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Environmental ArchaeologyThe Journal of Human Palaeoecology

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The utility of livestock dung for reconstructingrecent ethnological and environmental histories

Xavier Panadès i Blas, Jordi Bartolomé i Filella, Caroline Strömberg, IgnasiSoriano i Tomàs, Paul Buckland, Karen K. Serieyssol, Joan Bach i Plaza,Antonio Arillo Aranda, Francesca Lozar, Tony Stevenson, Lourdes Chamorro iLorenzo & Peter Ditchfield

To cite this article: Xavier Panadès i Blas, Jordi Bartolomé i Filella, Caroline Strömberg, IgnasiSoriano i Tomàs, Paul Buckland, Karen K. Serieyssol, Joan Bach i Plaza, Antonio Arillo Aranda,Francesca Lozar, Tony Stevenson, Lourdes Chamorro i Lorenzo & Peter Ditchfield (2016): Theutility of livestock dung for reconstructing recent ethnological and environmental histories,Environmental Archaeology

To link to this article: http://dx.doi.org/10.1080/14614103.2016.1142630

Published online: 31 Mar 2016.

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The utility of livestock dung for reconstructingrecent ethnological and environmentalhistoriesXavier Panadès i Blas1 , Jordi Bartolomé i Filella2 , Caroline Strömberg3,Ignasi Soriano i Tomàs4 , Paul Buckland5, Karen K. Serieyssol6, Joan Bach iPlaza7, Antonio Arillo Aranda8 , Francesca Lozar9, Tony Stevenson10, LourdesChamorro i Lorenzo4 , Peter Ditchfield11

1Departamento de Ecología, Genética y Microbiología, Facultad de Ciencias, Biológicas y Ambientales,Universidad de León, León, Spain, 2Departament de Ciència Animal i dels Aliments, Universitat Autònoma deBarcelona, Bellaterra, Barcelona, Catalonia, 3Department of Biology and Burke Museum of Natural History &Culture, University of Washington, Seattle, WA, USA, 4Departament de Biologia Vegetal, Universitat deBarcelona, Barcelona, Catalonia, 5BugsCEP, 20 Den Bank Close, Crosspool, Sheffield, S10 5PA, UK, 6DiatomResearch, Hery, France, 7Departament de Geologia, Edifici Ciències (Cs), Universitat Autònoma de Barcelona,Bellaterra, Barcelona, Catalonia, 8Departamento de Zoología y Antropología Física, Facultad de Biología,Universidad Complutense, Madrid, Spain, 9Dipartimento di Scienze della Terra, Università degli Studi diTorino, Torino, Italy, 10School of Geography, Politics and Sociology, Faculty of Humanities and Social Sciences,Newcastle University, UK, 11Research Laboratory for Archaeology and the History of Art, University of Oxford,Oxford, UK

Animal dung is evaluated here as a tool to reconstruct recent societal and environmental changes. Studiescompleted on the macro- and micro-contents from dung deposited in a mountain cave in Catalonia during the1970–1980s, preceding the socio-economic changes in the area, was supplemented by the testimony of thelast shepherd alive in the area. This information was also compared with evidence from aerial photographs ofthe area surrounding the cave taken in 1956 and 2009. Although taphonomic distortion of the dung shapeprecluded its assignment to its producers based on morphology, information from the organic remainsreflects the agro-pastoral activities in the mountainous area of the Catalan coast. A lack of calciumphosphate and abundant phytolith and epidermal remains from wild species indicates that the dung wasexcreted by ovicaprines, who sheltered in the cave during spells of adverse weather. These indicatorsalso signify that the animals fed only on natural vegetation. Insect remains in the dung include fragmentsof chafers, dung and ground beetles, bees and ants, and whilst the latter may have foraged in the cave,the others may have been accidentally ingested by herbivores. The nannofossils and diatoms found in thedung derive from tertiary crops 4 km away from the cave but within the range of the ovicaprine herd. Thelack of cereal and vegetable remains in the dung is consistent with the ethnographic record thatshepherds and farmers could not afford to supplement the herds’ diet with fodder crops or grain, aconsequence of the low productivity of the local subsistence agriculture. This study further discusses thesocio-economic impacts of the introduction of intensive farming, industrialisation and tourism in Cataloniaduring the latter part of the last century and how they are represented in the dung.

Keywords: Animal dung, Phytoliths, Diatoms, Insects, Karst, Catalonia

IntroductionWhere sufficiently well preserved, coprolites (fossilisedanimal dung) have been used successfully for archaeo-logical and vegetational reconstructions (e.g., Cabaneset al. 2009; Carrión et al. 2001, 2005; Charles 1998;González-Sampétriz et al. 2003; Scott 1987). This is

because animal and plant micro- and macro-remains,having passed through an animal’s gut, may surviveas a consequence of natural desiccation, partial miner-alisation, or both. The resultant coprolites may berepresentative of the prey species, local and regionalvegetation and agriculture (Linseele et al. 2010,2013; Wallace and Charles 2013). For example, arecent study, using seeds and pollen from dungCorrespondence to: Xavier Panadès i Blas, Flat 6, 35, Richmond Rd, KD6

1BX, Leeds, UK. E-mail: xpan100@gmail.com

© Association for Environmental Archaeology 2016DOI 10.1080/14614103.2016.1142630 Environmental Archaeology 2016 1

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excreted by modern cattle and sheep, provided a faith-ful representation of the local vegetation (Schepers andVan Haaster 2015). The subsistence economy in theNegev Highlands during the Iron Age, Byzantineand Early Islamic period has been elucidated fromthe macro- and micro-plant remains preserved fromthe dung of free-grazing livestock (Shahack-Grosset al. 2014). Pollen has also been extracted from carni-vore dung (Scott et al. 2003) and the composition ofthe late Pleistocene–Holocene vegetation of theCentral Zagros mountains has been reconstructedusing pollen from hyaena dung from Wezmeh cave inwestern Iran (Djamali et al. 2011).Despite this, the utility of animal dung as a tool to

reconstruct ethnological histories and environmentalchanges requires further evaluation. Such studieshave the potential to document the environmentaland social impacts, and vanishing ethnoarchaeologicalinformation from agro-pastoral regions, often as aconsequence of urbanisation since the middle of thelast century. Knowing the previous composition ofthe forest vegetation and the land-use of agro-pastoralsocieties would also help in the implementation ofcurrent managing and reforestation practices, aimedat preventing the devastating summer fires that regu-larly occur in Mediterranean areas (Moreno andOechel 2011; Trabaud and Lepart 1980; Whelan1995). In addition, comparing recent ethnographicrecords with coprolite remains can provide a blindtest for reconstruction of earlier periods.The recent discovery of animal dung in a cave in the

El Montmell area in the highlands of the Catalan coastprovides an opportunity to study recent environmentand social changes in the area (Fig. 1). The dunghad been deposited by livestock sheltering in the

cave. The contents are well preserved and the formeragro-pastoral practices and changes in vegetation areknown, based on the testimony of the last shepherdalive in the area, supplemented by aerial photographstaken between 1956 and 2009 (Figs. 2 and 3). This isan exceptional discovery because well-preservedanimal dung is rarely found in temperate/semi-aridregions, whilst dung may be the only source ofpalaeoecological information in arid environments(Carrión et al. 2001; Djamali et al. 2011; Scott 1987).

BackgroundDung rapidly decomposes under the warm and moistconditions of most temperate/arid environments butmay be recovered from cave and rock shelter sites,where stable, cooler conditions retard decompositionand facilitate the preservation of macro- and microfos-sil remains in the dung (Delhon et al. 2008; Mercuriet al. 2010; Shahack-Gross 2011). In Iberia, only afew unburnt dung remains have been unearthed incave deposits (Cabanes et al. 2009; Carrión et al.2001; González-Sampétriz et al., 2003). The Frenchterm fumier, referring to mixtures of dung andanimal litter, is also applied to sedimentary layers ofburnt or otherwise preserved dung in caves (e.g.,Allué et al. 2009; Badal 1999; Bergadà 1998). Theselayers have provided substantial information aboutpast societies, palaeoenvironment, palaeoclimate andthe diet of animals during the Quaternary. Forexample, dung, surface soil and sediments depositedin a cave in southeast Iberia have been used to recon-struct local and regional vegetation and the studyshowed that the animal dung yielded the highest con-centration and diversity of pollen taxa (Carrión 1999a,1999b; Carrión 2002; Carrión et al. 2005).

Carrión et al. (2007) provide reconstructions of thePleistocene landscapes of Central Iberia from pollenidentified in hyaena dung, which indicated a combi-nation of steppe-grassland areas with Asteraceae andChenopodiaceae, and wooded areas composed ofPinus and Juniperus species. The authors also detectedmixed oak forests, which were interpreted as refugia oftemperate and Mediterranean vegetation. In a similarstudy, hyaena coprolites have been used as part of amulti-proxy approach to reconstruct the glacial andlate glacial landscapes of northeast Iberia (González-Sampétriz et al. 2003). These identified steppe,Mediterranean shrubs, coniferous forest and refugesof meso-thermophilous taxa, which characterised thevaried climatic phases. However, it was concludedthat a multi-proxy approach, including pollen analysessourced from different traps, would provide moreaccurate reconstructions of past vegetation.

Similarly, Carrión et al. (2006) have explored thepotential of bat droppings (guano) as a tool forpalaeo-landscape reconstruction. They compared

Figure 1 Physical map of Catalonia placing the cave wherethe dung was uncovered.

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Figure 2 Aerial photo of the Urbanisation La Moixeta cave taken in 2009.

Figure 3 Aerial photo of the Urbanisation La Moixeta cave taken in 1956.

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the pollen spectra of surfaces with modern dungsamples from nine caves in southeastern Iberia andconcluded that guano could be a concise tool forenvironmental reconstructions, because it may effec-tively preserve pollen, with levels of taxon diversityand preservation superior to surface samples fromcaves.In the Balearic archipelago, diet and the causes of

the extinction of the cave goat, Myotragus balearicusBate, have been inferred from the contents of theirdung found in caves. DNA, pollen and macrofossilsdepict the animal as a mixed feeder, greatly dependenton box shrub (Buxus balearica) (Alcover et al. 1999;Bartolomé et al. 2011; Welker et al. 2014). Severalcauses seem to account for its extinction. Theseinclude increasing aridity changed the vegetation toopen maquis from the original mesophyllous cover ina short period of time, which induced the dramaticdecline of several species and in particular of box.The goat’s limited agility and lack of awareness to anew predator may also have made them easy preyfor the first humans that reached the archipelago.Being such a specialised feeder, it is believed thatthey could not adapt to such a rapid change in veg-etation (Alcover et al. 1999; Bartolomé et al. 2011;Welker et al. 2014). The climatic trend during thelate Holocene in the Balearic archipelago has beeninferred using stomatal density and indices of epider-mal remains of Buxus identified in the dung of thecave goat (González-Sampériz et al. 2003; Riveraet al. 2014). Stomata density can be directly associatedwith CO2 levels (Woodward 1987) which show that bythe Late Holocene CO2 levels were rising in theBalearic archipelago (Rivera et al. 2014).The Bronze Age environment and agricultural

practices have been reconstructed from pollen, phyto-lith and faecal spherulites from a fumier uncovered ina cave in the karst landscape of Atapuerca in centralIberia (Cabanes et al. 2009). Phytolith and faecalspherulites remains depict a range of land uses,including forest, grazing pastures and cultivatedfields, and the use of the cave for crop-processingand other agricultural activities. A pollen analyticalstudy of more recent badger (Meles meles) dung byCarrión et al. (2005) did not reveal any changes inthe anthropogenically modified landscape ofCentral Iberia over the last 300 years. The residuefrom a carnivore diet contained within the dung didnot bias their results.This paper attempts to test the utility of animal dung

as a tool to reconstruct recent environmental changesand human activities in the El Montmell region.This is achieved by comparing the information onagro-pastoral and vegetation composition providedby the dung remains, against contextual evidence,the shepherd’s testimony and aerial photographs.

MethodsThe samples examined were recovered from a cave inLa Moixeta in the El Montmell mountains, part ofthe Catalan Pre-coastal range, at an altitude of450 m a.s.l. (41°21′28.55′′N, 1°31′18.19′′E). The sub-strate of the Montmell range is a mixture of limestonesand other calcareous rocks ranging in age from theJurassic to the Miocene (Fig. 4). It has a graduallyundulating relief with small cliffs, canyons and numer-ous caves (Figs. 2 and 3) (Casas-Sainz and De Vicente2009; Enciclopèdia Catalana 2013; Masana 1994;Riba et al. 1976). The vegetation biome surroundingthe cave is Mediterranean, with some Euro-Siberianelements present on north-facing slopes and borderingwatercourses (Tables 1 and 2, Fig. 5). The dominantvegetation is composed of evergreen oak (Quercusilex) woodland with laurustinus (Viburnum tinus)forming a dense, richly bushy sclerophyllous under-storey and restricted growth of deciduous shrubby veg-etation on the stream’s banks, including olive willow(Salix elaeagnos subsp. angustifolia) and thickets ofMediterranean coriaria (Coriaria myrtifolia) andbramble (Rubus ulmifolius) (Tables 1 and 2, Fig. 5)(Quadrada 2007; Rovira i Gómez 1982). In therecent past, much of this would have been reduced

Figure 4 Geological map of the surroundings of the cave.Geologic map scale1:50·000, sheet number 419 (InstitutoGeológico y Minero de España, 2006). Legend: 1 – Triassic:marls–limestone: 2 – Jurassic–Early Cretaceous: sandstonesand clays, and clays and limestones; 3 – Lower Cretaceous:limestones and dolomite intercalations; 4 – Lower–UpperCretaceous: calcarenites, marls, clays; and sandstones; 5 –

Palaeogene–Palaeocene: clays; 6 – Palaeogene–Eocene:limestone and marls; 7 – Neogene–Miocene: conglomerates;8 – Quaternary: coarse sediments, gravel; 9 – Faults andmechanical contacts; 10 –Moixeta Cave; 11 –Boundary of theriver basin of Marmellar tributary; 12 – Drainage basin of theMarmellar tributary, as it is seen from the bridge.

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by grazing, creating an evergreen oak-dominatedwood pasture.The cave occurs alongside a gorge cut by a tributary

stream, the Marmellar. It is situated 15–20 m above

Table 2 List of vegetation identified along the Riera deMarmellar river bank (willow shrubby formations+Rubusulmifolius thicket + hygrophile rush and tall grassformations). C. 400 m a.s.l.

TreesFraxinus angustifolia

ShrubsSalix elaeagnos*Rubus ulmifolius*Coriaria myrtifoliaBupleurum fruticosumRosa caninaCrataegus monogynaPrunus spinosa

HerbsRumex obtusifolius*Mentha rotundifoliaPolygonum persicariaJuncus inflexusDipsacus fullonumCirsium monspelienseScirpus holoschoenusPrunella vulgaris

*means abundant species.

Figure 5 Habitats map of La Moixeta at 1:25·000 scale (Abriland Vallejo 1991; Cambra et al. 2006; Quadrada 2007). LaMoixeta Habitat map. Legend: 1 – Lowland holm-oak forests;2 – Aleppo pine (P. halepensis) forests with a thermo-Mediterranean woody understory; 3 – Aleppo pine (P.halepensis) forests with a calcicolous scrub layer; 4 – Kermesoak (Q. coccifera) garrigues, locally with Aleppo pineregeneration; 5 – Rosemary (Rosmarinus officinalis) scrubs,locally with Aleppo pine regeneration; 6 – Dry grasslandsusually dominated by Brachypodium phoenicoides; 7 – Riverbank vegetation complex: willow shrubby formations+Rubus ulmifolius thicket+ hygrophilous rush and tall grassformations; 8 – Vineyards; 9 –Urbanised areaswith patches ofnatural vegetation.

Table 1 Vegetation of La Moixeta relief: 1 Quercus ilex –

Q. faginea maquis (Viburno tini-Quercetum ilicis); 2 Aleppopine forest on Mediterranean scrub; 3 Kermes oak garrigue(Quercetum cocciferae)

[1] [2] [3]

Aspect NW W SSEInclination (°) 25 5 15Altitude (m) 440 430 460Total cover (%) 95 90 95Tree cover (%) – 60 5Tree height (m) 3 –

Shrub cover (%) – 50 95Shrub height (m) 2 1 1,5Herb cover (%) – 40 60Area (m2) 60 150 60Quercetea ilicis characteristicsQ. ilex 2.2 1.1 +Q. coccifera + 2.2 5.4Phillyrea latifolia 1.1 + +Rubia peregrina + + +Pistacia lentiscus + 1.1Asparagus acutifolius + +Lonicera implexa + +Rhamnus alaternus + 1.1Viburnum tinus 2.2Ruscus aculeatus +Smilax aspera +Viola alba +Clematis flammula +Querco-Fagetea characteristicsQ, faginea 4.3 2.2Buxus sempervirens 3.3Coronilla emerus 1.1Anemone hepatica +0.3 .Euphorbia amygdaloides +Rosmarinetea characteristicsRosmarinus officinalis 1.2 +Erica multiflora + +Bupleurum fruticescens +Stipa offneri +.2Argyrolobium zanonii +Euphorbia nicaeensis +Other speciesP. halepensis (+) 4.3 1.1Brachypodium retusum 2.2 2.3Bupleurum fruticosum 1.3 +Carex halleriana +0.2 +0.2Juniperus communis 1.2 +Juniperus oxycedrus + +Teucrium chamaedrys + +Ulex parviflorus 1.2 +Brachypodium phoenicoides 2.3Genista scorpius 1.1

Other species present only in one rélevéRel. 1: Aphyllanthes monspeliensis, Campanula sp., Carex gr.ornithopoda, Genista hispanica subsp. hispanica, Genistapatens, Hedera helix, Helleborus foetidus.Rel. 2: Carex flacca, Cistus albidus, Cistus salviifolius, Clematisvitalba, Conopodium majus subsp. ramosum, Crataegusmonogyna, Dorycnium hirsutum, Knautia sp., Lavandulalatifolia, Lonicera etrusca, Ranunculus sp., Rubus ulmifolius.Rel. 3: Centaurea linifolia, Dorycnium pentaphyllum, Fumanaericifolia, Galium lucidum, Ononis minutissima, Polygalarupestris, Rhamnus cf saxatilis, Sedum sediforme, Thymusvulgaris.Relief code: Country, county, community (Province), locality,UTM, survey date, authors.1 [2-010306]: El Montmell (Tarragona), La Moixeta, 31TCF37667983, 01/03/2006, I. Soriano, L. Chamorro, L. José-María2 [3-010306]: El Montmell (Tarragona), La Moixeta, 31TCF76417965, 01/03/2006, I. Soriano, L. Chamorro, L. José-María3 [1-010306]: El Montmell (Tarragona), La Moixeta, 31TCF76437977, 01/03/2006, I. Soriano, L. Chamorro, L. José-María

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the bank of the stream, where the bridge connects theroad towards La Moixeta. The cave has an outerchamber with a narrow, deep internal chamber andit is largely infilled with sandy sediment. Extensivedeposits largely of animal dung were found partlyburied in the surface sediments of both chambersand similar material occurred in small cavities in theroof of the cave, where several abandoned bird nestswere present.

The ethnographic and historical contextA few Pyrenean farming houses controlled agro-pas-toral activities in Catalonia and the resident ownersemployed low-paid sharecroppers and shepherds.Pastoralism was a large-scale enterprise that exerteda firm control over the economy and pastoralistswere the sole providers of meat for the main cities.In fact, butchers from the main cities in the southhad to travel to the north to buy livestock, whichwould be delivered during winter time as part of atranshumance cycle. The area sustained twice thecurrent population until the beginning of the 21stcentury, when transhumance in Catalonia finally col-lapsed (Llobet and Vilà Valentí 1951; Miralles andTutusaus 2005; Vilá Valentí 1973). The agro-pastoraleconomy of the mountainous area of the Catalancoast was substantially different from the Pyreneaneconomy. According to the nonagenarian shepherdAlfons Guasch i Martí, the harsh climatic conditionsof the El Montmell mountains, only allowed the devel-opment of what is essentially subsistence-based agri-culture, with small crops of barley and wheat,supplemented with a few vegetables, such as potatoes.Small farms only stocked a few pigs, chickens, a smallovicaprine flock and occasionally a mule, donkey or ahorse, but never cattle.Against this background, all resources were heavily

utilised. Dung from different livestock might beemployed selectively (cf. Charles 1998; Linseele et al.2013; Shahack-Gross 2011; Wallace and Charles2013). Whilst dung from ovicaprines might be burntas fuel or utilised for other purposes, Alfons confirmedthat in El Montmell area, ovicaprine, human, pig andavian dung was only used to fertilise horticultural andcereal crops and occasionally pastures. He pointed outthat burning dung was unnecessary owing to the greatabundance of wood in the area. However, he high-lighted that smoke from fires made from horse andmule dung had a specialist use: for the collection ofhoney. Smoke makes the bees docile to ensure thatthe honey was collected safely.Industrialisation, changes in agriculture practices

and the boom in tourism caused the demise of transhu-mance in Catalonia between 1920 and 1960 (Llobetand Vilà Valentí 1951; Miralles and Tutusaus 2005;Vilá Valentí 1973). The introduction of intensive

farming and the augmentation of irrigation systemsgenerated extra costs that the smaller farms couldnot absorb. It drastically reduced the grazing groundand consequently it raised the price of winter forageand reduced herd numbers. More intensive agriculturemeant that the herds were no longer needed to fertilisethe fields (Llobet and Vilà Valentí 1951; Miralles andTutusaus 2005; Vilá Valentí 1973). Industrialisationhad a more detrimental impact on the farms. Themassive growth of cities from 1920 onwards and thetourist boom of the 1960s created a high demand foremployment, which offered much better economicbenefits when compared to the farming houses. Thefinal development leading to the demise of transhu-mance was the construction of hydroelectric stationsin the Pyrenean region, stemming from the increasingenergy demands of the incipient industrialisation andthe expansion of urban areas. This attracted theremaining manpower available in the farming areasand resulted in a drastic depopulation of thePyrenees from which the region never recovered(Llobet and Vilà Valentí 1951; Miralles andTutusaus 2005; Vilá Valentí 1973). Furthermore, thewool industry also collapsed, owing to cheaperimports and more efficient production methods thatwere achieved using modern technology.

The disappearance of a pastoral herd-basedeconomy and the sharp depopulation of the centralmountainous area of the Catalan coast differed tothe changes evident in the Pyrenean region (Llobetand Vilà Valentí 1951; Miralles and Tutusaus 2005;Vilá Valentí 1973). El Montmell was a typical high-lands settlement in the central area of the Catalancoast. According to the former shepherd AlfonsGuasch i Martí, before small-scale urbanisation, thesettlement comprised a nucleus with scattered farm-houses in the surrounding mountains. Larger farmshad usually a resident owner who rented out smallerfarms to sharecroppers and the caves to shepherds.Owners and tenants also employed temporaryworkers at the busiest times of the year. It was oneof the main local centres of transhumance, takinglarge flocks of sheep from the lowlands to summerpasture in the uplands, and its economy was mainlybased on these herding activities, supplemented bysubsistence agriculture and forestry.

Unlike in the Pyrenees, Alfons noted that on theCatalan coast, subsistence and sedentary agro-pastoralactivities and exploitation of the forest only allowedthe subsistence of relatively small populations.However, socio-economic activities of the villagewere impermeable to changes occurring elsewhere inCatalonia until the 1940s. People in El Montmellhad a comfortable socio-economic standard andthere was less competition for local resources. Localswould cooperate and that jobs were fairly organised

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and local resources were shared amongst the localpopulations. Consequently, local populations werenot attracted by the opportunities offered by earlyindustrialisation. In fact, the population of ElMontmell remained at 529 inhabitants until the endof the 1950s (Institut d’Estadística de Catalunya2013). Alfons believes that the decline in subsistenceagro-pastoral and forest activities in the area of ElMontmell was a consequence of increasingly adverseweather conditions. Without these, local emigrationwould not have been so dramatic. Extreme snow-storms, frost and freezing temperatures between 1930and 1955 devastated local agriculture. Alfons remem-bers the frost of 1955 when most olive trees perishedand the harvest was so severely damaged that oliveswere unsuitable either for picking or for oil extraction.Later local wood production ceased because woodcut-ters could not compete with the much cheaper importsduring the mid-1980s. In addition, the introduction ofcheaper materials, such as plastic and silicone, pro-gressively replaced cork to make bottle stoppers(Espadalé i Reballí 2005; 2010; Rocas i Gutiérrezand Espadalé i Reballí 1987). Local producers in ElMontmell abandoned the industry. Groves of corkoak (Quercus suber) and other wood pasture were pro-gressively replaced by pine plantations (Pinus halepen-sis) and the ground flora became dominated byMediterranean heather (Erica multiflora), box (Buxussempervirens) and elements of the garrigue, mainlycomposed of kermes oak (Q. coccifera) (Espadalé iReballí 2005, 2010; Rocas i Gutiérrez and Espadalé iReballí 1987). According to Alfons, the farms sur-rounding the cave of La Moixeta and the nearbyhamlet of Marmellar were totally abandoned in the1960s and a proportion of the agricultural land wassold to developers. The population in El Montmelldwindled to 162 by 1981 (Institut d’Estadística deCatalunya 2013). Currently, the farm at Cal Turró isin ruins, surrounded by vineyards, whereas Cal LaMoixeta has been converted into a ‘rustic’ restaurantand hotel, and most of the agricultural land is urba-nised one (Figs. 2 and 3). This situation forcedpeople to migrate to Barcelona and other cities, tofind job opportunities offered by tourism and industri-alisation from the 1950 to the 1960s onwards. The areahas recovered since 1990 rising from 256 to 1175inhabitants in 2006. Alfons believes that this islargely as a result of improvements in the roadnetwork, restoration and gentrification of traditionalCatalan farm houses, small-scale urbanisation projectsand the expansion of rural tourism (EnciclopèdiaCatalana 2013).The depopulation and the disappearance of subsis-

tence pastoral activities in the area led to changes inthe vegetation, such as the disappearance of grazingand natural reforestation. Natural expansion of scrub

communities and conifers also occurred, particularlyin the central mountainous areas of the Catalancoast. Before the industrialisation and tourist boom,human intervention led to fragmentation of formeroak wood pasture with vineyards and afforestationwith extensive P. halepensis plantations. With theabandonment of grazing, the scrub community,including Brachypodium and Stipa sp., and shrubs,such as S. elaeagnos, R. ulmifolius, C. myrtifolia,Bupleurum fruticosum, Rosa canina, Crataegus mono-gyna, Prunus spinosa and Pinus sp. (Figs. 2 and 5)(Gordi et al. 1996; Otero et al 2013; Vila i Subirósand Gordi 2001), re-colonised or invaded abandonedagricultural land, hillsides formerly used for grazingand any deserted settlements. These changes can beseen by comparing the aerial photographs taken in1956 and 2009 (Figs. 2 and 3). The photo from 1956depicts the two rural houses and a landscape struc-tured with arable land and pasture. In contrast, by2009, it has been replaced by some urban developmentand natural re-forestation of Aleppo pinewoods withan understorey of heather or scrub.The golden age of transhumance in Catalonia

occurred between the 16th and 19th centuries(Llobet and Vilà Valentí 1951; Miralles andTutusaus 2005; Vilá Valentí, 1973). Transhumancecentres were located along the Pyrenees and wereorganised in three regions, which exhibited distinct tra-ditions and vocabulary, the cabanesa (western area),carrerada (eastern area) and lligalo (southern area).Shepherds from the cabanesa spent their summers inthe central area of the Catalan coast moving betweenthe La Noguera, Segrià, Urgell and Les Garrigues dis-tricts, whereas shepherds from the lligalo regionswould take their herds to the Maestrat district inValència and the mountains in Teruel province(Aragón). The shepherds from the carrerada regionmigrated to L’Empordà, Vallès counties and ElPenedès, where Alfons was located (Llobet and VilàValentí 1951; Miralles and Tutusaus 2005; ViláValentí 1973).The weather conditions in the Pyrenees are extreme

in winter and there is insufficient forage to feed ovi-caprine flocks. Pyrenean shepherds would conse-quently migrate southwards to access forage crops,which would grow in the milder conditions of thecoastal mountains (Llobet and Vilà Valentí 1951;Miralles and Tutusaus 2005; Vilá Valentí 1973).Northern transhumant shepherds would leave part oftheir herds with southern sedentary shepherds andtake the rest to the available pastures. In summertime,when the harsh hot and dry conditions of the southernmountains would deplete the fodder in the pastures,the southern shepherds would give part of theirherds to the Pyrenean ones, to be fed in the northernpasture, thereby completing the transhumance cycle

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(Llobet and Vilà Valentí 1951; Miralles andTutusaus 2005; Vilá Valentí 1973). Our informant,Alfons, always dealt with the same shepherd: JoanMuntanyès de Cal Miquel. He would travel132 km through mountain passages for 20 days,from La Pedra I La Coma (Sant Llorenç deMorunys, Solsonès County) in the carrerada trans-humance region, to Alfons’ farm (Fig. 6). Theywould mark their own individual animals beforeamalgamating their herds and Alfons would onlykeep those that would not survive the trip to thenorth, such as old and new born animals.Shepherds would pay for the service by donatingseveral of their animals.In the coastal mountains of Catalonia, sedentary

shepherds such as Alfons would feed their herds onnatural vegetation only and seek refuge in caves from

bad weather and during the heat of the day insummer. Occasionally, they would sleep in caves over-night. The caves were exclusively used by the shep-herds who paid a yearly rent to the owner. Nomadicshepherds never used the caves and rested undertrees or in designated stonemason refuges duringtranshumance periods. Alfons explained that the shep-herds from El Montmell never roamed for more than afew kilometres from the local farm and used threecaves in the area: ‘la Cova de la Plana Joana’ and ‘laCova dels Costers’, both located near the La Juncosadel Montmell, and ‘Cova de La Moixeta’. Shepherdsfrom the farms near the cave of La Moixeta, CalTurró and Cal La Moixeta (Figs. 2 and 3) had usedthe cave for at least four generations. Alfons purchasedthe herds from the shepherd in La Moixeta area whenthe latter retired.

Figure 6 Map showing the main transhumance routes in Catalonia. The route used by the Pyrenean shepherd is highlighted inred (modified from Fundació delMónRural and Institut Cartogràfic i Geològic de Catalunya 2010). A Sant Llorenç deMorunys andB La Juncosa del Montmell.

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Alfons stated that sheep defaecated and trampled ontheir dung in the caves during long spells of badweather. Dogs very seldom defaecated inside thecaves and wild animals avoided them. However, afterthe abandonment of agro-pastoral activities in thearea, the caves were used intermittently by wildanimals, in particular by badgers and foxes. Alfonsexplained that he and other locals would regularlyfind fresh dung on the cave floor when badgers usedthe cave as overnight shelters, setts and latrines. Henoted that badgers dug a hole at the cave entrance todefaecate in. Foxes would shelter in the caves whenthey were not occupied by badgers.

MethodsMorphological analysisTwelve dung samples were collected from the LaMoixeta cave and housed at the Museu Arqueològicdel Vendrell (Catalonia). They were organised intotwo groups MADV/561 (Figs. 7A–B and 8A–F) andMADV/562 (Fig. 9A–M). Four dung specimensfrom MADV/561 were used in this study(Fig. 7A–D). The surface of the dung was examinedusing a Nikon SMZ-10 binocular light microscope(10× to 60×). Dung size and morphology weremeasured following Chame (2003).

Radiocarbon datingIn the absence of any other dating evidence, the twosamples of animal dung from the group MADV/561(Figs. 7A and B) were dated at the OxfordRadiocarbon Accelerator (ORAU), and a furthersample from a dung specimen (Fig. 7A) of the groupMADV/561 (Fig. 7A) was AMS (Accelerator MassSpectrometry) dated at the Radiocarbon DatingLaboratory at the University of Waikato (Hamilton,New Zealand). AMS is used on samples smallerthan that needed for standard radiocarbon dating. Itcounts C14 atoms, rather than atoms which are decay-ing (Budzikiewicz and Grigsby 2006). The first usedthe standard methodology by McNichol et al.(2001), whereas the second applied the ConventionalAge or Percent Modern Carbon (pMC) method fol-lowing Stuiver and Polach (1977).

Geochemical analysisChemical composition of two specimens of animaldung from the group MADV/561 (Fig. 7A and B)were analysed using a Bruker D8 XRD with CuKα1radiation at the School of Earth and Environment atthe University of Leeds. The scans were processedusing the Bruker Eva with ICDD (InternationalCenter of Diffraction Data) Powder diffraction Filesand quantified using Bruker Topas software.

Figure 7 Specimens A–D used in this study, belonging to the group of dung from different producers reference MAVD-561stored at the Museu Arqueològic del Vendrell. A–C and B–D are photos of both sides of the same specimens. Notice striations indung A–C (arrows), and A–C and B–D (arrows) displaying dung beetle burrows.

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Epidermal and phytolith analysesIn order to examine plant content, the samples werewashed to remove extraneous material and lightlyground in a mortar to separate the epidermal frag-ments. Two subsamples weighing 10 g each were pre-pared. These were placed in a 5-ml test tube ofconcentrated HNO3 in a boiling water bath for 1minute. The samples were then diluted with 200 mlof water and sieved through 0.5 mm and 0.125 mmfilters. The 0.125–0.5 mm fraction was spread onglass microscope slides in a 50% aqueous solution ofglycerine. Cover slips (24×60 mm) were fixed ontothe slides with DPX microhistological varnish. Theslides were examined under a microscope at 100× to400× magnification to view traverses 2 mm wide and60 mm long with 3 mm between traverses.Vegetation percentages were registered by standardmicrohistological analyses of dung and compared todiet of current herbivores (Table 5) (cf. Bartoloméet al. 2011; Crocket 1959; García-González 1984;Stewart 1967).Phytolith analysis of the faeces of ca. 1 g of faecal

material was carried out. This was processed usingstandard methods, including chemical treatments toremove carbonates and organic matter centrifugationand density separation, and mounted for high magni-fication light microscopy and imaging (at 1000×, usingoil immersion) (Strömberg 2004). Over 200 (217) diag-nostic phytoliths were counted, resulting in an overallphytolith count of 375 (Table 3).The phytoliths were classified using a reference col-

lection from modern plants and grouped into ecologi-cally relevant classes (Strömberg 2003, 2004). Thoserelevant to this study are (1) Forest, diagnostic ofwoody and herbaceous dicotyledons, conifers andferns; (2) Closed-habitat grasses (Bambusoideae,Pharoideae, etc.); (3) Open-habitat grasses, dividedinto (a) Pooideae (C3), (b) the named PACMAD

clade (Duval et al. 2009; GPWG 2001), includingPanicoideae (mainly C4), and Chloridoideae (∼C4)and other PACMAD grasses (C3 or C4); (4) Grassshort cell phytoliths of unclear affinity; (5) Non-GSSC phytolith typical of grasses (e.g., wavy elongatesand trichomes) and (6) Ubiquitous or unknown phyto-liths. The relative abundances of phytoliths from theseclasses were used to determine the composition of thephytolith producing plants incorporated in the dung(Table 3). C3, C4 and CAM photosynthesis are thethree biochemical mechanisms are used by plants incarbon fixation (Sage et al. 2012). C3 are plants thatlive in cooler humid conditions, whereas C4 andCAM plants have devised alternative ways to survivehot conditions (Sage et al. 2012).

Modern vegetation types close to the cave containmoderate to low numbers of three grasses, Stipaoffneri, Brachypodium retusum and Brachypodiumphoenicoides. To investigate whether these grasses con-tributed to the dung assemblage, phytoliths wereextracted from leaf material of Brachypodium dista-chyum var. monostachyum, Brachypodium pinnatum,Brachypodium sylvaticum, Stipa tenuissima and Stipaneesiana, and analysed for comparison.

Pollen analysisPollen samples were subjected to conventional analyti-cal techniques of acetolysis and treatment with hydro-fluoric acid for the more siliceous samples (Mooreet al. 1991). Pollen counting was conducted at 400×,and 350 grains were counted for each sample. A2000 slide European and Mediterranean pollen refer-ence collection stored at Newcastle University wasused to aid uncertain identifications.

Diatom and nannofossil analysesFour diatom samples were exposed to acid accordingto the standard procedure established by Renberg(1990) and permanent slides were prepared withNaphrax®. Diatoms were identified under the lightmicroscope (Olympus BX40). Sample preparation ofnannofossils was conducted by spreading a fractionof dry sediment on to a cover glass together with adrop of distilled water. This procedure aimed toavoid laboratory contamination and to retain originalcomposition of the sample (see Monechi andThierstein 1985). The suspension was smeared on thecover glass with the aid of a wooden toothpick andthen dried on a hot plate. The cover glass was fixedon the slide with synthetic UV curing (NorlandOptical Adhesive). The slides were examined under alight microscope, both in transmitted and in polarisedlight, at a magnification of 1250×, scrutinising for cal-careous nannofossils (CNs).

Table 3 Sizes of the ovicaprine dung uncovered in the caveof La Moixeta.

Specimen number Length ×width (cm)

MADV/561A 1.62 × 0.67MADV/561B 1.16 × 0.70MADV/561C 2.05 × 0.69MADV/561D 1.86 × 1.37MADV/561E 1.16 × 0.88MADV/561F 1.42 × 0.71MADV/561G 2.37 × 1.03MADV/562A 7.49 × 4.42MADV/562B 7.35 × 5.38MADV/562C 3.16 × 1.95MADV/562D 1.95 × 1.83MADV/562E 1.88× 1.40MADV/562F 1.22 × 1.15MADV/562G 4.13 × 3.15MADV/562H 6.03 × 3.79

Note: Specimens followed numbering from Figs. 10 and 11.

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Entomological analysisTwo samples were examined for insect remains. Asample was rehydrated and gently disaggregated in dis-tilled water and the second was prepared without rehy-dration. The few insect fragments recovered werestored in 70% ethanol. Insect remains were identifiedunder a low power binocular microscope.

ResultsMorphological analysisThe dung specimens vary greatly in size and shape,and are very irregular, ranging from cylindrical tomore flattened specimens (Table 3, Figs. 7–11).Much of the material had clearly been disturbed bythe trampling of animals in the cave. The cylindricalspecimens display deep striations comparable tothose exhibited by current omnivore and herbivoremammal dung, such as that of badger, bear, wolf,dog and humans (Fig. 9C–F, I–K and 10A andB)(Martí Boada pers. comm.). Except for three well-compacted specimens, the rest were externally rough,dark brown and internally reddish and quite

Figure 9 Group of dung referenceMAVD-562 stored at the El Museu del Vendrell. A–D; B–E; C–F; G–L; H–J; and I–K are photos ofboth sides of the same specimens, and M is a single shot of a specimen.

Figure 8 Group of dung from different producers referenceMAVD-561 stored at the Museu Arqueològic del Vendrell. A–Dare photos of both sides of the same specimen.

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crumbly. Four show distinctive insect burrows(Figs. 7–11).

Radiocarbon datingSamples submitted to the Oxford RadiocarbonAccelerator Unit, contained insufficient carbon fordating, but other material sent to the RadiocarbonDating Laboratory at the University of Waikato(Hamilton, New Zealand) returned modern dates of30–50 years old (Table 4).

Geochemical analysisThe specimens are unlithified and composed of a cal-careous clay–silt matrix with scattered angular to sub-angular quartz grains. The presence of bone-derivedcalcium phosphate is used as a diagnosis for carnivoredung (Chin 2002; Pesquero et al. 2014) and this wasnot detected. The mineral composition of MADV/561 was slightly different from MADV/561(A)(Fig. 12A and B).

Epidermal and phytolith analysesThe phytolith assemblage in the dung is diverse interms of morphotypes. It mostly (∼57%) consists ofshort cells of grass silica (GSSC), with an additional∼22% non-diagnostic grass phytoliths (Fig. 13A–I).The remainder is almost all non-diagnostic morpho-types (Tables 5–6). This indicates that virtually all phy-toliths in the sample derive from grasses. The diet

represented by the dung therefore contained relativelyabundant grass. The GSSCs are >90% pooid, withpredominantly keeled rondels, conical rondels andseveral large crenates (Fig. 14A–D). PACCMADforms contribute only 3.8% of the GSSC assemblage,all of which are indicative of C3 PACCMAD grasses(e.g., Cortaderia selloana) (Kondo et al. 1994).Although C. selloana is originally from SouthAmerica, its presence in the phytolith assemblage isnot surprising. Known as Pampas grass in English, itwas introduced to France between 1775 and 1862(Bossard et al. 2000), and has spread over much ofEurope including Iberia. During the last 15 years, ithas become an invasive alien and has had a negativeimpact upon the native flora of northern Spain,especially in Cantabria (Sanz-Elorza et al. 2004).‘Gineri’ or ‘plomalls’, as it is known in Catalan, hasbeen widely used as an ornamental grass inCatalonia for the last century. It was planted in thepublic gardens when the agricultural land in LaMoixeta was urbanised in the 1960s, and has spreadin private gardens (Domènech et al. 2005; Linderet al. 2010). Thus, the concurrent presence ofCortaderia sp. pollen supports the age of the dung pro-vided by radiocarbon dating. It also suggests thatsome agro-pastoral activities were still being practisedwhile the urbanisation of the area was beingdeveloped.

The grass community that the dung producersgrazed upon was likely to have been almost 100%C3. Predominance of grass communities belonging toC3 are to be expected in arid mountainous areas,

Figure 10 Cylindrical dung displaying deep transverse andconcentric striations (arrows). A and B are photos of bothsides of the same specimen.

Figure 11 Dung displaying beetle burrows.

Table 4 Dating of a dung specimen (Fig. 7A–C) of the group MADV/561 (Fig. 7B–D), applying the Conventional Age or PercentModern Carbon (pMC) method following Stuiver and Polach (1977)

δ13C D14CC F14C%Higher calibrated age (68.2%probability)

Lower calibrated age (95.4%probability)

Total uncertaintyyears

−28.8± 143.4±

141.3±

1973–1977 calAD 1971–1980 calAD

±error 0.2‰ 4.6‰ 0.5%

Note: The isotopic fractionation, δ13C, is expressed as ‰ w.r.t. PDB.

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Figure 12 Graph showing themineral composition of the two specimens of dung analysed,MADV/561 (Figs. 11A and B). MADV/561(A) Aquartz 29.57%, calcite 46.76%, dolomite 9.63%, chlorite IIb 5.49%, muscovite 1M 8.54%, and orthoclase feldspar 0.00%,MADV/561(B) quartz 17.34%, calcite 37.93%, dolomite 12.61%, chlorite IIb 9.32%, muscovite 1M 12.87%, and orthoclasefeldspar 0.93%.

Figure 13 Some of the epidermal remains uncovered in the ovicaprine dung. A Trichomes of dicotyledon (herbaceous); Bepidermis of dicotyledon (herbaceous); C trichomes and epidermis of dicotyledon (herbaceous); D epidermis of bryophyte; Eepidermis of monocotyledon (graminid); F trichomes of leguminid (Trifolium?); G trichomes of Rosaceae (Rubus?); H trichomesand epidermis of dicotyledon (herbaceous); I epidermis and stoma of dicotyledon (herbaceous).

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because they are better adapted to cooler elevationsthat are exposed less often to very hot conditions(Sage et al. 2012). The high abundance of keeledrondels is consistent with, but by no means exclusiveto, Brachypodium sp. (Fig. 14A–D). In addition, asubset of the crenate forms is reminiscent of those pro-duced by this genus. However, the sample does notcontain the bilobates and polylobates that are abun-dant and typical of the Brachypodium leaf phytolithassemblages studied elsewhere (Strömberg, unpubl.data). Certain crenates are also similar to thosefound in Stipa and related grasses but the diagnosticstipoid bilobates are missing from the assemblage(Fig. 14A–D). The grasses consumed by the herbivoretherefore were mostly unknown pooid grasses and avery small proportion of Brachypodium and Stipa.The presence of the unknown pooid grasses reflectsthe sharp decline in pasture grasses and agriculturalland except for a few vineyards and orchards overthe last 50 years (Figures 2 and 3).

Pollen analysisThe specimens contained large quantities of pollen,dominated by 70% of Genista type. This suggests a

typical vegetation cover of Mediterranean garriguebut with some wet-loving taxa, as evidenced by thepresence of Salix (Table 6). There were also traceamounts of Pinus, Quercus, Ulmus, Mercurialis,Trifolium, Poaceae, Phillyrea, Galium, Rumex,Artemisia, Potentilla, Plantago, Geranium,Filipendula and spores of Filicales. This assemblagecorresponds to the extant flora in the immediate vicin-ity, except for Filipendula. Although F. vulgaris hasbeen recorded in nearby localities, it is locally a veryrare species (Table 6).

Diatoms and nannofossil analysesExamination of the dung revealed Nitzschia sp. and afurther species of diatoms (Fig. 15A-PO).Unfortunately, the identified genus includes bothfreshwater and marine species. Albeit relatively abun-dant, they were insufficient to run statistical analysis(Fig. 15A-O). Three species of CNs, all coccoliths,were obtained from the two samples analysed,

Table 6 Pollen percentage and number of vegetationfragments found in the dung specimens

Pollen type %Epidermalfragments Total

Genista type 70 P. halepensis 146Mentha type 11 Juniperus

oxycedrus12

Ericaceae 4.9 Ruscus aculeatus 9Salix sp. 3.4 Legumes 5Prunus type? 6.1 Other dicotyledons 332Poaceae, Phillyrea, Galium,

Rumex, Quercus,Artemisia, Potentilla,Plantago, Geranium,Ulmus, Filipendula andFilicales

4.6 Gramineae 318Briophyta 34Leguminosae 40Rubus sp. 36Floral fragments 22Fruit fragments 11Stem fragments 347Unidentifiable plant

fragments525

Animal fragments(insects)

38Figure 14 Phytoliths extracted from the ovicaprine dung. AStipa; B Chionochloid; C Keeled Rondels; D Brachypodium

Table 5 Phytolith percentage and number extracted from the dung specimens

Phytolith counts (%) of sample

Phytolith morphotypes* (%)

GSSC

FI CHTOT

POOID-D

POOID-ND

PACCMADgeneral

OtherGSSC

Non-diagnosticgrass

Other non-diagnosticgrass

Total phytolithcount

1.07 0.27 10.04 41.1 2.13 2.93 21.9 20.03 375

*GSSC, grass silica short cell; FI, forest indicator; phytoliths; CH TOT, closed habitats GSSC phytoliths; POOID-D, diagnostic pooidGSSCs; POOID-ND, non-diagnostic pooid GSSCs; PACCMAD general, C3/C4 PACCMAD clade (Panicoideae, Arundinoideae,Chloridoideae, Centothecoideae, Micrairoideae, Aristidoideae and Danthonioideae) GSSCs; Non-diagnostic grass, trichomes andwavy and spiny epidermal long cells and other morphotypes abundantly produced by grasses; Other non-diagnostic grass,phytoliths forms found in any types of plants.

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presumably having been ingested with mineral matter.The preservation of the specimens is very poor, withextensive overgrowth and/or dissolution, eventhough these taxa are resistant to diagenetic processes,in particular dissolution (Fig. 16A–E) (Thierstein1980). The three species from the dung are stratigra-phically distributed as follows: Watznaueria barnesiaefrom the Bajocian to the Maastrichtian (MiddleJurassic to Late Cretaceous); Cyclagelosphaera mar-gerelii, from the Bajocian to the Palaeocene and C.deflandrei, from the Tithonian to the Maastrichtian(Late Cretaceous) (Bown 1998); few specimens belong-ing to the genus Reticulofenestra sp., ranging from thePalaeocene to the Pleistocene, with a peak between theEocene and the late Miocene, were also found(Fig. 16A–E) (Wei and Wise 1990; Young 1998).Whilst there are no studies of the substrate microfossils

of the cave or those of the upper layers, the temporaldistribution of C. margerelii andW. barnesiae includesthe Jurassic-Lower Cretaceous limestones into whichthe cave is eroded, as well as the Cretaceous stratawhich outcrop in the area surrounding the cave (seeFigs. 4 and 16A–E).In contrast, the Cenozoic distribution of

Reticulofenestra sp. does not correspond to the ageof the rock present near the cave, and the originalsource of these nannofossils is likely to be in otherstrata in the catchment which also includesPalaeocene and Eocene rocks further north, ca.4.5 km from the La Moixeta cave (see Fig. 4). Eitherthe nannofossils were transported to the cave via thedrainage system, or they were incorporated in dungas a result of incidental ingestion during grazingsome distance from the cave (see Fig. 4).

Figure 15 A–C cysts; D unknown diatom; E part of a sponge spicule; F–G Diatom? – Aulocoseira?; H cyst; I Nitzshia sp.; J–Kunknown diatoms; L–M Nitzshia sp.; N–O spicules.

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Entomological analysisInsect remains from animal dung include not onlythose exploiting it as a primary habitat and their

predators (cf. Skidmore 1991), but also those acciden-tally ingested by the herbivore. The samples includedinsect remains of diverse sizes belonging to species cur-rently inhabiting the area (Fig. 17A–F). It is unfortu-nate that the fragments of pronotum and elytron ofGeotrupes cannot be identified beyond the speciespair, G. spiniger (Marsh.)/stercorarius (L.)(Fig. 17A–F). Both, however, occur in the dung of awide range of large herbivores (Koch 1989), althoughthe fragmented nature suggests the accidental con-sumption of a specimen by a grazer. The Europeanmole cricket, Gryllotalpa gryllotalpa (L.), is largelysubterranean, occurring in loose damp soils (Harantand Jarry 1982) and in wetlands (Marshall and Haes1988), although in the Mediterranean region it isalso recorded from dry, stony and sandy soils (Harris1982). Introduced to North America early lastcentury, Weiss and Dickerson’s (1918) comment thatadults may hibernate in horse manure and otherwarm places may be relevant to its appearance indung in a cave. Meadow spittlebugs (Philaenus cf. spu-marius) are numerous in grassland, depositing the fam-iliar ‘cuckoo spit’ to protect their larvae on stems, andlike the indeterminate weevils (Curculionidae), wereprobably accidentally ingested whilst grazing. Ants(Formicidae indet.) and the bumblebee, Bombus cf.terrestris, are probable accidental visitors to the caveand nothing can be added with regard to the indeter-minate Hymenoptera (Fig. 17A–F).

Figure 16 Nannofossils found in the dung. A, Watznaueriabarnesiae; B, Reticulofenestra sp.; C and D,Cyclagelosphaera deflandrei; E, Cyclagelosphaeramargerelii.

Figure 17 Insects uncovered in the dung. A, Gryllotalpa sp.; B, Gryllotalpa sp. head; C, Philaenus sp.; D, Curculionidae indet.; E,Messor sp.; F, Bombus sp.; G, Hymenoptera indet.

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DiscussionTaken overall, geochemical and biological materialsagree with the contextual evidence that the dung origi-nated from ovicaprine, rather than human, canine orother carnivore sources. The lack of calcium phos-phate removes carnivores as the possible dung produ-cers, and the included plant remains, pollen,epidermal and phytolith remains, and the calcareousclay–silt matrix with some quartz grains, suggest thatthe dung also contained sediment from the cave andother extraneous material via taphonomic processes,probably largely trampling by the animals, whichmay have destroyed any pelletal structure. The factthat pollen, epidermal and phytolith remains areentirely from natural vegetation concurs with the shep-herd’s comment that herds were fed entirely on naturalvegetation and pasture. The herds may have obtainedthe pre-Quaternary diatoms and nannofossils fromwater in the catchment. If this was the case, it wouldcertainly concord with the contextual evidence thatthe herds did not roam far from the cave.Alternatively, some could be contaminants from theCretaceous outcrops into which the cave is eroded;or could have been ingested incidentally duringgrazing from dust on vegetation. Dietary mineraldeficiencies may also lead to herbivores licking andchewing calcareous rocks (e.g., Chládeka andZapletal 2007; McDowell 1996). It is significant thatthe shape of the dung specimens was not diagnosticof the producers. These had been distorted and somesamples resembled carnivore (Figs. 9C–F, I–K and10A and B), cattle (Fig. 7A–D) and human(Fig. 10A–C and B–D), rather than ovicaprine dung.The dung also reflects the changes in vegetation as

described by the contextual evidence, and the presenceof unknown pooid grasses is the only difference fromcurrent vegetation. This reflects the sharp decline inpasture over the last 50 years, the abandonment ofagro-pastoral practices in the area, and the partialurbanisation of some of the agricultural land, exceptfor a few vineyards and orchards. It also registers theconsequent natural re-colonisation of the scrub com-munity and conifers, in the mountainous area of theCatalan coast.

ConclusionInterpreted in the light of the ethnographic evidence,the multidisciplinary study of the animal dung providesa reasonable picture of land use around the cave of ElMontmell, as well as identifying the source as ovica-prines. The mineral content and pollen, phytolith,and dermal contents of the dung allow the inferencethat the only producers of the dung were the localherds of sheep and/or goats that used the cave in therecent past as shelter. The lack of cereals and veg-etables remains also confirms that the herds were

entirely fed on natural vegetation, at least during epi-sodes when the animals utilised the cave. The shepherdwas able to confirm that because agricultural pro-ductivity was so the flock’s diet was not supplementedwith cereals and vegetables. Nannofossils and diatoms,sourced from outcrops 4 km away from the cave, mayindicate that herds roamed locally. The contents of thedung registered the existence of grazing pastures,which, as the aerial photographs confirm, have beenlost over the past 50 years to the rapid expansion offorest and scrub, although the paucity of currentgrazing means that this in itself is a new habitat. Asa caveat, it should be noted that the shape of thedung was not diagnostic of the producers and somespecimens resemble the coprolites of carnivores.Taphonomic processes, including animal tramplingand digging, have distorted the original shape, andmixed the dung with the cave’s sediment.More animal dung is expected to be unearthed in

the older layers of the cave sediment, since evidencefor agriculture and cattle farming has been recordedin the surrounding caves from at least the secondcentury AD (Burés et al. 1992). Thus, the cave of ElMontmell and perhaps other caves in the area havethe potential to provide a palaeoenvironmentalrecord for the central mountainous area of theCatalan coast. This preliminary study also serves tounderline the potential of a multidisciplinary studyof recent coprolites, backed by ethnographic research,in providing the background to the interpretation ofpast environments and societal changes.

AcknowledgementsWe would like to thank the two anonymous reviewersfor the very useful comments and suggestions whichhelp us improve the quality of our paper. The firstauthor would also like to give special thanks toGrandma Rosa for introducing him to the caves andother natural wonders of the area, and inducing alove for nature and study of the history of agro-pas-toral activities. We are deeply grateful to AlfonsGuash i Martí for the very helpful accounts of thefarming and herding practices in the area; transcrip-tions of his accounts have been deposited at theMuseu Etnològic de Barcelona. Thanks to AbelJulient i Vila (Institut Català d’Ornitologia), DrMartí Boada i Juncà (Institut de Ciència iTecnologia Ambiental (ICTA) de la UniversitatAutònoma de Barcelona), Professor Karen Chin(University of Colorado Boulder), Daniel Corominasi Muñoz, for having provided information on thenesting birds, carnivore dung, and the treatmentidentification of some of the phytoliths, respectively.Thanks are also due to the Fundació del Món Ruraland rgw Institut Cartogràfic i Geològic deCatalunya, 2010 for letting us use the map of the

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main transhumance routes in Catalonia; Dr AngelGalobart i Lorente (Institut Català de PaleontologiaMiquel Crusafont) for logistic and laboratorysupport; and Francesc Burjachs i Casas (ICREAResearch Professor at IPHES) for providing unpub-lished information about animal dung research inIberia; Dr Lesley Neve (University of Leeds) forrunning the geochemical analyses art the school ofEarth and Environment at the University of Leeds;and Dr A. Prieto i Márquez (University of Bristol)and Gary Morfitt (Leeds College of Art) for theirhelp in editing the images.

ORCIDXavier Panadès i Blas http://orcid.org/0000-0003-3812-4482Jordi Bartolomé i Filella http://orcid.org/0000-0002-3784-5248Soriano i Tomàs http://orcid.org/0000-0003-3875-7511Antonio Arillo Aranda http://orcid.org/0000-0002-4878-5797Lourdes Chamorro i Lorenzo http://orcid.org/0000-0002-5899-711X

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