Available online http://amq.aiqua.it ISSN (print): 2279-7327, ISSN (online): 2279-7335 Alpine and Mediterranean Quaternary, 34 (1), 2021, 89-108 PALEOECOLOGICAL FIRST RESULTS FROM THE MIDDLE PLEISTOCENE SEQUENCE OF COUDOULOUS I (QUERCY, LOT, FRANCE). Philippe Fernandez 1 , Marcel Jeannet † 1 , Jacques Jaubert 2 , Jean-Philip Brugal 1 1 CNRS, UMR 7269 LAMPEA, Aix Marseille Université, CNRS, Minist. Culture, Aix-en-Provence, France. 2 University of Bordeaux, CNRS-MC, UMR 5199 PACEA, Pessac, France. Corresponding author: Philippe Fernandez and Jean-Philip Brugal ABSTRACT: The Coudoulous I (Lot, France) sequence has yielded abundant fossil associations dated to the end of the Middle Pleistocene. The total sample consists of 102 taxa corresponding to 75 species of mammals (53 small and 22 large mammals), 11 species of amphibians and 16 species of reptiles distributed in five faunal units (FUs). After evaluating the sample quality with completeness indices (CI and CIbda), we established rarefaction curves that were discussed at length together with Shannon- Wiener diversity (H’), Margalef species richness (D), and the Sørensen-Dice similarity index. In order to better understand structu- ral changes in the mammal associations as well as the climate trends we built the body mass distribution with cenograms in the different FUs. Concurrently, we investigated the whole community using a more analytical approach with the “bioclimatic model” based on a climatic restriction index for each species to infer paleoclimatic proxies (i.e. temperatures, precipitation, aridity) and finally to assign each faunal unit to a dominant type of climate. Our results clearly indicate that the FUs could be systematically associated with a typical temperate climate with sub-Mediterranean temperate species for the largest part of the sequence, which is consistent with the dates obtained (U/Th, ESR/U-series, TT-OSL) and Marine Isotope Stage 7. Nevertheless, cold elements from a Boreal type climate component were also found in the sequence indicating possible climate oscillations in FUs. The upper part of the stratigraphy (FUII) is probably related to MIS 6 while the lowest part of the sequence (FUVII) could correspond to a cold phase of the MIS 7 or earlier (MIS 8?). Keywords: Vertebrate community, diversity, cenograms, bioclimatic model, climate reconstruction, MIS 6-7. 1. INTRODUCTION Less than a dozen sites in the French Mediterrane- an region are dated to the Middle Pleistocene, and per- haps only twenty in southern France (see fig. 2 in Brugal et al., 2020). Among them, the Coudoulous sites show one of the most well-constrained chronostratigraphical sequences, yielding very rich and diversified fossil verte- brate and invertebrate associations (Jaubert et al., 2005). The Quercy region, where Coudoulous is situat- ed, exhibits numerous and varied archeological and/or paleontological sites covering the last 0.3 Myr within a wide range of environmental and climatic contexts (Jaubert et al., 2013; Jeannet et al., 2013; Brugal et al., 2013). This region is located at the periphery of the Mediterranean ecological area, subject to the Atlantic conditions of the Aquitaine basin and bordered by the low mountainous area of the Massif Central. We pro- pose in this contribution to depict the succession of fos- sil associations throughout the complete sequence of Coudoulous I, attributed to the end of the Middle Pleis- tocene, with the aimof characterizing paleoecological and climatic dynamics. 2. THE COUDOULOUS SITES The Coudoulous (Coud) sites are located in the Quercy area (44°28'40"N, 1°39'50"E), not far from the Paleolithic cave art site of Pech-Merle (Cabrerets), a region of small Jurassic limestone plateaus divided by more or less narrow valleys and rivers. These sites are in a high position (280m asl) at the junction of the river Lot and its tributary, the Célé (Fig. 1), and correspond to two dis- tinct karstic cavities, namely Coud I and Coud II. First explored with a rescue excavation in 1978-1980 (Bonifay & Clottes, 1981; Jaubert, 1984), the sites were investigated between 1994 and 2003 by J. Jaubert (for Coud I) (Jaubert et al., 2005) and J.P. Brugal (for Coud https://doi.org/10.26382/AMQ.2021.12 90 Fernandez P. et al. into 4 main units/formations but only the lower and mid- dle units yield lithic artefact and fossil remains (Fig. 2). Several levels (from 3 at the top to 8 at the bottom) and sub-levels (at least 31) are distinguished within these units. The sequence is bracketed by two speleothems dated by U/Th (Fig. 2): the upper one (between levels 2 and 3) is assigned to MIS 5 (126[+20/-46] and 139[+13/- 11] kyr) and, the thick basal one (level 8g-e) at the limit of the method (229[+78/-42] and 202[+78/-39] kyr) is less accurate (Quinif in Jaubert et al., 2005; Couchoud, 2006). Two other stalagmitic floors are observed in the sequence (respectively levels 8a and 8c) and all are important periods of sedimentation with a pause in fossil II) (Brugal et al., 1998; Costamagno, 1999; Brugal, 2006). Here we focus on the Coudoulous I site, with a brief overview and synthesis of the different studies made of this site and its chronological sequence. 2.1. The Coudoulous I sequence The karstic cavity represents a pitfall (aven), with a main vertical entrance after the collapse of the roof above a large room (15-20 m diameter) presently com- pletely filled with detrital (clays, sands, angular limestone rocks) or carbonate (speleothems) deposits, to a total of ca.10 m in thickness (Jaubert et al., 2005). The se- quence is divided into 10 stratigraphic layers grouped Fig. 1 - Location of Coudoulous sites. Photo J. Jaubert and map reworked from Electronic reference (18). Fig. 2 - Coudoulous I, from left to right: Photo of the geological units (black arrow=2 m); Stratigraphic log with datations (grey bar=speleothems); Faunal Units; Distribution of non-identifiable bone remains (n total=80.452) throughout levels 5 to 8. 91 Paleoecologial first results from the Middle Pleistocene sequence of Coudoulous I (Quercy, Lot, France). deposition, and should be considered as natural limits or cut-offs. The sequence has been dated by U/Th, ESR/U-series and TT-OSL on teeth and bone and sedi- mentary quartz samples (Hernandez et al., 2015) giving correlations from MIS 7 to MIS 6. A very rich archaeological level (level 4) occurs at the top of the Coudoulous I sequence, and the levels (3 to 1) above it are sterile. Level 4 yielded abundant lithic artefacts (flint, quartzite), with Levallois core reduction, reported as Early Middle Paleolithic (EMP: stage 6.5) (Jaubert, 1995; Mourre, 1996), associated with a domi- nant species, the bison (98%), with very few horse and wolf remains. The bison record includes a minimum of 232 individuals, represented by all skeletal parts, but strongly affected by taphonomic processes, which have preferentially removed axial and cephalic bone ele- ments. Teeth are good zooarchaeological and paleocli- matic indicators and demonstrate a catastrophic mortali- ty pattern (juveniles and young adults dominant) and a late spring-early summer seasonality (Brugal, 1999a, 1999b; Brugal & Jaubert, 2006; Hélène Martin, pers. comm.). Oxygen isotope analysis of phosphate in tooth enamel (δ 18Op) shows contrasted seasons with a mini- mal annual temperature of 9±3 °C, which is about 4 °C lower than at present (Bernard et al., 2009). EMP or Lower Paleolithic artefacts are scarce and dispersed in the middle to lower units (levels 5a to 8d), and made on quartz, quartzite, metamorphic and volca- nic rocks. The techno-typological analysis places this industry within the variability of the Acheulean despite the absence of typically bifacial tools, but with some large cutting tools (LCTs) present (Jaubert, 1995; Jau- bert & Servelle, 1996). An interdisciplinary study has demonstrated that the cavity was used for marginal scavenging by Lower Paleolithic hominid groups and as a kill-butchery site during the EMP (Brugal & Jaubert, 1991, 2021; Jaubert et al., 2005). 2.2. Material This contribution is the first paleoecological analy- sis of the vertebrate associations from the middle and lower units (levels 5 to 8) of Coudoulous I (Fig. 2). This site represents one of the rare stratigraphic sequences of the Middle Pleistocene yielding a very rich and diver- sified fauna (NISP=9864). Moreover, a total of more than 80 000 splinters and bone shaft fragments (non- identifiable remains) were also collected during the ex- cavations. Following geological and stratigraphical ob- servations (Kervazo et al., 2021) and the distribution and abundance of faunal remains (Brugal, 2021), 6 faunal units (FU) were identified. According the strati- graphic log in Fig. 2, FUI includes only level 4. FUII and FUIII correspond to the middle units, with levels 5 and 6a to 7b, respectively. The two lower speleothems are used as limits in the distinction of the subsequent FUs (FUIV+V to FUVII). FUIV+V have been regrouped here and correspond to level 7c to 7f’. FUVI includes levels 8a to 8c and FUVII corresponds to level 8d. For the paleoecological purposes of this study the taxonomic determinations were realized at species level in five faunal units (FUII, FUIII, FUIV+V, FUVI, FUVII) thanks to detailed studies (Jeannet, 2021, for microvertebrates; Argant, 2021, for felids and ursids; Mallye, 2021, for mustelids; Boudadi-Maligne, 2021, for canids) and our own studies (equids, bovids, caprids, cervids, rhinoce- rotids, proboscids). The remains with marking errors or that might have been shuffled between different strati- graphic levels were not included. We took into account 9864 Number of Identified Specimens (NISP) from 102 species including small mammals such as rodents (n=35), lagomorphs (n=4), and chiropters (n=14), but also large mammals including ungulates (n=10) and carnivores from small to medium size (n=12), and re- spectively 16 and 11 species of reptiles and amphibians (Tab. 1). 3. METHODS Ever since the work of Leonard and Jones (1989) concerning the concept of “diversity”, methodological and theoretical approaches have continued to be topical in paleobiological contexts for studying animal commu- nities (rarefaction, richness, sample size, sample unit of measurement, appropriate diversity indices, etc.). Here we present the paleoecological analysis of the Cou- doulous I mammals, applying and sometimes cross- referencing different methods (concept, mathematical formulations, limits) to see if they lead to similar results in the different faunal units of this site. 3.1. Completeness indices (CI) and (CI bda) Natural taphonomic processes as well as carnivore activity and anthropogenic modifications are the key factors that explain biased samples from original depos- its. Nevertheless, completeness indices (CI) and (CIbda) (Maas & Krause, 1994; Maas et al., 1995) make it pos- sible to evaluate the quality of the sample in the faunal units, starting with the presence (1) absence (0) of spe- cies (Palombo et al., 2008). Completeness indices are based on the assumption of the range-through of the species represented, meaning that a given species, which is present before and after, but not during an interval of time is supposed to have been in continuous existence from its first to last appearance in the stratig- raphy. In this particular case, missing species probably result from taphonomic bias, poor sampling or bad preservation of the fossil remains. It should be added that between their first and last appearances, some missing species could have diffused to new landscapes or different habitats in response to strong climate changes. The completeness indices CI and CIbda are calcu- lated as follows (see details in Maas et al., 1995; Pal- ombo et al., 2008; Fernandez, 2009): Eq. (1) CI = [Nt / (Nt + Nrt)]*100 Eq. (2) CIbda = [Nbda/(Nbda+Nrt)]*100 Nt is the number of species known before, during and after the faunal unit considered, including taxa hav- ing their first or last occurrences in the interval. Nrt is the number of species known before and after, but not during the faunal unit interval. To calculate Nrt in the last and the first faunal units respectively FUII and FUVII, we used species’ chronological extension 92 Fernandez P. et al. Paleoecologial first results from the Middle Pleistocene sequence of Coudoulous I (Quercy, Lot, France). 93 from the works of Palombo et al. (2008) as well as Fer- nandez (2009). Because Nbda is the number of species only known before, during and after a faunal unit inter- val, CIbda excludes the first and last FU whereas CI al- lows us to take into account all of the faunal units (Maas et al., 1995; Palombo et al., 2008). 3.2. Rarefaction, richness (S), Margalef (D), Shannon -Wiener (H’) and the Sørensen-Dice index In this study we used the rarefaction method that was popularized by Sanders (1968) for marine inverte- brate communities and which has became a standard method. When applying Sanders's algorithm key as- sumptions (all met in Coudoulous I) are required such as a similar taxonomical rank between individuals col- lected from a similar place by using standardised proce- dures (Tipper, 1979). The brilliant idea of Sanders makes it possible to compare the number of species from different assemblages independently of the size of the initial samples and to see to what extent the sam- pling can induce a bias in the species richness (S) (i.e. area sampled, non-standardized sampling, catchability, etc.) (Raup, 1975; Tipper, 1979; Ricklefs & Miller, 1999; Gotelli & Colwell, 2001). Given the strong correlation between the number of identified specimens (NISP) and the minimum number of individuals (MNI) put forward by Grayson (1984) and later by Lyman (2008), rarefaction has been used in different paleobiological contexts, whether natural or anthropogenic (e.g. Adrain et al., 2000; Belmaker, 2006; Belmaker & Hovers, 2011; Po- pov & Marinska, 2007; Davis & Pyenson, 2007; Fernan- dez, 2009). In Coudoulous I we established rarefaction curves for each faunal unit from all the taxa present in the sequence and determined by the rank of the species (S=102). These curves were built with the "individual- based rarefaction" routine of the PAST program (Version 3.23) (Hammer et al., 2001; Hammer & Harper, 2006, see equations 6.10 to 6.13, p. 205). In addition to rarefaction, we used species diversity which measures species richness combined with even- ness, taking into account not only how many species are present but also how evenly distributed the numbers of each species are (see Eq. 3). It has been demonstrat- ed that diversity varies consistently according to large- scale geographical areas with δ and δ diversity (Rohde, 1992) but also in more restricted areas or habitats at a more local scale (δ diversity, Whittaker, 1972). Thus, there are many different indices and mathematical solu- tions to account for the diversity that can be defined in many ways (Hammer et Harper, 2006). These authors indicate that only the number of individuals should be used to estimate the diversity of a community. Conse- quently, in nearly all paleobiological contexts diversity indices will always be an approximation because MNI will never be the real number of individuals. As a result of the foregoing and the fact that we used NISP, which is strongly correlated to MNI, we focused on the Shan- non-Wiener diversity index (Shannon & Weaver, 1964) as it takes into account proportions for the frequency and it is completely adapted for local area (δ , the within assemblage diversity). We would like to stress that this index is sometimes erroneously labelled as the “Shannon-Weaver” index (see Spellerberg & Fedor, 2003 for this confusion). The formula implemented in Past (Version 3.23) that we used is as follows: Eq. (3) H’ = -∑pi ln pi Where (ln) is the logarithm to the base of e (sometime in the literature base two is used), and where pi is the proportion of individuals that belong to species i. Because species richness (S) increases with sam- ple size, we used Margalef’s species richness index (D) that attempts to compensate for this effect, as defined by the formula (Margalef, 1958): Eq. (4) D = (S-1) / ln n To go further in the analysis, we calculated the Sørensen-Dice index, which is a statistical indicator that measures the similarity between two samples and was developed independently in botany by Dice (1945) and Sørensen (1948). The Sørensen-Dice formula is as fol- lows: Eq. (5) Sørensen-Dice = 2J / (A+B) A and B represent the number of species in each sample and J the number of species common to the samples A and B. It is indexed from 0, a total absence of common elements, to 1 when two samples are identical (see Fig. 3). We do not go into further detail because the similarity coefficients are numerous and can only be explained by mathematical demonstrations (Shi, 1993). Comparing them to the most widely used such as Simp- son’s (1943) and Jaccard’s coefficients (1912), here we have chosen to put more weight on matches than on mismatches due to the multiplication of J by a factor of two (see Eq. 5). Furthermore, the Sørensen-Dice coeffi- cient normalizes with respect to the average rather than the total number of species in the two samples. Conse- quently, it is somewhat less sensitive to differences in sample size than Jaccard’s index. The Sørensen-Dice index is also different from Simpson’s coefficient, which is totally insensitive to the size of the larger sample and also does not take into account absences in the smaller sample of comparison (Hammer & Harper, 2006). 3.3. Body mass and cenograms The cenograms was proposed by Valverde (1964, 1967) in order to compare body size of non carnivore modern terrestrial mammals from different communities. Given that the weight distributions of species are stron- gly dependent on their environmental conditions in pre- dictable ways, Legendre (1986, 1989) and Gingerich (1989) extended the method into the fossil record to infer different paleoenvironmental characteristics as climatic < < < < < - - - - - - - Tab. 1 - Faunal units FUII to FUVII with species richness (S), number of identified specimens (NISP). For each species, log (ln) is given according to their body mass (see bibliographic references including Electronic references). Completeness indices (CI and CIbda) are calcu- lated from Eq. 1 & Eq. 2 in text. For predator prey ratio (PPR=n predator /n prey) see details in text. changes, namely, humidity or aridity, and types of landscapes, mainly open or closed. Although the initial work of Legendre was controversial on quantitative aspects (see Rodríguez, 1999) other quaternary Euro- pean paleoenvironmental studies have used the ceno- gram method (Montuire, 1994, 1999; Rodríguez et al., 1996; Montuire & Desclaux, 1997; Montuire & Marcolini, 2001; Valensi & Psathi, 2004; Hernández Fernández et al., 2006; Palombo & Giovinazzo, 2006; Fernandez, 2009). The initial graphic representation established by Legendre (1986) reports the body mass (ln on the y- axis) according to the rank of the species (on the x-axis from the heaviest to the lightest). Schematically, the plots of points are represented by segments with diffe- rent slopes (sometimes built by linear regression) and separated by breaks/gaps in the slope (see climatic trends in Fig. 4). Legendre’s model (1986, 1989) and Gingerich (1989) who quantified the slopes, gaps and breakpoints, took into account small-sized (500 g<), medium-sized (500 g> 250 kg<) and large-sized mam- mals (> 250 kg) with two breakpoints around 8 kg and 250 kg. There is a gap (500 g - 8 kg) with greater (steeper) slope for medium-sized species in an open environment (e.g. woodlands and savannas) whereas closed environments have a continuous distribution. In a typical humid environment the breakpoint around 250 kg marks the low regression slope for medium-sized species, with no break for closed landscapes and a gap around 500 g for a more open environment. Gingerich (1989) hypothesized that the relative scarcity of mam- mals of about 500 g in this type of environment is pro- bably related to the range of size, the density, and the diversity of insects and leaves, both of which are grea- test in forests. 3.4. Predator Prey Ratio (PPR) The PPR is simply the ratio between the number of predators and prey (PPR=n species of predators / n species of prey). For many Pleistocene sites, significant changes in PPR are difficult to interpret. Raia et al. (2007) found evidence for PPR inconstancy through time such as had been pointed out in previous studies (i.e. Van Valkenburgh & Janis, 1993). Here we assume an actualist approach whereby carnivores weighing up 21.5 kg feed mostly on prey that is 45% or less of their own mass (Carbone et al., 1999). Because carnivores above this limit feed mostly on prey that are greater than 45% of their own mass, we excluded from our cal- culation the mustelids of the sequence as well as Meles meles, Vulpes vulpes and Felis silvestris. 3.5. Bioclimatic analysis A reconstruction of the climate and the plant envi- ronment has been inferred for each Coudoulous I faunal unit from the so-called “bioclimatic model” developed by Hernández-Fernández (2001a, 2001b) and Hernández- Fernández & Peláez-Campomanes (2003, 2005). It has been very recently used on archaeological sites from the Middle Pleistocene with human remains (Lopez- Garcia et al., 2021), and adapted for insectivore and rodent communities from the Last Glacial Maximum to the Holocene (Royer et al., 2020). In the initial work of Hernández-Fernández (2001a, 2001b), 50 associations of present-day mammals were compiled in different latitudes and biotopes corresponding to 10 major biocli- matic components (BC) associated with different types of climates (Tab. 2). A typology adapted from previous works (Walter, 1970; Rivas-Martínez, 1994) made it possible to systematically identify different environmen- tal variables for each BC (e.g. type of vegetation, avera- ge annual temperatures, rain rate, thermal amplitu- de…). Thus, each mammalian species is associated with a CRIi (Climatic Restriction Index) value ranging from 0 to 1 which characterizes its climate dependence such that: Eq. (6) CRIi = 1/n With n the number of BCs where the species are represented and i the BC where the species appear. For the distribution of species by climatic types we used the work of Hernández-Fernández et al. (2007) and Jeannet (2010, 2021) for microfauna. For large mam- mals, and in rare cases their nearest living ecological analog species, we used the studies of Bhatnagar & Lovari (2008), Palombo (2015) and Lovari et al. (2016a, 2016b). For example, in Table 3, according to the data from Hernández-Fernández, (2001a, appendix 2), the CRIi value is equal to 1 for Ochotona pusilla, in BC VII, which means that all elements of this species are exclu- sively associated with this Arid-temperate climate. On the other hand, the CRIi of 0.33 for Microtus oecono- mus, which is associated with bioclimatic components VI, VIII and IX, indicates that this species is more eu- rioic than the previous one. It is therefore possible to calculate for several mammalian species a probability of association (BC) for each of the 10 bioclimatic compo- nents according to the following formula: 94 Fernandez P. et al. Tab. 2 - Bioclimatic components (BC) according to their type of climate and vegetation cover (from Hernàndez-Fernàndez, 2001a, table 2.1). Paleoecologial first results from the Middle Pleistocene sequence of Coudoulous I (Quercy, Lot, France). 95 Tab. 3 - Distribution of the mammalian species identified in FUII according to their Climatic Restriction Index (CRIi, see detailed Eq. 6 in text) and Bioclimatic Component (BCi, see detailed Eq. 7 in text) in accordance with Hernàndez-Fernàndez (2001a) for the CRIi. Eq. (7) BCi = (∑ CRIi)*100/S (S) being the number of species present in the faunal association (or species richness). Note also that the BCi value is equivalent to the cumulative rates of Andrews (1990) or to the % THI of Fernández-Jalvo et al. (1998). To end with our exam- ple, in Table 4, the highest probability BCi of the UFII mammalian association corresponds to BC VI (Arid- temperate climate) with 36.45%, followed by BC VIII with a probability of 18.66%, and so on. For each FU, we reported mammalian associa- tions and their most likely BC (Tab. 4). To go deeper into the climate reconstructions, in this table we calcu- lated some paleoclimatic proxies from the BCi values (Eq. 7) using the multiple linear regression coefficients from Hernández-Fernández (2001a, table 4.1 to 4.3). These indices and the parameters of their equations (b=intercept, a1 to a9=regression coefficients) are presented in the table 5 with the following compo- nents: - Mean Annual Temperature (°C) (MAT). - Mean Annual Thermal Amplitude (°C) (MATA). This expresses the difference (°C) between the mean temperature of the warmest month and that of the coldest month of the year. According Rivas-Martínez (1994), it is an index of continentality simple. - Annual Positive Temperatures Index (APTI). This provides a measure of the intensity of the summer heat. - Compensated Thermal Index (CTI). In extratropical zones, the CTI is designed to equilibrate the cold “excess” that occurs during winter in the continental climates (average temperature of the coldest month of the year minimum), or the excessively mild winter in the marked oceanic territories, so that these index values can be significantly compared (Rivas-Martínez et al., 2011). - Mean Annual Precipitation (mm) (MAP) - Mean Temperatures of the Coldest Month (°C) (MTCM) - Aridity Index (AI). We calculated this index from De Martonne (1926) which is still used with good results (Pellicone et al., 2019). It is simply the ratio between the mean annual precipitation (mm) (MAP) and the mean annual temperature (°C) (MAT) weighted by 10 such that: (AI) = MAP / MAT + 10. This index should be read carefully because the higher the AI, the lower the aridity. 4. RESULTS AND DISCUSSION 4.1. Sample quality In Coudoulous I, CI indicates a low-quality sample for FUII (52.88) and FUVII (28.43) whereas CIbda corre- sponds to a very slightly modified sample in FUIII, FUIV+V and FUVI respectively with 81.25, 91.66 and 92.85 (Tab. 1). An index value higher than 70 usually reflects very little disturbance in a faunal unit interval (Maas et al., 1995; Palombo et al., 2008). Biases are often related to lower values in shorter chronological intervals, which is clearly the case in FUVII with only one archaeological level (8d). The CI of this faunal unit (28.43) reflects the lowest quality sample in the whole sequence. On the other hand, the overlying FUVI shows a very high CIbda index (92.85). FUIV+V (CIbda=91.66) and FUIII (CIbda=81.25) also both indicate very good quality sam- ples mainly associated to the subdivisions of level 7, probably contemporary with MIS 7, which is supported by biomarkers (mammal species and/or evolutive stage) and absolute dates (e.g. Jaubert et al., 2005, 2013; Boudadi-Maligne, 2010; Hernandez et al., 2015). Final- ly, in the most recent FUII CI=52.88 indicates a poor- quality sample. This faunal unit, beneath level 4, is re- lated to the first part of MIS 6 (Jaubert et al., 2005; Her- nandez et al., 2015), a glacial period known for involv- ing dramatic drops in temperature (Mokeddem & McManus, 2016). The relatively low species richness of this faunal unit might coincide with this climate cooling. 4.2. Diversity (s.l.) In Coudoulous I, rarefaction curves illustrate the expected number of species whatever the initial sample size in each faunal unit (Fig. 3). The number of rarefied species in FUIV+V is higher than in the lower FUIII, FUVI, FUII and FUVII, respectively. In the same way the de- creasing values of the Margalef index (D) are totally in line with the rarefaction curves. Although the calcula- tions are quite different, both rarefaction and the Marga- lef index (D) reach the same result. The latter also com- pensates for the sample size effect by dividing the num- ber of species in a sample by the natural log of the number of individuals (see Eq. 4). Furthermore, the curve from FUVII does not reach the asymptotic plateau, indicating a significant bias in this sample (lowest CI), also shown by the weakness of the NISP and the spe- cies richness (NISP=118 and S=29) which correspond to the single archaeological level 8d. Interestingly enough, the species-rich communities may have a greater number of trophic levels than spe- cies-poor communities (Briand, 1983). It is in light of this evidence that Yachi & Loreau (1999) formulated the “insurance hypothesis” stating that high species rich- 96 Fernandez P. et al. Tab. 4 - Coudoulous I faunal units and their probabilities (P1 to P3) of association with a Bioclimatic Component (see Table 2 for BC de- scription). (S), species richness (excluding reptiles and amphibians). Paleoclimatic proxies with Mean Average T° (MAT), Mean Annual Thermal Amplitude (MATA), Annual Positive Temperatures Index (APTI), Compensated Thermal Index (CTI), Mean Annual Precipitations (MAP), Mean Temperatures of the Coldest Month (MTCM), Aridity Index (AI) (see text for details). Calculation for some paleoclimatic prox- ies from multiple linear regression coefficients provided in table 5. Tab. 5 - Paleoclimatic proxies with equation parameters (b=intercept, a1 to a9 = regression coefficients) from Hernández-Fernández (2001a, table 4.1 to 4.3) (see details in text). Paleoecologial first results from the Middle Pleistocene sequence of Coudoulous I (Quercy, Lot, France). 97 ness increases community stability with species that respond differently to environmental fluctuations (see also Ives et al., 2000). If we accept this hypothesis, the rarefaction curves associated with the highest Marga- lef indices could be good indicators of a stable com- munity in Coudoulous I (respectively FUIV+V, FUIII, FUVI, FUII, FUVII). In Figure 3, the Shannon-Wiener diversity index (H’) values are almost correlated to the species rich- ness (S) and Margalef’s values (D). Nevertheless, in FUVII the diversity is higher with H’=2.77 whereas the number of species is much less important (n=29) than in FUII (n=53) which has the lowest diversity H’=2.06. It means that independently of both species richness and the lowest CI (discussed above), the Shannon- Wiener index is very sensitive to the individual distribu- tion frequency among species in FUVII. In all faunal units the H’ values are comprised between 2.06 and 3.08. It is difficult to compare these values with other sites, knowing that diversity is stron- gly dependant on multiple local factors (e.g. climate, taxonomical groups, ecosystem involved, etc.). Never- theless, it is now widely accepted that high species diversity indicates a highly complex community with more species interactions (Whittaker, 1972). It is worth noting that in nature, whatever the taxonomic group studied, H’ often ranges between 1.5 and 3.5 accor- ding to Gaines et al. (1999). Another important consideration that should be emphasized is that H’ as well as D are completely independent of the temporal gradient in the different FUs whereas the Sørensen-Dice index almost always shows a greater similarity between the closest FUs through time. In other words, the more distant the FUs are, the less species they have in common. This should not be considered as a statistical artefact but most likely as a gradual turnover of species through time, probably due to paleoenvironmental changes. In southern Africa, Andrews & O’Brien (2000), combining 15 climatic variables with current mammal species (n=285), showed that woody plant species (n=1359) alone explained 75% of the variability in mammal species richness. Nevertheless, large mam- mals over 90 kg, scansorial, aquatic or fossorial mam- mals were not significantly correlated with woody plant species richness, unlike small-bodied arboreal frugivo- res and insectivores. This led us to test which species were most affected by possible climate and plant cover changes in the different faunal units of Coudoulous I. Therefore, we Fig. 3 - Rarefaction curves, Shannon-Wiener diversity (H’), Margalef species richness (D) and Sørensen-Dice similarity for each faunal unit (FUII to FUVII). NISP in Table 1 is used for calculation with Past 3.23, (Hammer et al., 2001). For rarefaction see details in text for “individual-based rarefaction" routine from PAST. For calculation of Shannon-Wiener (H’) see Eq. 3, for Margalef species richness (D) see Eq. 4 and for Sørensen-Dice similarity see Eq. 5 and Table 1. tested the proportions of species throughout the se- quence according to their size (microfauna vs macro- fauna) and their diet (herbivores vs carnivores). The results of the Chi² test indicate no significant differences between microfauna and macrofauna between the FUs. The same is true for the proportion of herbivores and carnivores (Table 6-a to 6-d). One possible explanation for these results is that regardless of the different types of climate in a small sampling area (e.g. within habitat), there is no signifi- cant difference between the richness of small and large size species within animal communities (Brown & Nico- letto, 1991). At larger spatial scales, the pattern is quite different with small species generally far exceeding the number of large species (Bakker & Kelt, 2000). 4.3. Cenograms The cenograms in Figure 4 illustrate the difference in total species richness (S) between the FUs as a func- tion of their body mass (y-axis): the higher the rank, the lightest the species (x-axis). In all FUs, a very clear break in slopes for species around 100 g can be ob- served, which has already started at about the limit of 500 g. Medium-sized species (500 g to 250 kg without carnivores) are less represented in all faunal units ex- cept in FUIV+V. Finally, the large-sized species (more than 250 kg) are scarce. According Legendre’s method, there are two general trends in Coudoulous I: either an open-arid landscape simultaneous with a decrease in small mammals as well as the total species richness (e.g. FUVII), or closed-arid landscapes with an increase in small mammals and highest species richness with possibly more humid climate trend (e.g. FUIV+V). These two opposing profiles should help inform our reading of the evolution of landscapes. For example, FUVII fits well with the results of Gingerich (1989) for open-arid envi- ronments. In these typical landscapes, the slope of the cenograms is more pronounced with a larger gap around the limit of 500 g, a scarcity of medium-sized species (500 g to 250 kg) and finally the lowest total species richness. Nevertheless, there are inconsisten- cies between the general shape of these two opposing cenograms and some paleoclimatic proxies such as MAP and AI (counterintuitively the higher the AI, the lower the aridity) (Tab. 4). Indeed, the latter show a more arid climatic trend in FUIV+V (AI=52.89 MAP = 972.20 mm) than in FUVII (AI=71.50 and MAP = 1236.79 mm). Furthermore, the range of values provided by De Martonne (1926) and reworked by Pellicone et al. (2019) for AI < 50 indicates a humid climate. The results obtained with the paleoclimatic proxies and the ceno- grams do not invalidate each other. However, they seem to indicate a wetter climate trend than the theoret- ical pattern of the model suggests, with more pro- nounced humidity in FUVII than in FUIV+V. Furthermore, we have to keep in mind that AI is a continuous com- plex gradient difficult to interpret, which spans several biomes (e.g. from tropical rainforest to tropical decidu- ous woodland, to savanna, to desert) (Hernández- Fernández et al., 2006). Gómez Cano et al. (2006) have shown that ceno- 98 Fernandez P. et al. Tab. 6 - a) Species richness (S) for microfauna and macrofauna in faunal units of Coudoulous I ; b) Chi² Test between species richness (microfauna vs macrofauna) in FUs; c) Species rich- ness (S) for herbivores and carnivores in FUs; d) Chi² Test between species richness (herbivores vs carnivores) in FUs. Fig. 4 - Cenograms according to the body mass (in g) for each faunal unit (FUII to FUVII) in Coudoulous I. For predator prey ratio (PPR=n predator /n prey) see details in text. Filled circles for bats, reptiles and amphibians. Red filled circles for carnivores. Paleoecologial first results from the Middle Pleistocene sequence of Coudoulous I (Quercy, Lot, France). 99 gram analyses work well with a random species loss of up to 60-70% in a fossil assemblage. The unequal sizes of the samples and the possible biases in representation of the species led us to calculate a confidence interval for the body mass of predators (excluding mustelids, M. meles, V. vulpes and F. silvestris), their prey (ungulates) and small mammals in each FU (Table 7-a). The boot- strap percentiles interval function in R (Version 3.2.1) makes it possible to provide the mean and the 2.5% and 97.5% percentile of the bootstrap distribution as confi- dence intervals (see details in Efron & Tibshirani, 1993). The predators, their prey and the small mammals showed relatively similar body mass confidence intervals in the different FUs (Table 7-a). To investigate this fur- ther we undertook a Kruskal-Wallis analysis, using the multiple pairwise comparisons option (XLSTAT Version 2017.1.1) to detect possible significant changes in their body mass (Table 7-b). Our results are mainly con- sistent with previous observations about a stable com- munity over time. Only the body mass values for small mammals from FUVII and FUVI are statistically different and should be explored to distinguish between evolu- tionary aspects (ecomorphology) or more likely sampling bias (s.l.) as discussed for FUVII. 4.4. Predation In present-day ecosytems, terrestrial carnivores are much rarer than most of their prey due to the competi- tion with predators of similar sizes to avoid the overcon- sumption of the resources on which they depend (Peters & Wassenberg, 1983; Hatton et al., 2015). As expected, the PPR calculated in all of the FUs are always lower than 1 (Fig. 4). They are relatively stable in the se- quence at around 0.6, with the exception of FUVII (PPR = 0.43) and FUII (PPR = 0.29). On a longer time scale, PPR ratio seems to be very dependent on the renewal of the Plio-Pleistocene fauna with the arrival or the extinction of certain predators (Raia et al., 2007; Croi- tor & Brugal, 2010). For now, it is difficult to infer pred- ators-prey interactions including man in Coudoulous I except that the greater PPR, the more important the CI in each FU. 4.5. Bioclimatic analysis In Coudoulous I, the mean annual temperatures (MAT) in the different faunal units are relatively similar to those established on the exclusive basis of the mi- crofauna by Jeannet (2021) (Tab. 4). Nevertheless, MAT alone does not permit the distinction between bioclimatic components, except in the case of very different ecozones (Rivas-Martínez et al., 1999). This is not the case with the mean annual thermal ampli- tude (MATA or IC in Rivas-Martinez et al., 1999) that constitutes an excellent indicator of continentality. The MATA values recorded in FUVII, FUIV+V and FUIII corre- spond to a semi-continental climate (IC 18 to 21 in Rivas-Martínez et al., 1999, p. 8) and to a sub- continental climate in FUVI and FUII (IC 21 to 28 ac- cording to the same authors). In both cases the win- ters were harsh and the summers hot and rainy. The annual positive temperature index (APTI) calculated at Coudoulous I indicates that the APTI values are all between around 900 and 1100 and correspond to the interval of the sub-Mediterranean temperate bioclimat- ic variant (800 1400, see Rivas-Martínez et al., 1999). Likewise, the compensated thermal indices Tab. 7 - a) Body mass (kg) bootstrap confidence interval distribution (95%) for prey (ungulates), predators (excluding all mustelids, Meles meles, Vulpes vulpes, Felis silvestris) and small mammals in FUs (calculation with the bootstrap percentiles interval function in R, Version 3.2.1). b) Kruskal-Wallis test for comparing the body mass (kg) of prey (ungulates), predators (excluding all mustelids, Meles meles, Vulpes vulpes, Felis silvestris) and small mammals in all FUs (Calculation from XLSTAT Version 2017.1.1, using multiple pairwise comparisons option for the recalculation of the ranks for each two-by-two combination of the samples, < 0.0001 is highly significant). of Coudoulous I all have values less than 180 which is consistent with this type of climate (see CTI in Rivas- Martínez et al., 2004). Finally, the most significant result concerns the faunal units, whose highest probabilities are all associated with the bioclimatic component VI, which corresponds to a typically temperate climate (Tab. 4). The vegetation cover of BC VI consists of a nemoral zone dominated by deciduous forests with broadleaf trees (e.g. maple, oak, hickory, beech, chest- nut) with several different kinds of plants like mountain laurel, azaleas and mosses. The bryophyte stratum and terrestrial lichens as well as various dwarf plants are very developed, while there are few herbaceous plants. Nevertheless, a poorly diversified or even mono-specific cover cannot be excluded in this type of environment (Hernández-Fernández, 2001a). This kind of vegetation cover is potentially located between the Boreal zone (coniferous forests) and the Mediterranean zone (sclerophyllous forests). In Table 4, FUII and FUVII, are both associated with BC VIII as second probabilities, with cold Boreal type climate elements. These results completely match the paleoenvironmental conclusions of Jeannet (2021) which indicate much more severe climatic conditions in FUII and FUVII. FUII could be related to MIS 6 with climat- ic degradation as mentioned previously, and FUVII to a cooler phase of MIS 7 or older (MIS 8?). The existence of large ice caps at the North Pole during ice ages, in conjunction with the Scandinavian anticyclone weather systems, likely generated continental surface winds coming from the East, and caused a decrease in precip- itation in Eurasia (COHMAP, 1988). For example, in France during these colder periods, there is a decrease in precipitation (down to a minimum of 600 mm during the Würm), unlike the hottest interglacial periods where precipitation can reach 1200 mm during the Eemian (Hernández-Fernández, 2001a). While it is very likely (Probability 1 and 2) that FUII, FUVII show climatic affini- ties with continental elements, the combination of aridity (AI) and rainfall pattern (MAP) related with thermal indi- ces (MAT, MTCM, APTI) seems to exclude a typical boreal climate in these faunal units, even in its mildest variant (sensu Rivas-Martínez et al., 1999, 2004) (see Tab. 4). However, in all of the FUs of Coudoulous I the as- sociation of some species present today but in environ- ments with very different environmental characteristics raises the question of their ecological plasticity and their contemporaneity. As an example, in FUIV+V we note the presence of Dicrostonyx torquatus and Sorex minutissi- mus which live today in very severe climatic conditions, unlike Oryctolagus cuniculus or to a lesser extent Micro- tus arvalis, which are generally associated with a much warmer climate. During the Late Pleistocene, the asso- ciation of D. torquatus and M. arvalis is not uncommon in southwestern France and reflects the ecological adaption of some taxa that evolved in very specific bi- omes (Royer et al., 2016). This kind of association is part of the “disharmonious communities” of animals and plants adapted to a climate that has no modern counter- part (Huggett, 2004). It could be explained by refuge areas forming a mosaic of settlements of small popula- tions dispersed in several habitats, known as the “patch -corridor-matrix model”, allowing their survival and co- habitation (Huggett & Cheesman, 2002). Regardless of the potential for climate adaptation of species and the plasticity of their ecological niche in the Pleistocene, there is no doubt that the temporal resolution of the faunal units of Coudoulous I corresponds to a grouping of different temporal phases. 5. CONCLUSION In current ecological studies, species richness has been demonstrated to vary consistently with large-scale geographical factors such as latitude and altitude. The sequence of Coudoulous I allows us to follow an entire animal community and its relationship to paleoenviron- mental changes through time at a local scale. First of all, it appears that FUII and FUVII are differ- ent from the other faunal units, with their low-quality samples, unbalanced and low PPR and the lowest spe- cies richness values. FUII, which is the most recent fau- nal unit, may belong to the first part of MIS 6, with new paleoenvironmental conditions. FUVII is the oldest faunal association and is very well individualized as it is com- posed of a single level (8d) that may correspond to a cooler phase of MIS 7 or older (MIS 8?). FUVII has the lowest species richness and above all an extremely disturbed assemblage as identified by the completeness index (CI). Nevertheless, FUVII has a greater diversity than FUII which provides a much larger number of spe- cies. This shows that the Shannon-Wiener (H’) performs well and is very sensitive to the individual distribution frequency even if there is a great imbalance of the spe- cies richness between samples. In the rest of the sequence, the faunal associations FUIII to FUVI show both greater diversity and species richness, thus indicating a greater number of trophic levels and a more stable community. The concept of the “insurance hypothesis”, which states that the greater the species richness, the more important are the interac- tions between species, is supported by the rarefaction curves that are systematically in line with Margalef (D). The number of species, according their body mass and their diet, displays no significant differences between all the FUs, as is usual in a small sampling area (e.g. with- in habitat). In addition, the Sørensen-Dice index almost always indicates a greater similarity of common species between the closest FUs throughout the sequence. This reveals a gradual turnover of species through time, probably due to paleoenvironmental changes. The predators, their prey and the small mammals showed no significant changes in their body mass through time, with the exception of the small mammals between FUVII and FUVI, which probably indicates the same pattern of evolution throughout the sequence. We showed clearly for FUII and FUVII that the lower the com- pleteness index, the lower the PPR. This ratio is practi- cally the same in the rest of the sequence but difficult to interpret from the small number of predators and their prey. Theoretically, body mass distribution analysis in the cenograms should be efficient with a random spe- 100 Fernandez P. et al. Paleoecologial first results from the Middle Pleistocene sequence of Coudoulous I (Quercy, Lot, France). 101 cies loss of up to 60-70% in a fossil assemblage. We established the cenograms in the different FUs and found them consistent with the pattern of a temperate climate corresponding to the bioclimatic component VI. According their theoretical slopes, breakpoints and species richness we identified two general trends cor- responding to an open-arid and more closed-arid cli- mate. Nevertheless, a cross reading with paleoclimatic proxies revealed a wetter climate trend than the theo- retical pattern arid-open landscape suggests. In addition to the cenograms, the bioclimatic model based on transfer functions from current mam- malian associations clearly showed that the paleoeco- logical requirements of Coudoulous I species were very similar between the different faunal units. While there were significant climatic changes, they remained limited enough to prevent any complete turnover of species. One of the key outputs is the maximum prob- abilities calculated for each FU that are all systemati- cally associated with the bioclimatic component VI. This component corresponds to a typical temperate climate with a predominance of deciduous forests that could potentially be located between the Boreal zone (coniferous forests) and the Mediterranean zone (sclerophyllous forests). At Coudoulous I, sub- Mediterranean temperate elements dominate through- out the sequence but faunal units such as FUVII, FUIV+V and FUIII show affinities with semi-continental or sub- continental conditions (i.e. FUVI, FUII). Finally, the paleoclimatic proxies from the bioclimatic model make it possible to exclude a boreal-type climate. ACKNOWLEDGEMENTS Several studies of interest to geology, dating, taxonomy and paleoenvironment have been carried out on this Coudoulous I sequence in recent years which will be soon published as part of a monograph 2021 under the direction of J. Jaubert. We want to warmly thank people of the team working on this site, especially faunal specialists A. Argant (Felids and Ursids), M. Boudadi-Maligne (Canids), F. Lacombat (Rhinocerotids), B. Labe (Proboscids), N. Lateur (Cervids), J.B. Mallye (Mustelids), and geo- stratigraphers B. Kervazo and colleagues. We thank the editor of AMQ as well as the two anony- mous reviewers for their unvaluable inputs and com- ments on previous draft. Finally, we are extremely grateful to Prof. Lucy Wilson (University of New Bruns- wick, Canada) for the corrected English version of the manuscript. REFERENCES Adrain J.M., Westrop S.R., Chatterton B.D.E., Ram- sköld L. 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