Cougars’ key to survival through the Late Pleistocene extinction: insights from dental microwear texture analysis

Cougars (Puma concolor) are one of only two large cats in North America to have survived the Late Pleistocene extinction (LPE), yet the specific key(s) to their relative success remains unknown. Here, we compare the dental microwear textures of Pleistocene cougars with sympatric felids from the La Brea Tar Pits in southern California that went extinct at the LPE (Panthera atrox and Smilodon fatalis), to clarify potential dietary factors that led to the cougar''s persistence through the LPE. We further assess whether the physical properties of food consumed have changed over time when compared with modern cougars in southern California. Using dental microwear texture analysis (DMTA), which quantifies surface features in three dimensions, we find that modern and Pleistocene cougars are not significantly different from modern African lions in any DMTA attributes, suggesting moderate durophagy (i.e. bone processing). Pleistocene cougars from La Brea have significantly greater complexity and textural fill volume than Panthera atrox (inferred to have primarily consumed flesh from fresh kills) and significantly greater variance in complexity values than S. fatalis. Ultimately, these results suggest that cougars already used or adopted a more generalized dietary strategy during the Pleistocene that may have been key to their subsequent success.     

Carnivores from Los Rincones,a leopard den in the highest mountain of the Iberian range (Moncayo,Zaragoza, Spain)

Carnivores are a common element in Pleistocene fossil assemblages. However, they are not so abundant in terms of the numbers of remains. Here, we present a palaeontological study of the carnivores from the Late Pleistocene (MIS 3) of Los Rincones (Spain), one of the few deposits accumulated by leopards. One hundred and ten leopard remains have been recovered. This carnivore is not the only inhabitant of the cave, and 175 remains belonging to Ursus arctos have also been recovered, making it one of the sites with the greatest number of brown bear remains in the Iberian Peninsula. The large number of leopard remains has allowed us to make a detailed morphological and biometrical study that has enabled us to classify the remains within the subspecies Panthera pardus spelaea. The European Ice Age leopard inhabited Europe during Upper Pleistocene and it presents some similarities with Panthera uncia. A study of the scarce remains of Canis lupus indicates that this was similar in size to Canis lupus maximus; the scarcity of the remains prevents us from assigning our remains to this subspecies. A study of the brown bear remains indicates that it is similar to other populations in the north of the Iberian Peninsula with this chronology.     

Radiographs Reveal Exceptional Forelimb Strength in the Sabertooth Cat,Smilodon fatalis

Background

The sabertooth cat, Smilodon fatalis, was an enigmatic predator without a true living analog. Their elongate canine teeth were more vulnerable to fracture than those of modern felids, making it imperative for them to immobilize prey with their forelimbs when making a kill. As a result, their need for heavily muscled forelimbs likely exceeded that of modern felids and thus should be reflected in their skeletons. Previous studies on forelimb bones of S. fatalis found them to be relatively robust but did not quantify their ability to withstand loading.

Methodology/Principal Findings

Using radiographs of the sabertooth cat, Smilodon fatalis, 28 extant felid species, and the larger, extinct American lion Panthera atrox, we measured cross-sectional properties of the humerus and femur to provide the first estimates of limb bone strength in bending and torsion. We found that the humeri of Smilodon were reinforced by cortical thickening to a greater degree than those observed in any living felid, or the much larger P. atrox. The femur of Smilodon also was thickened but not beyond the normal variation found in any other felid measured.

Conclusions/Significance

Based on the cross-sectional properties of its humerus, we interpret that Smilodon was a powerful predator that differed from extant felids in its greater ability to subdue prey using the forelimbs. This enhanced forelimb strength was part of an adaptive complex driven by the need to minimize the struggles of prey in order to protect the elongate canines from fracture and position the bite for a quick kill.     

Implications of Diet for the Extinction of Saber-Toothed Cats and American Lions

The saber-toothed cat, Smilodon fatalis, and American lion, Panthera atrox, were among the largest terrestrial carnivores that lived during the Pleistocene, going extinct along with other megafauna ∼12,000 years ago. Previous work suggests that times were difficult at La Brea (California) during the late Pleistocene, as nearly all carnivores have greater incidences of tooth breakage (used to infer greater carcass utilization) compared to today. As Dental Microwear Texture Analysis (DMTA) can differentiate between levels of bone consumption in extant carnivores, we use DMTA to clarify the dietary niches of extinct carnivorans from La Brea. Specifically, we test the hypothesis that times were tough at La Brea with carnivorous taxa utilizing more of the carcasses. Our results show no evidence of bone crushing by P. atrox, with DMTA attributes most similar to the extant cheetah, Acinonyx jubatus, which actively avoids bone. In contrast, S. fatalis has DMTA attributes most similar to the African lion Panthera leo, implying that S. fatalis did not avoid bone to the extent previously suggested by SEM microwear data. DMTA characters most indicative of bone consumption (i.e., complexity and textural fill volume) suggest that carcass utilization by the extinct carnivorans was not necessarily more complete during the Pleistocene at La Brea; thus, times may not have been “tougher” than the present. Additionally, minor to no significant differences in DMTA attributes from older (∼30–35 Ka) to younger (∼11.5 Ka) deposits offer little evidence that declining prey resources were a primary cause of extinction for these large cats.     

Phylogenetics of Panthera,including Panthera atrox,based on craniodental characters

The phylogenetic position of Panthera atrox within Felidae is still controversial despite many morphological and molecular studies addressing its relationships. This is in part due to the lack of consensus on a tree for Panthera. These inconsistencies suggest the need for further analysis and perhaps even different methodology to better understand pantherine evolution. Morphologic characters from the skull and dentary were analysed within Panthera to elucidate pantherine phylogeny. Extant taxa included Panthera leo (African lion), Panthera tigris (tiger), Panthera onca (jaguar), Panthera pardus (leopard), Uncia uncia (snow leopard) and Neofelis nebulosa (clouded leopard). Four outgroups were used: Crocuta crocuta (spotted hyena), Metailurus spp., Proailurus lemanensis and Pseudaelurus validus. Our study found a clade containing Panthera leo, Panthera tigris and Panthera atrox, suggesting that Panthera atrox is more closely related to the African lion and the tiger than the jaguar, in contrast to what has been recently proposed. Moreover, gross morphological similarities between Panthera atrox and Panthera onca are more likely the result of convergent hunting styles and/or prey selection, rather than phylogenetic affinity.     

On the status of “Urotherium antiquum” (Ameghino) (Xenarthra,Glyptodontidae)

Within the Late Neogene Glyptodontidae of the Pampean region of Argentina, “Urotherium antiquum” was described on the basis of some associated osteoderms of the dorsal carapace, which in addition include a partial skull and left hemimandible. The diagnostic characters are located on the exposed surface of the osteoderms of the dorsal carapace which somewhat resembles those of the Pleistocene genus Neuryurus. Although the relationship of “U. antiquum” to the remaining Glyptodontidae has never been clarified, some cladistic analyses suggest a close phylogenetic affinity with the clade composed of Plohophorus + (Glyptodon + Doedicurus). A careful comparison of “U. antiquum” to well-characterized taxa with similar stratigraphic and geographic provenance reveals that no significant skull differences are observed from Plohophorus figuratus Ameghino. It seems likely that the alleged ornamentation pattern that characterizes this species was produced by a taphonomic alteration of the exposed surface of the osteoderms, a process broadly distributed in glyptodonts having a “rosette” ornamentation pattern. Finally, some characters present in the osteoderms of the type specimen of “U. antiquum” suggest that it may be a juvenile specimen. In summary, “U. antiquum” should be considered a junior synonym of P. figuratus.     

Evolutionary changes in craniomandibular shape in the great cats (Neofelis Griffith and Panthera Oken)

Evolutionary shape changes in skull and mandibular anatomy was analysed in 223 specimens of pantherine felids (Neofelis nebulosa, Panthera leo, Panthera onca, Panthera pardus, Panthera tigris, Panthera uncia) compared to a small‐felid outgroup, consisting of 86 specimens of nine different species, using digital surface morphometry on 25 (skull) and 17 (mandible) landmarks. Shape evolution in the pantherine species is complex and nonlinear, and involves both large‐scale and small‐scale shape changes. Shape changes frequently differ among the ingroup species, but the four large Panthera species (leo, onca, pardus, tigris) bear some resemblance to each other. The leopard and jaguar bear the closest resemblance to each other, and several shape changes are common to the lion and tiger, but have probably evolved convergently as a result of large size. The lion has undergone the largest and most numerous shape changes from a small‐felid outgroup. Certain shape changes in the skull and, in some respects, the mandible of the clouded leopard bear resemblance to those in the four large Panthera species. The snow leopard is often regarded as the most primitive of the extant Panthera, and skull and mandibular shape changes often diverge markedly from those observed in the other five ingroup taxa; its overall skull shape is rather similar to the small‐felid outgroup. This indicates that the shape changes in the clouded leopard are convergent with those of the four large Panthera species. Landmark integration showed no significant correlation with molecular phylogeny, chiefly owing to the snow leopard being placed among the four large Panthera species. A traditional phylogenetic topology with the snow leopard as the basal‐most species of Panthera yielded a weak but nonsignificant phylogenetic signal. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 766–778.     

Stephanorhinus hundsheimensis (Mammalia,Rhinocerotidae) from the late early Pleistocene deposits of the Denizli Basin (Anatolia,Turkey)

Pleistocene rhinoceroses are poorly documented in Turkey where they have been reported only from the late early Pleistocene (1.3–1.1 Ma) travertine deposits of the Denizli Basin. In this work, new rhinoceros remains collected from this basin are assigned to a relatively large-sized Stephanorhinus hundsheimensis on the basis of their morphology and morphometry. The first Turkish record of this species is approximately coeval with the first appearance of S. hundsheimensis in Europe, chronologically referred to the late early Pleistocene, ca. 1.2 Ma. During that time, S. etruscus still survived in Iberian Peninsula, central Italy and Dacian Basin. The presence of two successive evolutionary morphs for S. hundsheimensis during the Pleistocene is not confirmed.     

Biogeography and chronology of the Eurasian extinct rhinoceros Stephanorhinus etruscus (Mammalia,Rhinocerotidae)

Stephanorhinus etruscus is one of the most abundantly recorded and better known Eurasian Early Pleistocene rhinoceroses. Nevertheless, the first and last appearances of this species, as well as its paleogeographic distribution, are controversial and debated in literature. S. etruscus is documented since the latest Pliocene in Spain (Las Higueruelas), Italy (Montopoli and Castelnuovo di Barardenga), France (Perrier–Les Étouaires) and Romania (Iar??–Cariera Veche). During the Early Pleistocene, S. etruscus occurred in several Spanish, French and Italian localities, as well as in The Netherlands (e.g., Tegelen), Germany (e.g., Thiede), Greece (e.g., Aivaliki) and Israel (e.g., Ubeidiya). The last appearance of S. etruscus in Eurasia is debatable. Etruscan rhino populations survived till the Jaramillo subchrone (around 1.1 Ma) in France (Bois-de-Riquet), Romania (Betfia XII) and Hungary (Osztramos 2 and 8), and close to the early–middle Pleistocene transition in Spain (Cueva Victoria, Huéscar 1, Atapuerca TD4, TD6 and TD8), and Italy (Monte delle Piche).     

Stephanorhinus jeanvireti (Mammalia,Rhinocerotidae) from the early Pleistocene of Colțești (southwestern Romania)

The Rhinocerotidae material from the early Pleistocene Tetoiu Formation of Colțești (southwestern Romania) is here described for the first time. The rhinoceros is documented by calcaneus and second, third, and fourth metatarsals, probably belonging to the same individual. The morphology and the dimensions of these specimens enable us to record the presence of Stephanorhinus jeanvireti, a relatively rare rhinoceros usually reported from late Pliocene European localities and recently considered a junior synonym of S. elatus. Nevertheless, the taxon Rhinoceros elatus is here regarded as a nomen dubium and the name S. jeanvireti is retained in order to maintain nomenclatural stability, being it based on much more diagnostic material. In Romania, S. jeanvireti has been listed within a few late Pliocene faunal assemblages, but the rhinoceros remains are fragmentary and isolated bones. The Colțești find is among the best documented records of S. jeanvireti in Romania. The biochronological distribution of S. jeanvireti is mainly confined within the MNQ16 (early and early middle Villafranchian), and its first occurrence is doubtfully reported in late MNQ15 faunas. The record of Colțești (MNQ17/MNQ18) represents, instead, the last occurrence of this taxon in Europe.     

Cold-climate rodent indicators for the Late Pleistocene of Central Iberia: New data from the Buena Pinta Cave (Pinilla del Valle,Madrid Region,Spain)

In the Iberian Peninsula, the Late Pleistocene record of small mammal indicators of cold climates is largely restricted to two sets of sites at the eastern and western ends of the Pyrenees. Some assemblages from other sites at the Peninsular centre have, however, recently yielded such taxa. This work describes the remains of three such rodent species from the Buena Pinta Cave, a site in the Sierra de Guadarrama mountains in the Spanish Central System. Excavation campaigns in the cave have taken place every summer since 2003. Thermoluminescence analyses of sediments from levels 2–5 of the site suggest an age corresponding to the middle of the Late Pleistocene, within Marine Isotope Stage 4 or the beginning of Marine Isotope Stage 3. Sieve-washing and picking out of the small fossils contained in the sediments of levels 2–5 yielded several thousand small mammal teeth and other remains, dominated by Microtus arvalis. Smaller numbers of remains belonging to other rodents typical of cold climates were also identified, such as Microtus oeconomus, Microtus gregalis and Chionomys nivalis. Thus, the small mammal record of the Buena Pinta Cave shows that rodent indicators of cold climates reached the centre of the Iberian Peninsula during the mid-Late Pleistocene, i.e., well before the Last Glacial Maximum. These findings represent one of the southernmost Pleistocene records for M. oeconomus in Europe, and the most southerly for M. gregalis.     

Comparative Biomechanical Modeling of Metatherian and Placental Saber-Tooths: A Different Kind of Bite for an Extreme Pouched Predator

Questions surrounding the dramatic morphology of saber-tooths, and the presumably deadly purpose to which it was put, have long excited scholarly and popular attention. Among saber-toothed species, the iconic North American placental, Smilodon fatalis, and the bizarre South American sparassodont, Thylacosmilus atrox, represent extreme forms commonly forwarded as examples of convergent evolution. For S. fatalis, some consensus has been reached on the question of killing behaviour, with most researchers accepting the canine-shear bite hypothesis, wherein both head-depressing and jaw closing musculatures played a role in delivery of the fatal bite. However, whether, or to what degree, T. atrox may have applied a similar approach remains an open question. Here we apply a three-dimensional computational approach to examine convergence in mechanical performance between the two species. We find that, in many respects, the placental S. fatalis (a true felid) was more similar to the metatherian T. atrox than to a conical-toothed cat. In modeling of both saber-tooths we found that jaw-adductor-driven bite forces were low, but that simulations invoking neck musculature revealed less cranio-mandibular stress than in a conical-toothed cat. However, our study also revealed differences between the two saber-tooths likely reflected in the modus operandi of the kill. Jaw-adductor-driven bite forces were extremely weak in T. atrox, and its skull was even better-adapted to resist stress induced by head-depressors. Considered together with the fact that the center of the arc described by the canines was closer to the jaw-joint in Smilodon, our results are consistent with both jaw-closing and neck musculature playing a role in prey dispatch for the placental, as has been previously suggested. However, for T. atrox, we conclude that the jaw-adductors probably played no major part in the killing bite. We propose that the metatherian presents a more complete commitment to the already extreme saber-tooth ‘lifestyle’.     

Les ostracodes plio-pléistocènes de la côte orientale de la Péninsule du Cap Bon (Tunisie)

This article concerns the study of the ostracods of the Upper Pliocene-Lower Pleistocene sedimentary deposits cropping out along the south-eastern coast of the Cape Bon Peninsula (Tunisia, Hammamet region, Sidi Khelas section). Fourteen samples were taken along a thirty-two-meters thick sedimentary silico-clastic series. Seventy species of ostracods belonging to twenty-four genera were found. The autochthonous association, which originally lived in an environment located in the infralittoral and upper circalittoral stages of the benthic marine domain, is cyclically affected by inputs from the more superficial environments due to the presence of relatively well-developed lagoon and/or estuary environments in the surrounding coastal areas. Within the ostracod association, the presence of Bythocythere turgida – a true “northern guest” – is particularly significant. This species indicates that these sediments were deposited during the cold phases of the Early Pleistocene. Among the ostracods found, those belonging to the families Trachyleberididae, Hemicytheridae and Bythocytheridae, which are dominant and include particularly significant species from the environmental and paleoclimatic point of view, are systematically recorded and commented on. Some species that are probably new have been described and photographed, but are currently left in open nomenclature because of how few specimens have been found.     

A new whale barnacle from the early Pleistocene of Italy suggests an ancient right whale breeding ground in the Mediterranean

The fragmentary fossil history of whale barnacles (Cirripedia: Coronulidae) is mainly constituted by remains of Coronula spp. in Plio-Pleistocene deposits found along extant humpback whale migration routes, and especially in occurrence of breeding areas. Here we report the presence of a single compartment of Cetopirus along with remains of Coronula bifida in Lower Pleistocene open shelf deposits of Salento (South Italy). This is the first occurrence of the genus Cetopirus prior to the Late Glacial period (ca. 15–10 ky before Present), and the specimen here studied is designated as the holotype of the new fossil species Cetopirus fragilis. Since Cetopirus is currently found exclusively inhabiting the skin of the right whales (Eubalaena spp.), the fossil material here studied suggests the persistence of Balaenidae in the Mediterranean Basin during the Early Pleistocene and supports the existence of a baleen whale migratory route active between the central Mediterranean and the North Atlantic during the Plio-Pleistocene.     

The Xujiayao 14 Mandibular Ramus and Pleistocene Homo Mandibular Variation

The earlier Late Pleistocene mandibular ramus from Xujiayao (northern China) preserves traits that vary distributionally among western Old World Pleistocene Homo samples and between Early/Middle Pleistocene archaic humans and Late Pleistocene modern humans in eastern Eurasia. Xujiayao 14 presents a lateral mandibular notch crest, an open mandibular foramen, a wide ramus, an asymmetrical mandibular notch, an enlarged superior medial pterygoid tubercle, (probably) a retromolar space, and gonial eversion, as well as an unusual depression in the planum triangulare. The first two traits appear ancestral for Later Pleistocene and recent Homo and are dominant among modern humans. The second two traits largely separate Xujiayao 14 and archaic Homo from modern humans. The next two traits are found in the highest frequency among the Neandertals, although gonial eversion contrasts with Late Pleistocene Neandertals. Xujiayao 14, in the context of Pleistocene and recent Homo samples and the other Xujiayao human remains, therefore provides a morphological mosaic, highlighting regional variation through the Pleistocene.     

Inner structural organization of the distal humerus in Paranthropus and Homo

The taxonomic attribution of isolated hominin distal humeri has been a matter of uncertainty and disagreement notwithstanding their relative abundance in the fossil record. Four taxonomically-based morphotypes, respectively representing P. boisei, P. robustus, non-erectus early Homo and H. erectus, have been identified based on the cross-sectional outer shape variation of an assemblage of Plio-Pleistocene eastern and southern African specimens (Lague, 2015). However, the existence of possible differences between Paranthropus and Homo in the inner structural organisation at this skeletal site remains unexplored. We used noninvasive imaging techniques to tentatively characterize the endostructural organization of five early Pleistocene distal humeri from South Africa (TM 1517g, SK 24600, SKX 10924, SKX 34805) and Ethiopia (Gombore IB), which have been variably attributed to Paranthropus or Homo. While the investigated specimens reveal diverse degrees of inner preservation related to their taphonomic and diagenetic history, in all but SK 24600 from Swartkrans we could comparatively assess some geometric properties at the most distal cross-sectional level (%CA, I_x/I_y, I_max/I_min) and quantify cortical bone thickness topographic variation across the preserved shaft portions by means of a 2-3D Relative Cortical Thickness index. Whenever possible, we also provided details about the site-specific organization of the cancellous network and measured the same parameters in a comparative sample of twelve adult extant humans. For most features, our results indicate two main patterns: the first includes the specimens TM 1517g, SKX 10924 and SKX 34805, while the second endostructural morphotype sets apart the robust Homo aff. erectus Gombore IB specimen from Melka Kunture, which more closely resembles the condition displayed by our comparative human sample. Notably, marked differences in the amount and pattern of proximodistal cortical bone distribution have been detected between Gombore IB and SKX 34805 from Swartkrans. Given its discordant outer and inner signatures, we conclude that the taxonomic status of SKX 34805 deserves further investigations.     

Western European Quaternary lions: new working hypotheses

Cave lions (Panthera spelaea), which spread throughout Western Europe for several thousand years, disappeared approximately 14 000–14 500 years ago. They were supposedly replaced by modern lions (Panthera leo) approximately 8000 years ago. Modern lions reached the steppes of Ukraine and Hungary, without penetrating the forests of Central Europe. The present study focuses on Italian and Spanish findings that possibly bridge the alleged absence of these big cats from Europe for 6000 years. Fossil lion remains from reliably radiocarbon‐dated levels have been plotted against the δ¹⁸O curve and mapped. The accumulated evidence indicates that lions inhabited Western Europe uninterruptedly from the early Middle Pleistocene up to the Early Holocene. Moreover, all of the latest Pleistocene/early Holocene lion‐bearing localities do not range farther than the 44th parallel north and are located at relatively high altitudes. Two working hypotheses are formulated: one, which is less likely because it is not supported by palaeontological evidence indicating earlier migrations of lions from Africa, suggests that modern lions entered Western Europe prior to 8000 years ago; the second, which is more probable, suggests that P. spelaea (or an advanced offspring of the species) survived up until the latest Pleistocene. Panthera leo accessed Eastern Europe between 6000–6500 and 8000 years ago but was prevented from penetrating further west, probably because of the intrusive presence of their indigenous European relatives, and/or the increasing encroachment of modern human populations. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 66–77.     

Snake venomics and antivenomics of Bothrops atrox venoms from Colombia and the Amazon regions of Brazil,Perú and Ecuador suggest the occurrence of geographic variation of venom phenotype by a trend towards paedomorphism

The venom proteomes of Bothrops atrox from Colombia, Brazil, Ecuador, and Perú were characterized using venomic and antivenomic strategies. Our results evidence the existence of two geographically differentiated venom phenotypes. The venom from Colombia comprises at least 26 different proteins belonging to 9 different groups of toxins. PI-metalloproteinases and K49-PLA₂ molecules represent the most abundant toxins. On the other hand, the venoms from Brazilian, Ecuadorian, and Peruvian B. atrox contain predominantly PIII-metalloproteinases. These toxin profiles correlate with the venom phenotypes of adult and juvenile B. asper from Costa Rica, respectively, suggesting that paedomorphism represented a selective trend during the trans-Amazonian southward expansion of B. atrox through the Andean Corridor. The high degree of crossreactivity of a Costa Rican polyvalent (Bothrops asper, Lachesis stenophrys, Crotalus simus) antivenom against B. atrox venoms further evidenced the close evolutionary kinship between B. asper and B. atrox. This antivenom was more efficient immunodepleting proteins from the venoms of B. atrox from Brazil, Ecuador, and Perú than from Colombia. Such behaviour may be rationalized taking into account the lower content of poorly immunogenic toxins, such as PLA₂ molecules and PI-SVMPs in the paedomorphic venoms. The immunological profile of the Costa Rican antivenom strongly suggests the possibility of using this antivenom for the management of snakebites by B. atrox in Colombia and the Amazon regions of Ecuador, Perú and Brazil.     

Small and large mammals from the Ciota Ciara cave (Borgosesia,Vercelli, Italy): An Isotope Stage 5 assemblage

The Ciota Ciara cave is located within the Monte Fenera karst system (Borgosesia, Vercelli, Italy) at 670 m a.s.l. The cave entrance presents a deposit with Mousterian quartz and flint industry. The faunal remains from Stratigraphic Units 13, 14 and 103 are the subject of this work and are presented here as a whole for the first time. The large mammal assemblage is dominated by Ursus spelaeus. In addition, a few remains of carnivores such as Panthera leo spelaea, Panthera pardus, Meles meles have been found together with ungulates such as Cervus elaphus and Rupicapra rupicapra. The small mammal assemblage is characterized by a high biodiversity, especially in bats, by the dominance of Clethrionomys glareolus and by a relatively large number of Pliomys coronensis, a species that seems to disappear from the Italian Peninsula at the end of Marine Isotope Stage 5 or during the beginning of MIS 4. The changes in frequency of the small and large mammals between the two S.U. suggest a change from a relatively cold-humid (S.U. 14) to a warmer-still humid climate (S.U. 13). Although no radiometric dates are available yet, the small and large mammal assemblages, the presence of Pliomys coronensis and the climate change inferred by the variation of the small mammal frequencies allow us to correlate these two units of the Ciota Ciara cave to a relatively warm moment of Marine Isotope Stage 5.