C/3 29La Riera 21e26 713La Pila IV 296Morin 2 9El Pielago IeII 5 8Rascaño 2B 5La Garma A NeO 4168La Fragua 4 355El Perro 2C/3 977El Horno 3e1 23El Miron 106e107 3Goikolau VIeV 187Santimamiñe ALMP þ SLNC 3Antoliña LANC 10Erralla II 2Laminak II II 286AzilianLa Paloma 2 2Oscura de Ania 1e2 6Los Azules 3 ent 12Los Canes 4 1La Riera 27 27e28 8393La Pila III 6525La Lastrilla VII 534La Fragua 3 328El Perro 2A/B 14,227Arenaza III 1Santimamiñe ARCP 4Laminak II I 340Abbitaga VeVI 21Anton Koba VIII 2Ekain VeII 311MesolithicMazaculos 1e3/A3 8322La Llana 1 6704El Penincial e 1621Balmori E1eC1 3208Coberizas B1 3070La Riera 29-30-B 4272Arnero A 255Bricia A 1529Fonfria e 20Vidiago e 124Lledias B 502Los Canes 6e10 627Pozal’egua 2e1 1601La Pila IeII 8827Cofresnedo V0 4El Perro 1 15,242La Fragua 1 11,900La Chora mustr.92 159La Trecha e 1505Kobeaga II AMCK-AMK 2882Santimamiñe H-SLN 36Marizulo IVeIII 246

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2004; Marín-Arroyo and González-Morales, 2009). Adaptationistshave interpreted these changes as the logical response adopted byhunteregatherer groups facing a progressive reforestation and arising sea level, which modified the availability of previously un-wanted resources such as marine shellfish (Bailey, 1983; Arias,1991; Craighead, 1995; Bailey and Craighead, 2003, 2004). How-ever, other researchers have claimed that the intensification ofmarine resource exploitation was a direct consequence of humandemographic growth (Clark and Lerner, 1980; Straus et al., 1980,Straus, 1986; Clark, 1987, 2004; Marín-Arroyo, 2009b). The abun-dant population would then need more resources as terrestrialmammals would not be sufficient to provide them with therequired energy. Ungulates would then suffer some hunting stress(Straus, 1979; Estévez, 2005) and diet would widen to ensure foodprocurement from other sources (Marín-Arroyo and González-Morales, 2009).

Traditionally, intensification in the use of coastal environmentsduring the Cantabrian Mesolithic has been investigated in terms ofthe evolution in shell size (Ortea, 1986; Straus and Clark, 1986;Bailey and Craighead, 2003; González Morales et al., 2004;Gutierrez-Zugasti, 2011) or the effect of sea level rise on the pres-ence and visibility of sites (Bailey, 1978, 1983; González-Morales,1982; Fano, 1996, 2004; Bailey and Craighead, 2004; GarciaMoreno, 2010). However, little attention has been paid to ungu-late faunas, the low abundance of which has prevented an accurateeconomic interpretation. It is nonetheless the scarcity of ungulateremains which makes them so interesting for understanding theeconomic changes that took place during that period, particularlyas ungulates made up the bulk of the diet during the Middle andUpper Palaeolithic.

Given this, a reappraisal of available faunal information,including from recent excavations, has been conducted as a com-plementary study to the numerous available malacological studies.A new zooarchaeological and taphonomic approach has beenapplied with a special focus on the possible identification ofhunting pressure. The results reinforce the developmental theory,as a demographic crisis due to population growth would be morecoherent with the available data.

2. Materials and methods

2.1. The Cantabrian Pleistocene/Holocene transition

The Cantabrian Coast (northern Spain) is a strip of land,approximately 350 km long and 30e50 km wide, located betweenthe Atlantic Ocean to the north and the CantabrianMountains (withpeaks of about 1500e2600 m above sea level) to the south. Thisshort latitudinal distance is covered by fast-flowing rivers that runessentially perpendicular to the coast, carving a series of steepvalleys that to some extent limit the communication routes throughthe region. Whilst in the eastern provinces of Asturias and Canta-bria small coastal plains exist, in the western area of the BasqueCountry the mountain ridges often reach the coastline (Straus,1992). The region benefits from the presence of warm watersbrought by the North Atlantic Drift of the Gulf Stream, which leadsto relatively mild climatic conditions for this northern latitude. As aresult of these conditions, the area holds a rich archaeological re-cord, above all during the Upper Palaeolithic, when it acted as arefugium for European animal and human populations duringsome periods (Consuegra et al., 2002; Achilli et al., 2004; Pereiraet al., 2005; Pardiñas et al., 2012; Meiri et al., 2013).

The Pleistocene/Holocene transition produced notable changesin the life of the Cantabrian hunteregatherer groups. Environ-mental changes such as sea level rise (Mary, 1992), the end of thecold period and subsequent extensive reforestation (García

Moreno, 2007) affected the social and economic organisation ofthe human populations. Flora and fauna became adapted to moretemperate climatic conditions (Harrison and Digerfeldt, 1993)whilst cold-adapted species disappeared (Altuna, 1992). The avail-able zooarchaeological record for this period reveals a shift inhunting preferences, which now included some ungulates scarcelyexploited before, such as wild boar and roe deer, in similar pro-portions to the previously dominating red deer and ibex. However,the most significant dietary transformation of that time was un-doubtedly the intense and unprecedented exploitation of marineresources. Limpets, top shells and mussels started to be consumed

Table 2MNI values of ungulate species in the Cantabrian Pleistocene/Holocene transition.BOS: Bos primigenius/Bison sp.; EQUUS: Equus sp.; CEEL: Cervus elaphus; CPCP:Capreolus capreolus; RATA: Rangifer tarandus; SUSC: Sus scrofa; CPPY: Capra pyr-enaica; RURU: Rupicapra rupicapra.

Sites and levels BOS EQUUS CEEL CPCP RATA SUSC CPPY RURU

Late Magdalenian MNIPaloma 4 1 1 30 1 3 3Sofoxo 3 1 8 4 1 4 3Coimbre 1 4 8 560 31 1 722 57Tito Bustillo Ic 2 3 17 3 1Riera 21e23 3 16 3 2 4Riera 24 2 11 3 2 11 2Castillo 6e9 14 78 114 4 7 23Castillo Dari 3 4 41 3 1Pendo II 3 3 35 1 2 5 1Morin 2 1 3 9 2 1 2 3 3Rascaño 2 1 4 1 1 15Pielago 5e6El Perro 2c 7 6 9 1 1 9La Fragua 4 2 8 3 3 5 1El Horno 1 1 2 4 1El Horno 2 1 4 1 1 12 1El Miron 106 1 6 1 1 9 3El Miron 106.1 4 1 4 2El Miron 12 1 2 2 3El Miron 307 4 1 1 2 3El Miron 308 3 2 2El Miron 103 2 1El Miron 104 1 3 3 1El Miron 104.2 1 1Atxeta E 2 5 1 2Santimamiñe VI 5 7 31 4 1 8 5 5Lezetxiki I 1 1 1 1 1 1Lamiñak II 1 2 7 5 3 3Urtiaga D 3 2 37 13 7 3 20 15Ekain VI 1 2 1 1 4 1Ermitia 1 1 3 1 2 2 13 2Erralla IIIeI 1 1 4 2 1 8 4Azilian MNILa Paloma 2 1 1 11 3 1 1 1Riera 27 2 3 28 6 4 7 2Cueto de la Mina 1 1 9 1 1 1Pendo I 1 1 11 1 1 1 1Morin I 2 1 5 1 1 1Rascaño 1 1 9 1 26 3El Perro 2Ae2B 5 3 6 1 1 5La Fragua 3 1 2 2 1El Miron 11 4 3 1 2 1El Miron 11.1 2 1 4 1El Miron 11.2 2 1 1 1El Miron 305 1 2 1 1 2El Miron 306 1 1 5 3 1 3 2El Miron 102 5 3 1 3 2El Miron 102.1 2 2 1 2 1El Miron 102.2 3 1 1 1Atxeta CeD 1 5 2 2Urtiaga C 16 9 5 3 4Ekain IIeV 3 1 16 5 1 10 3Ermitia 1 3 2 6 1Aitzbitarte IV I 2 2 6 2 1 2 3Mesolithic MNILa Riera 29 5 1 1 1Poza L’Egua 45 4 10 4 1Mazaculos 3 7 12 5 4 1Mazaculos 1e2 9 3 3La Fragua 1 2 2 3 4 3El Perro 1 1El Miron 10.1 2 2 1Cubio Redondo 4 2 3 1 2Urratxa IIIKobeaga II 2 2 2

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widely (Gutierrez-Zugasti, 2011). In fact, the main feature of theCantabrian Mesolithic (ca 8900e4900 cal BP) was the accumula-tion of vast shell middens along the coast in the western part ofCantabrian Spain, establishing the so-called Asturian culture(González-Morales, 1982; Clark, 1983). In the eastern part of Can-tabria, shell accumulations appearmostly within the archaeologicallevels. In both areas, these sites were located along the coastalmargin, no more than 5 km far from the littoral zone (Bailey andCraighead, 2003; Fano, 2004), contrasting with the apparentabandonment of the interior. Seasonality data confirm that some ofthese sites were more than temporary settlements, acting as resi-dential locations where the last hunteregatherer groups livedthroughout the year, exploiting marine and river resources as wellas ungulates in the adjacent coastal plains, as is verified for examplein Mazaculos II Cave (Gonzalez Morales, 1982; Marín-Arroyo andGonzalez-Morales, 2009).

2.2. The zooarchaeological record

In order to investigate the evolution of subsistence strategyduring the Cantabrian Pleistocene/Holocene transition, dataregarding mollusc and ungulate exploitation have been compiled.In the first case, the Minimum Number of Individuals (MNI) hasbeen gathered corresponding to main marine species (bivalves andgastropods) from Gutiérrez-Zugasti (2009), Álvarez-Fernández(2011) and references therein. Secondly, MNI values as defined inKlein and Cruz-Uribe (1984) for the principal ungulate species havebeen retrieved from Marín-Arroyo (2010), Yravedra (2001) andreferences therein. These MNI data are presented in Tables 1 and 2respectively and have been classified in the 3 cultural periods thatprevailed during the Pleistocene/Holocene transition, the so-calledLate Magdalenian, Azilian and Asturian/Mesolithic. Fig. 1 shows thelocations of the studied sites. When available, age profiles, estab-lishing the number of adult and juvenile individuals, have been alsorecorded. Finally, some of the zooarchaeological studies provideskeletal profiles based both on the Number of Identified Specimens(NISP as defined in Payne, 1975) or the Minimum Number of Ele-ments (MNE as defined inMarín-Arroyo, 2009a). Given the need forhomogeneous comparison and the correlation between NISP andMNE indices (Lyman, 2004), NISP values for the followinganatomical parts have been obtained: mandible, vertebrae, ribs,sacrum, scapula, humerus, radiuseulna, carpals, pelvis, meta-carpals, femur, tibia, tarsals, astragalus, calcaneus, metatarsals andphalanges.

2.3. Evidence of hunting pressure

The available zooarchaeological record during the Pleistocene/Holocene transition on the Cantabrian Coast constitutes, in general,an excellent opportunity to evaluate the evolution of the humanstrategy of subsistence, and in particular to verify if there weresigns of an economic crisis. There is some evidence that can beinvoked to do this:

� First, if previously exploited species were depleted, dietarybreath would widen to incorporate new lower-ranked species(MacArthur and Pianka, 1966; Charnov, 1973; Smith, 1983;Waguespack and Surovell, 2003). This could be tested bycalculating average figures for the inverse of Simpson’s Indexand for the high to low-ranked species ratio.

� Second, an imbalance between required food and available re-sources would decrease the age at death of hunted prey. Ingeneral, adult individuals are more productive and thus wouldrank higher than juveniles (Stiner, 1994: 377); If they sufferedfrom overhunting, juvenile individuals would start to be

Fig. 1. Location of archaeological sites considered in this analysis: 1. La Paloma, 2. Oscura de Ania, 3. Las Caldas, 4. Sofoxo, 5. Los Azules, 6. Tito Bustillo, 7. El Penincial, 8. Coberizas, 9.Poza L’Egua, 10. Arnero, 11. Bricia, 12. Cueto de la Mina., 13. La Riera, 14. Lledias, 15. Fonfría, 16. Balmori, 17. Los Canes, 18. La Llana, 19. Coimbre 1, 20. Vidiago, 21. Mazaculos, 22. La Pila,23. Castillo, 24. Pendo, 25. Morín, 26. El Piélago IeII, 27. Rascaño, 28. La Garma A, 29. Cubio Redondo, 30. Cofresnedo, 31. La Chora, 32. La Fragua, 33. El Perro, 34. El Mirón, 35. ElHorno, 36. La Trecha, 37. La Lastrilla, 38. Urratxa III, 39. Arenaza, 40. Atxeta, 41. Antoliña, 42. Santimamiñe, 43. Urtiaga, 44. Kobeaga II, 45. Abbitaga, 46. Laminak II, 47. Goikolau, 48.Ermitia, 49. Lezetxiki, 50. Anton Koba, 51. Ekain, 52. Erralla.

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exploited more intensively (Speth, 1983, 2004; Broughton, 1994,2002; Grayson, 2001; Munro, 2004). Consequently, adult to ju-venile ratios would decrease.

� Third, when resources were scarce in the vicinity of a certain siteand changing residency was not likely to improve the situation,logistic mobility and related catchment areas would have toincrease. As travelling times became longer, differential trans-port of carcasses would intensify in order to optimize the energycontributed to the base camp (Cannon, 2003; Marín-Arroyo,2009a). Munro (2009) and Speth (2004) argue that this wouldresult in lower cranial to post-cranial anatomical ratios in theassemblages. A more detailed analysis can be made bycomparing red deer skeletal profiles in several archaeologicalsites belonging to different cultural periods, with theoreticalprofiles obtained by assuming certain hypotheses of bothcarcass processing at the kill site and post-depositional attrition.The degree of resemblance can be tested by means of a corre-spondence analysis. The possible ways human groups mightprocess carcasses at the kill-site are summarized in Table 3. Onceelements are discarded and deposited at the base camp, theyundergo attritional processes of variable intensity, characterisedby a parameter b that can vary between 0 and 8. Thus, the

Table 3Proposed strategies of carcass processing at the kill-site and transport to the basecamp. P can take value of 0, 0.2, 0.4, 0.6, 0.8 and 1. Head is represented bymandibles;axial skeleton by vertebrae, ribs, pelvis and sacrum; forelimb by scapula, humerus,radius/ulna, metacarpal, carpals and phalanges; and hind limb by femur, tibia,metatarsal, astragalus, calcaneum, tarsals and phalanges.

Strategy Probability Transport to the site

Head Axial Forelimb Hind limb

Complete contribution 1 Yes Yes Yes YesAxial processing at the

kill-siteP Yes Yes Yes Yes(1-P)/2 No No Yes Yes(1-P)/2 No No No Yes

Appendicular processingat kill-site

P Yes Yes Yes Yes(1-P)/2 No Yes No Yes(1-P)/2 No Yes No No

likelihood of survival of one element, in the absence of betterestimates, is obtained as an exponential function of the bonedensity (Rogers, 2000). A value of b ¼ 1 implies that only 50% ofthe bones survive, whereas if b ¼ 2, the survival percentage isreduced to 25%.

� Fourth, if there were a collapse of ungulate populations due tohunting pressure, human groups would be forced to rely onother types of resources, thus reducing the overall intake frommammalian faunas. This can be verified by estimating the evo-lution of average biomass per archaeological level, which can beobtained by multiplying MNI values per edible weight. If thisungulate population collapse provoked a human demographiccrisis, the alternative resources that would have been consumed(molluscs and other marine species in the case of the CantabrianMesolithic) would not be able to substitute the energy lost.

3. Results

3.1. Shellfish versus ungulate productivity

The change in the diet of Cantabrian Late Glacial populations,although notable by the number of remains recovered, is betterdiscerned when a homogeneous comparison is made, and thus,Table 4 shows the relative abundance of ungulate and mollusc in-dividuals in the archaeological record of each cultural period. It isclear, from this, that there was a shift towards a more intenseexploitation of marine resources and a reduction in mammalianfaunal consumption. Whether this trend could also indicate areduction in the energetic procurement obtained from the envi-ronment and therefore a decrease in the potential human popula-tion numbers that could be sustained upon it, would depend on theability of molluscs to substitute the nutrients previously providedby ungulates.

In order to test this, the amount of energy per level associatedwith each type of resource has been calculated (see Table 5) bytaking into account the edible weight of each individual accordingto Garrard (1998) in the case of ungulates and Dupont and Gruet(2002) and Dupont (2003) in the case of molluscs. To transform

Table 4Relative importance of ungulates and molluscs in the human diet as shown by MNIvalues.

Cultural period Ungulates Molluscs

TotalMNI

Totallevels

MNI/level

TotalMNI

Totallevels

MNI/level

Late Magdalenian 2332 32 72.9 11,530 29 397.6Azilian 372 21 18 30,707 25 1228.3Mesolithic 154 10 14.8 72,656 32 2270.5

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this weight into energy, the nutrient database of the US Depart-ment of Agriculture (USDA) provides estimations for rawmeat fromdifferent game prey including the main ungulate species (red deerand ibex) hunted during the Cantabrian Pleistocene/Holocenetransition, these taxa resulting in an average energy content of1328 � 359 kcal/kg, whilst marine molluscs yield 831 � 233 kcal/kg. When the energy provided per level is compared for each cul-tural period, the energy difference between ungulates and marinemolluscs is considerable. It is apparent, at least as far as the avail-able archaeological record is concerned, that the intensification inthe exploitation of littoral biotopes during the Azilian and theMesolithic could not have replaced the consumption of ungulates atthe rates they were hunted during the Upper Palaeolithic and LateMagdalenian. There would then have been an abrupt fall in theenergy obtained from the environment that would have beeninsufficient if human populations remained equal in size.

Gutiérrez-Zugasti (2009) argued that this type of calculationmight be biased towards ungulates, as MNI determination wouldtend to overestimate the real number of hunted individuals.However, although the recovery of one single bone from anarchaeological site would lead to the assumption of all edible partsof the animal being eaten, this scenario is usually rare and normallythe number of bone remains that account for an individual is high(for example in the case of El Mirón Cave the NISP/MNI ratio for theLate Magdalenian levels is 8.4; while in the case of the Mesolithiclevels of Mazaculos the ratio is 16.7), which together with the factthat defleshing at the kill-site is a common procedure, would meanthat the possible overestimation would certainly not explain theaforementioned difference between the energetic procurement ofeach type of resource. Other scholars state that the rise in sea levelmight have hidden many Late Glacial sites (Ortea, 1986; Álvarez-Fernández, 2007; Bicho and Haws, 2008; Gutiérrez-Zugasti,2009), which would mean that the available record is incomplete.While this could be true, the average value per level would not beaffected by this fact, and the conclusions drawn from them can bemaintained.

Nevertheless, there may still be doubts about a possible bias,and therefore, the question of potential productivity should beaddressed in order to understand the implications of the intensi-fication of marine food procurement observed in the Azilian andMesolithic. In this sense, the role of molluscs and other coastalresources had been traditionally regarded as secondary, and char-acterized as low-ranking in energetic yield (Osborn, 1977; Yesner,1987), although their utilization as key resources at times of

Table 5Relative energy contributed by ungulates and molluscs per archaeological level and cult

Cultural period Ungulates

Total weight (kg) kg/level kca

Late Magdalenian 159,899 4998 6,6Azilian 28,340 1350 1,7Mesolithic 11,945 1327 1,7

climatic and environmental change and/or during demographiccrises has been also vindicated (Stiner,1999; Aura et al., 2002, 2009;Stringer et al., 2008). Defenders of Optimal Foraging Theory (OFT)in general and the prey-choicemodel in particular have long arguedthat only when abundance of high-ranking prey (i.e. Pleistocenemegafauna) decreased, due to either overhunting or environmentalchange, would human diet broaden by including less productiveitems (Smith, 1983; Hawkes and O’Connell, 1992; Bird andO’Connell, 2006).

Following the OFT, molluscs cannot be compared with largegame when the overall energy yield needs to be optimised. How-ever, coastal resources can also provide energy together with pro-teins and some fats, although in small packages in comparisonwithterrestrial mammals (Wing and Brown, 1979; Erlandson, 1988).With a steadily increasing number of archaeological examplesshowing diet diversification within scenarios that defy OFT postu-lates (Jones, 2006; Bicho and Haws, 2008; Atici, 2009; Starkovichand Stiner, 2009; Lee, 2011), the OFT framework is challenged. Apossible explanation for this apparent contradiction could be theapplication of new harvesting or processing techniques(Winterhalder and Goland, 1997; Madsen and Schmitt, 1998) andthe adoption of storage or scheduling practices (Hill et al., 1987;Lupo, 2007), which tend to reduce searching and handling timeand therefore to increase productivity. Coastal resource yields caneasily be improved by these kinds of changes. Additionally, theirlocation is much more predictable and their exploitation far easierthan for ungulates, which makes them low-risk resources that canbe harvested by women, children and elderly people (Meehan,1982) and could represent, together with plants and small game,the bulk of the daily diet (Hawkes et al., 1991, 2001). The con-sumption of marine resources might be then regarded as a way ofreducing starvation risks, an economic strategy that can be pursuedby hunteregatherers groups in addition to the maximisation ofenergy intake (Winterhalder and Smith, 1981; Bettinger, 1991;Kelly, 1995; Gremillion, 1996). Finally, coastal resources can alsobe the source of some nutrients such as iron, calcium and Vitamin Dthat are absent or present only in small quantities in terrestrialmammals but that are relevant in maintaining human health(Hockett and Haws, 2003).

As far as average energy content is concerned, the values pro-vided by the USDA confirm the early assumptions of the OFT hy-pothesis, although hunting/foraging costs should be taken intoaccount in order to estimate the final productivity. In this context,the focus will be on red deer as the most represented ungulate inCantabrian mammalian faunas. The average weight of a genericindividual can be established as 93.5 kg (Carranza, 2004), withedible parts (meat plus fat) at 36.7 kg according to Binford’s (1978)observations on caribou. Given the calorific content provided by theUSDA (1110 kcal/kg), the average gross energy related to a red deerindividual will be in the region of 40,778 Kcal.

On the other hand, hunting costs will equal the sum of travel-ling, searching and handling expenditures. Encounter rates (in-dividuals/h) are highly dependent on prey abundance althoughhunters’ territorial knowledge can increase their numbers some-what. However, on average prey abundance can be reasonably

ural period.

Molluscs

l/level Total weight (kg) kg/level kcal/level

35,788 15 0.5 42892,136 40 1.6 132262,477 94 3.0 2444

Table 6Evolution of ungulates consumption along the Cantabrian Pleistocene to Holocenetransition.

Cultural period Inverse ofSimpson’s index

High/lowranked species

Adults/juveniles

Late Magdalenian 1.72 19.0 1.74Azilian 2.13 4.2 1.33Mesolithic 2.37 2.0 1.05

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related to searching times. Based on experimental values calculatedduring extermination events of feral goat populations in NewZealand, Brennan et al. (1993) provide a good mathematical cor-relation that can be applied to the present analysis. But beforesearching time can be obtained, the typical density of red deerpopulations must be estimated. The use of contemporary obser-vations here is problematic as they can be influenced by particularecological characteristics such as climate, the availability of plantresources and the presence of predators. Consequently, the range ofpotential values is large and imprecise. Hence, general experi-mental trends in the form of allometric relationships are bettersuited. Of the available equations, Nowak’s formula (1999) has beenchosen as it appears to better fit modern-day observations of thestudy region, which results in a typical density of 8.05 individuals/km2 and a typical searching time of 1 h. Likewise, handling time,including pursuit, hunting and processing time, for white-taileddeer (Odocoileus virginianus) has been estimated at about 2 h bySimms (1987). Finally, optimal travel time for red deer exploitationhas been estimated by Marín-Arroyo (2009a), based on the appli-cation of Cannon’s (2003) Central Place Forager Prey Choice andMadrigal and Holt’s (2002) experimental data of white tail deerbutchering times, as 2.15 h in each direction.

As a whole, the total expedition time would be around 8 h,which would be consistent with the ethnographic observationsincluded in Binford (2001). Consequently, the productivity of reddeer huntingwould be around 5.000 kcal/h. Obviously these figurescould be reduced considering that not all expeditions are successfuland that each expedition group could be formed by more than onehunter. In addition, human hunting pressure can directly affect preyabundance, thus reducing encounter rates. Marín-Arroyo (2009b)has shown that an exploitation of more than 15% of red deer in-dividuals in the Cantabrian niche could easily lead to their extinc-tion, whereas an exploitation of 8% would reduce their numbers byhalf. However, in the latter case, this would mean that more than300 individuals per year could still be consumed in a typicalhunting territory of 523 km2 (the area being located 2.15 h travel-ling distance from a certain settlement, given an average travellingspeed of 6 km/h), which could sustain a fairly substantial humanpopulation.

In the case of molluscs, there is some ethnographic informationregarding their productivity. For example, Bird et al. (2001) reporton encounter return rates of 1492� 173 kcal/h and 575� 56 kcal/h,for reef flat collecting and rocky shore harvesting of shellfishrespectively, amongMeriampopulations of the Torres Strait Islands.However, Thomas (2007) conducted a study in the atolls ofWesternKiribati, obtaining significantly lower return rates for reef flat andreef crest foraging (72 � 72 kcal/h and 12�6 kcal/h respectively) bytaking into account both searching and handling times. Conse-quently, if total foraging times are considered, these types of re-sources seem to have productivity values an order of magnitudelower than those of ungulates, even if ungulate hunting successrates are noticeably small. In addition, molluscs, being a commonresource usually with unrestricted access, are prone to over-exploitation (Hardin, 1968). In fact, there are several studies thatshow how relatively small-scale human foraging could negativelyaffect the structure of biological communities in intertidal zones(e.g. Okera, 1976; Siegfried, 1994; McLachlan et al., 1996; Castilla,1999). Although it has generally been assumed that marine re-sources are abundant and self-renewing, recent ecological viewsconsiders human foraging as a serious potential threat for thesespecies (Roberts andHawkins,1999;Wolff, 2000). In fact, in order toprevent a collapse of coastal ecosystems, current Cantabrian lawsforbid the extraction of more than 2 kg of shellfish per authorizedperson fromeach tide during the open season. As a result of this typeof policy, shellfishing has become a sustainable economic activity on

the Cantabrian Coast and currently roughly 130,000 kg of shellfishare gathered each year in the provinces of Asturias and Cantabria(Gobierno de Asturias, 2010; Gobierno de Cantabria, 2010), 41% ofwhich are obtained on foot, both in the intertidal zone and on therocky shores (MAGRAMA, 2009). Given the length in kilometres ofthe related coastal region, this means a production ratio of 98 kg/km/yr or 81,438 kcal/km/yr. Returning to the aforementionedtypical hunting territory and assuming an associated coast length of24 km (the quotient between coast length and drainage area in theCantabrian region is 0.046 km�1 according to the Spanish Ministryof Environment), the expected energy extraction from shellfishwould be 1,954,512 kcal/yr if current standards were met. This en-ergy corresponds to the consumption of only 48 red deer in-dividuals. In such a case, there should be no reason to think thatmarine molluscs could have replaced terrestrial mammals duringthe Mesolithic if Palaeolithic human population levels were main-tained. The role of shellfish as a complementary and seasonalresource would then be strengthened.

3.2. Intensification of ungulate exploitation

A decrease in the numbers of hunted ungulates is not evidenceof exploitation pressure but a sign of lower overall energetic pro-curement that, if not complemented by other types of resources,would mean a reduction of the human population as sustained bytheir consumption. The Cantabrian Pleistocene/Holocene transitionis also characterized, however, by a modification in hunting pref-erences, and thus diet diversity widened to include an increasingnumber of species. This diversification, shown in Table 6 by highervalues of the inverse of Simpson’s Index, was due to the inclusion ofpreviously less-frequently hunted animals such as wild boar androe deer and a much lower predominance of red deer and ibex.

This trend has been related in the past to a progressive refor-estation of the region and amore temperate climate that could havevaried the abundance of each species. However, modern ethologicalstudies (Alados and Escós, 2003; Carranza, 2004; Pérez Barberíaand García González, 2004; Mateos Quesada, 2005) state that thehabitats of red deer and roe deer on the one hand, and ibex on theother, often overlap, so that environmental changes would affectthem in the same way. In addition, red deer are more likely to beinfluenced by the amount of food present than by climate itself(Mariezkurrena, 1983; Bugalho and Milne, 2003), while forestsnormally act as a refuge area for this species during light hours(Carranza et al., 1991) providing complementary resources tograzing, such as tender branches and fruits (Rodríguez-Berrocal,1978). However, a shift towards milder weather conditions wouldnot have reduced ibex populations either according to severalbiological studies (Sæter et al., 2002; Jacobson et al., 2004; Grøtanet al., 2008) and it should be noted that its altitudinal range canvary from200 to 3300m a.s.l. (Granados et al., 2002). Consequently,if the abundance of red deer and ibex was not modified by themilder Holocene climate, both these species comprising the bulk ofhuman diet during Late Pleistocene, there is no reasonwhy huntingpreference would have shifted towards the consumption of moredangerous or less productive animals. Nonetheless, as shown inTable 6, the quotient between high and low ranked species (the

Fig. 2. Correspondence analysis between recorded and theoretical skeletal profiles for red deer. Axis 1 accounts for the transport strategy with more appendicular processing at thekill-site to the right. Axis 2 accounts for the degree of attrition with less conservation to the top. Late Magdalenian sites: 1. Paloma 4, 2. La Riera 21e23, 3. La Riera 24, 4. Morín 2, 5.Santimamiñe VI, 6. Lamiñak II, 7. Urtiaga D-E, 8. La Fragua 4, 9. El Mirón 104, 10. El Mirón 106. Azilian sites: 11. Paloma 2, 12. La Riera 27, 13. El Perro, 14. Ekain IIIeV, 15. El Mirón 11,16. El Mirón 306, 17. El Mirón 102. Mesolithic sites: 18. La Riera 29, 19. Cubio Redondo, 20. Mazaculos 3. 21. Marizulo III.

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latter represented by roe deer and wild boar) decreased sharplywith the arrival of the Holocene, a fact that could only be explainedby an overexploitation of high-ranked prey due to demographicpressure (Marín-Arroyo, 2009a,b).

Interestingly enough, human groups not only started to huntlow productivity species on a large scale but also to consume lowproductivity individuals within a species. Table 6 presents thechange in the adult to juvenile ratio taken during the periodstudied, showing a tendency to consumemore young individuals inthe Azilian and Mesolithic, which not only would mean that theaveragemeat weight consumedwould have decreased but also thatthe very survivorship of ungulate populations would have beenthreatened as less individuals reached reproductive age. This un-sustainable strategy is a clear sign of hunting pressure and re-inforces the hypothesis of the existence of a demographic crisis.

3.3. Expansion of hunting territories

A final piece of evidence for the occurrence of overhuntingduring the Cantabrian Late Glacial comes from the expansion ofcatchment areas. It is clear that lower prey abundances around acertain settlement due to a previous intense exploitation wouldforce human groups to travel longer distances to hunt their prey. Inorder to optimize the amount of energy contributed to the basecamp (Cannon, 2003; Marín-Arroyo, 2009a) this would lead tohigher carcass processing at the kill-site and differential transportof anatomical parts. For example, Speth (2004) and Munro (2009)argue that larger hunting territories would reduce the proportionof cranial bone remains at the residential sites due to the lowerproductivity of that part of the carcass. This hypothesis is confirmedin the Cantabrian Late Glacial zooarchaeological records, with acranial to post-cranial ratio decreasing from 0.07 during the LateMagdalenian to 0.06 in the Azilian and 0.05 in the Mesolithic.

A more integral approach is presented in Fig. 2 based on theskeletal profiles of red deer available in the literature (MNE values

were rarely obtained; NISP values were used instead given the highcorrelation existing between these indices according to Lyman,2004). The transport strategy carried out for each site and cul-tural period is discriminated by extracting the potential attritionand comparing the original anatomical representation with sometheoretical strategies.

As can be seen, there is some variation that might be related tosome extent to the specific topographic location of each site and therecovery procedures and analytical methods of quantification.However, Late Magdalenian sites tend in general to be character-ized by a nearly complete carcass contribution or by a moderateappendicular processing at the kill-site. This would be consistentwith fairly small catchment areas and short travelling times. AzilianandMesolithic sites, in turn, have awider dispersion, some of themwith an intense processing at the kill-site either of the axial orappendicular skeleton. This pattern would be the result of an in-crease in travelling times, forcing hunters to reduce load weightsbefore returning to the base camp in order to optimize the ener-getic yield. This scenario is in agreement with a decrease in un-gulate encounter rates due to lower animal population densities.

4. Discussion

Since Binford (1968) and Flannery (1969) placed the origin ofNeolithic behaviour in the diversification of human diets and pro-curement strategies that had occurred at the end of the Pleistocenedue to unprecedented demographic crowding in certain regions ofthe world, the so-called developmental theory spread amongstarchaeological researchers (Cohen, 1977; Keeley, 1988; Bar-Yosefand Meadow, 1995; Stiner et al., 1999). On the Cantabrian Coast,in the north of the Iberian Peninsula, the diversification in subsis-tence observed in Late Upper Palaeolithic sites, with an increasinguse of marine resources (estuarine and rocky shore molluscs, fish,urchins and crabs) was also seen by Straus (1979) as a result ofpopulation pressure. Based on the archaeological record of La Riera,

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as well as other Cantabrian sites, Clark and Straus (1986) furtherdeveloped this model by including intensification in the con-sumption of terrestrial faunas and a significant exploitation ofbirds. For them, these economic changes were clearly not linked toclimatic variations, apart from the increase of wild boar and roedeer, but could also have been caused by overexploitation of pre-viously hunted prey, and a decrease in Littorina littorea consump-tion (Straus, 1992; Clark, 2004).

As a counter argument, adaptationist theories have been alsobeen invoked to explain the Cantabrian Pleistocene to Holocenetransition. Remarkably, Bailey (1978, 1983) has been continuouslyarguing that climatic shifts and variations in sea level could havebeen plausible causes of the changes observed in the subsistenceeconomy. Craighead (1995) came to support this view too, basednot only on information from Patella vulgata but also from themammal faunas of La Riera. Both authors have claimed that the sortof evidence normally used to support demographic pressure as thelikely explanation (i.e, changes in limpet size andmortality profiles)was in fact better correlated to the effect of sea level rise and cli-matic oscillations on the habitats of mollusc communities (Baileyand Craighead, 2003, 2004, but see responses in; GonzálezMorales et al., 2004; Gutierrez-Zugasti, 2011). Bailey (2004a,2004b) even emphasized the potential of coastal environments ashighly productive areas with abundant and diverse kinds of re-sources, being suitable to support large human populations.Furthermore, the exploitation of coastal environments during theIberian Upper Palaeolithic, along with small terrestrial game (i.e.rabbits) and plant resources, could have been underestimated inthe earlier Palaeolithic, the evidence now being inaccessible asmany existing coastal sites become submerged (Ortea, 1986;Álvarez-Fernández, 2007; Bicho and Haws, 2008; Gutiérrez-Zugasti, 2009; Bicho et al., 2010).

Most of the previous debate has made use of marine data ingeneral, and limpet sizes in particular, as the main focus of analysis.Surprisingly enough, the use of terrestrial faunas in analysis, whichwere widely studied during the Palaeolithic period, becomesomewhat reduced in Mesolithic studies. This is easily under-standable given the overwhelming presence of shell middens inCantabrian Mesolithic sites, a remarkable sign of the intensificationin the exploitation of littoral habitats (González Morales et al.,2004). However, terrestrial faunas, scarce as they are, can stillprovide valuable information on the likely cause of the observedeconomic shift. In this sense, the results of the present analysis haveshown that:

1. The Pleistocene to Holocene transition was characterized on theCantabrian Coast by a progressive widening of human dietbreadth with the inclusion of less productive ungulates, con-sumption of which could only be explained if the populationdensity of previously-hunted species, mainly red deer and ibex,had decreased significantly.

2. The intensification in the exploitation of littoral environmentsseen during the Azilian and the Mesolithic should not beregarded as an alternative strategy of subsistence if humanpopulations remained constant. The amount of energy providedby marine resources is not sufficient to account for the decreasein the exploitation of mammalian faunas, at least as far as theavailable archaeological record is concerned. In addition, overallproductivity of molluscs cannot be compared to that of un-gulates, being an order of magnitude lower.

3. There is clear evidence of ungulate overhunting during theAzilian and Mesolithic such as the increase in the consumptionof juvenile individuals, the noticeable reduction of overall foodprocurement provided by ungulates, and the enlargement ofcatchment areas as available resources became depleted.

Given this, the increasing importance of marine resourcesderived from the archaeological record at the end of the Pleistocenecould potentially be considered as having been due to the exploi-tation of secondary, more predictable resources if not for the dra-matic fall in the number of hunted ungulates. This, along with thedecrease in the total energy extracted from the environment, pointstowards the occurrence of a demographic crisis, as the humanpopulation that existed in the Late Magdalenian could no longerhave been sustained. Nonetheless, Straus (1979, 1992) doubted thatthis ungulate under-representation was a real reflection of thestrategy of subsistence of that time, arguing that shell middenswere only part of the archaeological record left by hunteregatherergroups. Based on ethnographic observations, Straus claimed thatintensive shellfish collection could have only been a winter com-plement when other resources such as plants and ungulates werelimited, and that open-air camps would have been discovered hadevidence of them not been lost over time. However, studies on theintentionality of site location (Fano, 1998), the limited success infinding other types of settlements despite intensive research con-ducted in the last three decades in the Cantabrian area, and therelatively limited sea level change from that of Late Upper Palae-olithic times (Bailey and Craighead, 2004) have progressivelyweakened this hypothesis. In this scenario, the limited exploitationof ungulates should be regarded as a typical trend of the Mesolithiceconomy, and therefore, interpreted accordingly.

Recently, Estévez (2005) and Marín-Arroyo and González-Morales (2009) have suggested the possibility that a rapiddecrease in temperatures during the Preboreal oscillations, causedby the sudden interruption of the North Atlantic oceanic circulationfrom 11 to 8 ka BP (Clark et al., 2001; Teller et al., 2002), might haveprovoked a catastrophic reduction of terrestrial resources thatcould have dramatically affected the Cantabrian human pop-ulations. Based on pollen reconstructions, Davis et al. (2003) esti-mated that, for example, the 8.2 ka BP event could have caused adecrease in annual temperature for south-western Europe of about2 �C. As proposed by Marín-Arroyo (2009b), this kind of climaticimbalance could easily have provoked the collapse of ungulatepopulations that were simultaneously suffering from a significantlevel of hunting. In fact, a succession of several harsh winters withheavy snow might be enough to lead to the extinction of the reddeer populations in a particular area, as occurred in the Erregerenamountains (Navarra) in 1979 (Mariezkurrena, 1983). This kind ofrapid resource depletion would have forced human groups tomigrate to other less affected areas in order to survive.

In summary, a demographic crisis could explain the abrupteconomic and technological changes detected in the region duringthe Pleistocene to Holocene transition more effectively than theadaptationist postulates, which may be coherent with an increasein the exploitation of marine environments as complementary re-sources but clearly fail to address the issue of the decrease in un-gulate consumption.

Acknowledgements

This research has been funded by the Ministerio de Economía yCompetitividad with a Ramon y Cajal Research contract (RYC-2011-00695) at the Universidad de Cantabria. I would like to thank MarkLewis for his comments and for reviewing the English of thismanuscript. I am also grateful to David Ocio for his kindly support.

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