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Biochronological data for the middle pleistocene site of Grotte de la Carrière (Lachambre karstic complex, Corneilla-De-Conflent, Eastern Pyrenees, Southern France) inferred from the small-mammal assemblage

Données biochronologiques pour le site pléistocène moyen de la Grotte de la Carrière (complexe karstique Lachambre, Corneilla-de-Conflent, Pyrenées-Orientales, Sud de la France) à partir de l’association de micromammifères
Juan Manuel López‑García, Elisa Luzi, Marc Furio, Ivette Susanna, Manel Llenas et Joan Madurell‑Malapeira
p. 133-138

Résumés

La montagne d’Embullà, située entre les municipalités de Ria et Corneilla-de-Conflent (Pyrénées-Orientales) possède un grand complexe karstique avec plus de 20 grottes disposées sur plusieurs niveaux, la Grotte de Lachambre étant la plus grande avec plus de 26 km de galeries. Les recherches faites par les auteurs depuis 2011 ont permis la découverte de plusieurs sites du Pléistocène moyen et supérieur. La Grotte de la Carrière est un de ces nouveaux sites. La première reconnaissance menée en mai 2015 a permis de collecter plus de 2000 restes de grands et petits vertébrés dans huit couches stratigraphiques différentes. L’assemblage de petits mammifères du niveau le plus riche (couche 4) comprend au moins deux insectivores (Talpa europaea et Sorex sp.) et huit rongeurs (Microtus arvalis, Microtus agrestis jansoni, Iberomys brecciensis, Clethrionomys glareolus, Pliomys coronensis, Apodemus gr. sylvaticus-flavicollis, Allocricetus bursae et Sciurus sp.). Cette association est très similaire à celles des autres sites du Pléistocène moyen du sud de la France, comme la Caune de l’Arago ou la Grotte du Lazaret, et est aussi similaire à celles d’autres sites ibériques du Pléistocène moyen, comme Gran Dolina (TD10), Galeria ou Sima del Elefante (TE18‑19) dans la Sierra d’Atapuerca. Ces données suggèrent un âge approximatif pour la Grotte de la Carrière - couche 4 entre 220 et 450 ka . De plus, la chronologie proposée à partir de l’assemblage des petits mammifères coïncide avec celle des grands mammifères découverts dans ce niveau, qui incluent principalement les espèces Ursus deningeri, Panthera fossilis et Canis mosbachensis.

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Texte intégral

We want to thank the editor Dr. Nicole Limondin-Lozouet and the two reviewers, Dr. Emmanuel Desclaux and Prof. Gloria Cuenca-Bescós, for their comments that improve the final version of the manuscript. This paper has been funded by the Spanish Ministerio de Economía y Competitividad (CGL2014-54373-P) and the Generalitat de Catalunya (2014 SGR 416 GRC and 2014-SGR900. E. Luzi is the beneficiary of a PhD scholarship funded under the Erasmus Mundus Programme - International Doctorate in Quaternary and Prehistory. The authors thank Jean Louis Perez, Caroline Guérineau, Philippe Galant, Jérôme Durbet, Miquel Nebot, Teresa Hernández and Julià González for various kinds of support. We would also like to thank Dr. Hugues-Alexandre Blain and Rupert Glasgow for French and English revisions, respectively.

1 - Introduction

1Rodents are one of the stratigraphically most significant groups of mammals in the European Pleistocene, because they have diversied more than other taxa over this relatively short geological time span. Arvicolines (voles) are particularly useful for biochronological correlation, because of the abundance of their remains and their rapid evolutionary rates. In addition, some voles undertook rapid, long-range migrations and may have had extensive geographical distributions, making interregional correlations possible (Minwer-Barakat et al., 2011). For these reasons, arvicolines have been used to establish the biostratigraphic subdivisions of the Quaternary (Chaline, 1972; van der Meulen, 1973; Agustí, 1986; Sala & Masini, 2007; Cuenca-Bescós et al., 2010; Agustí et al., 2015; among many others).

2Grotte de la Carrière is a newly discovered site in the Embullà Mountain, located between the municipalities of Ria and Corneilla-de-Conflent (Eastern Pyrenees) (fig. 1A). This mountain chain possesses a large karstic complex with more than 20 caves situated at different altitudinal levels. Réseau Lachambre Cave, with more than 26 km of galleries, is the largest of these. The research carried out by the authors since 2011 has led to the discovery of several Middle to Late Pleistocene sites (Madurell-Malapeira et al., 2015). The first paleontological survey of Grotte de la Carrière, undertaken in May 2015, resulted in the unearthing of more than 2000 large- and small-vertebrate remains in eight different stratigraphical layers. The small-mammal material recovered comes mainly from layer 4, a five-centimetre-thick layer composed of brown clays with boulders some centimetres in size comprising quartzes and calcareous schists.

3This paper aims to provide a study of the small-mammal (insectivore and rodent) faunas from Grotte de la Carrière. Particularly interesting is the correlation and chronological position of Grotte de la Carrière in relation to other Middle Pleistocene sites from southern France, Spain and Italy, based mainly on a study of their vole species. This study thus offers a good opportunity for adding to our knowledge of the small-mammal assemblages of the Middle Pleistocene of Western Europe.

Fig. 1: A/ Location of Grotte de la Carrière and the main Middle Pleistocene sites mentioned in the text. B/ Nomenclature and measuring methods used for the first lower molars in the description of Arvicolinae. C/ Measurements and indices of m1 of Microtus agrestis jansoni and Iberomys brecciensis from Grotte de la Carrière. Linear data (L, W, a) are in mm. Abbreviations for B: a. length of the anteroconid complex; ACC. anteroconid complex; AC

Fig. 1: A/ Location of Grotte de la Carrière and the main Middle Pleistocene sites mentioned in the text. B/ Nomenclature and measuring methods used for the first lower molars in the description of Arvicolinae. C/ Measurements and indices of m1 of Microtus agrestis jansoni and Iberomys brecciensis from Grotte de la Carrière. Linear data (L, W, a) are in mm. Abbreviations for B: a. length of the anteroconid complex; ACC. anteroconid complex; AC

2 - Material and methods

4During the excavation of Grotte de la Carrière, sediments from each layer were screen -washed in order to obtain the small -vertebrate fossil remains. Most of the material comes from layer 4. The specimens under study were sorted at the Institut Català de Paleontologia Miquel Crusafont (ICP, Barcelona, Spain). The Grotte de la Carrière - layer 4 assemblage includes a total of 91 identied small mammals corresponding to a minimum number of 42 individuals, representing at least ten taxa: Talpa europaea, Sorex sp., Microtus arvalis, Microtus agrestis jansoni, Iberomys brecciensis ( = mediterraneus; priority discussed by Cuenca-Bescós et al. 2014), Clethrionomys glareolus, Pliomys coronensis ( = lenki; priority discussed by Terzea, 1983), Apodemus gr. sylvaticus-flavicollis, Allocricetus bursae and Sciurus sp.

5The nomenclature for the description of the arvicoline teeth (only rst lower molars are considered) used for the chronological inferences is that from van der Meulen (1973) and Martin (1987). Length, width, and parameter a are those proposed by van der Meulen (1973), and parameters Li and La are those proposed by Cuenca-Bescós et al. (1995). Index La/Li is the ratio between parameters Li and La (fig. 1B & C).

3 - Biochronological remarks

6Small mammals, and particularly arvicolines, evolved rapidly during the Quaternary, so their record provides an important basis for stratigraphic divisions and correlations between distant areas. In particular, the quick evolutionary radiation of the ‘Microtus group’ (which includes forms ascribed to Microtus, Allophaiomys, Stenocranius, Terricola, Iberomys and Pallasiinus) constitutes a very useful tool for correlating European Pleistocene faunas. In our case, the most diagnostic species from a chronological point of view are the voles Microtus agrestis jansoni and Iberomys brecciensis (fig. 2A).

3.1 - Microtus agrestis jansoni (Chaline, 1972)

7Microtus agrestis morphotype jansoni was first described by Chaline (1972) on the basis of material from Grotte de l’Escale (Saint-Estève-Janson, Bouches-du-Rhône, France) as a vole with agrestis-like morphology: m1 that presents a posterior loop, 5 closed triangles with a clear labio-lingual asymmetry, alternatingly confluent T6 and T7, and an asymmetric anterior cap. In particular, the jansoni-type is characterized by a middle-large size, a deep LRA4, the presence of T9 in the ACC and a broad LRA5. In some cases, the populations of the late Middle Pleistocene develop a small BRA4 on T6, resulting in a T8 that recalls the mimomyan-fold. This archaic character is rare in Late Pleistocene populations (1/1000). Chaline (1972) and Cuenca-Bescós et al. (1999) suggest a trend toward a decrease in size for this sub-species during the Middle Pleistocene.

8Other populations of M. agrestis jansoni have been identified in the Middle and Late Pleistocene sites of Caune de l’Arago (Brunet-Lecomte & Paunesco, 2004), La Fage (Mourer-Chauvire et al., 2003), Grotte des Cèdres (Defleur et al., 1990) and Combe Grenal (Chaline, 1972) in France, and in the Middle Pleistocene sites of Galeria, Zarpazos, Gran Dolina TD10, Sima del Elefante TE18-19 (Cuenca-Bescós et al., 1999, 2010; Galindo-Pellicena et al., 2011; López-García et al., 2011), Mollet (López-García et al., 2014) and Valdavara-3 in Spain.

9The specimens from Grotte de la Carrière identified as Microtus agrestis jansoni present all the morphological characters proper to this sub-species. One specimen also shows a T8 which, taking into account the number of remains recovered, suggests a Middle Pleistocene age. From a biometric point of view, the mean length of m1 (2.796 mm) is lower than those from the type-population of L’Escale (2.87 mm) and from TE 18-19 (2.886 mm) and Zarpazos (2.849 mm), and it is close to those from Mollet (2.775 mm), TD10 (2.776 mm) and Valdavara-3 (2.801 mm). The labio-lingual asymmetry is more pronounced in Grotte de la Carrière (La/Li = 57) than in the samples from Mollet, TD10, Zarpazos, Valdavara-3 and TE 18-19 (La/Li = 41.5, 51.5, 51.6, 52.5 and 54 respectively). The population of M. agrestis from layer IIb of Teixoneres Cave (López-García et al., 2012) (Late Pleistocene, Spain) is smaller and more symmetrical (L = 2.662 mm; La/Li = 64.1) than those of the Middle Pleistocene (fig. 2B). Thus, a morphometric comparison between these populations indicates that Grotte de la Carrière has an age close to Valdavara-3 and TD10 and younger than Zarpazos (Falguères et al., 2013; Demuro et al., 2014), ranging from 360 to 250 ka.

Fig 2: A/ Some small-mammal remains identified from layer 4 of Grotte de la Carrière. B/ Distribution of the species Microtus agrestis jansoni (above) and Iberomys brecciensis (below) from several Middle Pleistocene sites and some Late Pleistocene sites with Microtus agrestis (Tx) and Iberomys cabrerae (Abric Romaní, Gegant and Gorh), including Grotte de la Carrière, for the mean of variables La/Li and L. C/ Chronological position for Grotte de la Carrière in relation to the other Middle Pleistocene sites cited in the text and the oxygen isotope curve for this chronological period.

Fig 2: A/ Some small-mammal remains identified from layer 4 of Grotte de la Carrière. B/ Distribution of the species Microtus agrestis jansoni (above) and Iberomys brecciensis (below) from several Middle Pleistocene sites and some Late Pleistocene sites with Microtus agrestis (Tx) and Iberomys cabrerae (Abric Romaní, Gegant and Gorh), including Grotte de la Carrière, for the mean of variables La/Li and L. C/ Chronological position for Grotte de la Carrière in relation to the other Middle Pleistocene sites cited in the text and the oxygen isotope curve for this chronological period.

A/ 1-2. right and left m1 Iberomys brecciensis (C1-N4-D6-8 and C4-N4-D6-23); 3-4. left m1 Clethrionomys glareolus (C2-N4-D6-15 and C5-N4-D6-29); 5-6. right and left m1 Microtus agrestis jansoni (C2-N4-D6-12 and C2-N4-D6-13); 7-8. left and right m1 Microtus arvalis (C2-N4-D6-10 and C1-N4-D6-1); 9. left m1 Pliomys coronensis (C1-N4-D6-4); 10. left m1 Apodemus gr. sylvaticus-flavicollis (C6-N4-D6-31/1); 11. right m1 Allocricetus bursae (C1-N4-D6-9). All teeth are oriented in occlusal view. Scale 1 mm. B/ For abbreviations of the ratios see Material and Methods. TE:Atapuerca Sima del Elefante layers TE18-TE19; TD: Atapuerca Gran Dolina layer TD10; TZG: Atapuerca Trinchera Zarpazos-Galeria; MSII : Montagnola Senese II; Val-3: Valdavara-3; Gorh: Gorham’s cave; Tx : Teixoneres cave.

3.2 - Iberomys brecciensis (= Mediterraneus) (Giebel, 1847)

10Microtus (Iberomys) brecciensis was described by Chaline in 1972 as a subgenus of Microtus species. Chaline proposed a specimen drawn by Cuvier (1823) as the holotype, and Sète as the type locality, a karst fissure of imprecise location where lagomorphs have been found.

11The distinctive morphological traits of I. brecciensis are the T4 and T5 that are closed in all specimens. In some specimens, BRA4 forms a reduced re-entrant angle, and in most m1, T6 is clearly separated from the anterior cusp due to the strong development of the re-entrant angle LRA4. These characteristics allow our material to be attributed to the species I. brecciensis, thus distinguishing it, moreover, from the present-day species I. cabrerae and from the fossil species from the Early Pleistocene I. huescarensis.

12Other populations of I. brecciensis have been identified in Middle Pleistocene sites such as Caune de l’Arago, Orgnac and Le Lazaret (Hanquet, 2011; Hanquet & Desclaux, 2011) in France, Valdemino or Montagnola Senese II in Italy (López-García et al., 2015), and Galeria, Zarpazos, Gran Dolina TD10, Sima del Elefante TE18-19 (Cuenca-Bescós et al., 1999, 2010; Galindo-Pellicena et al., 2011; López-García et al., 2011) and Mollet (López-García et al., 2014) in Spain.

13The specimens from Grotte de la Carrière identified as Iberomys brecciensis present all the morphological characters proper to this species, suggesting a Middle Pleistocene age for layer 4. From the biometric point of view, m1 (mean length 2.830 mm) is longer than those from the sites of Valdemino (2.69 mm) and Mollet (2.7 mm), shorter than those from Orgnac (2.96 mm), Lazaret (3.15 mm), TE 18-19 (2.91 mm) and Galeria (3.12 mm), and similar to those from Arago I (2.89 mm), L’Escale (2.89 mm), Montagnola Senese II (2.83 mm) and TD10 (2.88 mm). The labio-lingual asymmetry is less pronounced in Grotte de la Carrière (La/Li = 47.6) than in Galeria (La/Li = 48.2), but more pronounced than in Valdemino, Montagnola Senese II, TD10, TE18-19 and Mollet (36.8, 41.8, 42.7, 45.2 and 45.9 respectively). The populations of I. cabrerae from the Late Pleistocene sites of Cova del Gegant (L = 3.31 mm; La/Li = 54.7), Abric Romaní (L = 3.35 mm; La/Li = 56.9) and Gorham’s cave (L = 3.40 mm; La/Li = 55.07) (López-García et al., 2008; López-García, 2011) are larger and more asymmetrical than those of the Middle Pleistocene (fig. 2B). Thus, morphological and biometric comparisons between these populations indicate that Grotte de la Carrière has an age close to TD10, Galeria and TE18-19, older than Mollet, and younger than Valdemino and Montagnola Senese II (López-García et al., 2011; Maroto et al., 2012; Falguères et al., 2013; Demuro et al., 2014; ; de Lombera-Hermida et al., 2015; López-García et al., 2015), ranging from 450 to 215 ka.

14In summary, the small-mammal association from layer 4 of Grotte de la Carrière fits within the range of typical associations from the Middle Pleistocene faunas of the southwestern Mediterranean and can be included in FU6 (ranging from 600 to 125 ka) as defined by Cuenca-Bescós et al. (2010), where the characteristic species of this Faunal Unit are I. brecciensis, M. arvalis, M. agrestis jansoni, Pliomys coronensis and Allocricetus bursae. Thus, the Grotte de la Carrière association is also very similar to other Middle Pleistocene sites from the south of France, such as Caune del Arago or Grotte du Lazaret (Hanquet, 2011; Hanquet & Desclaux, 2011), and Spain, such as Gran Dolina (TD10), Galeria and Sima del Elefante (TE18-19) from the Sierra de Atapuerca (Cuenca-Bescós et al., 1999, 2010; Galindo-Pellicena et al., 2011; López-García et al., 2011). All these data, together with the morphological and biometric studies of the species Iberomys brecciensis and Microtus agrestis jansoni, suggest a rough age for layer 4 of between 450 and 220 ka (fig. 2C). The chronology proposed on the basis of the micromammal assemblage is in accordance with the large mammals recovered and identified from this layer, Ursus deningeri, Panthera fossilis and Canis mosbachensis being the most characteristic species.

4 - Conclusions

15Our study of the small-mammal assemblage from layer 4 of Grotte de la Carrière allows the following conclusions to be drawn:

  • -1/ Ten small-mammal (insectivore and rodent) species are currently known to be present in Grotte de la Carrière - layer 4: Talpa europaea, Sorex sp., Microtus arvalis, Microtus agrestis jansoni, Iberomys brecciensis, Clethrionomys glareolus, Pliomys coronensis, Apodemus gr. sylvaticus-flavicollis, Allocricetus bursae and Sciurus sp.

  • -2/ Within the context of the small- and large-mammal assemblages from layer 4 of Grotte de la Carrière, the evolutionary states of the vole species Microtus agrestis jansoni and Iberomys brecciensis are indicative of a chronological age within the range of 450 and 220 ka .

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Table des illustrations

Titre Fig. 1: A/ Location of Grotte de la Carrière and the main Middle Pleistocene sites mentioned in the text. B/ Nomenclature and measuring methods used for the first lower molars in the description of Arvicolinae. C/ Measurements and indices of m1 of Microtus agrestis jansoni and Iberomys brecciensis from Grotte de la Carrière. Linear data (L, W, a) are in mm. Abbreviations for B: a. length of the anteroconid complex; ACC. anteroconid complex; AC
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Titre Fig 2: A/ Some small-mammal remains identified from layer 4 of Grotte de la Carrière. B/ Distribution of the species Microtus agrestis jansoni (above) and Iberomys brecciensis (below) from several Middle Pleistocene sites and some Late Pleistocene sites with Microtus agrestis (Tx) and Iberomys cabrerae (Abric Romaní, Gegant and Gorh), including Grotte de la Carrière, for the mean of variables La/Li and L. C/ Chronological position for Grotte de la Carrière in relation to the other Middle Pleistocene sites cited in the text and the oxygen isotope curve for this chronological period.
Légende A/ 1-2. right and left m1 Iberomys brecciensis (C1-N4-D6-8 and C4-N4-D6-23); 3-4. left m1 Clethrionomys glareolus (C2-N4-D6-15 and C5-N4-D6-29); 5-6. right and left m1 Microtus agrestis jansoni (C2-N4-D6-12 and C2-N4-D6-13); 7-8. left and right m1 Microtus arvalis (C2-N4-D6-10 and C1-N4-D6-1); 9. left m1 Pliomys coronensis (C1-N4-D6-4); 10. left m1 Apodemus gr. sylvaticus-flavicollis (C6-N4-D6-31/1); 11. right m1 Allocricetus bursae (C1-N4-D6-9). All teeth are oriented in occlusal view. Scale 1 mm. B/ For abbreviations of the ratios see Material and Methods. TE:Atapuerca Sima del Elefante layers TE18-TE19; TD: Atapuerca Gran Dolina layer TD10; TZG: Atapuerca Trinchera Zarpazos-Galeria; MSII : Montagnola Senese II; Val-3: Valdavara-3; Gorh: Gorham’s cave; Tx : Teixoneres cave.
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Juan Manuel López‑García, Elisa Luzi, Marc Furio, Ivette Susanna, Manel Llenas et Joan Madurell‑Malapeira, « Biochronological data for the middle pleistocene site of Grotte de la Carrière (Lachambre karstic complex, Corneilla-De-Conflent, Eastern Pyrenees, Southern France) inferred from the small-mammal assemblage », Quaternaire, vol. 27/2 | 2016, 133-138.

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Juan Manuel López‑García, Elisa Luzi, Marc Furio, Ivette Susanna, Manel Llenas et Joan Madurell‑Malapeira, « Biochronological data for the middle pleistocene site of Grotte de la Carrière (Lachambre karstic complex, Corneilla-De-Conflent, Eastern Pyrenees, Southern France) inferred from the small-mammal assemblage », Quaternaire [En ligne], vol. 27/2 | 2016, mis en ligne le 01 juin 2018, consulté le 19 juin 2018. URL : http://journals.openedition.org/quaternaire/7601 ; DOI : 10.4000/quaternaire.7601

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Auteurs

Juan Manuel López‑García

 IPHES, Institut Català de Paleoecologia Humana i Evolució Social, Campus Sescelades URV, edifici W3, ES-43007 TARRAGONA. Email: jmlopez@iphes.cat; Àrea de Prehistòria, Universitat Rovira i Virgili (URV), Avinguda de Catalunya 35, ES-43002 TARRAGONA. Email: jlopez@prehistoria.urv.cat

Articles du même auteur

Elisa Luzi

Àrea de Prehistòria, Universitat Rovira i Virgili (URV), Avinguda de Catalunya 35, ES-43002 TARRAGONA. Email: elisa.luzi@urv.cat

Marc Furio

Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, Edifici ICTA-ICP, Carrer de les columnes s/n, ES-08193 CERDANYOLA DEL VALLÈS, BARCELONA. Email : marc.furio@icp.cat

Ivette Susanna

Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, Edifici ICTA-ICP, Carrer de les columnes s/n, ES-08193 CERDANYOLA DEL VALLÈS, BARCELONA. Email: ivette.susanna@icp.cat

Manel Llenas

Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, Edifici ICTA-ICP, Carrer de les columnes s/n, ES-08193 CERDANYOLA DEL VALLÈS, BARCELONA. Email: manel.llenas@icp.cat

Joan Madurell‑Malapeira

Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, Edifici ICTA-ICP, Carrer de les columnes s/n, ES-08193 CERDANYOLA DEL VALLÈS, BARCELONA. Email: joan.madurell@icp.cat

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