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A Late Pleistocene (MIS3) ungulate mammalassemblage (Los Rincones, Zaragoza, Spain) in theEurosiberian–Mediterranean boundaryVíctor Sauquéa, Ricardo García-Gonzálezb & Gloria Cuenca-Bescósa

a Grupo Aragosaurus-IUCA, Paleontología, Facultad de Ciencias, Universidad de Zaragoza,C/Pedro Cerbuna 12, E-50009 Zaragoza, Spainb Instituto Pirenaico de Ecología (IPE-CSIC), Avda. Nuestra Señora de la Victoria s/n, 22700Jaca, SpainPublished online: 03 Sep 2014.

To cite this article: Víctor Sauqué, Ricardo García-González & Gloria Cuenca-Bescós (2014): A Late Pleistocene (MIS3)ungulate mammal assemblage (Los Rincones, Zaragoza, Spain) in the Eurosiberian–Mediterranean boundary, HistoricalBiology: An International Journal of Paleobiology, DOI: 10.1080/08912963.2014.945926

To link to this article: http://dx.doi.org/10.1080/08912963.2014.945926

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A Late Pleistocene (MIS3) ungulate mammal assemblage (Los Rincones, Zaragoza, Spain) in theEurosiberian–Mediterranean boundary

Vıctor Sauquea*, Ricardo Garcıa-Gonzalezb1 and Gloria Cuenca-Bescosa2

aGrupo Aragosaurus-IUCA, Paleontologıa, Facultad de Ciencias, Universidad de Zaragoza, C/Pedro Cerbuna 12, E-50009 Zaragoza,Spain; bInstituto Pirenaico de Ecologıa (IPE-CSIC), Avda. Nuestra Senora de la Victoria s/n, 22700 Jaca, Spain

(Received 29 March 2014; accepted 15 July 2014)

The Late Pleistocene archaeo-palaeontological sites in the Iberian Peninsula are located mainly on the coasts. Here, wepresent for the first time a palaeoenvironmental proxy for Upper Pleistocene locality (Marine Isotope Stage 3 MIS3) that isin the interior peninsular, in the Moncayo massif (Zaragoza). This is actually the boundary between Mediterranean andEurosiberian climatic regions. This study is based in the site ungulates: Capra pyrenaica, which is larger in size than thecurrent and fossil Capra from the Mediterranean area of the Iberian Peninsula. The horses have a small size, which is similarto that of the horses from Fontainhas and Casares. Significantly lesser in the number of specimens are the roe deer, thesouthern chamois and the auroch. The study and comparison of the faunal assemblage of the locality of Los Rincones withother sites of the Iberian MIS3 show a cluster of sites in the southern Peninsular, though separated, due to the abundance ofCapra pyrenaica. The association of Los Rincones is similar to the southern peninsular sites such as Nerja, Gorham Cave,Cova Beneito and Zafarraya. The ungulate assemblage of Los Rincones represents a landscape with temperate climate,presence of steppe and patches of forest, similar to the current landscape surrounding the cavity today.

Keywords: ungulates; Iberian Peninsula; Late Pleistocene; Capra pyrenaica; Equus ferus

1. Introduction

The Iberian Peninsula is an exceptional area for studying

the faunal associations of the Late Pleistocene, because on

account of its geographical configuration it displays two

distinct climatic ranges: the Atlantic and the Mediterra-

nean (Schmidt et al. 2012). Furthermore, the Iberian

Peninsula contains the southwestern limit to the cold

faunas that make up what is also known as the

‘Mammuthus–Coelodonta faunal complex’ (Kahlke

1999, Kahlke 2014). These faunas generally occur on a

sporadic basis, being most frequent during Marine Isotope

Stage 3 (MIS3) (Alvarez-Lao and Garcıa 2010, 2011).

Although there are many Late Pleistocene (MIS3) sites in

the Iberian Peninsula, most are concentrated in coastal

areas or in the Pyrenean region, and there are few

references to inland sites with faunas from the period in

question (d’Errico and Sanchez Goni 2003; Stewart 2007;

Schmidt et al. 2012).

The objective of this article is to undertake a

taxonomic and biometric study of the ungulates found in

the cave of Los Rincones (Zaragoza, Spain) and to carry

out a faunal analysis of the association present at the site,

comparing it with the bibliographical data. Through a

comparative study of the fauna, the aim is to define the

landscape of the inland area of the Iberian Peninsula

during the Late Pleistocene (MIS3), a landscape that was

inhabited by large predators such as the leopard and Homo

neanderthalensis.

2. Description of the study area

The cave of Los Rincones is located in the Sierra del

Moncayo, in the central part of the Iberian Range in the

north of the Iberian Peninsula (Figure 1(a)). On account of

the isolated relief it presents between the boundary of the

Ebro Valley and the Castilian Meseta as well as its high

altitude (2315m), its special orientation and its geological

diversity, the Moncayo massif is endowed with special

characteristics that are unique within its geographical

setting. It represents a confluence of three biogeographical

subregions (Uribe-Echebarrıa 2002). The northern slope of

the Moncayo is subject to the wet, cold influence of fronts

from the Atlantic, harbouring plant taxa of a northern

character, such as Quercus robur (Gomez Garcıa et al.

2003). By contrast, the surrounding land shows a marked

Mediterranean character, with vegetation dominated by

evergreen Quercus such as Quercus ilex and Quercus

coccifera. A few kilometres to the east lies the Ebro Valley,

with its notable aridity, where species characteristic of

semi-arid steppes such as Lygeum spartum can be found

(Hernandez and Valle 1989; Uribe-Echebarrıa 2002).

The climatological data from Agramonte de Moncayo

(Zaragoza), situated at an altitude of 1090m, show a mean

annual precipitation of 927mm and a mean annual

temperature of 9.48C (Hernandez and Valle 1989). The

pronounced altitude gradient and the NW–SE climatic

dissymmetry result in a great diversity of plant

environments, with three vegetation levels at present:

q 2014 Taylor & Francis

*Corresponding author. Email: vsauque@unizar.es

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meso-, supra- and oro-Mediterranean, with the existence

of a fourth level under discussion, the cryoro-Mediterra-

nean, which would be the equivalent of an Alpine level.

The only comparable reference as regards the predominant

vegetation during the period to which the fossil remains

from Los Rincones correspond (MIS3) would be Gabasa

Cave. A pollen study carried out on hyena coprolites

dating to between 50 and 40 ka reflects a mosaic glacial

landscape that includes Pinus and Juniperus woodlands

and steppes with Chenopodiaceae, Poaceae, Artemisia

and Asteraceae, suggesting an arid and cold/cool climate.

The presence of mesophilous and thermophilous tree and

shrub taxa (Quercus ilex, Quercus coccifera) also suggests

the existence of temperate Mediterranean refugia during

the period in question (Gonzalez-Samperiz et al. 2003,

2005).

The relief around the cave of Los Rincones is abrupt,

with the site surrounded by steep cliffs (Figure 1(b)),

which would foster the presence of rupicolous species

such as Rupicapra pyrenaica and Capra pyrenaica.

However, the ravine of Los Rincones opens into a broad

valley that could have hosted open-environment species

such as Equus and Bos primigenius and woodland species

such as Capreolus capreolus and Cervus elaphus.

Figure 1. (a) Geographical location of Los Rincones cave. (b) Panoramic view of the Los Rincones ravine. (c) Panoramic view of themain entrance of the cave. (d) Plan view of the cave. (e) Elevation view of the cave.

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2.1. The site of Los Rincones: origin of theaccumulation

The cave of Los Rincones is situated at the head of the

ravine of Los Rincones (with a SW orientation), in the

municipality of Purujosa (Zaragoza, Spain). It opens at an

altitude of 1010m (see diagram of geographical location)

(Figure 1(c)). The cave is divided into a number of

chambers. The remains under study in this article appeared

at the surface of the ‘Ursus gallery’ and the ‘Leopard

gallery’, which are connected and are considered a single

site (Figure 1(d),(e)). The sample presents a similar pattern

of fragmentation, with similar tooth and cut marks, and

breaking pattern due to collapse of celling blocks. Also the

faunal composition, the skeletal survival profiles and

the degree of preservation of the remains, as well as the

distribution of fragments of the same anatomical element

in distinct galleries of the site, indicate that the process of

accumulation was similar: bones, with other clastic

sediments, were carried in from the surface (allogenic

transport) to the Ursus gallery, until the cone blocked the

mouth of the cave (Sauque et al. 2014). The faunal

association under study in this article date at least to the

Late Pleistocene, probably MIS3. We sent samples to a

radiocarbon laboratory (Beta Analytics, Miami, FL, USA)

but unfortunately we could not obtain radiometric ages by

the 14C method due to lack of collagen in the bones. Also

during the early stages of cleaning the gallery, a piece of

Mousterian industry was also discovered in the cave of Los

Rincones.The upper part of the sedimentary cone that

closes off the original entrance to the cave has been dated

on the basis of a microfaunal association consisting of

Microtus, Iberomys and Pliomys lenki. Such an association

is characteristic of the Late Pleistocene. The species

Pliomys lenki disappears between 50 ka and 40 ka from the

centre of the Iberian Peninsula, where it is only found at

Mousterian localities (Cuenca-Bescos et al. 2010; Sauque

and Cuenca-Bescos 2013).

Late Pleistocene faunal accumulations are generally

produced by the activity of humans or carnivores (Cruz-

Uribe 1991; Stiner 1991; Diedrich 2011; Enloe 2012).

Noteworthy among the latter are the accumulations

generated by hyenas, which are much more frequent

than those generated by leopards (Yravedra 2006;

Diedrich 2013; Sauque et al. 2014). Both hyenas and

humans tend to break bones in order to extract the marrow,

so the presence of whole bones is very rare at such sites.

The site of Los Rincones is one of the few sites where the

accumulation is attributed to the activity of a leopard as the

main accumulator (Sauque and Cuenca-Bescos 2013;

Sauque et al. 2014). Nonetheless, the site also testifies to

the sporadic presence of Neanderthals, who left their mark

in the form of cut marks in some bones, as well as remains

of lithic industry (Sauque et al. 2014). As a result, the

accumulation is composed mainly of small-sized herbi-

vores that fall within the range of leopard consumption.

The accumulation consists primarily of remains of Capra

pyrenaica (Table 1; Figure 2), thus providing a great

opportunity to carry out a biometric study of this

taxonomically controversial taxon (Garrido 2008;

Garcıa-Gonzalez 2011).

3. Materials and methods

For taxonomic identification, the following authors have

been followed: Bibikova (1958), Pales and Lambert

(1971), Eisenmann (1986), Gee (1993), Lister (1996) and

Fernandez (2001). Use has also been made of the reference

collections of the UZ (Universidad de Zaragoza) and the

Table 1. Composition of the Los Rincones ungulateassemblage.

Taxa NISP %NISP MNI %MNI

Cervus elaphus 13 2.11 2 6.25Capreolus capreolus 26 4.23 2 6.25Rupicapra pyrenaica 29 4.72 3 9.38Capra pyrenaica 528 85.85 20 62.5Bos primigenius 4 0.65 1 3.13Equus ferus 12 1.95 3 9.38Equus hydruntinus 2 0.33 1 3.13Total 615 100 32 100

Figure 2. Pie chart of ungulates from Los Rincones.

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IPE (Instituto Pirenaico de Ecologıa). In order to assess the

skeletal representation of the assemblage from Los

Rincones, we have used the number of remains, the

number of identified specimens (NISP) and the minimum

number of individuals (MNI), which have been calculated

in accordance with Brain (1981) and Lyman (1994a).

In order to determine the age of death, dental

replacement and the degree of dental eruption have been

used (Morris 1978; Hillson 1992), as well as the degree of

fusion of the epiphyses in long bones (Morris 1972).

Furthermore, for each species the criteria proposed by

other specialists have been followed: e.g. those from

Perez-Ripoll (1988) and Vigal and Marchordom (1985) for

Capra pyrenaica; those from Tome and Vigne (2003) for

Capreolus capreolus; those from Perez-Barberıa (1994)

for Rupicapra pyrenaica and those from Aitken (1975),

Azorit et al. (2002), D’Errico and Vanhaeren (2002) and

Mariezkurrena (1983) for Cervus elaphus. The measure-

ments were taken with a digital calliper (Digimatic Caliper

CD-800CX) in accordance with von den Driesch (1976) and

Bibikova (1958). For the remains from Equus, the

methodology proposed by Eisenmann (1986) has been

followed.

All measurements are given in millimetres. The

morphometric data have been compared with data

collections from western Europe published by a variety

of authors (see citations in Tables 2–7), as well as with our

own data gathered directly from the collections of the

Museo Nacional de Ciencias Naturales in Madrid

(MNCN), the Museu de Prehistoria de Valencia (MPV),

the Instituto Pirenaico Ecologıa (IPE), the Zoologische

Staatssammlung Munchen (ZSM), the Museo Paleontolo-

gico de Zaragoza (MPZ), the Museo Arqueologico

Provincial de Huesca (MAPH), the Instituto Alaves de la

Naturaleza (IAN), Institut fur Palaoanatomie, Domestika-

tionsforschung und Geschichte der Tiermedizin (Ludwig

Maximilians university Munchen) (LMU) and the

Naturhistorisches Museum Basel (NMB).

To provide a statistical comparison of the site of Los

Rincones with the faunal associations of other sites, we

Table 2. Selected comparative measurements on Cervus elaphus postcranial elements.

Site (mm) Tib. DW Mtt. PW 1st Pha. PW 1st Pha. DW 1st Pha. L 2nd Pha. PW 2nd Pha. L

Los Rinconesa Mean (n) 48.7 (1) 41. 6 (1) 20.3 (2) 18.8 (2) 57.4 (2) 21.0 (1) 37.7 (1)Min–max 19.7–20.9 18.8–18.9 56.5–58.2

Cova Negrab Mean (n) 43.9 (4) 19.77 (4) 18.2 (4) 54.4 (4) 18.6 (8) 38.6 (8)Min–max 41–48.6 16.3–21.8 14.5–21 50.8–60 16–20.9 33.7–42.2

Gabasac Mean (n) 46.7 (1) 34 (2) 20.8 (14) 20.2 (3) 52.9 (2) 20.24 (9) 37.2 (2)Min–max 31–37 18–23.8 17.7–22.5 50.3–55.4 18.1–23.4 36.7–37.6

Camino Caved Mean (n) 34.47 (4)Min–max 32.5–34.4

Lumentxa (Mag–Sol)e Mean (n) 51.2 (3) 45.8 (7) 22.2 (25) 21.1 (44) 58.3 (8) 21.6 (16) 41.7 (6)Min–max 48.5–53 41.5–54 19–25 18.4–24.5 53.5–64 18.4–23.5 38.5–44.5

Lumentxa n. VII (Aur)e Mean (n) 46.3 (4) 23.2 (8) 21.9 (8) 58.6 (4) 22.5 (5) 42.8 (4)Min–max 40–52.5 21.5–25.5 19.5–25 55–64 22–24 41–45

Santimamine nVI (Mag)e Mean (n) 55.6 (8) 45.5 (18) 21.7 (13) 57 (1) 22.5 (21) 44.7 (9)Min–max 52–61.5 42–50 18.8–24.5 19.6–25.5 40.5–46.5

Santimamine nVIII (Aur)e Mean (n) 53.9 (5) 44.4 (7) 22.8 (3) 22.5 (10) 62.3 (2) 23.1 (4) 47.3 (2)Min–max 51.5–60 43–45.5 21.5–23.5 20–24.5 60–64.5 21.5–24 46.5–48

Urtiaga Df Mean (n) 55.8 (3) 39 (5) 21.5 (4) 20.6 (11) 58.7 (10) 22.5 (12) 44.6 (4)Min–max 53.5–59 35.6–44 18.4–23 18.3–23 50.6–63 21–26 44–46.7

Labeko Kobag Mean (n) 54.4 (7) 41.4 (11) 23.2 (38) 22.3 (36) 58 (34) 23.1 (44) 43.8 (44)Min–max 50.5–60 37–45 22–25 20.5–24.5 54–63.5 21.5–25 39–47

La Paloma (Mag)h Mean (n) 50.5 (42) 43.6 (8) 21.9 (17) 19.9 (58) 54.6 (10) 21.8 (52) 41.7 (17)Min–max 45–57 42–46 19–25.5 18.6–22.5 51–61 19–24.5 37.5–47

El Mironi Mean (n) 52.2 (1) 23 (9) 20.5 (12) 60.4 (1) 21.5 (8) 38.9 (3)Min–max 21.3–27.1 17.4–24.4 17.2–24.3

Nerjaj Mean (n) 19.5 (2) 18.5 (2) 55.5 (1) 18.5 (1) 40 (1)Min–max 18.5–20.5 17–19.5

Parpallok Mean (n) 45.7 (19)Min–max 43.1–50

Morinl Mean (n) 52.4 (4) 48.3 (1) 22.2 21.8 (19) 54.8 21.8 (7)Min–max 49.1–52.8 21.4–24.2 19.4–24.8 53.5–56.1 20.7–22.5

Abauntzm Mean (n) 52 (3) 22 (3) 22 (2) 58.5 (3) 24.8 (2) 46.5 (2)Min–max 50–55.5 20–24 20–24 55–59.5 24–25.5 45–48

Notes: Tib., tibia; Mtt., metatarsus; 1st Pha., first phalanx; 2nd Pha., second phalanx; L, maximum length; DW, distal width; PW, proximal width. Mag,Magdalenian; Sol, Solutrean; Aur, Aurignacian.aThis study; bPerez-Ripoll (1977); cBlasco-Sancho (1995); dAlvarez-Lao et al. (2013); eCastanos (1986); fAltuna (1972); gAltuna and Mariezkurrena(2000); hMariezkurrena and Altuna (1983); iMarın Arroyo (2010); jMorales and Martın (1995); kDavidson (1989); lAltuna (1971); mAltuna et al. (2002).

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have undertaken correspondence analysis (CA) using

PAST v.2.14 (Hammer et al. 2001). CA is a way of

projecting a multivariate database onto two or three

dimensions in order to visualise the cluster patterns. It is

thus a good method for discovering geographical group-

ings and patterns and for bringing to light environmental

gradients (Greenacre 1984; Jongman et al. 1995; Legendre

and Legendre 1998).

The algorithms used by CA present two problems. The

first is the tendency to compress the end of the ordinate

axis, squeezing together the samples and taxa, which is not

useful. The second is the arch effect, which occurs when

the environmental gradient presents ‘losses’ in the first and

second ordination axes instead of being linked only with

the first axis as it should be. To avoid these problems, we

have used a detrended correspondence analysis (DCA) for

our comparison of faunal associations. First, this rescales

itself, removing the compression at the ends of the axes;

afterwards, it carries out the detrending, stretching and

bending the arch until it becomes straight. This type of

analysis is of great value for the ordination of ecological

databases (Hill and Gauch 1980).

4. Taxonomy

The ungulates from Los Rincones show a good state of

preservation due to the non-anthropic origin of the

accumulation (Sauque et al. 2014). Thanks to the low

levels of modification, it has been possible to identify

taxonomically a total of 615 ungulate remains (Table 1).

These remains correspond to the following taxa: Cervus

elaphus (red deer), Capreolus capreolus (roe deer), Capra

pyrenaica (Iberian wild goat), R. pyrenaica (Pyrenean

chamois), Equus hydruntinus (European ass), Equus ferus

(horse) and Bos primigenius (aurochs) (Table 1; Figure 2).

The main ungulate is Capra pyrenaica, constituting

86% of the total ungulate remains. The carnivores, which

are not included in this study, are Ursus arctos (brown

Table 3. Selected comparative measurements on Capreolus capreolus cranial and postcranial elements.

Site (mm) m3. L Hum. DW Ra.PW Mtcp. DW Tib. DW

Los Rinconesa Mean (n) 15.1 (1) 27.1 (1) 29.5 (1) 22.2 (1) 27.4 (1)Min–max

Gabasab Mean (n) 27.3 22.2 (1) 25.5 (3)Min–max 27–27.6 24.9–26

Jou Puertac Mean (n) 17.2 (2) 29.8 (5) 21.6 (1) 27.7 (4)Min–max 16.3–18.1 28.2–31.4 21–22.5 26.5–28.7

Urtiagad Mean (n) 16.5 (24) 31.1 (12) 24.4 (13) 30.7 (2)Min–max 14.5–17.6 28.2–33.4 21.3–27.9 29.9–31.5

Santimaminee Mean (n) 16.1 (27) 29.8 (13) 29.2 (3) 23.9 (8)Min–max 14.2–17.6 28–32.5 27–32.5 23–25

La Rieraf Mean (n) 16.9 (8) 29.4 (5) 24 (2)Min–max 15.5–18 22.5–32 23.5–24.5

Lumentxag Mean (n) 27 (1) 25 (2)Min–max 24–26

Notes: m3 L., lower third molar; Hum. humerus; Ra., radius; Mtcp., metacarpus; Tib., tibia; L, maximum length; DW, distal width; PW, proximal width.aThis study; bBlasco-Sancho (1995); cAlvarez-Lao (2014); dAltuna (1972); eCastanos 1984; fAltuna (1986); gCastanos (1986).

Table 4. Values of the index of lateral articular facet and distal trochlea astragalus of Bos and Bison according to different authors.

Astragalus

Lateral articular facet index Distal trochlea index

Reference Genus n Mean Variability N Mean Variability

Bibikova (1958) Bos .58 93–98.7Bison 50–54 73.3–89.2

Stampfli (1963) Bos 11 54.3 46.4–62.2 16 90 82–122.7Bison 12 45.9 41.6–56.6 12 81.7 76–100

Altuna (1972) Bos 3 52 46.5–55.8 8 89.7 81.2–95.5Buitrago-Villaplana (1992) Bos 8 53 43.3–69.4 8 95 83–108.5Sala et al. (2010) Bos 1 61.81 1 93.3

Bison 1 40.95 2 88.43 82.95–93.91Perez-Ripoll (1977) Bos 1 61.1 97.1Los Rincones Bos 1 60 1 110

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Table

5.

SelectedcomparativemeasurementsonCapra

pyrenaicacranialandpostcranialelem

ents.

Site

(mm)

LP2–P4

LM1–M3

m3.L

Hum.DW

Ra.PW

Ra.DW

Mtcp.PW

Mtcp.DW

Mtcp.L

Tib.DW

Ast.L

Ast.W

Mtt.L

Mtt.PW

Mtt.DW

LosRincones

aMean(n)

23.1

(1)

46.1

(1)

27.2

(11)

39.3

(10)

39.9

(6)

42.4

(4)

33.7

(12)

38.1

(8)

152.6

(6)

34.2

(8)

36(15)

23.9

(16)

150.4

(1)

26.5

(4)

32(4)

Min–max

25.2–29.3

34.8–45.9

38.5–41.5

40.2–45.3

28.5–37.8

31.6–40.7

145.6–158.3

29.8–37.9

34.2–40.6

21.9–28.2

24.1–30.2

29.6–35.6

JouPuertab

Mean(n)

24.6

(1)

53.6

(1)

25.5

(2)

34.2

(4)

37.6

(3)

38.3

(2)

Min–max

24.54–26.5

27–36.3

36.7–38.1

37.7–38.8

TitoBustillo

cMean(n)

22.85(2)

40.8

(5)

42.5

(1)

35.5

(1)

35.8

(2)

33.5

(1)

35.7

(7)

23.4

(7)

32.2

(3)

Min–max

21.7–24

34.5–45

34–37.5

32–40

21–27

31–33.5

LaRiera

dMean(n)

24.5

(2)

23.8

(23)

29(2)

35.7

(3)

31(6)

Min–max

23.5–25.5

19.5–28.5

28–30

30–39

29.5–34.5

UrtiagaDe

Mean(n)

23.7

(7)

55.9

(3)

27(23)

43.7

(13)

41.5

(1)

34.9(6)

145(1)

36.5

(1)

37(29)

24.2

(27)

Min–max

22.1–26

50.3–62

24–29.3

39.5–48.5

33.3–39

34.3–40.5

21.5–28

Chaves

(Mag)f

Mean(n)

24(1)

54(1)

23.6

(29)

36.7

(5)

42.3

(4)

31.25(2)

35(1)

34.3

(5)

32(9)

34.1

(5)

147.5

(1)

28.8

(3)

Min–max

21–29.5

31–41.5

36.5–45

27.5–35

28.5–41.5

27–36.5

31–35.5

27.5–30

Gabasag

Mean(n)

29(58)

43(11)

40(8)

37.4

(3)

31.3

(12)

35.2

(5)

142.9

(5)

35.3

(12)

37.6

(34)

24.5

(30)

148.1

(4)

28(10)

32.2

(5)

Min–max

25.8–32.2

38.2–48.6

37.8–42.8

37–37.9

24.7–37.4

27.4–41.4

129–156

31.8–39.1

32.2–41.7

21.5–27.5

143–151.1

25.7–29.9

29.7–37

Santimam

ineh

Mean(n)

25.5

(4)

48.6

(4)

26.8

(8)

40.8

(2)

30.5

(1)

35.3(5)

35.3

(3)

37.8

(12)

23.5

(11)

179(1)

25.5

(1)

31(1)

Min–max

22.5–31.5

44.5–54

21.5–32.5

37–44.5

30.5–41

33.5–38.5

30–40.5

19.4–27.5

Bolinkobai

Mean(n)

27(1)

26(37)

38.8

(2)

29.7

(3)

39.5

(3)

36.6

(31)

23.9

(28)

Min–max

22–28.5

38.5–39

29.5–30

33–43.5

33.5–40.5

21–27.5

Lumentxah

Mean(n)

21.5

(1)

25.5

(1)

35.5

(3)

43(1)

40(1)

40.5

(2)

156.5

(1)

29.3

(2)

32.1

(8)

21.1

(9)

32.3

(2)

Min–max

32.5–41

35–46

29–29.5

30.5–33.5

18.5–23

30.5–36

Errallaj

Mean(n)

25.6

(4)

48.6

(4)

25.9

(87)

39.6

(7)

39.4

(6)

41(1)

34.7

(3)

37.5

(4)

149.7

(3)

34.5

(8)

38.8

(6)

26.2

(5)

152.3

(2)

29.4

(4)

35.2

(3)

Min–max

23.5–29

44.5–54

23–29

37–42.5

32–48

33.4–35.5

33–41

141–154

29–39

37.5–40.5

24.5–28

150–154.5

27–31.5

32.5–39

LosCasares

kMean(n)

51.1

(2)

28.2(4)

42(1)

Min–max

50.2–52

25.5–31.4

CovaNegra

(MIS

5)l

Mean(n)

22.1

(1)

51.1

(1)

37.2

(1)

43.4

(3)

41(2)

31.6

(8)

34.3

(7)

32.7

(4)

37.3

(6)

23.8

(5)

Min–max

41.1–48

37.5–44

29–37

31.6–42.5

31.3–35

32.2–41

20.3–26.4

Parpallo

(UPa)

m,n

Mean(n)

26.8

(144)

34.9

(259)

36(37)

38.4

(71)

29.5

(49)

33.9

(113)

137.6

(29)

31.7

(63)

149.4

(29)

22.5

(23)

30.1

(19)

Min–max

23.5–30.4

28.9–39.6

36.5–41

31.9–45.9

25.4–32.1

26–37.1

123.8–143.4

27.9–35.2

133–160

21–27.5

27–34.5

Recolduc(Sol-Gra)n

Mean(n)

27.3

(9)

30.6

(9)

128.6

(6)

143.4

(5)

24(5)

28.3

(5)

Min–max

26.3–28.9

27.2

234.7

123.4–143.1

132.2–157.5

21.6–26.6

26.7–31.6

Malladetes

(UPa)

nMean(n)

28.4

(6)

32.3

(7)

131.1

(1)

137.1

(2)

23.9

(2)

29.5

(8)

Min–max

25.7–30.1

31.6–33.8

133.4–140.9

23.1–24.7

27.9–31

Nerja

(Mag)o,n

Min–max

27.3

(1)

34.5

(3)

36(1)

23(1)

28.6

(1)

Mean(n)

33.1–37.2

Capra

pyrenaicaextantn

Mean(n)

28.1

(131)

33.9

(131)

125.4

(131)

32.5

(136)

20.7

(135)

133.6

(135)

23.5

(135)

27.4

(135)

Min–max

23.2–33.3

25.9–38

104–144.2

27.1–36.4

18–24.8

105.77–155.48

18.7–26.1

21.1–33

Notes:P2–P4,upper

premolarseries;M1–M1,upper

molarseries;m3L.,lower

thirdmolar;Hum.,humerus;Ra.,radius;Mtcp.,metacarpus;Tib.,tibia;Ast.,astragalus;Mtt.,metatarsus;L,maxim

um

length;W,maxim

um

width;DW,distalwidth;PW,proxim

alwidth.Mag,Magdalenian;Sol,Solutrean;Gra,Gravettian;Aur,Aurignacian;UPa,Upper

Palaeolihitc.

aThisstudy;bAlvarez-Lao

(2014);

cAltuna(1976);

dAltuna(1986);

eAltuna(1972);

f Castanos(1993);

gBlasco-Sancho(1995);

hCastanos(1984);

i Castanos(1986);

j AltunaandMariezkurrena(1985);

kAltuna

(1973);

l Perez

Ripoll(1977);

mDavidson(1989);

nSauque(unpublished

data);oMoralesandMartın(1995).

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Table

6.

SelectedcomparativemeasurementsonRupicapra

pyrenaicacranialandpostcranialelem

ents.

Site

(mm)

m3.L

Hum.DW

Ra.PW

Tib.DW

Ast.L

Ast.W

1stPha.L

1stPha.PW

2ndPha.L

2ndPha.PW

LosRincones

aMean(n)

17.1

(3)

28(2)

28.8

(1)

27.1

(1)

30.4

(4)

19.9

(4)

41.8

(1)

13.3

(1)

27.5

(4)

13.7

(4)

Min–max

16.9–17.4

26.4–29.6

29.6–31.2

19.1–20.8

25.8–28.5

12.8–14.2

Abauntzb

Mean(n)

18.2

(6)

32.6

(4)

28.5

(2)

31.3

(5)

20.4

(5)

46.3

(5)

14.2

(5)

30.4

(6)

13.8

(6)

Min–max

17.1–8.7

31.5–33.5

28–29

30.5–32

19–22

41.5–50

13.8–14.8

29–33

12.5–15.5

Santimam

inenIII–

V(M

ag-Sol)c

Mean(n)

18.5

(8)

31.3

(2)

33.5

(2)

25(1)

30.4

(4)

19.5

(3)

42.3

(6)

12.8

(7)

26(2)

11.1

(2)

Min–max

17–19.7

30.5–32

32.5–34.5

28–31.5

18.2–21

37.5–47.5

11.6–14.5

22–30

9.4–12.7

Gabasad

Mean(n)

19.2

(9)

27.8

(3)

28.7

(1)

31.2

20.5

(3)

46.6

(7)

13.6

(9)

28(4)

12.7

(4)

Min–max

17.8–21.5

27.6–28.1

29–32.3

20–20.9

44.9–49.1

12.8–14.3

23.8–32.3

10.4–13.9

CovaNegra

eMean(n)

29.5

(1)

29.1

(2)

31.8

(1)

21.1

(1)

44.2

(1)

12.2

(1)

Min–max

29–29.2

Chaves

fMean(n)

19(4)

31(2)

29(1)

31(1)

20.15(2)

13.13(3)

28(1)

13.3

(1)

Min–max

18.4–19.6

30.5–31.5

30.5–31.5

19.8–20.5

12.3–14

Bolinkoban.III–

VI(M

ag-G

ra)g

Mean(n)

18.8

(2)

29.2

(3)

19.8

(1)

40.8

(5)

12.7

(7)

28.5

(5)

12.7

(5)

Min–max

18.3–19.3

28–30.5

31–47

11.9–13.2

26–31.5

12–13.5

LumentxanIV

.V.VI(M

ag-Sol)g

Mean(n)

17.4

(1)

29.7

(3)

26.8

(2)

22.5

(1)

10.7

(1)

Min–max

29–30.5

25.5–28

CuevaMillanh

Mean(n)

17.8

(7)

30.4

(1)

20.5

(1)

12.1

(1)

27.3

(3)

12.1(4)

Min–max

17–19.2

25.7–29

11.6–13

Ekaini

Mean(n)

17.6

(8)

31(1)

31.5

(3)

31.5

(5)

20.3

(3)

46.8

(10)

13.7

(10)

30.5

(3)

13.2

(3)

Min–max

17.3–18.3

30.5–33

30–34

19.7–21.2

42.5–49.5

13–14

30–31

13–13.5

JouPuertaj

Mean(n)

19.3

(1)

32.2

(2)

27.22(3)

Min–max

31–33.4

26.4–28.9

Naranco

jMean(n)

17.46(4)

30.42(7)

26.3

(10)

Min–max

16.3–18.9

29.5–33.4

25.36–27.92

Lezetxikik

Mean(n)

18(19)

33.3

(2)

27.9

(6)

Min–max

16.4–19

32.1–34.4

26.3–29.5

Urtiagak

Mean(n)

18.9

(16)

29.9

(8)

Min–max

15.3–20.6

27.6–32.8

Amaldal

Mean(n)

17.8

(3)

28.75(2)

Min–max

17.3–18.5

28.5–29

LaPalomam

Mean(n)

16.7

(2)

31.5

(3)

28.5

(1)

Min–max

14.7–18.7

31–32

Valdegoban

Mean(n)

30.7

(8)

31.8

(19)

26.3

(27)

Min–max

28.8–33

24.6–38.9

23.2–28.5

Notes:m3.L,lower

thirdmolar;Hum.humerus;Ra.,radius;Tib.,tibia;Ast.,astragalus;1stPha.,firstphalanx;2ndPha.,secondphalanx;L,maxim

um

length;W,maxim

um

width;DW,distalwidth;PW,

proxim

alwidth.Mag,Magdalenian;Sol,Solutrean;Gra,Gravettian.

aThisstudy;bAltunaetal.(2002);

cCastanos(1984);

dBlasco-Sancho(1995);

ePerez-Ripoll(1977);

f Castanos(1993);

gCastanos(1986);

hPerez

LegidoandCerdeno(1992);

i AltunaandMariezkurrena(1984);

j Alvarez-Lao

(2014);

kAltuna(1972);

l Altuna(1990);

mCastanos(1980);

nArceredillo

etal.(2011).

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Table

7.

SelectedcomparativemeasurementsonEquuspostcranialelem

ents.

Site

Taxa

(mm)

1stPha.L

1stPha.PW

1stPha.DW

2ndPha.L

2ndPha.PW

2ndPha.DW

3rd

Pha.LM

3rd

Pha.W

LosRincones

aEquusferus

Mean(n)

78.8

(4)

49.3

(3)

44.1

(3)

45.8

(5)

49.9

(4)

44.9

(5)

58.8

(1)

70.7

(1)

Min–max

75.2–84.1

47.4–50.3

43.7–44.7

41.9–48.1

48.8–51.6

41.4–48.7

Cuetodela

Minab

Equusferus

Mean(n)

84.3

(2)

58(1)

47(2)

50.2

(5)

54.4

(4)

48.9

(4)

48.25(4)

83.5

(1)

Min–max

80–88.5

45–49

46–51

50.5–58

47–53.5

44–54

Santimam

inec

Equuscaballus

Mean(n)

87.8

(3)

55.1

(4)

46.3

(3)

50.6

(5)

55.3

(5)

50(4)

Min–max

83.9–93

54.5–59.4

44–48.6

50–51.2

53.6–58.6

48.6–51.9

Lumentxac

Equuscaballus

Mean(n)

88(2)

59.8

(2)

50.3

(2)

50.3

(2)

56.3

(2)

53(2)

68.5

(2)

84.5

(2)

Min–max

84.5–91.5

59–60.5

50–50.5

48.5–52

54.5–58

52–54

68–69

83.5–84.5

Atxuric

Equuscaballus

Mean(n)

82(1)

54.5

(1)

47(1)

Min–max

Bolinkobac

Equuscaballus

Mean(n)

42(1)

49.1

(4)

56.3

(4)

48.8

(4)

66(1)

78.5

(1)

Min–max

47.5–50

54–58.5

43.5–51

San

Lorenzo

IIc

Equuscaballus

Mean(n)

42.5

(1)

47(1)

42.5

(1)

Min–max

AbricRomanıd

Equuscaballus

Mean(n)

85.8

(1)

52.6

(1)

43.3

(1)

Min–max

LosCasares

eEquuscaballuscasarensis

Mean(n)

77.3

(2)

46(1)

37(1)

Min–max

76.5–78

E.caballusgallicuse

Equuscaballusgallicus

Mean(n)

84.7

(25)

60.1

(25)

50(25)

Min–max

79–89

57.6–66

46.5–53

E.cfgallicuse

Equuscf

gallicus

Mean(n)

81.6

(47)

57.1

(47)

47.4

(47)

Min–max

74.5–91

53.5–65.5

42–53.5

UrtiagaDf

Equuscaballus

Mean(n)

82(1)

54.5

(1)

45(1)

50.65(2)

57(2)

51.75(2)

50.5

(2)

77(2)

Min–max

49–52.3

55–59

49.5–54

50–51

76–78

Aitzbartef

Equuscaballus

Mean(n)

55(1)

75.7

(1)

Min–max

Gruta

deFontainhas

gEquuscaballusantunensis

Mean(n)

79.6

(4)

52.8

(3)

41.7

(3)

46.2

(1)

48(1)

43(1)

50.5

(2)

70.75(2)

Min–max

78.5–82

52–54

41–42

47–57

70–71.5

Algar

deJoao

Ram

osg

Equuscaballusantunensis

Mean(n)

82(2)

52.75(2)

42.5

(1)

49.5

(1)

54(1)

45(1)

53.5

(2)

70.5

(2)

Min–max

82

52.5–53

42.5

53–54

69–72

PdreiradeSalem

asg

Equuscaballusantunensis

Mean(n)

87(3)

50.85(2)

42(2)

46.5

(1)

50(1)

47(1)

Min–max

84.5–89

50–51.7

41–43

Parpallo

hEquuscaballus

Mean(n)

83(12)

50.4

(11)

40.8

(20)

47.1

(29)

49(28)

44.5

(30)

Min–max

75–89

47–51

37.4–44.1

44–50.5

43–51.5

40.5–49.2

CovaNegra

iEquuscaballus

Mean(n)

51.6

(2)

55.1

(2)

53(2)

50(1)

74.8

(1)

Min–max

49.2–54

51.3–58.9

52.1–53.8

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bear), Canis lupus (grey wolf), Panthera pardus spelaea

(Ice Age leopard) and Lynx sp.

Abundant microfaunal taxa are also present, including

the insectivoreCrocidura sp., the rodentsMicrotus arvalis,

Microtus agrestis, Chionomys nivalis, Arvicola terrestris,

Terricola sp., Pliomys lenki and Apodemus sp., and the

lagomorph Lagomorpha indet., as well as Aves, squamates,

the tortoise Testudo hermanni and fish. This small-

vertebrate association is typical of the Late Pleistocene of

the Iberian Peninsula (Cuenca-Bescos et al. 2010; Sauque

and Cuenca-Bescos 2013; Sauque et al. 2014).

4.1. Order Artiodactyla Owen, 1848

4.1.1. Family Cervidae Goldfuss, 1820

Cervids are scarce at Los Rincones, representing just 6.3%

of the ungulates. Although they only form a small

proportion of the total, their presence is interesting as it

indicates a woody environment (Table 1; Figure 2).

4.1.1.1. Cervus elaphus Linnaeus, 1758. Thirteen

remains were recovered, representing 2.1% of the

ungulates. The dental remains are scarce; only two incisors

were recovered. The most abundant remains are the

phalanges (six), two first phalanges, two second and two

third phalanges; also there are one metatarsus and one tibia

(Figure 3).

The metatarsus (Ri10/O-13/172) presents a clear

separation between grand cuneiform facet and escafocu-

boides facet such as in Cervus elaphus while in Dama

dama they appear to meet (Lister 1996). Also, Ri10/O-

13/172 shows a single large foramen typical of Cervus

elaphus while Dama dama presents a network of pores

(Lister 1996). First phalanx (Ri10/P13/1, Ri10/O-13/81) of

Los Rincones shows a posterior articular facet with a mid-

groove which is typical of Cervus elaphus (Lister 1996).

The dimensions of the postcranial material that was

found correspond to a medium-sized deer, one larger than

the deer from Cueva Camino and Cova Negra (MIS5), but

somewhat smaller than those from the north of the Iberian

Peninsula (the sites of Lumentxa, Santimamine, Urtiaga D,

Labeko Koba, La Paloma, El Miron, Morın and Abauntz)

that correspond to the last cold stage (MIS3 and MIS2).

The deer from Los Rincones is similar in size to deer from

Gabasa, Parpallo and Nerja (MIS3 and MIS2), which are

found in a Mediterranean climatic environment (Table 2).

The larger size of these deer from the cold stages was

already pointed out by Mariezkurrena (1983). This greater

size could be related to Bergmann’s rule (cold climates

select for the survival of large-sized animals), or it could

be due to differences in food quality, the high quality of the

vegetation during the cold periods allowing ruminants

such as deer to attain large sizes (Guthrie 1990).Gabasaj

Equuscaballus

Mean(n)

82.4

(4)

51(3)

41.5

(6)

48.3

(2)

49.1

(2)

46(2)

68.4

(1)

78.8

(1)

Min–max

80–88

46.4–54.6

39–43.9

47.5–49

46.6–51.5

43–49

Abauntzk

Equusferuscf.gallicus

Mean(n)

82.5

(3)

56.5

(1)

48.5

(2)

49(1)

54.5

(1)

52(1)

64(2)

77(1)

Min–max

78–85

48–49

63–65

Hornosdela

Penal

Equuscaballus

Mean(n)

80(1)

55(1)

42(1)

47(1)

57(1)

53(1)

Min–max

TitoBustillo

mEquusferusgallicus

Mean(n)

83(2)

55(2)

47.5

(2)

49.5

(1)

57(1)

49(1)

Min–max

53–57

42–53

Equusferusprzew

alskiiextantn

Equusferusprzew

alskii

Mean(n)

78.1

(27)

49.6

(27)

43.8

(27)

44.4

(85)

49(85)

43.3

(85)

64.6

(47)

70.7

(47)

Min–max

74–84

46–55

40.5–47.1

40–50

45–54

39.5–52

56–78

63.5–78

Notes:1stPha.,firstphalanx;2ndPha.,secondphalanx;3rd

Pha.,thirdphalanx;L,maxim

um

length;W,maxim

um

width;DW,distalwidth;PW,proxim

alwidth.

aThisstudy;bCastanos(1982);

cCastanos(1986);

dSanchez

(1990);

eAltuna(1973);

f Altuna(1972);

gCardoso

andEisenmann(1989);

hDavidson(1989);

i Perez-Ripoll(1977);

j Blasco-Sancho(1995);

kAltuna

etal.(2002);

l Yravedra

(2010);

mAltuna(1976);

nEisenmannonlinedata.

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4.1.1.2. Capreolus capreolus Linnaeus, 1758. Twenty-

six remains were recovered, representing 4.2% of the

ungulates. Cranial and postcranial remains were found

(Figure 3). The minimum number of individuals is two,

one adult and one subadult of roughly 19–20 months.

Among the remains recovered, an antler (Ri10/

O13/230) is particularly notable for its taxonomic value.

This is composed of a central branch that presents a tip that

grows forwards and upwards in its upper part. The rear tip

projecting backwards situated a third of the way along the

central branch is not observed, since the individual is

juvenile. It presents great development of the rosettes, with

pearled patterning. These characteristics are typical of

Capreolus capreolus (Sempere et al. 1996; Mateos-

Quesada 2011) (Figure 3).

The remains that could be measured have been

compared with the measurements from the bibliography

(Table 3). The remains recovered from Los Rincones are

similar in size to the roe deer from Gabasa (MIS3) but

smaller than those from the north of the Iberian Peninsula

(the sites of Jou Puerta, Urtiaga, La Riera, Santimamine

and Lumentxa) that correspond to the last cold stage

(MIS3 and MIS2) (Table 3).

The presence of Capreolus capreolus is interesting

since the species does not occur frequently in the sites of the

Quaternary of western Europe (Altuna 1972; Lister 1986;

Figure 3. (a) Right tibia Cervus elaphus (Ri10/O-13/14). (b) Right metatarsus Cervus elaphus (Ri10/O-13/172). (c) Left radiusCapreolus capreolus (Ri10/O-13/235). (d) Metacarpus Capreolus capreolus (Ri10/O-14/87). (e) Left antler Capreolus capreolus (Ri10/O-13/4; Ri10/O-14/45). (f) Right antler Capreolus capreolus (Ri10/O-13/230). (g) Right tibia Capreolus capreolus (Ri10/O-13/75). (h)Left mandible m2–m3 Capreolus capreolus (Ri10/O-13/73). (i) Right humerus Rupicapra pyrenaica (Ri10/N10/34). (j) Left tibiaRupicapra pyrenaica (Ri10/N10/188). (k) Left lower third molar Rupicapra pyrenaica (Ri10/N10/72). (l) Right lower third molarRupicapra pyrenaica (Ri10/N10/71).

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Alvarez-Lao 2014). In the Iberian Peninsula, it has mainly

been recovered in northern regions, though its presence

never represents more than 20% of the NISP of the

ungulates of the sites in question (see Appendix 1 %NISP).

4.1.2. Family Bovidae Gray, 1821

Bovids are predominant in the ungulate association from

Los Rincones, constituting 91.2% of the NISP of ungulates.

The most abundant taxon is the Iberian wild goat (Capra

pyrenaica), followed by much lower percentages of the

Pyrenean chamois (Rupicapra pyrenaica) and the aurochs

(Bos primigenius) (Figure 2; Table 1).

4.1.2.1. Bos primigenius Bojanus, 1827. The remains of

Bos primigenius are scarce, representing just 0.7% of the

NISP of ungulates.Moreover, they seem to belong to a single

individual, since theywere collectedwithin a very small area,

are similar in size and articulate with one another. The

remains recovered belong to the postcranial skeleton: a talus

bone, a metatarsus, a scaphocuboid and a third phalanx

(Figure 4).

The distinction between Bos and Bison is complex, and

the absence of the cranial or dental remains that are best

for distinguishing them obliged us to use a variety of

morphological and metric criteria. In spite of the

complexity, the distinction is relevant because the taxa

Figure 4. (a), (b) Right metatarsus Bos primigenius (Ri10/O-13/147). (c) Right astragalus Bos primigenius (Ri10/N10/14). (d) Firstphalanx Equus ferus (Ri10/N10/31). (e) First phalanx Equus ferus (Ri10/N11/1) (f) First phalanx Equus ferus (Ri10/O-13/71). (g) Thirdphalanx Equus ferus (Ri10/N10/64). (h) Second phalanx Equus ferus (Ri10/O-14/53). (i) Second phalanx Equus ferus (Ri10/N10/98). (j)Second phalanx Equus ferus (Ri10/N10/150). (k) Third phalanx Equus hydruntinus (Ri10/M10/10). (l) Third phalanx Equus hydruntinus(Ri10/N10/57).

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indicate different environments: Bison priscus is associ-

ated with steppe regions (Brugal 1985), whereas Bos

primigenius is found both in open and woodland areas

(Ekstrom 1993).

Only the proximal part of the metatarsus is preserved,

where the proximal articular facets corresponding to the great

cuneiform and the scaphocuboid can be seen to be slightly

separated, which is a trait characteristic of Bos, whereas in

Bison they are joined (Brugal 1985; Gee 1993). Moreover,

these facets are asymmetrical,which differentiates them from

Bison (Alvarez-Lao and Garcıa-Garcıa 2006). Another

criterion that supports an ascription of the fragment to Bos

is the lack of the medial tubercle (Gee 1993).

With the talus bone, we have followed morphological

and metric criteria. In the case of Bos, the morphology of

the plantar groove separating the articular surfaces of the

calcaneus and the scaphocuboid displays an angle of 908both in the plantar and medial area of the bone and on the

lateral margin, whereas in Bison it follows a gently arching

course that ends in an open angle (Altuna 1972; Buitrago-

Villaplana 1992; Gee 1993; Sala et al. 2010). Specimen

Ri10/N10/14 clearly presents straight angles; furthermore,

it does not display an articular facet for the scaphocuboid,

which is a character typical of Bison (Bibikova 1958;

Altuna 1972; Buitrago-Villaplana 1992; Gee 1993; Sala

et al. 2010). The metric criteria applied to the talus are the

index of the lateral articular facet and that of the distal

trochlea (Bibikova 1958). Table 4 presents the values for

these indices, as given by various authors. It can be seen

that there is an overlap between the two species in these

values (Sala et al. 2010), though values above 56.6 for the

articular facet index have only been published for Bos

(Stampfli 1963). For the distal trochlea index, the overlap

is substantial, but values below 81.2 correspond to Bison

(Altuna 1972), whereas values above 100 correspond to

Bos (Stampfli 1963). The value of the lateral articular facet

index for specimen Ri10/N10/14 is 60, and the distal

trochlea index has a value of 110. Both these values place

it clearly within the range of variability of Bos and exclude

it from that of Bison. In short, metric and morphological

characteristics enable us to assign the large-sized bovid

remains from Los Rincones to the species Bos primigen-

ius, which inhabited Europe until historical times

(Degerbøl and Fredskild 1970).

4.1.2.2. Capra pyrenaica (Schinz, 1838). The Iberian

wild goat is thepredominant taxonat the site ofLosRincones;

528 remains were recovered, which represent 85.9% of the

totalNISPofungulates (Table 1). Theage ofdeath reveals the

presence of 20 individuals at the site. Grouped by age, these

are one neonate individual, three juvenile individuals, seven

subadults, five adults and four old individuals.

The sample includes almost all the anatomical

elements, especially bones of the extremities, but also

cranial remains, mainly isolated teeth, axial elements,

scapulae and pelvis (Figures 5–7).

The taxonomy of the genus Capra is based principally

on the morphological characteristics of the horns. In the

absence of these elements, M3 has been one of the most-

used elements, in particular the presence or absence of the

metastylar wing and the width of the interstylar surface

(Cregut-Bonnoure 1992). On the basis of these characters,

Cregut-Bonnoure (1992, 2006) proposed that Capra

pyrenaica is derived from a common ancestor with the

Caucasian goat (Capra caucasica praepyrenaica), which

would have arrived at the Massif Central in France in the

course of the Eemian. Once there, it would have evolved

into Capra pyrenaica, moving towards the south of France

and reaching the Pyrenees in the Magdalenian. On this

theory, from the Pyrenees it would have subsequently

colonised the Iberian Peninsula. Currently, in the light of

the presence of Capra in Iberia for more than 100 kyr

(Altuna 1992), the high variability of the distinguishing

morphological characters proposed by Cregut-Bonnoure

(Magniez 2009) and the results of molecular studies

showing the kinship of Capra ibex and Capra pyrenaica

(Pidancier et al. 2006; Urena et al. 2011), the hypothesis of

the Caucasian origin of Capra pyrenaica has rather lost

plausibility (Garcıa-Gonzalez 2011). Although the exact

taxonomic classification of Iberian goats is far from being

resolved, however, it does seem very likely that Capra

pyrenaica was already present and differentiated in the

Late Pleistocene. Accordingly, it would make sense to

ascribe the remains from Los Rincones to this taxon,

irrespective of possible hybridisations with Capra ibex that

might have taken place in the northeast of the Iberian

Peninsula and south of France during this period (Garcıa-

Gonzalez 2012).

The comparative data for Capra pyrenaica presented

in Table 5 suggest the existence of two groups (clades?) in

the Late Pleistocene of the Iberian Peninsula, separable by

their dimensions: one northern group, characterised by

their large size, and another southern group that is smaller

in size (Table 5; Figure 8). The Capra pyrenaica remains

from Los Rincones would fall within the range of variation

of the first group. This size gradient would have been

reproduced in the Holocene (Castanos 2004) and at

present, for this and other characteristics have been

recognised in the present-day subspecies (Cabrera 1911).

Do the differences in size among Iberian goats in the Late

Pleistocene correspond to phylogenetic differences? The

current subspecific differentiation of Capra pyrenaica is a

matter of controversy (Acevedo and Cassinello 2009), and

the systematics should probably be revised in the near

future, even at the specific level (Garcıa-Gonzalez 2011).

The north–south size gradient of the Iberian goats does not

seem to be a temporary case of Bergmann’s rule, but seems

rather to be due to the possible abundance of trophic

resources in two contrasting environments: the Eurosiber-

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ian (temperate and fertile) and the Mediterranean (arid and

poor). The increase in size that took place in Capra

pyrenaica in the interglacial period (MIS5) at Cova Negra

(Perez-Ripoll 1977; Table 5; Figure 8) or during the

Holocene (Garcıa-Gonzalez 2012) would support the

hypothesis of the abundance of resources.

4.1.2.3. Rupicapra pyrenaica Linnaeus, 1858. As

regards the Pyrenean chamois, 29 remains were recovered,

representing 4.7% of the NISP of the ungulates at the site

(Table 1; Figure 2). Both appendicular and cranial

elements were found. Among the cranial material, three

m3, one m2, one M2 and one M3 were recovered (Figure 3).

The m3 presents an external wall with a metastyle and

parastyle and an oval talonid with a practically polygonal

section. The distal edge is straight and not convex (Prat

1966).

The talus bones, with four elements recovered, and the

phalanges, with six elements recovered, are the most

numerous elements of Rupicapra pyrenaica at the site, and

their measurements have been compared with a sample of

Rupicapra pyrenaica from the Pleistocene of the Iberian

Peninsula (Table 6). The chamois from Los Rincones is

similar in size to that from Cueva Millan and smaller than

those from the sites in the north of the Iberian Peninsula

(Santimamine, Abauntz, Bolinkoba, Lumentxa, Ekain,

Naranco, Aitzbitatre, Lezetxiki, Urtiaga, Amalda,

La Paloma and Valdegoba) that correspond to the last

cold stage (MIS3 and MIS2). Furthermore, it is smaller

than those from the Ebro Valley sites of Gabasa and

Chaves (MIS3 and MIS2, respectively) (Table 6).

The genus Rupicapra (chamois) is a small-sized

mountain goat-antelope that inhabits Europe and western

Asia. The genus comprises two different species:

Rupicapra rupicapra, present in the Alps, the Balkans

and the Caucasus; and Rupicapra pyrenaica, present in the

Iberian Peninsula and Italy (Lovari and Scala 1980;

Nascetti et al. 1985; Perez et al. 2002).

The time of divergence between Rupicapra rupicapra

and Rupicapra pyrenaica has been estimated on the basis

of their genetic distance, but different methods yield

different ages. Molecular clocks show a separation that

took place at ca. 1.7Ma (Rodrıguez et al. 2010), whereas

phylogenetic analysis based on Y chromosomes has given

a more recent age of 655 ka (Perez et al. 2011). These

genetic studies imply that the chamois recovered from the

Late Pleistocene of the Iberian Peninsula belong to the

species Rupicapra pyrenaica (Alvarez-Lao 2014).

4.2. Order Perissodactyla Owen, 1848

4.2.1. Family Equidae Gray, 1821

The systematics of horses is under revision. To underline

the difference between domestic and wild species, the

Figure 5. Remains of Capra pyrenaica. (a) Left maxilla P2–M3 (Ri10/N10/41). (b) Right mandible of a juvenile p2-p4 (Ri10/O-13/92).(c) Left mandible p3–m3 (Ri10/N10/117). (d) Left M3 (Ri10/O-13/126). (e) Right M3 (Ri10/N10/83). (f) Right M3 (Ri10/GL9/9). (g)Right M3 (Ri10/O-13/201). (h) Right M3 (Ri10/O-13/203). (i) Right mandible p3–m3 (Ri10/O-13/27). (j) Atlas (Ri10/O-13/118). (k)Atlas (Ri10/N11/2). (l) Atlas (Ri10/O-13/82).

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International Commission on Zoological Nomenclature in

2003 proposed that the name of a wild species – in the

case of the horse Equus ferus Boddaert, 1785 – should

take precedence over the name of the domesticated

species. However, the name of the domesticated species,

Equus caballus Linnaeus, 1758, continues to be valid

(Gentry et al. 2004; Conti et al. 2010). Accordingly, we

here use the designation Equus ferus to refer to the horses

of the Pleistocene even though they may previously have

been classified as Equus caballus.

It is difficult to distinguish between the different

species of equids on the basis of morphological characters

alone (Eisenmann 1986; Dive and Eisenmann 1991;

Orlando et al. 2009). In the present case, this is particularly

difficult due to the lack of preserved cranial remains,

which are those that provide most systematic information.

Figure 6. Remains of Capra pyrenaica. (a) Right humerus (Ri10/N11/41). (b) Right humerus (Ri10/O-14/12). (c) Right humerus (Ri10/J10/1). (d) Left humerus (Ri10/O-13/179). (e) Left radius (Ri10/O-13/165). (f) Left radius (Ri10/N10/208).(g) Right metacarpus (Ri10/N10/195). (h) Right metacarpus (Ri10/O-13/199). (i) Left metacarpus (Ri10/N19/151). (j) Left metacarpus (Ri10/M9/17). (k) Leftmetacarpus (Ri10/GL7/19; Ri10/GL7/2; Ri10/GL7/5). (l) Right metacarpus (Ri10/O-13/184).

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For this reason, the remains have been classified by means

of a biometric study.

4.2.1.1. Equus ferus Linnaeus, 1758. A fragment of

metapod and phalanges were recovered from the site: four

first phalanges, three of which are posterior, six second

phalanges and a third phalanx (Figure 4). The phalanges

were distinguished as posterior or anterior in accordance

with the criteria proposed by Prat (1957), Eisenmann and

De Giuli (1974) and Dive and Eisenmann (1991) (Figures 9

and 10).

As a whole, the horse from Los Rincones is small in

size compared with the other horses of the Late

Pleistocene of the Iberian Peninsula, and especially

compared with the horses of the Upper Palaeolithic of

the Basque Country from sites such as Atxuri, Santima-

mine and Lumentxa (Castanos 1984) (Table 7; Figure 11).

Its size is similar to that of the horse from Fontainhas,

which forms part of the subspecies Equus ferus antunesi,

characterised by its small dimensions (Cardoso and

Eisenmann 1989), and it is also close in size to the

subspecies Equus ferus casarensis described at Los

Casares (Altuna 1973). The measurements of the horse

from Los Rincones are very similar to the current

population of Equus ferus przewalskii (Table 7). The third

phalanx from Los Rincones presents a small maximum

width, which may be related to the type of land over which

Figure 7. Remains ofCapra pyrenaica. (a) Lumbar vertebras and sacrum (Ri10/O-13/90, Ri10/O-13/93). (b) Left scapula (Ri10/GL8/5).(c) Left scapula (Ri10/O-13/219). (d) Left tibia (Ri10/GL9/12). (e) Left tibia Ri10/M9/19). (f) Right tibia (Ri10/O-13/102). (g) Right tibia(Ri10/M10/3). (h) Right tibia (Ri10/P13/5). (i) Right femur (Ri10/O-13/218; Ri10/O-13/234; Ri10/O-14/15).

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the animals moved; large ungual phalanges are associated

with soft ground, whereas narrow ones indicate an adaptation

for movement over craggy reliefs (Eisenmann 1984).

Despite its small size, the horse from Los Rincones

does not fall within the range of variation for the

measurements of Equus hydruntinus; as can be seen in

Figure 9. It lies within the range of variation for Equus

ferus przewalskii as well as for the Equus ferus of the Late

Pleistocene.

4.2.1.2. Equus hydruntinus Regalia, 1907. Two remains

of Equus hydruntinuswere recovered, which correspond to

two third phalanges (Figure 4). This taxon is an indicator

for conditions of aridity and open spaces (Burke et al.

2003; Orlando et al. 2006; Feranec et al. 2010).

Equus hydruntinus was described by Regalia (1907) in

the Grotta Romanelli. As it showed characteristics similar

to the onager, he designated as Equus (Asinus) hydruntinus

(Conti et al. 2010). Subsequent studies of the systematic

Figure 8. Graph representing the maximum value, minimum and the mean of Capra pyrenaica from different Pleistocene sites andextant animals. Mtcp., metacarpus; Mtt., metatarsus; Tib., tibia; Hum., humerus; L, maximum length; DW, distal width.

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position of this taxon classified it as an ass (Stehlin and

Graziosi 1935; Gromova 1949), a zebra (Davis 1980) and a

stenonian form (Forsten 1999). In recent years, however,

the study of material from Crimea has brought to light the

phylogenetic affinity between Equus hydruntinus and

Equus hemionus (Burke et al. 2003), confirmed by DNA

studies on remains from Crimea and Iran (Orlando et al.

2006). Equus hydruntinus was the last species of the

Pleistocene megafauna to go extinct in the Iberian

Peninsula, as there is evidence of its presence as late as

the 16th century (Quesada and von Lettow-Vorbeck 1992).

As there are no available biometric data for the third

phalanges of Equus hydruntinus and these display

similarly proportioned bones of the extremities to onagers

(Eisenmann and Baryshnikov 1995; Forsten 1999; Burke

et al. 2003; Orlando et al. 2006), we have used the

measurements from modern-day onagers taken from

Eisenmann’s collection (online data at http://www.

vera-eisenmann.com) for the purpose of biometric

comparison.

The bones of the extremities of Equus hydruntinus are

slender, and the third phalanges are pointed (Davis 2002).

Figure 9. Graph representing greatest length versus the proximal width of the first phalanx of fossil and extant Equus species. Speciesdata points and corresponding regression lines are plotted together.

Figure 10. Graph representing greatest length versus the greatest width of the third phalanx of fossil and extant Equus species. Speciesdata points and corresponding regression lines are plotted together.

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The third phalanges from Los Rincones with a size similar

to Equus hemionus (Figure 10) and a sharp-pointed end

belong to Equus hydruntinus.

5. Faunal composition

5.1. Analysis of the faunal composition in theEuropean context of MIS3–MIS2

The faunal composition, expressed as the percentage of

each herbivore species at the site of Los Rincones, was

compared with that from 89 European sites, with a total of

127 levels with different chronologies, but all of them

belonging to the isotope stages MIS3 and MIS2. Of these

sites, 56 are located in the Iberian Peninsula, and 33 in the

rest of Europe, both in western and central parts (France,

United Kingdom, Germany, the Czech Republic and

Switzerland; see Appendix 1). To compare the faunal

associations present at each site, the NISP was calculated

for the herbivores in each case. This parameter is

considered the most reliable (Lyman 1994b), as there is

no taphonomic bias towards any of the species in the

assemblages (Alvarez-Lao 2014). In order to minimise

the taphonomic biases that might be produced by the

accumulators of the remains, this study has taken account

Figure 11. Graph representing maximum value, minimum and the mean of Equus from different Upper Pleistocene sites and extantanimals. 1st Pha., first phalanx; 2nd Pha., second phalanx; L, maximum length; PW, proximal width.

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both of sites where the accumulators are hunter-gatherers

(Chaves, Grotte de la Vache, Gabasa, Abric Romanı,

Santimamine and Nerja) and of those where they are

carnivores (Zafarraya, Los Rincones, Zarzamora and Las

Caldas). It has only used assemblages with an NISP

greater than 100, in accordance with the methodology

proposed by Alvarez-Lao (2014). Moreover, certain taxa

have been grouped together, as in the case of Capra

pyrenaica and Capra ibex (C.p/i) and Rupicapra pyrenaica

and Rupicapra rupicapra (R.r/p). We follow Deng et al.

(2011) in the taxonomic nomenclature of Stephanorhinus

hemitoechus.

The CA has allowed us to place the sites with a similar

taxonomic composition close to one another. At the same

time, the taxa with a similar distribution throughout the

sample should also be placed close to one another.

The DCA shows a clear separation of the sites of

central and Western Europe in relation to those of the

Iberian Peninsula. The sites of central and western Europe

(black dots) are situated on the right of the graph and are

characterised by the presence of the faunas of the

‘Mammuthus-Coelodonta faunal complex’, comprising

Mammuthus primigenius, Coelodonta antiquitatis, Mega-

loceros giganteus, Rangifer tarandus, Saiga tartarica,

Equus ferus and bovids (Bison priscus/Bos primigenius)

(Guthrie 1990; Kahlke 1999; West 2000). The mammoth

and the woolly rhinoceros have been recovered in the

Iberian Peninsula at 25 and 23 sites, respectively (Alvarez-

Lao and Garcıa 2011, 2012). However, none of the Iberian

sites presents this fauna at percentages that place it within

the area of sites with the typical faunal association of the

Mammuthus–Coelodonta complex. An exception among

the sites in the west is La Grotte de la Vache in the French

Pyrenees. Its position on the left of the graph is due to the

abundance of Capra pyrenaica, which represents 88% of

the NISP of the ungulates (Pailhaugue 1995) (Figure 12).

The sites of the Iberian Peninsula (grey dots) are

located on the left of the graph, with the exception of

Urtiagako Leizea. This site is situated on the right due to

its high numbers of reindeer, Rangifer tarandus (Altuna

1984), which bring it closer to the French sites of the

Massif Central, where the reindeer is very abundant

(Delpech 1983, 1990). The sites of the Iberian Peninsula

are characterised by the presence of taxa associated with

more temperate environments than the cold European

faunas. Such taxa are Capra pyrenaica, Cervus elaphus,

Figure 12. Ordination of palaeontological sites and species along the first two axes of the DCA on the basis of their ungulatecomposition. Grey, Iberian Peninsula sites; black, Western and Middle European sites; star, Los Rincones. Taxa abbreviations: R.t.,Rangifer tarandus; C.e., Cervus elaphus; M.g.,Megaloceros giganteus; C.c., Capreolus capreolus; C.i./p., Capra ibex/pyrenaica; R.r./p.,Rupicapra rupicapra/pyrenaica; S.t., Saiga tatarica; Bov., Bovinae indet.; S.s., Sus scrofa; E.f., Equus ferus; E.h., Equus hydruntinus; C.a., Coelodonta antiquitatis; S.h., Stephanorhinus hemitoechus; M.p., Mammuthus primigenius. Site abbreviations: Zaf, Zafarraya; Ne,Nerja; Vg, Vanguard Cave; Go, Gorham’s Cave; G.V, Grotte de la Vache; Ab R, Abric Romanı; Ge, Cova del Gegant; Mo, Cueva Morın;La, Labeko Koba; Lez, Lezika; Ar, Arbreda; Zar, Zarzamora; Du, L’abri Duruthy; Mor, Morancourt; Srb, Srbsko Chulum-Komin Cave; H.S, Hohle Stein; Pe, Perick Cave; Vo, Vogelherd (IV–V); Ly, Lynford; Gei, Geissenklosterle; Hor, L’Hortu; U.L, Urtiagako Leizea; Fac,Abri du Facteur; Com, Roc de Combe; Lau, Laugerie Haute Ouest.

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Capreolus capreolus, Rupicapra pyrenaica, Sus scrofa and

Stephanorhinus hemitoechus (steppe rhinoceros)

(Figure 12).

The DCA shows an oblique general gradient of

increasing latitude and decreasing temperature, running

from the upper right to the lower left corner. In the upper

right corner are the sites of Lynford (UK) and

Geissenklosterle, Hohle Stein, Vogelherd and Perick

Cave (Germany), sites characterised by the presence of

larger-sized taxa of the Mammuthus–Coelodonta com-

plex, such as Mammuthus primigenius, Coelodonta

antiquitatis and Megaloceros giganteus. In the lower

right corner of the graph, we find the sites located in the

Massif Central in France, such as Abri Pataud, Roc de

Combe, Laugerie Haute-Ouest and Abri Facteur among

others, which are characterised by a predominance of

Rangifer tarandus and Saiga tatarica. On the far right of

the horizontal ordination of the graph, we find both French

sites (Morancourt, Duruthy and Hortus) and Middle

European sites (Srbsko Chulum-Komin Cave from Czech

Republic), characterised by the presence of Equus ferus

and large bovids. Close to these sites are Iberian sites such

as l’Arbreda, Abric Romanı and Cova del Gegant

(Catalonia), and Cueva Morın, Labeko Koba and Lezika

(Basque-Cantabrian region). Both the northern part of

Catalonia and the Basque Country functioned as faunal

corridors in the Pleistocene (Alvarez-Lao and Garcıa

2011), so it is hardly surprising that these areas are the

most similar to European ones. Despite their similarity to

European sites, these sites exhibit a greater proportion of

Equus hydruntinus, associated with arid environments

(Burke et al. 2003; Feranec et al. 2010). The sites of the

Iberian Peninsula hardly show any variation with respect

to axis 2 (horizontal alignment), whereas their distribution

in relation to axis 1 marks a transition from the

Mediterranean conditions of the sites in the south (Nerja,

Zafarraya, Gorham’s Cave and Vanguard Cave) on the left

of the graph, with a high proportion of Capra pyrenaica,

towards more Atlantic, mid-European conditions, exem-

plified by the sites of the Basque Country and Catalonia on

Figure 13. Ordination of palaeontological sites and species of the Iberian Peninsula along the first two axes of theDCAon the basis of theirungulate composition. White: Iberian Peninsula sites; Star: Los Rincones. Taxa abbreviations: C.e., Cervus elaphus; C.c., Capreoluscapreolus; C.i./p., Capra ibex/pyrenaica; R.i/p., Rupicapra rupicapra/pyrenaica; Bov., Bovinae; S.s., Sus scrofa; E.f., Equus ferus; E.h.,Equus hydruntinus; S.h., Stephanorhinus hemitoechus. Site abbreviations: Zar, Zarzamora; Bu, Cueva del Buho; Mo, Cueva Morın; Ge,Cova del Gegant; La, Labeko Koba; Fi, Figueira Brava (2–3); Ar, Arbreda; Es, Escoural; Ab R, Abric Romanı; Pe, Cueva del Pendo; Mi,Cueva Millan; Le, Lezetxiki (IIIa); HP, Hornos de la Pena; Ax, Axlor; Am, Amalda; Va, Valdegoba; Ai, Aizbiarte IV; Ab, Abauntz; Er,Cueva Ermita; JP, JouPuerta; Arr, Arrillor; PD, Pego doDiablo; GC,Grota Nova da Columbeira; Co, Covalejos, At, Atxagakoa; Li, Linares;Sa, Santimamine; Lu, Lumentxa n VII; Ca, Los Casares; Cal, Las Caldas; Mie, Pena Miel; Ga, Gabasa 1; Be, Cova Beneito; Vg, VanguardCave; Ne, Nerja; Zaf, Zafarraya.

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the far right of the graph (with the exception of La Grotte

de la Vache). In spite of being located in an area that is

climatically at the boundary between Atlantic and

Mediterranean environments, the site of Los Rincones is

closer to the sites of the south in faunal terms (Figure 12).

5.2. The site of Los Rincones in the Iberian context of

MIS3

The herbivore association present at the site of Los

Rincones was compared with those at 37 sites in the Iberian

Peninsulawith a chronology close toMIS3 (Figure 13). The

CA shows that the sites with taxa associated with open

environments such as Equus hydruntinus, Equus ferus,

Stephanorhinus hemitoechus and the large bovids (Brugal

1985; Burke et al. 2003; Feranec et al. 2010; Sala et al.

2011) are situated in the left part of the graph. These sites

are Cueva de la Zarzamora, CuevaMorın, Cova del Gegant,

Labeko Koba and Figueira Brava. On the right, by contrast,

are the sites of the south of the Iberian Peninsula such as

Zafarraya, Nerja, Vanguard Cave and Cova Beneito, which

are characterised by a great abundance of Capra pyrenaica

associated with rocky areas. Furthermore, it should be

borne in mind that this graph shows a comparison of sites

from MIS3, at which time human hunter-gatherers had not

yet specialised in hunting Capra. This phenomenon

appeared in the Upper Palaeolithic (MIS2) (Freeman

1973; Straus 1987; Gamble 1995). As such, the high

percentages of Capra in the Mediterranean sites cannot be

attributed to the presence of specialised hunters, but rather

to the abundance of these taxa in the area of the

accumulation. In the central part of the graph, we find

mainly Basque–Cantabrian sites under Atlantic influence.

Axis 2 separates sites with woodland taxa such as Cervus

elaphus, Capreolus capreolus and Sus scrofa (Covalejos,

Linares, Lumentxa and Aitbiarte) in the lower part from

sites in the upper central part (Amalda, Valdegoba and

Hornos de la Pena) characterised by the presence of taxa

associated with rocky environments such as the chamois,

but with a more Atlantic climatic affinity than Capra

pyrenaica (Feranec et al. 2010).

Despite currently being located in a geographical

situation close to the boundary between the Mediterranean

and Atlantic climatic regions, in MIS3 the site of Los

Rincones was close to sites such as Zafarraya, Nerja, Cova

Beneito and Vanguard Cave situated in the Mediterranean

climatic region in the south of the Iberian Peninsula and

characterised by a strong presence of Capra pyrenaica, a

species with a high drought-tolerance (Feranec et al.

2010).

The faunal association from los Rincones is similar

to southern Iberian sites (Zafarraya, Nerja and

Vanguard). This fact may reflect similar environmental

conditions, but it can also be due to a bias produced by

the main accumulator agent. In Los Rincones, most of

the ungulates recovered were transported there by Ice

Age leopards, which used the cave as a refuge for

protecting their kills (Sauque et al. 2014). This predator,

as his relative the Panthera uncia (snow leopard), shows

a predilection by preys of small–medium size, especially

goats (see references in Diedrich 2013). This phenom-

enon, added to the placement of the cave in a rocky

wall, may produce an overrepresentation of Capra

pyrenaica. Anyway, although leopards focus mainly on

prey ranging from 20 to 80 kg in weight (Mills and

Harvey 2001), they are not selective hunters and have

the broadest diet of the larger predators with 92 prey

species recorded in sub-Saharan Africa (Mills and

Harvey 2001; Hayward et al. 2006), which may indicate

that a high percentage of Capra pyrenaica is related to

its great abundance in the landscape which surrounds the

cave. However, the low percentage of equids, rhinoceros

and bovids (Bos primigenius and Bison) found in Los

Rincones could not be related to a low abundance of

these taxa in the environment, but to the impossibility of

leopards to drag these preys to their dens because they

weigh more than the range the leopard could hunt.

Therefore, the abundance of these might not reflect the

‘real’ situation surrounding the cave and subsequently

the faunal association would be more similar to the

Iberian sites placed in the inland such as Zarzamora,

Cueva del Buho and Los Casares.

To sum up, the analysis of faunal associations is a great

tool to analyse the environmental patterns, though we have

to take these results carefully, especially when the

accumulator may generate bias in the association.

6. Palaeoenvironmental context

The herbivores at Los Rincones represent various types of

landscape. The horses (Equus ferus and Equus hydrunti-

nus) indicate open environments (Nowak 1999; Blasco

et al. 2011), and Equus hydruntinus is also indicative of

conditions of semi-aridity (Burke et al. 2003; Feranec et al.

2010) (Figure 14). However, Cervus elaphus and

Capreolus capreolus suggest woodland habitats (Delpech

and Prat 1980; Carranza et al. 1991; Mateos-Quesada

2011; Carranza 2011), whereas Bos primigenius can

inhabit both woody environments and plains (Ekstrom

1993). However, the best-represented herbivores both in

terms of NISP and MNI are those associated with high or

medium mountain areas with abrupt reliefs, such as Capra

pyrenaica and Rupicapra pyrenaica (Alados and Escos

1996; Granados et al. 2001; Perez-Barberıa et al. 2010)

(Figure 15). In this context, it is interesting to bear in mind

that the site has also yielded a peripheral bone from the

shell of a Hermann’s tortoise (Testudo hermanni), which is

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Figure 14. In the forefront, a herd of horses (Equus ferus). Behind, to the right, three Equus hydruntinus might be seen in a steppesurrounding Los Rincones cave. In the background, the Moncayo massive (2315m a.s.l) (Illustration Gianfranco Mensi).

Figure 15. Ice Age leopard (Panthera pardus spelaea) chasing a herd of Spanish wild goat (Capra pyrenaica) in the ravine of LosRincones. Behind, the mouth of the cave (1000m a.s.l) (Illustration Gianfranco Mensi).

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a good indicator of Mediterranean conditions (Morales

Perez and Serra 2009).

The great abundance of Capra pyrenaica and the

presence of Rupicapra pyrenaica indicate that in the

vicinity of the site there were rocky mountains. This

landscape would be similar to what we currently find in the

area around the cave, the mouth of which opens at an

altitude of some 1010m in a ravine with rocky walls, ideal

for both species (Figure 1).

7. Conclusions

The ungulate association from Los Rincones provides us

with palaeoenvironmental information about the region

around the Moncayo during MIS3. In spite of its current

importance as a boundary between Mediterranean and

Eurosiberian climatic regions, until now there were no

data available for a reconstruction of its landscape during

the Late Pleistocene.

The great diversity of ungulates present at the site of Los

Rincones, with seven species – Cervus elaphus,Capreolus

capreolus, Rupicapra pyrenaica, Capra pyrenaica, Bos

primigenius, Equus ferus and Equus hydruntinus –

associated with different environments, indicates a high

taxonomic and environmental diversity corresponding both

to the situation of the massif as an ecotone and to the

particular location of the site, not very different from

present-day conditions. Moreover, both Equus hydruntinus

andBos primigenius are species that are now extinct but that

survived into historical times in Europe.

The ungulate association from Los Rincones is

dominated by Capra pyrenaica. Furthermore, this taxon

is similar in size to the goats from the sites in the north of the

Iberian Peninsula, which are larger than present-day goats

and the fossil goats of the south of the Iberian Peninsula.

The horses present at Los Rincones are smaller in size than

those recovered from the Basque–Cantabrian region.

The faunal associations of the Iberian Peninsula during

MIS3 and MIS2 are clearly distinct from those present at

the sites of western and central Europe. The statistical

ordination analysis shows a latitudinal/climatic gradient

from sites with a greater presence of faunas of the

Mammuthus–Coelodonta complex towards sites with a

predominance of species with a Mediterranean affinity

(Capra pyrenaica, Sus scrofa, Cervus elaphus and

Stephanorhinus hemitoechus). At a midpoint are the sites

characterised by the presence of Equus and large bovids

associated with steppe environments.

Our study of faunal associations in the context of the

Iberian Peninsula during MIS3 indicates a cluster of sites

in the south of the peninsula that are separate from the

other sites in the peninsula due to their high presence of

Capra pyrenaica. These sites are Nerja, Gorham’s Cave,

Cueva Beneito and Zafarraya, as well as the site of Los

Rincones, which is included within this group.

The ungulate association of Los Rincones indicates a

landscape with a temperate climate characterised by the

presence of steppes, woodland and rocky areas in

the vicinity of the cave. This landscape would be similar

to the present-day landscape of the ravine of Los Rincones,

where the cave of the same name is located.

Acknowledgements

We thank the Centro Espeleologico de Aragon (SpeleologicalCentre of Aragon), in particular Mario Gisbert, for the discoveryand topography of the cave. The Government of Aragon, theCEA (Centro de Espeleologıa de Aragon) and the University ofZaragoza have partially subsidised the palaeontological activitiesat Los Rincones. This study forms part of the projects ‘GruposConsolidados H54’ of the Government of Aragon and IUCA(Instituto Universitario de Ciencias Ambientales) and CGL2012-38434-C03-01. The first author (VS) also thanks Alfred Sanchis(MPV), Josefina Barreiro and Patricia Perez (MNCN), JulioRamon (MAPH), Mario Laurino (IAN) Jose Ignacio Canudo(MPZ), Henriette Obermaier (LMU) and Caroline Lang (ZSM)for allowing him to study the material for comparison, as well asLourdes Montes, Pillar Utrilla and Mª Fernanda Blasco forgranting access to unpublished data on Gabasa as well as theirvaluable comments. He also thanks Raquel Rabal and CristinaSola for their comments and discussion on taphonomy; VeraEisenmann for her discussions and clarifications on horses; JorgeColmenar for his help with the photographs, and FranciscoGutierrez and Ira Sasowsky for their valuable comments aboutkarst and geology. Rupert Glasgow revised the English grammar.A particular thanks to palaeoartist Gianfranco Mensi for hisprecise reconstruction. The authors also thank Joan MadurellMalapeira and anonymous reviewer for the useful suggestionsthat greatly improved the manuscript. We are especially gratefulfor the comments and improvements of the editor Gareth Dyke.Finally, many thanks go to all the members of the excavationteam of the 2009 and 2010 field campaigns at Los Rincones.

Notes

1. Email: rgarciag@ipe.csic.es2. Email: cuencag@unizar.es

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