Bats (Chiroptera, Mammalia) are a valuable and independent source of palaeoenvironmental information in Quaternary sites because bat assemblages are not necessarily produced by the same processes as those of other small mammals. Here, we report a small collection of bats (MNI 15) from a cave infill of the Aguilón P7 (AGP-7) site (Zaragoza, Spain) dated Late Pleistocene (MIS 3). This is one of the rare localities south of the Ebro River (right bank) with Neanderthal tracks; therefore, the palaeoenvironmental data provided here may help us to understand Neanderthal history. Nine bat taxa were identified, viz. Rhinolophus ferumequinum, R. hipposideros, Myotis gr. myotis/blythii, M. cf. bechsteinii, M. emarginatus, M. daubentonii, Plecotus gr. auritus/austriacus, Hipsugo savii vel. Pipistrellus kuhlii/nathusii, and Miniopterus schreibersii. The presence of a high number of yearlings of the former four species suggests the presence of breeding colonies. The surprisingly high diversity of the sample indicates a variegated vegetation cover and relatively high humidity for the MIS 3 stage in the region.
Les chauves-souris (Chiroptera, Mammalia) retrouvées dans les sites fossiles quaternaires sont une source indépendante précieuse d’information paléoenvironnementale, puisque les assemblages de chiroptères ne proviennent pas nécessairement des mêmes processus que celles d’autres petits mammifères. Ici, nous décrivons pour la première fois une petite association relativement diversifiée de chauves-souris, trouvée dans une localité d’âge Pléistocène supérieur (MIS 3), le site d’Aguilón P7 (Saragosse, Espagne). C’est l’une des rares localités du Sud de l’Èbre (rive droite) ayant des traces des Néandertaliens ; par conséquent, les données paléoenvironnementales fournies ici peuvent aider à comprendre leur histoire. Neuf taxons de chiroptères ont été identifiés : Rhinolophus ferrumequinum, Rhinolophus hipposideros, Myotis gr. myotis/blythii, M. cf. bechsteinii, M. emarginatus, M. daubentonii, Plecotus gr. auritus/austriacus, Hipsugo savii vel. Pipistrellus kuhlii/nathusii et Miniopterus schreibersii. La présence d’un nombre élevé de jeunes individus des quatre premières espèces suggère la présence de colonies de reproduction. L’importante et surprenante diversité de l’échantillon indiquent pour le MIS 3, dans la région, une couverture végétale variée et une humidité relativement élevée.
Chiroptera, Taxonomy, Taphonomy, Palaeoenvironment, Late PleistoceneChiroptera, Taxonomie, Taphonomie, Paléoenvironnement, Pléistocène supérieurpresentedHandled by Lars Vanden Hoek OstendeIntroduction
Bat fossil remains (Chiroptera, Mammalia) are interesting palaeoenvironmental indicators in Quaternary sites mainly due to their specific habitat preferences. Bats are only occasionally preyed upon by owls and other predators, such as carnivorous mammals, snakes or lizards (García et al., 2005, Kowalski, 1995, Rosina and Shokhrin, 2011 and Sommer et al., 2009), but their habit of forming roosting colonies in caves sometimes allows the formation of exceptionally well-preserved assemblages (López-García and Sevilla, 2012). Because bat assemblages are not necessarily produced by the same processes as other small mammal assemblages, they provide a valuable independent source of palaeoenvironmental information.
Here, we describe for the first time a relatively diverse association of bats found in Aguilón P7 Cave (AGP-7), a new locality dating to the Late Pleistocene (MIS 3) in the central Ebro Valley. AGP-7 is one of the rare localities of the south of the Ebro River with Neanderthal tracks, and palaeoenvironmental data provided by this site may help understanding Neanderthal history (Cuenca-Bescós et al., 2010 and Sauqué et al., 2014a). We focus on the taxonomic identification of the bat remains, the origin of the accumulation, the palaeoenvironment, and the comparison with other chiropteran associations from the Iberian Late Pleistocene.
The localityGeographical and geological location
AGP-7 is a palaeontological cave-site located in the north face of the Iberian Range to the south of the River Ebro. It is situated on a small hill named Cerro Pezón; the mouth of the cave is open at 683 m above sea level (masl) on the left bank of the Valdeaguilón ravine, close to the town of Aguilón and 55 km from the main city of Zaragoza (Fig. 1) (Gisbert and Pastor, 2009). The P7 cave is part of a karst system in Upper Jurassic (Malm) marine limestones that opens at the contact between the two main formations from this age in the Iberian Range: the marls and clays of Loriguilla (Kimmeridgian) and the thick limestones of Higueruelas (Tithonian) (Cuenca-Bescós et al., 2010). As with other cases of speleogenesis in the Iberian Peninsula, the karst system is expected to have developed during the Pliocene–Early Pleistocene (Ortega et al., 2013). Its geometry, orientation and karst-cavity filling were conditioned by the incision of the main rivers, such as the River Ebro, and the development and incision of their hydrological basins. In the light of a preliminary geological survey made in the area in May 2010, the drainage of the karst system presumably changed from an original exit towards the River Huerva, the main river in the area, to the current discharge towards the Valdeaguilón Ravine, a secondary incision in the hydrological system of the area. The P7 cave is rather shallow and consists of an outer entrance hall and an inner gallery (Fig. 2). The cavity filling sediment of the outer entrance hall, where the main palaeontological excavation took place, consists of a single sedimentary level, 85 cm thick, comprising heterometric limestone ridges in a reddish clay matrix, probably corresponding to a single deposition event, while sediments in the inner gallery were likely deposited during a previous stage (Cuenca-Bescós et al., 2010).
Dating the sedimentary fill of the AGP-7
A bone fragment that came out of a sample of fossil remains from the surface cleaning, almost certainly from the top of the section, was dated by radiocarbon resulting in 34,760 ± 360 BP–34,860 ± 360 BP (Beta – 222732), providing a minimum age of the reservoir in the area's known slope (P in Fig. 2). In recent years, a second radiocarbon date was provided by a bone from a roe deer Capreolus capreolus exhumed from the base of the section. This gave an age of just under 46.3 ka BP (OxA-27902; for the methodology employed see Bronk Ramsey et al., 2004). Thus, the faunal and lithic assemblages point to a Late Pleistocene biostratigraphic age, likely from the beginning of Marine Isotope Stage 3 (MIS3). MIS 3 was a period characterized by rapid climate changes (Sánchez-Goñi and D’Errico, 2005), supporting the existence of Pleistocene communities with no present-day analogy, and that shared distributions that are currently disjunct (López-García et al., 2010, Sauqué et al., 2014b and Sommer and Nadachowski, 2006).
The faunal assemblage
Even though only a small part of AGP-7 cave has been excavated, the species richness and the number of remains are notable. The faunal list is as follows (Cuenca-Bescós et al., 2010, Galán, 2013 and Núñez-Lahuerta et al., 2015): small mammals—Crocidura sp., Lagomorpha indet., Apodemus sp., Microtus sp., Microtus (Terricola) sp., Iberomys cabrerae and Chionomys nivalis; large mammals—Cuon alpinus, Canis lupus, Crocuta crocuta, Panthera pardus, Vulpes vulpes, Ursus arctos, Lynx pardinus, Felis silvestris, Meles meles, Cervus elaphus, Capra pyrenaica, Capreolus capreolus, Rupicapra pyrenaica and Equus ferus; birds—Galliformes indet., Lagopus sp., Aquila chrysaetos, Gyps fulvus, Passeridae indet., Anthus trivialis, Prunella modularis, Sturnus cf. unicolor, and Corvus monedula. The cave may have been used as a den by large carnivores, yet some Mousterian tools have been recovered, including a scraper, and tool marks in some fossils have been observed, suggesting that the cave was visited by Neanderthals, even though no human remains have been found.
Material and methodsAbbreviations
Institutional – MPZ: Palaeontological Museum of Zaragoza; EBD: Doñana Biological Station-CSIC; MNCN: National Museum of Natural Science-CSIC, Madrid; GA: Aragosaurus Group-IUCA, Zaragoza; CIAMA: International Centre for Water and Environment-La Alfranca, Zaragoza.
Dental terminology (Fig. 3) – I: upper incisor; C: upper canine; P: upper premolars; M: upper molars; the teeth from the lower dental series are indicated in the same way but using lowercase letters. The anatomical parts of molars are abbreviated as follows, PA: paracone; ME: metacone; PR: protocone; PAC: paracrista; MEC: metacrista; PRC: protocrista; PAS: parastyle; MSS: mesostyle; MES: metastyle; HY: hypocone; PL: paraconule; ML: metaconule; PAD: paraconid; MED: metaconid; PRD: protoconid; END: entoconid; HYD: hypoconid; HLD: hypoconulid; ENCD: entocristid; PALD: paralophid; tg: trigonid; td: talonid.
Anatomical elements – max: maxilla; md: mandible; hu: humerus; d. e.: distal epiphysis; diap.: diaphysis; fr: fragment.
Sample processing and fossil sorting
The bat fossil remains studied here consist of disarticulated bone fragments collected by water-screening the sediment excavated at the Cave P7 during the years 2005, 2009 and 2010. Superimposed screens of 0.2 cm and 0.05 cm mesh were used. The palaeontological excavation was divided into six different sectors (Fig. 2): Entrada (E), Pendiente (P), Centro (C), Galería (G), Fondo (F) and Hiedra (H); in situ material was extracted from sector F and from sectors E, P, C and H. The fossils were picked out from the residues, and then the bat remains were separated and identified. They are all kept in the MPZ. To evaluate the bat representation, the number of identified specimens (NISP) and the minimum number of individuals (MNI) were calculated for each taxon in accordance with Lyman (1994). For taxonomic identification these authors have been followed: Dupuis, 1986 and Felten et al., 1973, Menu and Popelard (1987), Sevilla (1988). The reference collections of the EBD, MNCN, GA and CIAMA were used.
Palaeoenvironmental approach
We used the habitat weighting method (see Andrews, 2006 and Evans et al., 1981) to infer palaeoenvironmental characteristics. Five types of habitat are defined according to concrete environmental features: “open dry (OD)”, meadows under seasonal climate change; “open humid (OH)”, evergreen meadows with dense pastures and suitable topsoil; “woodland (Wo)”, mature forest including woodland margins and forest patches with moderate ground cover; “rocky (Ro)”, areas with a suitable rocky or stony substratum; and “water (Wa)”, areas along streams, lakes and ponds. Bat remains from sector F (Fig. 2) were not taken into account for the palaeoenvironmental approach as they correspond to a previous accumulation event (Cuenca-Bescós et al., 2010), but they were identified and measured.
Recorded taxa
Order CHIROPTERA Blumenbach, 1779
Family RHINOLOPHIDAE Gray, 1825
Genus RhinolophusLacépède, 1799
Rhinolophus ferrumequinum (Schreber, 1774) – Greater horseshoe bat (present habitat and distribution in Table 1).
MATERIAL (Fig. 4A, H; Appendix A): 1 max (fr) with M1, M2; 1 max (fr) with M3; 3C; 1 P4; 1 md with p4, m1; 1 md with p3, p4, m1, m2, m3; 1 md (fr) with p4; 1 c; 2 m1; 2 m2; 2 m3; 1 m1 (fr).
DESCRIPTION: R. ferrumequinum is the most abundant species in AGP-7 and one of the largest. The C has a trapezoidal occlusal outline with a strong crown with three sides and a distally bent apex. The lingual side is the widest; it is flat and presents a small heel. The cingulum thickness is variable and has a zigzag morphology, inflexions pointing alternately to apex and root. The P4 has a rectangular occlusal outline, a wide, well-developed heel and a narrow cingulum, which disappears distally. The upper molars present a parastyle forming an angle, though only the M1 has a (straight) metastyle. The M1 and M2 have a rectangular occlusal outline and a well-developed parastyle and heel. The protocone is on the mesial margin and its base lacks a cingulum. The M3 has a subtriangular occlusal outline and presents a small parastyle. The metacone is smaller than the protocone, and there is a slightly developed metaconule on the post-protocrista. The maxilla fragments present the infra-orbital foramen located over the alveolus of the M2, and the alveolus of the M3 is located on the zygomatic arch root. The c has a semicircular occlusal outline in the base but the cusp has two sides, its apex and root are aligned and the cingulum is narrow and forms a small projection at posterior side. The p3 is very small – sometimes even absent – and appears out of the dental series. The p4 has a trapezoidal occlusal outline with three sides and a narrow cingulum, which is slightly thicker at the distal margin. The lower molars are of the nyctalodont type and present a narrow and regular labial cingulum. The m1 and m2 have an open trigonid, which is narrower in m2, and their lingual cusplets are not aligned, the hypoconulid having a labial position. The m3 presents a narrow trigonid and a reduced talonid due to the labial migration of the entoconid. The mandibles have a long and flat horizontal body and an almost rectangular mandibular ramus with flat and low coronoid process. The mental protuberance appears well-developed; the mental foramen is circular and located under the p2 alveolus, or under the junction of the alveoli of c and p2.
DISCUSSION: The morphological features and the size (Appendix A) of the teeth and bones of this fossil material conform to the specimens of R. ferrumequinum described in Sevilla (1988).
Rhinolophus hipposideros (Bechstein, 1800) – Lesser horseshoe bat (present habitat and distribution in Table 1).
MATERIAL (Fig. 4E; Appendix A): 1 max (fr) with P4, M1, M2.
DESCRIPTION: This is the smallest species of this genus in the Palaearctic region. The P4 has a rectangular occlusal outline and a wide, well-developed heel. The thickness of the cingulum is constant along the heel. The upper molars present a parastyle forming an angle, a straight metastyle and a comparatively wide heel. The M1 has a square occlusal outline, while the M2 is narrower. The base of the protocone lacks a cingulum and in the M1, it advances beyond the heel. Although the studied maxilla is fragmented, the bulging-out nasal region can be appreciated, and also the huge palatine incision caused by the disarticulation of the premaxilla, typical in this genus. The posterior rim of the horizontal plate of the palatine presents a square outline, a well-developed posterior nasal spine and a choanal line anteriorly displaced in comparison with other members of the genus.
DISCUSSION: The wide heel of the upper teeth and the size of the remains (Appendix A) indicate that the remains belong to R. hipposideros according to Sevilla (1988). The features of the palatine posterior rim have been also observed in the R. hipposideros specimens from the EBD collection.
Family VESPERTILIONIDAE Gray, 1821
Genus MyotisKaup, 1829
Myotis myotis (Borkhausen, 1797) – Greater mouse-eared bat (present habitat and distribution in Table 1).
Myotis blythii (Tomes, 1857) – Lesser mouse-eared bat (present habitat and distribution in Table 1).
Myotis gr. myotis/blythii
MATERIAL (Fig. 4F, I; Appendix A): 1 max (fr) with P4, M1; 1 P2; 2 P4; 1 M1; 2 c; 3 p4; 5 m1 (fr); 1 m2 (fr).
DESCRIPTION: The P2 has an oval occlusal outline, while P4 is triangular with a very strong crown; both have a thick cingulum. The M1 has a square occlusal outline. It presents a well-developed parastyle forming an angle, and a smaller bent metastyle. The cingulum is thick, but it is not present at the base of the protocone and the lingual. There is a metaconule. The maxilla fragment presents the infra-orbital foramen located above the junction of the P4 and M1 alveoli; the bone bar between the foramen and the orbit is rather thick. The c has a semicircular occlusal outline in the base but the cusp has three sides, apex and root being aligned. In the distal side, the thick cingulum is slightly projected backwards and also forms an inflexion upwards; it disappears in the anterior side. The p4 has a rectangular occlusal outline and a three-faced crown; the distal side is strongly concave. The cingulum is thick, irregular and continuous. The lower molars are of the myotodont type, with a thick and slightly irregular cingulum. The trigonid is rather closed in m1 and m2, and narrower in the latter. The lingual cusplets are not aligned; the hypoconulid has a more labial position. The entocristid is almost straight.
DISCUSSION: These two species (M. myotis and M. blythii) are difficult to distinguish from one another as their bones and teeth are very similar in morphology and size, although M. myotis is in general terms slightly larger than M. blythii (López-García, 2008 and Sevilla, 1988). The morphological features and the size (Appendix A) of the remains indicate that they belong to a large species of the genus Myotis (M. myotis or M. blythii) based on the data provided by Sevilla (1988). Some biometric methods allow the two species to be distinguished, but these are hard to use when the sample is composed of isolated teeth. The measurements of the m3 are a good diagnostic element (Sevilla, 1988), but this molar has not been found in the assemblage. A biometric analysis has been performed on the M1 specimens (after López-García, 2008), comparing them with current specimens from the collections at the EBD, GA, MNCN and CIAMA suggests that they probably belong to M. myotis. However, as the remaining elements cannot be assigned to either of the two species either by biometric or morphological criteria, the whole sample has been classified as M. gr. myotis/blythii.
Myotis bechsteinii (Kuhl, 1817) – Bechstein's bat (present habitat and distribution in Table 1).
Myotis cf. bechsteinii
MATERIAL (Fig. 5A; Appendix A): 1 hu (d. e. + diap.).
DESCRIPTION: The distal humeral epiphysis presents a rather shallow antecubital fossa. In external view, the trochlea and the condyle dip slightly to the epicondyle and there is a shallow furrow between them. The distal ends of the trochlea and the condyle have a similar height. The posterior epicondyle outline presents a concave–convex morphology with a pointed but not sharp edge.
DISCUSSION: The specimen presents typical Myotis features – a reduced epitrochlea and a styloid process. The morphology coincides with the fossil specimens of M. bechsteinii reported by Sevilla (1990); the epiphysis length (Appendix A) agrees with data for that species provided by Felten et al. (1973). However, a single humeral fragment does not allow unequivocal identification in the case of M. bechsteinii so we determined it as M. cf. bechsteinii.
Myotis emarginatus (Geoffroy, 1806). Geoffroy's bat (present habitat and distribution in Table 1).
MATERIAL (Fig. 4B; Appendix A): 1C; 1 m3; 2 m2.
DESCRIPTION: The C has an oval occlusal outline with two sides; apex and root are aligned. The labial side presents a longitudinal furrow in the distal margin. The cingulum is thicker in the anterolabial margin. The m2 has a closed trigonid and its cingulum is thick and almost regular. The lingual cusplets are almost aligned, except for the hypoconulid, which is slightly labial. The paraconid and metaconid cusplets are similar in height, while the entoconid is slightly lower. The paralophid is concave and the entocristid is almost straight. The m3 has a closed trigonid and a medium-thick cingulum, which is thinner in the mesial and distal margins. The paralophid is concave and not angular. The talonid appears reduced due to the labial displacement of the hypoconulid.
DISCUSSION: The main morphological features and the size (Appendix A) of the specimens described above coincide with the data reported by Sevilla (1990) for M. emarginatus. According to this author, the occlusal outline and the furrow in the C, and the disposition of the talonid cusplets in the lower molars, are diagnostic elements for distinguishing this taxon from other similarly sized species of Myotis, such as M. nattereri. The development of the entoconid in the m2 differs somewhat from that observed in the material used for comparison, where this is the highest and strongest labial cusp. However, this was not considered reason enough to assign the specimens to another taxon, but was regarded as intraspecific variability.
Myotis daubentonii (Kuhl, 1817) – Daubenton's bat (present habitat and distribution in Table 1).
MATERIAL (Fig. 4J; Appendix A): 1 M1; 1 md (fr) with m2; 1 m1.
DESCRIPTION: M. daubentonii is small in comparison with other members of the genus Myotis. The M1 has a subtriangular occlusal outline. It presents a well-developed parastyle forming an angle, and a slightly bent metastyle. The metacone is markedly stronger than the paracone and somewhat higher. The base of the protocone presents a strong cingulum in its distal margin. There is a well-developed metaconule and also a paraconule is present. The mandible fragment shows a circular mental foramen located under the p2 alveolus. The lower molars present a thick and irregular cingulum. The trigonid is rather closed in m1 and m2, being narrower in m2. The lingual cusplets are almost aligned except for the hypoconulid, which is slightly labial. It is notable that the entoconid is the strongest lingual cusp, as high as the metaconid. The paralophid has a shallow concavity.
DISCUSSION: The size (Appendix A) and morphological features of the described specimens agree with data for Myotis daubentonii from Sevilla (1990). The presence of a paraconule in the M1 allows distinguishing this taxon from M. mystacinus according to Menu and Popelard (1987). Regarding the lower molars, the entoconid in lingual view is remarkably strong in both teeth, which is a feature that only has been observed in extant M. daubentonii from the reference collection but not in other small Myotis, such as M. mystacinus.
Myotis emarginatus/daubentonii
MATERIAL (Fig. 5B; Appendix A): 1 hu (d. e. + diap.).
DESCRIPTION: The distal humeral epiphysis presents a well-marked antecubital fossa. In external view, the trochlea and the condyle dip slightly to the epitrochlea and there is a shallow but somewhat angular furrow between them. The distal end of the trochlea is higher than the condyle. The epitrochlea has an angular contour.
DISCUSSION: The size (Appendix A) as well as the morphology of the specimens agrees with M. emarginatus, M. nattereri and M. daubentonii (Dupuis, 1986) (Felten et al., 1973). The length of the epitrochlea is shorter than in M. nattereri, which is a character indicated by Dupuis (1986) for distinguishing this taxon from M. emarginatus and M. daubentonii. Another diagnostic character is the size of the styloid process, but this part of the bone is broken in the studied specimen.
Genus PlecotusGeoffroy, 1818
Plecotus auritus (Linnaeus, 1758) – Brown long-eared bat (present habitat and distribution in Table 1).
Plecotus austriacus (Fischer, 1829) – Grey long-eared bat (present habitat and distribution in Table 1).
DESCRIPTION: The C has an almost circular occlusal outline. The crown has two sides (labial and lingual), both of which have a longitudinal furrow in the distal half, the junction between the two sides forming a blunt edge. The cingulum is thick and almost regular. The M2 has an elongated, rectangular occlusal outline, and the base of the protocone has a circular outline and presents a thick cingulum, which continues along the distal and mesial sides. The M2 has a well-developed parastyle forming an angle, and a smaller bent metastyle. The metacone is markedly stronger and higher than the paracone. The lower molars are of the myotodont type and have a thick and very irregular cingulum. The m1 trigonid is open, and the lingual cusplets are almost aligned except for the hypoconulid, which is slightly labial. The paraconid, metaconid and entoconid have a similar height and are somewhat short and wide. The union between the paraconid and metaconid shows a deep depression.
DISCUSSION: The material has been assigned to the genus Plecotus on the basis of the occlusal outline of the M2, the geometry and disposition of the trigonid and hypoconulid of the lower molars, and the occlusal outline and furrow of C, which are diagnostic characters of the two Iberian species of the genus according to the fossil and present-day specimens described by Sevilla, 1988 and Sevilla, 1990. However, distinguishing between P. auritus and P. austriacus is hard when the sample is composed of isolated teeth. According to Sevilla (1988), C and P4 are diagnostic elements. The junction between the two sides of the upper canines forming a blunted edge is characteristic of P. auritus, as this edge is sharper and the cusp is longer in P. austriacus. The upper canine probably belongs to P. auritus, but as there are no clear criteria applicable for distinguishing the remaining specimens, the whole sample has been classified as P. gr. auritus/austriacus.
Genus HypsugoKolenati, 1856 vel Genus PipistrellusKaup, 1829
Hypsugo savii (Bonaparte, 1837) – Savi's pipistrelle (present habitat and distribution in Table 1).
P. kuhlii (Kuhl, 1817) – Kuhl's pipistrelle (present habitat and distribution in Table 1).
P. nathusii (Keyserling. and Blasius, 1839) – Nathusius's pipistrelle (present habitat and distribution in Table 1).
Hypsugo savii vel Pipistrellus gr. kuhlii/nathusii
MATERIAL (Fig. 5C; Appendix A): 1 hu (d. e. + diap.).
DESCRIPTION: The distal humeral epiphysis has an internally projected delta-shaped styloid process. The antecubital fossa is rather deep. In external view, the trochlea and the condyle dip slightly to the epitrochlea and there is a shallow furrow between them; the epicondyle is hidden by the condyle. The posterior epicondyle outline presents a concave–convex morphology with a bulge at the edge. The distal margin of the trochlea goes slightly beyond that of the condyle, and the distal margin of the condyle goes slightly beyond that of the epicondyle.
DISCUSSION: Based on the general morphology and the size (Appendix A) of the specimen, it can belong to the taxa H. savii, P. kuhlii or P. nathusii based on Dupuis (1986) and Felten et al. (1973).
Genus MiniopterusBonaparte, 1837
Miniopterus schreibersii (Kuhl, 1817) – Schreibers’ bat (present habitat and distribution in Table 1).
MATERIAL (Fig. 4D; Fig. 5D; Appendix A): 1 hu. (d. e. + diap.); 1 C; 1 m2.
DESCRIPTION: This is a medium-sized bat in relation to other cave-dwelling bats. The C has a nearly oval occlusal outline. The apex and the root are aligned. The cingulum is continuous, slightly irregular and not very thick. The crown has four deep longitudinal furrows, one of them in the distolabial margin, the others on the lingual side. The lower molars are of the nyctalodont type and present a thin and regular cingulum. The m2 trigonid is open and shows a small lingual cingulum. The lingual cusplets are almost aligned except for the hypoconulid, which has a labial position. The metaconid is the highest lingual cusp; the unions between the cusplets are deep depressions. The entocristid forms a marked angle with the entoconid. The paralophid is concave. The distal humeral epiphysis presents a long, flattened blade-shaped styloid process. The antecubital fossa is well-developed. In external view, the condyle dips slightly to the trochlea. The furrows separating the condyle, trochlea and epicondyle are rather deep, the furrow between the latter two being the deepest.
DISCUSSION: According to Sevilla (1988), the deep longitudinal furrows of the C and the morphology and disposition of the m2 cusplets described above are diagnostic for the species M. schreibersii, as is the distal humeral epiphysis, which presents a blade-shaped styloid process and deep external furrows.
Results and discussion
All chiropteran fossil remains were recovered from the AGP-7 assemblage totaling 51 specimens. Every specimen has been described and assigned to a taxon. Nine taxa of the order Chiroptera have been identified (Table 2).
Taphonomy. The origin of the accumulation
The post-cranial remains in the bat fossil assemblage of AGP-7 represent less than 8% of the total, the dominant elements are isolated teeth while complete bones are scarce (Table 3). Teeth show different colours from white to dark brown (e.g., Fig. 4C, I). Clear evidence of digestion is only observed in one humerus affecting its olecranon fossa (Fig. 5C). Around 25% of the remains are partially affected by manganese oxide mineralization (e.g., Fig. 4A, F; Fig. 5D), which is a common post-burial alteration in cave-sediments (Bennàsar-Serra, 2010); only one isolated tooth is affected by carbonate concretion. Notably, 85% of dental remains belonged to yearlings showing no teeth-wear; only a few dental specimens of R. ferrumequinum (e.g., Fig. 4H) and M. gr. myotis/blythii belonged adult individuals.
The abundance of such typical cave-dwelling bats as R. ferrumequinum and M. gr. myotis/blythii suggests that both taxa were accumulated by in situ death within the cave. However, the scarce complete bones, the differential loss of post-cranial remains and the dominance of isolated teeth indicate some grade of transport. We propose a process of dragging-out of the remains from an inner hall of the cave where the in situ accumulation actually took place. Both taxa show a mixed age composition with yearlings and old individuals, which is the typical pattern of hibernating colonies thanatocenosis (López-García and Sevilla, 2012). Some minority taxa in AGP-7 assemblage are also typical cave-dwelling species, such as R. hipposideros, M. bechsteinii, M. emarginatus or Miniopterus schreibersii. On the other hand, some others rarely roost in Iberian caves: M. daubentonii, P. gr. auritus/austriacus, H. avii and P. gr. kuhlii/nathusii (Palomo et al., 2007), from which at least one specimen shows marks of digestion. A relatively high diversity of the bat assemblage together with a low proportion of bat fossils in the entire small vertebrate sample (bat remains represent less of the 5% of the total recovered remains of small vertebrates) is the characteristic pattern of an owl-accumulated assemblage (Kowalski, 1995 and López-García and Sevilla, 2012). Our proposal for this group is that probably both processes (owl-predation and death within the cave) contributed to the accumulation. The near-exclusive presence of yearlings could be explained as the typical pattern of nursery colonies thanatocenosis (López-García and Sevilla, 2012) or as a higher vulnerability to predation for young bats (Petrželková et al., 2004 and Rosina and Shokhrin, 2011).
Palaeoenvironmental approach
The results of our habitat weighting analysis suggest a patched landscape where woodland and transitional woodland areas would dominate, alternating with seasonal meadows (Fig. 6). The occurrence of the typical cave-bats R. ferrumequinum, R. hipposideros, M. gr. myotis/blythii and M. schreibersii indicates the presence of exposed rock areas with cavities and/or fissures where they would roost. The existence of some surface water bodies is a limiting factor for the taxa M. daubentonii and R. hipposideros. Although all the species identified in the bat assemblage from AGP-7 currently inhabit the Iberian Peninsula, no similar community has been reported in the central Ebro Valley. However, similar communities are known in other Iberian regions: the eastern Cantabrian margin, the Sierra de Guadarrama (central Iberia) and the Sierra de Francia (central-western Iberia) (Palomo et al., 2007); all these localities are under the influence of an oceanic climate. However, the lack of studies of present-day bat communities in the area of the central Ebro Valley must also be taken into account. Our proposal, based on the analysis of the bat assemblage from AGP-7, is that the area surrounding the locality was characterized by a mosaic of forest and meadows habitats with transitional areas, where the water supply was great enough to support a rich vegetation cover and to maintain a generally wetter, more oceanic-influenced climate than nowadays in the area.
The other fossil mammals from AGP-7 consist of taxa with highly diverse habitat preferences, which is consistent with the patched landscape obtained with bats. The dominant small mammal at AGP-7 is Iberomys cabrerae, an endemic Iberian species adapted to a Mediterranean climate and evergreen meadows (Cuenca-Bescós et al., 2010 and López-García and Cuenca-Bescós, 2012); other small mammals with a lower presence show preferences for woodland (Apodemus sp.) or rocky, snow areas, such as Chionomys nivalis. The large mammals are eurythermal species with preference for temperate climate. The herbivorous mammals of the association, of which Cervus elaphus is the dominant taxon, points to a landscape characterized by a mosaic of forest and open habitats with transitional areas as the chiropteran association does.
Comparison with other Iberian Late Pleistocene bat assemblages
Several Late Pleistocene bat assemblages have been studied all along Spain (Appendix B) although this is the first report from the Ebro Valley region. The dominant taxon in AGP-7 site is R. ferrumequinum, which is a common species in Pleistocene Iberian sites; regarding the two following taxa in terms of abundance, M. gr. myotis/blythii is also a common species, but P. gr. auritus/austriacus is rather uncommon (López-García, 2008 and Sevilla, 1988). Besides, other common species in Quaternary sites, such as R. gr. euryale/mehelyi and M. schreibersii are respectively absent or poorly represented in AGP-7; both of them are markedly thermophilic (Sevilla, 1988). The assemblage includes a rare species in the Iberian Pleistocene: M. daubentonii, which has only been reported in one other Iberian locality: Quibas (Lower Pleistocene, Murcia, Spain), a cool-period site (Agustí et al., 2009). Finally, a remarkable characteristic of the AGP-7 site is its high bat diversity despite of the small amount of remains recovered. Considering other Iberian sites aged MIS 3 (Appendix B), only La Carihuela (Granada) has a more diverse bat association than AGP-7, having been identified 14 taxa in there (Sevilla, 1988) among which are all the taxa identified in AGP-7. Another relatively diverse site is El Reguerillo (Madrid) where seven bat taxa have been identified although this assemblage rather differs from the one in Aguilón in terms of taxonomic composition (Sevilla, 1988; Appendix B).
Conclusions
The following species were identified in a small set of bat remains obtained from AGP-7 (MNI = 15): R. ferrumequinum, R. hipposideros, M. gr. myotis/blythii, M. cf. bechsteinii, M. emarginatus, M. daubentonii, P. gr. auritus/austriacus, H. savii vel. P. gr. kuhlii/nathusii and M. schreibersii. All the identified bat taxa from AGP-7 presently inhabit the Iberian Peninsula, but the association is more typical of regions with an oceanic climate and woodland environments than the extant conditions in the Aguilón surrounding area. Our conclusion is that the area surrounding the AGP-7 locality presented a heterogeneous vegetation structure composed of an alternation of forest and meadow zones, where the water supply was great enough to support a rich vegetation cover and to maintain generally wetter, rather more oceanic-influenced climate conditions than nowadays. Further studies on Late Pleistocene sites in the region of the Ebro Valley will be of interest in order to obtain a complete overview of the state of chiropters in this area and period; for the moment only one Iberian site with this age, La Carihuela, is comparable with AGP-7 in terms of diversity and taxonomic composition.
Acknowledgments
The first author is the recipient of a Ph. D. fellowship from the Gobierno de Aragón. This work is a contribution to the project “Grupos Consolidados H54” of the Gobierno de Aragón and the European Social Fund. Arturo Hernandez discovered AGP-7, and the CEA undertook the topography of the cave. C. Nuñez is the recipient of a Ph. D. fellowship from the Gobierno de Aragón and J.M. López-García is the recipient of a Beatriu de Pinós postdoctoral fellowship (2011BP-A00272) from the Generalitat de Catalunya co-funded by the EU through the Marie Curie Actions of the 7th Framework Program for R+D. We would like to thank the excavation and sieving-screening team (J. Rofes, R. Rabal, D. Ramón and J. Ruiz). I. Sasowsky, University of Akron, and F. Gutierrez, University of Zaragoza, give us information about the speleogenesys of the cave. We would also like to thank the staff of the Centro de Recuperación de Fauna Silvestre-La Alfranca for their help during the taxonomic analysis. J. Arroyo Cabrales collaborated making the photographs of the EBD collection, and C. Ibañez from the EBD, kindly loaned the comparative specimens. This work has also been partially supported by MICINN project CGL2009-12703-C03-03, CGL2012-38434-C03-01, and the Fundación Ancestros. R. Glasgow revised the English grammar and H.-A. Blain translated the French texts. We want to thank the Associate editor Prof. Lars van den Hoek Ostende and the two anonymous reviewers for their comments that improve the final version of the manuscript.
Measurements of bat specimens from Aguilón P7 site. The measures are expressed in millimeters.
Iberian Late Pleistocene (MIS 3) bat assemblages. Data obtained from <xref rid="bib0050" ref-type="bibr">Burjachs et al. (2012)</xref>, <xref rid="bib0155" ref-type="bibr">López-García (2008)</xref>, <xref rid="bib0175" ref-type="bibr">López-García et al., 2014</xref>, <xref rid="bib0180" ref-type="bibr">López-García et al., 2012a</xref> and <xref rid="bib0185" ref-type="bibr">López-García et al., 2012b</xref>, <xref rid="bib0245" ref-type="bibr">Sevilla (1988)</xref>
Rhinolophus ferreumequinumRhinolophus mehelyiRhinolophus euryaleRhinolophus euryale-mehelyiRhinolophus hipposiderosRhinolophus sp.Myotis myotisMyotis bythii (oxignathus)Myotis myotis blythiiMyotis bechsteiniMyotis nattereriMyotis emarginatusAguilónxxxcf.xNorthCueva de CovalejosxLezetxikixLabeko KobaxCenterCueva MillánxEl ReguerilloxxxxxxCueva de la Buena Pinta (U2)xxxxNorth-eastGabasaxxxxCova de l’Arbreda (G-H)xCova de l’Arbreda (I)xxCova de Teixoneres (II-III)xxxOlopte BxxxxGalls CarbonersxxAbric Romaní (E-O)xCova dels Xaragalls (C3-C8)xxE.Cova NegraxSouthLa CarihuelaxxxxxxxxxxIbex Cavecf.ZafarrayaMyotis daubentoniiMyotis capacciniiMyotis sp.Eptesicus serotinusEptesicus sp.Plecotus auritusPlecotus austriacusPlecotus auritus-austriacusBarbastella barbastellusNyctalus lasiopterusNycatlus leisleriPipistrellus pipistrellusPipistrellus sp.Hypsugo savii – Pipistrellus kuhlii-nathusiiMiniopterus schreibersiiMiniopterus sp.Tadarida teniotisAguilónxxxxNorthCueva de CovalejosLezetxikiLabeko KobaCenterCueva MillánEl ReguerilloxCueva de la Buena Pinta (U2)xNorth-eastGabasaCova de l’Arbreda (G-H)xxCova de l’Arbreda (I)Cova de Teixoneres (II-III)xxOlopte BxGalls CarbonersxxxAbric Romaní (E-O)xxxCova dels Xaragalls (C3-C8)xxxE.Cova NegraxSouthLa CarihuelaxxxxIbex CavexZafarraya
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Geographical and geological location of the Aguilón P7 site, modified from Cortés-Gracia and Casas-Sainz (1996).
Situation géographique et géologique du site d’Aguilón P7, modifié d’après Cortés-Gracia and Casas-Sainz (1996).
Plan view of the Aguilón P7 cave and the excavation grid. The following sectors were distinguished: E: entrada, P: pendiente, C: centro and H: hiedra in the outer entrance hall; and sectors G: galería and F: fondo in the inner gallery. Modified from Gisbert and Pastor (2009); Cuenca-Bescós et al. (2010).
Plan de la grotte P7 d’Aguilón et du carroyage de fouille. Les secteurs suivants ont été distingués E : entrada, P : pendiente, C : centro et H : hiedra dans le hall d’entrée extérieur et les secteurs G : galería et F : fondo dans la galerie intérieure. Modifié d’après Gisbert et Pastor (2009) ; Cuenca-Bescós et al. (2010).
Anatomy of bat cranial skeleton. A. Lateral view of skull and hemimandible, and ventral view of skull, modified from Benzal and de Paz (1990). B. Occlusal view of upper molar (left) and lower molar (right), modified from Sevilla (1988). Abbreviated names are explained in the paragraph “Abbreviations”.
Anatomie du crâne d’une chauve-souris. A. Vue latérale du crâne et de la mandibule, et vue ventrale du crâne, modifié d’après Benzal et de Paz (1990). B. Vue occlusale de la molaire supérieure (à gauche) et de la molaire inférieure (à droite), modifié d’après Sevilla (1988). Les abréviations sont expliquées dans le paragraphe « Abréviations ».
Fossil cranial bat remains from the Aguilón P7 site. A. Right C of Rhinolophus ferrumequinum (MPZ 2014/257). B. Left C of Myotis emarginatus (MPZ 2014/278). C. Left C of Plecotus gr. auritus/austriacus (MPZ 2014/286). D. Right C of Miniopterus schreibersii (MPZ 2014/290). E. Fragment of right maxilla of Rhinolophus hipposideros (MPZ 2014/240). F. Fragment of left maxilla of Myotis gr. myotis/blythii (MPZ 2014/271). G. Right M2 of Plecotus gr. auritus/austriacus (MPZ 2014/285). H. Left mandible of Rhinolophus ferrumequinum (MPZ 2014/241). I. Left p4 of Myotis gr. myotis/blythii (MPZ 2014/265). J. Fragment of left mandible of Myotis daubentonii (MPZ 2014/279).
Restes crâniens fossiles de chauve-souris du site d’Aguilón P7. A. C droite de Rhinolophus ferrumequinum (MPZ 2014/257). B. C gauche de Myotis emarginatus (MPZ 2014/278). C. C gauche de Plecotus gr. auritus/austriacus (MPZ 2014/286). D. C droite de Miniopterus schreibersii (MPZ 2014/290). E. Fragment de maxillaire droit de Rhinolophus hipposideros (MPZ 2014/240). F. Fragment de maxillaire gauche de Myotis gr. myotis/blythii (MPZ 2014/271). G. M2 droite de Plecotus gr. auritus/austriacus (MPZ 2014/285). H. Mandibule gauche de Rhinolophus ferrumequinum (MPZ 2014/241). I. p4 gauche de Myotis gr. myotis/blythii (MPZ 2014/265). J. Fragment du mandibule gauche de Myotis daubentonii (MPZ 2014/279).
Distal humeral epiphysis of fossil bat remains from the Aguilón P7 site. A. Left distal humeral epiphysis of M. cf. bechsteinii (MPZ 2014/282). B. Left distal humeral epiphysis of M. emarginatus/daubentonii (MPZ 2014/281). C. Left distal humeral epiphysis of H. savii vel. P. gr. kuhlii/nathusii (MPZ 2014/287). D. Left distal humeral epiphysis of M. schreibersii (MPZ 2014/288).
Restes postcrâniens fossiles de chauve-souris du site d’Aguilón P7. A. Épiphyse distale gauche de M. cf. bechsteinii (MPZ 2014/282). B. Épiphyse distale gauche de M. emarginatus/daubentonii (MPZ 2014/281). C. Épiphyse distale gauche de H. savii vel. P. gr. kuhlii/nathusii (MPZ 2014/287). D. Épiphyse distale gauche de M. schreibersii (MPZ 2014/288).
Habitat composition percentages inferred from the bat fossil assemblage of Aguilón P7 (excluding the material from sector F). OD: open dry; OH: open humid; Wo: woodland/woodland margins; Ro: rocky; Wa: water. The most abundant taxa in the assemblage are A: R. ferrumequinum; B: M. gr. myotis/blythii; C: P. gr. auritus/austriacus.
Pourcentages de composition par habitat, déduits de l’assemblage fossile de chiroptères d’Aguilón P7 excepté le matériel du secteur F. OD : ouvert et sec ; OH : ouvert et humide ; Wo : forêt/bordures de forêt ; Ro : rocheux ; Wa : aquatique. Les taxons les plus abondants dans l’assemblage sont A: R. ferrumequinum; B: M. gr. myotis/blythii; C: P. gr. auritus/austriacus.
Synthetic notes on current distribution, habitat preferences and roosting habits for the bat species considered in this work. Masl: meters above sea level. Data obtained from Palomo et al. (2007).
Notes synthétiques sur la distribution actuelle, préférences d’habitat et modes de gite pour les espèces de chiroptères considérées dans ce travail. Masl : mètres au-dessus du niveau de la mer. Les données ont été extraites de Palomo et al. (2007).
TaxonGeographic range in EuropeDistribution (Spain)Alt. range (Spain)Habitat preferences (Spain)Roosting habits (Spain)Rhinolophus ferrumequinumFrom England to the Mediterranean peninsulasAll Spain (with the Baleares); scarcer in the internal areas0–1600 maslUbiquitous; mainly prefer transitional woodland areasMainly caves. Small coloniesRhinolophus hipposiderosFrom the southern Germany and Poland to the Medit. peninsulasAll Spain (with the Baleares); scarcer in the internal areas0–2000 maslWoodland and transitional woodland; water bodiesCaves. Nursery colonies; solitary hibernationMyotis myotisWestern, central and southern EuropeAll Spain (with the Baleares); scarcer in the central Ebro Valley and the South Submeseta0–2060 maslWoodland, transitional woodland and woodland marginsCaves. Massive nursery colonies; smaller hibernating coloniesMyotis blythiiSouthern Europe and southern parts of central EuropeAll Spain (with the Baleares), except the Northwest margin; very scarce0–100 maslOpen areas, seasonal meadowsCaves. Massive nursery colonies; smaller hibernating coloniesMyotis bechsteiniiAcross central and southern Europe (including islands)Mainly in the Cantabrian margen and the Central Range; very scarce0–500 maslWoodlandMainly caves. Small nursery colonies; solitary hibernationMyotis emarginatusFrom the South of the Netherlands and Germany across southern EuropeAll Spain (with the Baleares); very scarce, especially in the most internal areas0–1700 maslWoodlands; fre-quently avoiding the most closed areasCaves. Nursery coloniesMyotis daubentoniiAlmost all Eurpe; scarcer at the southern margins of the Medit. peninsulasAcross the north-western Peninsula, sporadically in the Medit. margin0–1600 maslAlways linked to water bodiesCaves, tree-holes. Small nursery colonies; solitary hibernationPlecotus auritusAll the palaearctic regionMountain ranges of the northern Peninsula0–1600 maslMature woodlandsCaves, tree-holes. Nursery colonies; solitary hibernationPlecotus austriacusMiddle latitudes, from England to southern EuropeAll Spain (with the Baleares); not very abundant but frequent0–2100 maslWoodland, transitional woodland and dry, seasonal meadowsCaves, tree-holes. Nursery colonies; solitary hibernationPipistrellus kuhliiAcross southern EuropeAll Spain (with the Baleares and the Canaries); except north-western margin0–1500 maslMainly open woodland; but high synanthropyTree-holes and crevices. Small to medium nursery coloniesPipistrellus nathusiiAll Europe from Scandinavia to the Medit. peninsulasMainly limited to northern Peninsula; very scarce0–900 maslWoodlandTree-holes, crevices. Small to medium nursery coloniesHypsugo saviiAcross the Medit. EuropeAlmost all Spain (with the Canaries); scarce0–3300 maslAll type of habitatsTree-holes, crevices. Small to medium nursery coloniesMiniopterus schreibersiiAcross the southern EuropeAll Spain (with the Baleares)0–1400 maslOpen and woodland areas; rocky habitats where caves or crevices occurCaves, crevices. Massive nursery and hibernation colonies
Quantitative presence indices for the taxa making up the total bat assemblage of Aguilón P7, Zaragoza, Spain, and distribution by habitat (according to Palomo et al., 2007). NISP: number of identified specimens; MNI: minimum number of individuals; OD: open dry; OH: open humid; Wo: woodland/woodland margins; Ro: rocky; Wa: water.
Indices de présence quantitative pour les taxons de la totalité de l’assemblage de chiroptères d’Aguilón P7, Saragosse, Espagne, et distribution par habitat (selon Palomo et al., 2007). NISP : nombre de spécimens identifiés ; MNI : nombre minimum d’individus ; OD : ouvert et sec ; OH : ouvert et humide ; Wo : forêt/bordures de forêt ; Ro : rocheux ; Wa : aquatique.
Quantitative data for the osteological composition of the total bat assemblage of Aguilón P7 (Zaragoza, Spain). n: number of elements; %: percentage of the whole bat assemblage constituted by each element.
Données quantitatives pour la composition ostéologique de l’assemblage total de chiroptères d’Aguilón P7 (Saragosse, Espagne). n : nombre d’éléments ; % : pourcentage de chaque élément dans tout l’assemblage de chauves-souris.
Elementn%max (fr)35.9M47.8C611.8P212P435.9md23.9md (fr)23.9c35.9p435.9m1223.5m (fr)815.7hu (d. e. + diap.)47.8Total51100