Riparo Mochi is considered a key site for the Middle–Upper Palaeolithic chrono-cultural sequence in the Mediterranean area. Climatic and environmental conditions have been reconstructed after the study of the small mammal assemblage by means of linear regression method and the Habitat Weighting method. The climate proxies for the late Mousterian and Proto-Aurignacian (cultural units I to G) indicate an environment with a relative high percentage of forest component during two cold and one relative warm oscillations. Open environments with low temperatures are registered for the Gravettian (cultural Unit D) allowing a possible relation to the Heinrich 3 Event. The final Gravettian (cultural Unit C) shows a relative warm peak, tentatively related to Greenland Interstadial 2. The late Epigravettian (cultural Unit A) is characterized by a high percentage of forest-related environment suggesting that this horizon formed during the Bølling–Allerød Interstadial. Finally, the Riparo Mochi sequence contributes to reconstruct the processes of faunal changes along the northern Tyrrhenian coast in a period of climatic instability, especially during the end of Marine Isotope Stage 3 when foraging groups adopted different mobility strategies between southern France and the Italian peninsula.
Le Riparo Mochi est un site clé pour la séquence chrono-culturelle Paléolithique moyen–supérieur de la région méditerranéenne. Les conditions climatiques et environnementales ont été reconstruites à partir de l’étude de l’assemblage de petits mammifères par les méthodes de moyennes de régression linéaire et Habitat Weighting. Les données climatiques pour le Moustérien final et le Proto-Aurignacien (unités culturelles I à G) indiquent un environnement composé, d’une part relativement importante de forêt au cours de deux oscillations froides et une oscillation relativement chaude. Un environnement ouvert avec des températures basses est enregistré pour le Gravettien (unité culturelle D), permettant une corrélation possible avec l’évènement Heinrich 3. Le Gravettien final (unité culturelle C) présente des pics chauds, pouvant être reliés au Greenland Interstadial 2. L’Épigravettien final (unité culturelle A) est caractérisé par une proportion importante d’environnement forestier, suggérant que cet horizon s’est formé pendant l’interglaciaire Bølling–Allerød. La séquence du Riparo Mochi contribue à la reconstruction des processus de changement faunique le long de la côte Nord-Tyrrhénienne pendant une période d’instabilité climatique, particulièrement à la fin du MIS 3, lorsque les groupes de chasseurs-cueilleurs adoptent différentes stratégies de mobilité entre le Sud de la France et la péninsule Italienne.
Upper Pleistocene, Paleoclimatology, Late Glacial, Gravettian, Insectivores, RodentsPléistocène supérieur, Paléoclimatologie, Dernier Glaciaire, Gravettien, Insectivores, RongeurspresentedHandled by Lars W. van den Hoek OstendeIntroduction
The environment and climate reconstructions of MIS 3 and 2 of central and southern Europe have been widely discussed (Barron and Pollard, 2002, Monegato et al., 2011 and Pini et al., 2010a among others). Most research focuses on the possible climatic causes that led to the Neanderthal extinction and that could have driven the diffusion of Anatomically Modern Humans in Europe (e.g., Fedele et al., 2002, Higham et al., 2009 and López-García et al., 2015).
Thirteen continuous and repeated climate cycles, such as Dansgaard–Oeschger (DO = GI) and Heinrich Events (H), have been recognized between 50,000 and 20,000 years BP in Greenland and North Atlantic cores (Andersen et al., 2006, Bond et al., 1993, Dansgaard et al., 1993 and Rasmussen et al., 2014, among others). By contrast, continental records supporting these events are rare and usually incomplete, mainly because of hiatuses in sedimentation and/or poor dating (Spötl and Mangini, 2002). Nevertheless, several studies allowed us to correlate the MIS 3 climate cycles to changes in the small mammal communities found throughout the stratigraphic sequences, especially in the Mediterranean area (e.g., Berto et al., 2017a, Burjachs et al., 2012, López-García et al., 2014b, López-García et al., 2012, López-García et al., 2015 and Royer et al., 2013) contributing to reconstruct the climate conditions in which prehistoric hunter-gatherers lived.
In this context, little is known about small mammal variations in northwestern Italy during MIS 3 and 2, especially in the coastal region (Berto, 2013). This area has been recently recognized as key to better understand the land use strategies of the last Neanderthals as well as the diffusion of the Upper Palaeolithic technocomplexes such as the Protoaurignacian, Aurignacian, and Gravettian (Grimaldi and Santaniello, 2014 and Grimaldi et al., 2014, Santaniello, 2016). The northern Tyrrhenian coast, the so-called Liguro-Provençal Arc (Fig. 1a), forms a narrow littoral corridor, 400 km long and a few kilometres wide, delimited by the Apennines and the western Alps to the north, and the Tyrrhenian Sea to the south, further linking central Tyrrhenian Italy to the Rhône Valley in France. During the MIS 3, the landscape of the Liguro-Provençal Arc was not very different from today's (Arobba and Caramiello, 2009, Pons-Branchu et al., 2010 and Watts et al., 2000). This region was a narrow corridor even during glacial time according to the bathymetry of the sea bottom (IBCM, 2014). Consequently, from an archaeological perspective, the Liguro-Provençal Arc should have been a natural axis channelling the exchange of both humans and animals between central Italy and southern France (Porraz et al., 2010).
Here, we present for the first time a study of the entire collection of small mammals (insectivores and rodents) from the Riparo Mochi site (Balzi Rossi, Ventimiglia, Italy). This site holds one of the most important sections in the Tyrrhenian area, spanning from Mousterian to Epigravettian, and it is considered a key site for the investigations on the Middle-Upper Palaeolithic boundary (Douka et al., 2012 and Higham et al., 2014). This paper aims to reconstruct the local environment and climate changes based on the small mammals. The reconstruction will be discussed taking in account other northern Italy small mammal sequences – such as Grotta del Broion (Colamussi, 2002) and Grotta di Fumane (López-García et al., 2015) – as well as pollen archives from southern France and northern Italy with their correlation to Oxygen Isotope records.
Riparo Mochi
Riparo Mochi (Mochi rock shelter) is a broad, shallow rock shelter situated in the Balzi Rossi archaeological complex, in western Liguria on the border between Italy and France (43°47′3.66″N, 7°32′4.18″E, 30 m a.s.l.). It is one of several prehistoric sites, also known as “Grimaldi caves” (de Villeneuve et al., 1919) that provided archaeological deposits ranging from the early Middle to the final Upper Palaeolithic (Fig. 1a).
The site was discovered in 1938 by A.C. Blanc and L. Cardini of the Italian Institute of Human Palaeontology (IsIPU). They performed first a small-scale testing over three trenches (A, B, C) (Blanc, 1938); trench A, later called “Central Trench”, was enlarged and excavated systematically in 1941, 1942 and 1949. In 1959, another excavation carried out by Cardini yielded the Upper Palaeolithic layers located at the eastern and western side of the Central Trench. For nearly four decades the site was abandoned, but from 1995 to 2005, A. Bietti (La Sapienza University and IIPU) resumed the excavations. Since 2007, the University of Trento, in collaboration with the former Soprintendenza Archeologica della Liguria started new research on this site, which is still ongoing. Thanks to this project, new radiocarbon dates of the lower part of the Upper Palaeolithic sequence have been published (Douka et al., 2012). Based on this evidence, the bottom of the cultural Unit G (Protoaurignacian) of Riparo Mochi, dating to around 37–36.5 ka BP, is the oldest directly-dated Upper Palaeolithic assemblage in Italy.
The chronocultural sequence, approximately 10 m deep, consists of nine units, mainly inferred in retrospect upon the characteristics of the embedded cultural remains (Grimaldi and Santaniello, 2014, Grimaldi et al., 2014, Laplace, 1977 and Tomasso et al., 2014 among others), and named A to I, from top to bottom (Fig. 1b). Each cultural unit was excavated in 10 to 5 cm spits. Geoarchaeological observations on the upper 5.5 m of the sequence (Unit C–H) allow us to subdivide it into 3 facies. The first (Facies 1) consists of thick layers composed of limestone rubble in a sandy silty loam matrix; the second (Facies 2) is composed of silty clay loam units with hearts interbedded between them (Facies 3) (Douka et al., 2012). The lithological characteristics of the sequence in association with the site chronology, already described in Douka et al. (2012), show that the sequence was deposited mainly between MIS 3 and 2. Sixteen radiocarbon dates are available from the top of the cultural Unit I (Mousterian) to the bottom of cultural Unit D (Noaillian–Gravettian) providing a chronological range between 41,630–40,410 cal. BP and 29,380–28,510 years cal. BP. The upper part of the sequence has not been dated yet and the relative chronology is inferred by lithic evidence: the cultural Unit A yielded late Epigravettian tools while the cultural Unit C, covered by the sterile Unit B, has been attributed to Final Gravettian.
Previous paleoenvironment analyses
Several paleoenvironment analyses have been previously carried out on Riparo Mochi. These studies have focused on the pollen sequence (Renault-Miskovsky, 1972), large mammals (Alhaique, 2000, Arellano, 2004, Arellano, 2009 and Zeppieri, 2009, summarised in Tagliacozzo et al., 2012), and a sample of small mammals (Abbassi, 1999).
Twelve pollen samples collected from cultural Units G and F–E (1959 Cardini sequence, East sector) allowed us to reconstruct the environment of the region at the beginning of the Upper Palaeolithic (Renault-Miskovsky, 1972). The identified pollen essences came from plants currently living in the reliefs next to the site (starting from 600 m a.s.l.), such as conifers and deciduous trees, and from typical Mediterranean plants such as maritime pine and Oleaceae, nowadays present along the coast. Four environment oscillations were detected: a change from an open environment (Arboreal Pollen = 5%), with a regional cold and dry climate, to an increase of wooden areas (AP = 40–65%) with termophilic essences (Oleaceae 10–30%), suggesting a warmer phase related to the Arcy interstadial. Other two oscillations, testified only by two samples in Units F and E, show a cold and arid climate followed by a warmer phase.
The faunal evidence coming from Riparo Mochi was only partially studied by several authors. Arellano (2009) reported the large mammal species for the Mousterian sequence (cultural Unit I) without providing any detailed paleoenvironment and paleoclimatic reconstruction; however, three environmental changes were observed. In the lower spits, cervids are dominant and wild boar (Sus scrofa) is very abundant while the presence of roe deer suggests a relatively humid climate. This condition changes in the medium part of the unit where mammals such as Rupicapra rupicapra and Capra ibex increase and wild boar–among the forest ungulates, the most sensitive one to climatic changes–shows an abrupt decline, suggesting a colder phase in association with more arid conditions. In the upper part of the unit (i.e. final Mousterian), the forest fauna becomes dominant in association with Dama dama remains.
Alhaique (2000) analysed the faunal assemblage coming from the 1996–1996 Bietti's excavation (G to C Units). The Upper Palaeolithic sequence, although poor in material, provided some information. Red deer and roe deer are present in the whole sequence, but roe deer shows a higher frequency in unit G. Marmota marmota is also present in Units C and D; according to the author, the only individual found in Unit F might be intrusive.
A more detailed study of the large mammals evidence coming from Unit D (Gravettian, 1959 Cardini's excavation) is presented in Zeppieri (2009), later summarized in Tagliacozzo et al. (2012). The faunal remains from Unit D were divided into phases (from IV to I, from the bottom to top of the Unit) that have been associated with seasonality of several occupations; this subdivision should no longer be considered correct as it was established following the artificial stratigraphy defined by Cardini himself without any viable correlation with the geo-archaeological evidence (compare with Santaniello, 2016). Broadly speaking, at the base of the Unit D (Phase IV), the faunal spectrum indicates a wooded environment, with conifers and deciduous trees, characterized by a temperate to cool, climate. Such an ecosystem changed due to colder temperatures and a dry climate in the central phase of Unit D (Phase III and II), where the upper limit of the forest left space for alpine meadows. Finally, on top of Unit D (Phase I), the forest coverage was thicker, with temperate to cool and more humid climatic conditions. However, in all periods, there is a contemporaneous presence of temperate and forest fauna (Cervus elaphus, Capreolus capreolus, Sus scrofa) that probably lived in the immediate surroundings of the site, together with cold and open environment species (Capra ibex, Rupicapra rupicapra, Marmota marmota) which were present in mountain areas above the tree line.
A sample of small mammal fauna, collected by the “Laboratoire départemental de Préhistoire du Lazaret” in 1993 (Paunesco et al., 2010), was studied by Abbassi (1999). The sample was divided in cultural units following Blanc's (1954) stratigraphy with the only exception for the so called “Unit J”, probably representing the base of the Unit I. The rodents identified along the sequence are Marmota marmota, Eliomys quercinus, Muscardinus avellanarius, Microtus arvalis, Microtus (Terricola) multiplex, Chionomys nivalis, Arvicola amphibius, Arvicola sapidus, Clethrionomys glareolus, and Apodemus flavicollis, while the specimen identified as Microtus (Terricola) savii was later revised as Microtus (Terricola) multiplex (Paunesco et al., 2010). Abbassi (1999) suggested a Mediterranean climate, similar to present conditions, at the base of the sequence (possibly Unit “J”), which changes toward a colder and continental climate because of the presence of Chionomys nivalis and Microtus (Terricola) multiplex. The evidence found at the Middle–Upper Palaeolithic transition indicates an arid climate which becomes more continental in the upper Units G to C.
Material and methods
The small mammal remains from Riparo Mochi are represented by disarticulated bone fragments collected by water-screening using 1 mm mesh sieves during the Cardini (1938–1949, 1959) and Bietti (1995–2005) campaigns. The small mammal assemblage includes a total of 1244 remains, corresponding to a minimum number of 795 individuals (Table 1, Fig. 2). The specific attribution of this material was mainly based on the best diagnostic elements: mandible, maxilla and isolated teeth for rodents, mandible and maxilla for shrews, mandible, maxilla, isolated teeth and postcranial bones for Talpidae.
The taxonomic classification follows Wilson and Reeder (2005), except for Clethrionomys glareolus (for the priority over Myodes, see Tesakov et al., 2010). Data on the distribution and habitat of the species were taken from Amori et al. (2008), Boitani et al. (2003) and Mitchell-Jones et al. (1999).
We calculated the palaeodiversity using the Simpson index of Evenness = 1–∑(pi2), where pi is the proportion of individuals in the ith species (Harper, 2005 and Magurran, 2004). The evenness index is constrained between 0 and 1. The index has been calculated using PAST 3.04 avoiding redundant determinations (i.e. for Microtus agrestis, the individuals determined as Microtus cf. agrestis have not been included in the Simpson index calculation) (Hammer et al., 2001). A Rattus sp. first lower molar found on top of Unit D has been considered intrusive from Holocene sediments, thus this species has been not taken in account for paleoenvironment and paleoclimate reconstructions. Finally, despite the criticisms outlined above, a comparison with the seasonal phases described by Zeppieri in Tagliacozzo et al. (2012) is also discussed.
Paleoenvironment reconstruction
In order to reconstruct the paleoenvironment at Riparo Mochi, we used the Habitat Weighting method (Andrews, 2006 and Evans et al., 1981), assigning each small mammal taxon to habitat(s) where it can be found today in Europe. To this purpose, habitats were divided up into six types (Cuenca-Bescós et al., 2010, Cuenca-Bescós et al., 2009 and López-García et al., 2014a): open land with either dry or wet meadows (OD and OH, respectively); woodland environments, divided into open woodland, woodland margins and forest patches (OW) and woodland and mature forest habitat (W); water, areas along streams, lakes and ponds (Wa); and habitats with a suitable rocky or stony substratum (R) (Table 1).
Climate reconstruction
Paleoclimatic data from Riparo Mochi have been calculated using the bioclimatic model described by Hernández Fernández, 2001a and Hernández Fernández, 2001b. First, the mammal assemblage was assigned to the climatic types described in Hernández Fernández (2001b) and in Walter (1970), following the values established in Hernández Fernández and Peláez-Campomanes (2005): IV–subtropical with winter rains and summer droughts; VI–typical temperate; VII–arid-temperate; VIII–cold-temperate (boreal); IX–polar. The faunal assemblage was analysed using the Climatic Restriction Index (CRIi = 1/n, where n is the number of climatic zones inhabited by the species and i is the climatic zone where the species appears).
Afterwards, the Bioclimatic Component (BC) was calculated using the following formula: BCi = (Σ CRIi) × 100/S, where S is the number of species by unit at Riparo Mochi. From the BC, climatic parameters were estimated with multiple linear regression method using the values given by Hernández Fernández and Peláez-Campomanes (2005). Mean Annual Temperatures (MAT), Mean Temperatures of the Coldest and Warmest month (MTC and MTW respectively) and Mean Annual Precipitation (MAP) were obtained. The data were compared with the present conditions registered at Sanremo meteorological station (Imperia, Liguria, 43°47’N, 7°46’E, 9 m a.s.l.), approximately 20 km east from the site. The current data are: MAT = 16.4 °C, MTW = 23.5 °C, MTC = 9.5 °C and MAP = 773 mm.
Results and discussionThe Riparo Mochi sequence
The small mammal assemblage of Riparo Mochi is characterized by high percentage of open environment-related arvicolids such as Microtus arvalis and Microtus (Terricola) gr. multiplex-subterraneus. Species that live in forest or in forest margins (Apodemus gr. sylvaticus-flavicollis, Clethrionomys glareolus) reach relative high percentages only in Unit I, F, and A.
Arvicola amphibius, a species that in the Italian peninsula lives near water streams, lakes and ponds, is present throughout the sequence. All Arvicola specimens have positive or Microtus-type enamel (Fig. 2, number 1 and 2). Thus, the presence of Arvicola sapidus, a species characterized by a negative or Mimomys-type enamel, signalled by Abbassi (1999) at Riparo Mochi is not confirmed.
The BCi values (Table 2), related only to presence/absence of species, are dominated by component VI because most the rodents found at Riparo Mochi currently live in temperate climate areas (Mitchell-Jones et al., 1999). Nevertheless, the Habitat Weighting variations, connected to the abundant or poor presence of insectivores and rodents in the assemblage, indicate that several environment changes happened during the formation of the sequence.
As the Units from I to E (Table 1) are characterized by a low MNI, conclusions on environment and climate changes during the transition from the Mousterian to the Aurignacian must be considered with caution.
In Unit I (NMI = 45), the biodiversity is high and equally distributed (Simpson index = 0.865; Table 2) and the presence of forest-related species is relatively high (OW = 22.68%; W = 30.56). The precipitation value is the lowest of the entire sequence (MAP = 872 mm), while temperature proxies indicate the presence of very cold conditions (MAT = 3.9 °C). Such climate data are directly related to the high percentage of Chionomys nivalis (8.9%), the absence of forest species such as Glis glis, the low percentage of Erinaceus europaeus, and the high percentage of Microtus arvalis and Microtus (Terricola) gr. multiplex-subterraneus. Thus, it is possible to suggest the presence of an environment characterized by tree patches in an open and rocky area rather than a dense forested landscape.
The cold climate evolves to slightly warmer conditions through the sequence (Units H and G) reflected in a more open environment. The rise in temperature and open environment may be related to a different precipitation distribution that caused the loss of biodiversity and the forest environment reduction (Fig. 3).
Unit F (early Aurignacian) is the most reliable layer of this portion of the sequence (NMI = 75): it is dominated by Microtus arvalis (32%), but the genus Apodemus is well represented (Apodemus sp. and Apodemus gr. sylvaticus-flavicollis > 20%). The environment was probably made up of sparse wooden areas without bush and open meadows. This reconstruction can be implemented thanks to the palynological analyses carried out at the Riparo Bombrini, a rock shelter located few tens of meters eastward to Riparo Mochi, which yielded layers related to Middle-Upper Palaeolithic transition. The authors suggested the presence of space pinewood, mixed oakwood, and thermophilous taxa related to the coastal area in the final Mousterian layers; warmer temperatures with a reduced tree covering were recorded during the deposition of the early Upper Palaeolithic layers (Arobba and Caramiello, 2009).
The loss of equality in the small mammal assemblage distribution continues in Unit E of Riparo Mochi. Here, few small mammals have been recovered (NMI = 23) to reach the lowest values in Unit D. Phases D IV, III, and II are dominated by one species, Microtus arvalis; the biodiversity is relatively low when compared to the lowermost phase (1-D values from 0.598 to 0.714). The forest components decrease in favour of the one indicative of more open environments due to a cold and dry climate. From phase IV to II, a slight climate improvement is registered; its peak is reached during a relative temperate cycle which characterizes the phase I (top of Unit D) as well as the Unit C, even if the general context is dominated by cold climate conditions. In these units, the arise of a water component represented mainly by Arvicola amphibius suggests the presence of a fluvial or a water-related environment, probably due to the presence of the San Luigi river, about 450 m to the west of the site.
After the thin and sterile Unit B, the Riparo Mochi sequence ends with the cultural Unit A (Epigravettian). This Unit also has a low NMI but the high percentage of forest-related indicators and the relative warm temperatures suggest that the unit formed in an environment similar to the present one.
Correlation between Riparo Mochi sequence and other climate proxies
The relatively high percentage of forest related species in Units I to F suggests that, in the period of time between DO (=GI) 12 and H 4 (following Douka et al., 2012), the western Liguria environment was conditioned by the Mediterranean Sea influence. This is also confirmed by pollen data indicating the presence of Mediterranean plants (Oleaceae) through the Riparo Mochi sequence (Renault-Miskovsky, 1972). In contrast, the percentage decrease of Microtus (Terricola) multiplex-subterraneus and Clethrionomys glareolus suggests an environment modification from a forest with shrubs and a well-formed substratum (Unit I, Mousterian) to an open forest (Unit F, early Aurignacian). The change is probably due to a climate deterioration related to a stadial episode of MIS 3. Unfortunately, it is not possible to argue whether this change is related only to H 4 rather or any other stadials (GS 12, 11, 10, and 9).
Riparo Mochi Units from I to F can be compared to layers SU A5 + A6 to A1 of the Fumane Cave (Verona, Italy). There, a decrease in percentage of woodland coverage is registered and this reduction culminates in layers A3-A1 that are attributed to Heinrich Event 4 (López-García et al., 2015). Differences between the two sequences must be noticed: at the Riparo Mochi, the biodiversity is lower than in the Fumane cave and boreal/continental climate indicators, as well as eastern Europe species such as Dinaromys bogdanovi, Microtus oeconomus, Sicista betulina, and Cricetus cricetus are absent. The composition of the Riparo Mochi assemblage is better related to southwestern Italian peninsula coastal sites, at Grotta della Cala and Grotta della Serratura (Bambini, 1996 and Bertolini et al., 1996), where such cold climate related species are absent, rather than to northern Italian Peninsula sites (Berto, 2013; Bon and Boscato, 1996, López-García et al., 2014a and Masini and Abbazzi, 1997).
Three short and not yet chronologically determined interstadials characterized by sparse tree cover are also found in two French site pollen records: Les Échets, zone G (Beaulieu and Reille, 2008) and Bouchet (Reille et al., 2000). Finally, the Riparo Mochi cultural Units I to F can be related to the palaeoenvironmental reconstructions suggested for Fimon Lake (FP 17a, 17b, and 17c) and for Azzano Decimo (AZ 65-67), in the southern Alps. The southern Alps pollen records show a thick tree coverage and, even during stadial periods because the environment was dominated by Pinus and Betula probably due to orographic precipitations (Pini et al., 2010b, 2009). Available dates (Douka et al., 2012) allow us to relate the base of Unit D of Riparo Mochi (Zeppieri's phase IV) to Heinrich Event 3. The effects of the cold and arid climate are clearly visible on the small mammal assemblage–especially when phases IV and III of Unit D are considered – and they can be related to Les Échets (zone O), Fimon Lake (FP 18), and Azzano Decimo (AZ 68).
The Unit D (phase III) temperature increase followed by an open environment decrease visible at the top of Unit D of Riparo Mochi can be related to episodes following the H3. No abrupt changes are visible in the transition from the top of Unit D to Unit C, although the latter is attributed to the final Gravettian technocomplex, chronologically related to Last Glacial Maximum (LGM) and DO 2 (Berto, 2013; Douka et al., 2012 and Palma di Cesnola, 1993). No radiometric dates are available yet, but two hypotheses can be proposed assuming that the open environment decrease in Unit C is related to the DO 2: A) the upper part of Unit D can be related to DO 4-3, two cycles chronologically close and hardly distinguishable without radiometric datings in a continental sequence, a hiatus in sedimentation is also present between LGM and DO 2; B) the deposition of Unit D took place in period between H3 and LGM.
Italian small mammal sequences related to H3 and following DO cycles are very scarce; the climatic deterioration related to this event in northern Italy is visible only at Grotta del Broion (layers G2–F), where a decrease of forest related species is registered (Colamussi, 2002). In southern Italy, the forest indicators decrease at Grotta della Cala (3m-1) (Bambini, 1996) and the increase in percentage of Microtus arvalis at Grotta Paglicci (22b to 21c) can be related to H3 (Berto et al., 2017a).
The late Epigravettian Unit A of Riparo Mochi is characterized by the high percentage of forest related species. The Unit probably accumulated after the turnover registered during the Late Glacial in the small mammal communities. In southwestern France, cold-climate species such as Microtus oeconomus, Microtus gregalis and Spermophilus sp. were replaced by forest-related species (Royer et al., 2016) while in the southern Alps, species related to cold-continental climates such as Dinaromys bogdanovi, Microtus oeconomus and Cricetus cricetus disappeared (i.e. at Riparo Tagliente, Berto et al., 2017b). Finally, in southern Italy the small mammal communities, dominated during the MIS 3 and 2 by Microtus arvalis, changed to forest related assemblages (dominated by Glis glis) or to a strictly Mediterranean-related community dominated by Microtus (Terricola) savii (Berto et al., 2017a and López-García et al., 2014a). The changes are related to Bølling–Allerød Interstadial afforestation registered also in pollen sequences (Berto et al., 2017b and Ravazzi et al., 2007).
Conclusions
The small mammal analysis from Riparo Mochi allows us to reconstruct the environment and its changes in the region surrounding the site from the final Middle Palaeolithic to the Upper Palaeolithic. The changes are due to D/O cycles and Heinrich Events registered during MIS 3 and 2 and detected in numerous climate proxies of European continental sequences.
Due to its particular position, the small mammal community of Riparo Mochi has been influenced by the Mediterranean climate and, even during the coldest stadial period of MIS 3, species related to boreal regions usually documented in northern Italy did not reach this area.
During the Middle to Upper Palaeolithic transition, two cold and one slightly warm oscillations were detected depicting a forest and bush reduction near the shelter. It was probably due to stadial moments not highlighted throughout the low small mammals MNI registered in the collection.
During the Gravettian, the environment changed from one dominated mainly by open grasslands, related to a cold and dry climate, to one characterized by open forests with an increase of water streams, the latter probably due to the flow rate of the near San Luigi river. The radiometric date at the base of Unit D and the presence of lithic tools related to final Gravettian in Unit C indicate that this portion of the sequence formed between H3 and D/O (GI) 2.
The small mammal assemblage of the uppermost Unit A, related to final Epigravettian, indicates the presence of a forested environment. This Unit probably formed after the faunal turnover registered in southern Europe during the Late Glacial.
The sequence of small mammal assemblages at Riparo Mochi fills a gap between southern Italy and southern France small mammal sequences, confirming the sequence as a reference one not only for the Middle to Upper Palaeolithic transition but also for the transition from MIS 3 and MIS 2.
Acknowledgements
The Dipartimento di Lettere e Filosofia, Università degli studi di Trento (Italia) partially supported this study. The former Soprintendenza per i Beni Archeologici della Liguria granted the permission for the study; in particular, we thank Dr. Angiolo Del Lucchese and Dr. Elisabetta Starnini. We are grateful to many people and friends who helped us during the fieldwork, especially to M. A. Segré and the staff of the Museo Preistorico dei Balzi Rossi. We want to thank the editorial secretary Fathia Lemhemdi and the reviewers Juan Manuel López-García and Lars van den Hoek Ostende for their comments that improved the final version of the manuscript. The authors also thank Benedetto Sala and Elisa Luzi for their useful suggestions and Julie Arnaud for the French spelling check. Finally, we are in debt with A. Segre who generously gave us direct information and original field documentation about the Riparo Mochi excavation.
Supplementary data
BCi computed in each layer (BCi = (Σ CRIi) × 100/S); linear regression values (Rodentia) used for MAP, MAT, MTC and MTW calculation (Hernández Fernández and Peláez-Campomanes, 2005); species distribution by climatic preferences (Hernández Fernández, 2001a and Hernández Fernández, 2001b): I: equatorial; II: tropical with summer rains; II/III: transition tropical semi-arid; III: sub-tropical arid; IV: winter rain and summer drought; V: warm-temperate; VI: typical temperate; VII: arid-temperate; VIII: cold-temperate (boreal); IX: Arctic.
BCi calculés pour chaque niveau (BCi = (Σ CRIi) × 100/S) ; valeurs de régression linéaire (Rodentia) utilisées pour le calcul des MAP, MAT, MTC et MTW (Hernández Fernández and Peláez-Campomanes, 2005) ; distribution des espèces par préférence climatique (Hernández Fernández, 2001a and Hernández Fernández, 2001b) : I : équatoriale ; II : tropicale avec mousson estive ; II/III : transition tropicale semi-aride ; III : subtropicale aride ; IV : pluie hivernale et sécheresse estivale ; V : tempéré chaud ; VI : tempéré typique ; VII : tempéré aride ; VIII : tempéré froid (boréal) ; IX : arctique.
AbbassiM.Les rongeurs du sud-est de la France et de Ligurie : implications systématiques, biostratigraphiques et paleoenvironnementales (PhD thesis)1999Museum national d’histoire naturelleParisAlhaiqueF.Risultati preliminari dell’analisi dei resti faunistici rinvenuti nei livelli del Paleolitico superiore di Riparo Mochi (Balzi Rossi): scavi 1995-1996MalerbaG.CilliC.GiacobiniG.Atti Del 2°Convegno Nazionale Di Archeozoologia (Asti, 1997)2000Abaco EdizioniLa Spezia125–130AmoriG.ContoliL.NappiA.Fauna d’Italia: mammalia II. Erinaceomorpha, Soricomorpha, Lagomorpha, Rodentia2008CalderiniBolognaAndersenK.K.SvenssonA.JohnsenS.J.RasmussenS.O.BiglerM.RöthlisbergerR.RuthU.Siggaard-AndersenM.-L.Peder SteffensenJ.Dahl-JensenD.The Greenland Ice Core Chronology 2005, 15–42ka. Part 1: constructing the time scaleQuat. Sci. Rev.2520063246–325710.1016/j.quascirev.2006.08.002AndrewsP.Taphonomic effects of faunal impoverishment and faunal mixingPalaeogeogr. Palaeoclimatol. Palaeoecol.2412006572–589ArellanoA.Les grands mammifères des niveaux moustériens de l’abri Mochi (grottes de Grimaldi, Vintimille, Italie). Fouilles de 1949Quat. Nov. VII2004139–157ArellanoA.Les grands mammifères des niveaux moustériens de l’abri Mochi (grottes de Grimaldi, Vintimille, Italie). Fouilles de 1949Bull. Mus. Anthr. Préhist. Monaco49200929–39ArobbaD.CaramielloR.Analisi paleobotanica sui sedimenti del Riparo Bombrini (Balzi Rossi, Ventimiglia)Bull. musée Anthropol. Prehist. Monaco49200941–48BambiniI.Microteriofauna del Pleniglaciale superiore di Grotta della Cala (Marina di Camerota, Salerno) (Master thesis)1996Università di FerraraItalyBarronE.PollardD.High-Resolution Climate Simulations of Oxygen Isotope Stage 3 in EuropeQuat. Res582002296–30910.1006/qres.2002.2374BeaulieuJ.-L. deReilleM.A long Upper Pleistocene pollen record from Les Echets, near Lyon, FranceBoreas132008111–13210.1111/j.1502-3885.1984.tb00066.xBertoC.BoscatoP.BoschinF.LuziE.RonchitelliA.Paleoenvironmental and paleoclimatic context during the Upper Paleolithic (late Upper Pleistocene) in the Italian Peninsula. The small mammal record from Grotta Paglicci (Rignano Garganico, Foggia, Southern Italy)Quat. Sci. Rev.168201730–4110.1016/j.quascirev.2017.05.004BertoC.LuziE.Montanari CaniniG.GuerreschiA.FontanaF.Climate and landscape in Italy during Late Epigravettian. The Late Glacial small mammal sequence of Riparo Tagliente (Stallavena di Grezzana, Verona, Italy)Quat. Sci. Rev.1842017132–14210.1016/j.quascirev.2017.07.022BertoC.Distribuzione ed evoluzione delle associazioni a piccoli mammiferi della penisola italiana durante il Pleistocene superiore (PhD thesis)2013Università di FerraraBertoliniM.FedozziS.MartiniF.SalaB.Late Glacial and Holocene climatic oscillations inferred from the variations in the micromammal associations at Grotta della Serratura (Marina di Camerota, Salerno, S Italy)Il Quaternario - Ital. J. Quat. Sci.91996561–566BlancA.C.Nuovo giacimento Paleolitico e Mesolitico ai Balzi Rossi di GrimaldiRend. R. Acc. LinceiVI1938107–113BlancA.C.CardiniL.SegreA.G.Il riparo Mochi ai Balzi Rossi di Grimaldi. Scavi 1938-1949Paleontogr. Ital.5019541–43BoitaniL.LovariS.Vigna TagliantiA.Fauna d’Italia. Mammalia III. Carnivora, Artiodactylavol. 382003EdagricoleMilanoBonM.BoscatoP.Associazioni faunistiche dell’Olocene antico nell’Italia del sud: confronto tra un sito adriatico (Grotta delle Mura, Monopoli, Bari) ed uno tirrenico (Grotta della Serratura, Marina di Camerota, Salerno)Quat. - Ital. J. Quat. Sci.91996567–572BondG.BroeckerW.S.JohnsenS.McManusJ.LabeyrieL.JouzelJ.BonaniG.Correlations between climate records from North Atlantic sediments and Greenland iceNature3651993143–147BurjachsF.López-GarcíaJ.M.AlluéE.BlainH.-A.RivalsF.BennàsarM.ExpósitoI.Palaeoecology of Neanderthals during Dansgaard–Oeschger cycles in northeastern Iberia (Abric Romaní): From regional to global scaleQuat. Int.247201226–3710.1016/j.quaint.2011.01.035ColamussiV.Studi Climatici sul Quaternario mediante l’uso dei Micromammiferi (PhD thesis)2002Università di Ferrara e FirenzeCuenca-BescósG.StrausL.G.García-PimientaJ.C.MoralesM.R.G.López-GarcíaJ.M.Late Quaternary small mammal turnover in the Cantabrian Region: The extinction of Pliomys lenki (RodentiaMammalia). Quat. Int.2122010129–13610.1016/j.quaint.2009.06.006Cuenca-BescósG.StrausL.G.González MoralesM.R.García PimientaJ.C.The reconstruction of past environments through small mammals: from the Mousterian to the Bronze Age in El Mirón Cave (Cantabria, Spain)J. Archaeol. Sci.362009947–95510.1016/j.jas.2008.09.025DansgaardW.JohnsenS.J.ClausenH.B.Dahl-JensenD.GundestrupN.S.HammerC.U.HvidbergC.S.SteffensenJ.P.SveinbjörnsdóttirA.E.JouzelJ.BondG.Evidence for general instability of past climate from a 250-kyr ice-core recordNature3641993218–220de VilleneuveL.BouleM.VerneauR.CartailacE.Les grottes de Grimaldi (Baousse-Rousse): Geologie et paleontologie. 2 vols1919Imprimerie de MonacoMonte CarloDoukaK.GrimaldiS.BoschianG.del LuccheseA.HighamT.F.G.A new chronostratigraphic framework for the Upper Paleolithic of Riparo Mochi (Italy)J. Hum. Evol.622012286–29910.1016/j.jhevol.2011.11.009EvansE.M.N.Van CouveringJ.A.H.AndrewsP.Palaeoecology of Miocene sites in Western KenyaJ. Hum. Evol.10198199–116FedeleF.G.GiaccioB.IsaiaR.OrsiG.Ecosystem Impact of the Campanian Ignimbrite Eruption in Late Pleistocene EuropeQuat. Res.572002420–42410.1006/qres.2002.2331GrimaldiS.SantanielloF.New insights into Final Mousterian lithic production in western ItalyQuat. Int.3502014116–129GrimaldiS.PorrazG.SantanielloF.Raw material procurement and land use in the northern Mediterranean Arc: insight from the first Proto-Aurignacian of Riparo Mochi (Balzi Rossi, Italy)Quartar612014113–127HammerØ.HarperD.A.T.RyanP.D.Past: Paleontological Statistics Software package for Education and data analysisPalaeontol. Electron.420011–9HarperD.A.T.Paleontological Data Analysis2005Blackwell PublishingMalden, MA, USA10.1002/9780470750711Hernández FernándezM.Bioclimatic discriminant capacity of terrestrial mammal faunasGlob. Ecol. Biogeogr.102001189–20410.1046/j.1466-822x.2001.00218.xHernández FernándezM.Análisis paleoecológico y paleoclimático de las sucesiones de mamíferos del Plio-Pleistoceno ibérico (PhD thesis)2001Universidad Complutense de MadridHernández FernándezM.Peláez-CampomanesP.Quantitative palaeoclimatic inference based on terrestrial mammal faunasGlob. Ecol. Biogeogr.14200539–5610.1111/j.1466-822X.2004.00125.xHighamT.BrockF.PeresaniM.BroglioA.WoodR.DoukaK.Problems with radiocarbon dating the Middle to Upper Paleolithic transition in ItalyQuat. Sci. Rev.2820091257–126710.1016/j.quascirev.2008.12.018HighamT.DoukaK.WoodR.RamseyC.B.BrockF.BasellL.CampsM.ArrizabalagaA.BaenaJ.Barroso-RuízC.BergmanC.BoitardC.BoscatoP.CaparrósM.ConardN.J.DrailyC.FromentA.GalvánB.GambassiniP.Garcia-MorenoA.GrimaldiS.HaesaertsP.HoltB.Iriarte-ChiapussoM.-J.JelinekA.Jordá PardoJ.F.Maíllo-FernándezJ.-M.MaromA.MarotoJ.MenéndezM.MetzL.MorinE.MoroniA.NegrinoF.PanagopoulouE.PeresaniM.PirsonS.de la RasillaM.Riel-SalvatoreJ.RonchitelliA.SantamariaD.SemalP.SlimakL.SolerJ.SolerN.VillaluengaA.PinhasiR.JacobiR.The timing and spatiotemporal patterning of Neanderthal disappearanceNature5122014306–30910.1038/nature13621IBCM - International Bathymetric Chart of the Mediterranean, 2014. Ocean Mapping Project of the Inter-governmental Oceanographic Commission. http://www.ngdc.noaa.gov/mgg/ibcm/ibcm.html. (accessed 13 November 2017).LaplaceG.Il Riparo Mochi ai Balzi Rossi di Grimaldi (Scavi 1939–1949). Les industries leptolithiquesRiv. di Sci. Preist.XXXII19773–131López-GarcíaJ.M.BertoC.ColamussiV.ValleC.D.Lo VetroD.LuziE.MalavasiG.MartiniF.SalaB.Paaleoenvironment and palaeoclimatic reconstruction of the latest Pleistocene–Holocene sequence from Grotta del Romito (Calabria, southern Italy) using the small-mammal assemblagesPalaeogeogr. Palaeoclimatol. Palaeoecol.4092014169–17910.1016/j.palaeo.2014.05.017López-GarcíaJ.M.BlainH.-A.BennàsarM.Fernández-GarcíaM.Environmental and climatic context of Neanderthal occupation in southwestern Europe during MIS3 inferred from the small-vertebrate assemblagesQuat. Int.3262014319–32810.1016/j.quaint.2013.09.010López-GarcíaJ.M.BlainH.-A.BurjachsF.BallesterosA.AlluéE.Cuevas-RuizG.E.RivalsF.BlascoR.MoralesJ.I.HidalgoA.R.CarbonellE.SerratD.RosellJ.A multidisciplinary approach to reconstructing the chronology and environment of southwestern European Neanderthals: the contribution of Teixoneres cave (Moià, Barcelona, Spain)Quat. Sci. Rev.43201233–4410.1016/j.quascirev.2012.04.008López-GarcíaJ.M.dalla ValleC.CremaschiM.PeresaniM.Reconstruction of the Neanderthal and Modern Human landscape and climate from the Fumane cave sequence (Verona, Italy) using small-mammal assemblagesQuat. Sci. Rev.12820151–1310.1016/j.quascirev.2015.09.013MagurranA.E.Measuring Biological Diversity2004Blackwell Science LtdOxfordMasiniF.AbbazziL.L’associazione di mammiferi della Grotta di CastelcivitaGambassiniP.Il Paleolitico Di Castelcivita, Culture Ambiente1997ElectaNapoli33–59Mitchell-JonesA.J.AmoriG.BogdanowiczW.KrystufekB.ReijindersP.J.H.SpitzenbergerF.StubbeM.ThissenJ.B.M.VohralikV.ZimaJ.The atlas of European mammals1999Academic PressLondonMonegatoG.PiniR.RavazziC.ReimerP.J.WickL.Correlating Alpine glaciation with Adriatic sea-level changes through lake and alluvial stratigraphyJ. Quat. Sci.262011791–80410.1002/jqs.1502Palma di CesnolaA.Il Paleolitico superiore in Italia1993Garlatti e Razzai EditoriFirenzePaunescoA.C.Brunet-LecomteP.AbbassiM.Révision spécifique des campagnols de sous-genre Microtus (Terricola) (Mammalia, Rodentia) de l’Abri Mochi (Ligurie, Italie)Bull. Mus. Anthr. Préhist. Monaco50201031–38PiniR.RavazziC.DoneganaM.Pollen stratigraphy, vegetation and climate history of the last 215 ka in the Azzano Decimo core (plain of Friuli, north-eastern Italy)Quat. Sci. Rev.2820091268–1290PiniR.RavazziC.ReimerP.J.The vegetation and climate history of the last glacial cycle in a new pollen record from Lake Fimon (southern Alpine foreland, N-Italy)Quat. Sci. Rev.2920103115–313710.1016/j.quascirev.2010.06.040PiniR.RavazziC.ReimerP.J.The vegetation and climate history of the last glacial cycle in a new pollen record from Lake Fimon (southern Alpine foreland, N-Italy)Quat. Sci. Rev.2920103115–313710.1016/j.quascirev.2010.06.040Pons-BranchuE.HamelinB.LossonB.JailletS.BrulheteJ.Speleothem evidence of warm episodes in northeast France during Marine Oxygen Isotope Stage 3 and implications for permafrost distribution in northern EuropeQuat. Res.742010246–251PorrazG.SimonP.PasquiniA.Identité technique et comportements économiques des groupes protoaurignaciens à la grotte de l’Observatoire (Principauté de Monaco)Gallia Préhist.52201033–59RasmussenS.O.BiglerM.BlockleyS.P.BlunierT.BuchardtS.L.ClausenH.B.CvijanovicI.Dahl-JensenD.JohnsenS.J.FischerH.GkinisV.GuillevicM.HoekW.Z.LoweJ.J.PedroJ.B.PoppT.SeierstadI.K.SteffensenJ.P.SvenssonA.M.VallelongaP.VintherB.M.WalkerM.J.C.WheatleyJ.J.WinstrupM.A stratigraphic framework for abrupt climatic changes during the Last Glacial period based on three synchronized Greenland ice-core records: refining and extending the INTIMATE event stratigraphyQuat. Sci. Rev.106201414–2810.1016/j.quascirev.2014.09.007RavazziC.PeresaniM.PiniR.VescoviE.Il Tardoglaciale nelle Alpi Italiane e in Pianura Padana. Evoluzione stratigrafica, storia della vegetazione e del popolamento antropicoIl Quaternario2022007163–184ReilleM.BeaulieuJ.-L. DeSvobodovaH.Andrieu-PonelV.GoeuryC.Pollen analytical biostratigraphy of the last five climatic cycles from a long continental sequence from the Velay region (Massif Central, France)J. Quat. Sci.152000665–685doi:10.1002/1099-1417(200010)15:7<665:AID-JQS560>3.0.CO;2-GRenault-MiskovskyJ.Contribution a la paléoclimatologie du midi Méditerranéen pendant la dernière glaciation et le post-glaciaire, d’après l’étude palynologique du remplissage des grottes et abris sous rocheBull. Mus. Anthr. préhist. Monaco181972145–210RoyerA.LécuyerC.MontuireS.EscarguelG.FourelF.MannA.MaureilleB.Late Pleistocene (MIS 3-4) climate inferred from micromammal communities and δ18O of rodents from Les Pradelles, FranceQuat. Res.802013113–12410.1016/j.yqres.2013.03.007RoyerA.MontuireS.LegendreS.DiscampsE.JeannetM.LécuyerC.Investigating the Influence of Climate Changes on Rodent Communities at a Regional-Scale (MIS 1-3, Southwestern France)PLoS One1112016e014560010.1371/journal.pone.0145600SantanielloF.Il Gravettiano dell’Italia tirrenica nel contesto mediterraneo: definizione delle strategie di insediamento e mobilità attraverso lo studio delle materie prime e delle industrie litiche (PhD Thesis)2016Università degli studi di Trento - Université de Nice/Sophia AntipolisSpötlC.ManginiA.Stalagmite from the Austrian Alps reveals Dansgaard–Oeschger events during isotope stage 3: Implications for the absolute chronology of Greenland ice coresEarth Planet. Sci. Lett.2032002507–51810.1016/S0012-821X(02)00837-3TagliacozzoA.ZeppieriF.FioreI.SpinapoliceE.Del LuccheseA.Archaeozoological evidence of subsistence strategies during the Gravettian at Riparo Mochi (Balzi Rossi, Ventimiglia, Imperia - Italy)Quat. Int.2522012142–15410.1016/j.quaint.2011.03.059TesakovA.S.LebedevV.S.BannikovaA.A.AbramsonN.I.Clethrionomys Tilesius, 1850 is the valid generic name for red-backed voles and Myodes Pallas, 1811 is a junior synonym of Lemmus Link, 1795Russ. J. Theriol.9201083–86TomassoA.NaudinotN.BinderD.GrimaldiS.Unité et diversité dans l’Épigravettien récent de l’arc liguro-provençalLanglaisM.NaudinotN.PeresaniM.Les groupes culturels de la transition Pléistocène-Holocène entre Atlantique et Adriatique. Actes de la Sance de la Sociéte préhistorique française de Bordeaux (24-25 mai 2012)2014Séances de la Société Préhistorique Française 3Paris155–184WalterH.Vegetationszonen und Klima1970UlmerStuttgartWattsW.A.AllenJ.R.M.HuntleyB.Palaeoecology of three interstadial events during oxygen-isotope Stages 3 and 4: a lacustrine record from Lago Grande di Monticchio, southern ItalyPalaeogeogr. Palaeoclimatol. Palaeoecol.155200083–93WilsonD.E.ReederD.M.third ed.Mammal Species of the World. A Taxonomic and Geographic Referencevol. 22005Johns Hopkins University PressBaltimoraZeppieriF.I Macromammiferi del livello Gravettiano “D” di Riparo Mochi (Balzi Rossi,, Ventimiglia), scavi Blanc e Cardini 1938 e 1959 (Master Thesis)2009Università degli Studi di Ferrara
A: Location of Riparo Mochi (red square); location of the most complete Italian Peninsula small mammal sequences (black squares); location of cited pollen records (blue dots). B: Riparo Mochi stratigraphy and selected radiocarbon dates cal BP (original drawing from Segre and modified from Blanc, 1954; for a complete framework about radiocarbon dates see Douka et al., 2012).
Localisation du Riparo Mochi (carré rouge) ; localisation des séquences de petits mammifères les plus complètes de la péninsule Italienne (carrés noirs) ; localisation des données palynologiques citées (points bleus). B : Stratigraphie du Riparo Mochi (dessin original de Segre modifié par Blanc, 1954 ; Douka et al., 2012).
Some identified small mammals from Riparo Mochi, all scales are 1 mm long. 1 and 2: Arvicola amphibius, left and right m1; 3 and 4: Chionomys nivalis, right m1; 5, 6, and 7: Microtus arvalis, one right m1 and two left m1; 8: Microtus (Terricola) gr. multiplex-subterraneus, right m1; 9 and 10: Clethrionomys glareolus, left and right m1; 11: Apodemus sylvaticus, left m1; 12: Apodemus flavicollis, right m1; 13: Eliomys quercinus, left m1, m2, and m3; 14: Sorex gr. araneus, left mandible; 15: Neomys sp., mandibular condyle; 16: Erinaceus europaeus, right m1.
Exemples de petits mammifères identifiés au Riparo Mochi ; toutes les échelles sont de 1 mm de long. 1 et 2 : Arvicola amphibius, m1 gauche et droite ; 3 et 4 : Chionomys nivalis, m1 droite ; 5, 6 et 7 : Microtus arvalis, une m1 droite et deux m1 gauches ; 8 : Microtus (Terricola) gr. multiplex-subterraneus, m1 droite ; 9 et 10 : Clethrionomys glareolus, m1 gauche et droite ; 11 : Apodemus sylvaticus, m1 gauche ; 12 : Apodemus flavicollis, m1 droite ; 13 : Eliomys quercinus, m1, m2, m3 gauches ; 14 : Sorex gr. araneus, mandibule gauche ; 15 : Neomys sp., condyle mandibulaire ; 16 : Erinaceus europaeus, m1 droite.
Representation of the small mammal Simpson diversity index (1-D), the mean annual precipitation (MAP), the mean temperature of the coldest month (MTC), the mean annual temperature (MAT), the mean temperature of the warmest month (MTW) and the landscape percentages (OD: open meadows; OH: open humid meadows; W: woodlands; R: rocky areas; Wa: areas along streams, lakes and ponds). GI: Greenland Interstadial; GS: Greenland Stadial.
Représentation de l’indice de diversité de Simpson des petits mammifères (1-D), moyenne des précipitations annuelles (MAP), moyenne des températures du mois le plus froid (MTC), moyenne annuelle des températures (MAT), moyenne des températures du mois le plus chaud (MTW) et pourcentages de paysage (OD : prairies ouvertes ; OH : prairies humides ouvertes ; W : bois ; R : zones rocheuses ; Wa : zones le long de ruisseaux, lacs et étangs). GI : Interstade groenlandais. GS : Stade groenlandais.
Small mammal percentages of Minimum Number of Individuals (MNI) and small mammal distribution by habitat.
Pourcentages de nombre minimum d’individus (NMI) de petits mammifères et distribution des petits mammifères par habitat.
Table 1ACD ID IID IIID IVEFGHIODOHOWWRWaArvicola amphibius16.122.726.910.28.78.64.310.71331.48.91Chionomys nivalis7.65.86.84.64.32.74.35.78.91Microtus arvalis19.437.930.848.96056.260.93243.528.615.61Microtus agrestis12.95.73.14.32.20.50.5Microtus cf. agrestis1.91.10.50.5Microtus (Terricola) gr. multiplex-subterraneus6.56.13.86.88.210.58.79.34.311.4200.50.5Clethrionomys glareolus0.64.32.915.60.250.75Apodemus sp.16.16.15.84.51.53.113.018.78.75.711.11Apodemus sylvaticus1.51.90.52.74.41Apodemus flavicollis3.21.52.30.51.22.78.72.91Rattus sp.1.9--Eliomys quercinus3.21.50.61.32.20.750.25Talpa sp.1.91.50.60.50.5Talpa europaea22.612.115.411.48.76.28.71617.411.46.70.50.5Erinaceus europaeus3.82.32.542.20.250.75Sorex gr. araneus2.11.22.20.750.25Sorex sp.0.50.750.25Neomys sp.3.00.750.5Total MNI316652881951622375233545Total NISP521147916030624440105285066
Biodiversity, Climate and Landscape values. Percentage representation of small mammal taxa associated with open dry meadows (OD), open humid meadows (OH), open woodland environments (OW), woodland environments (W), rocky environments (R) and landscapes constituted by river, lakes and ponds (Wa); bioclimatic components (BCi); temperature and precipitation reconstruction for Riparo Mochi sequence: MAT: Mean Annual Temperature, MTW: Mean Temperature of the Warmest month, MTC: Mean Temperature of the Coldest month, MAP: Mean Annual Precipitation. Number of taxa in each layer; values obtained for evenness: Simpson diversity index = 1 – ∑(pi2).
Valeurs de biodiversité, climat et paysage. Représentation des pourcentages des taxons de petits mammifères associés à une prairie sèche ouverte (OD), une prairie humide ouverte (OH), un environnement boisé ouvert (OW), un environnement boisé (W), un environnement rocheux (R) et des paysages constitués de rivières, lacs et étangs (Wa) ; composants bioclimatiques (BCi) ; reconstruction des températures et des précipitations de la séquence du Riparo Mochi : MAT, moyenne des températures annuelles, MTW, moyenne des températures du mois le plus chaud, MTC, moyenne des températures du mois le plus froid, MAP, moyenne des précipitations annuelles. Nombre de taxons pour chaque niveau ; valeurs obtenues pour la régularité des espèces : indice de diversité de Simpson = 1–∑(pi2).