Paleoecology of human interaction with associated faunas

The article presents the application of K-means clustering algorithm to sorting female and male pedicles of reindeer (Rangifer tarandus) from the Paleolithic site of Cosăuţi from Soroca District (the Republic of Moldova). According to the obtained results, 29.8% of frontal bones with pedicles belong to males, while the rest of 70.2% of specimens belong to females. The obtained value of the sex ratio of the reindeer pedicle sample confirms the earlier study that obtained the same results based on sexual dimorphism of limb bones. The volume of pedicles represents the most secure parameter that allows to distinguish female pedicles from male pedicles. The distribution of pedicle volume size in the samples of males and females corresponds to the age profiles of the samples.

The paper discusses the geographic origin and the cultural context of pearl-shaped beads/pendants from the Late Paleolithic site of Cosăuți. The morphological and biometric characteristics of the pendants reveal the fact that they are made of the upper canines of red deer Cervus elaphus, a species adapted to mild climate conditions. The osteological material from Cosăuți belongs exclusively to representatives of the cold-adapted fauna and does not contain other skeletal remains of red deer. The artifacts made of red deer upper canines from Cosăuți suggest cultural contacts between the Paleolithic populations from the Prut-Dniester area and the southern part of Balkans where Cervus elaphus persisted during the Last Glacial Maximum.

The new archaeozoological and taphonomical data identify the Middle Paleolithic site of Buzdujeni 1 as a periodic cave hyena (Crocuta crocuta spelaea) den. The most complete taphonomic evidence on hyena den come from the recently excavated squares K-5 and L-5 of layer 3, but the presence of hyena at Buzdujeni 1 site is also recorded in layers 4, 5, and 6. The evidence of cave hyena presence at Buzdujeni 1 include steak-like bone fragments, nibbled and chewed stick-like bone fragments, gnawed bones of large prey species (Bison priscus, Coelodonta antiquitatis), and skeletal remains of juvenile individuals. The composition of fauna from Buzdujeni 1 indicates generally open landscape in the environments of the site when Paleolithic layers were accumulated.

The article presents a description of cervid remains from the Middle Pleistocene Acheulean site of Gruta da Aroeira (= Galerias Pesadas) in central Portugal. The assemblage comprises the remains of four deer species: Cervus elaphus, Praedama cf. savini, Haploidoceros mediterraneus, and Dama cf. vallonnetensis, making Gruta da Aroeira the first site in the Iberian Peninsula at which the genus Haploidoceros has been documented in the Middle Pleistocene. Virtually all the cervids documented at the site demonstrate a degree of endemism, including evolutionary modifications of skull, antlers and dentition or a reduction in body size. The unusual richness of the cervid community at Gruta da Aroeira may be related to the specific biogeographic conditions of the Middle Pleistocene in Iberia. Thus, while the Iberian Peninsula’s geographical link with the temperate west Eurasian zone facilitated the dispersal of cervids of palearctic origin into Iberia, it impeded the dispersal of ruminants from warmer, more arid areas. The endemic character of the Middle Pleistocene cervids and the biodiversity of the Iberian Peninsula should shed some light on the paleobiogeography of Iberian hominins and their role in hominin hunting or their economic strategies.

The article attests the presence of the genus Rucervus in the paleontological record of Europe and presents the description of new species of large-sized deer Rucervus radulescui sp. nov. from the Early Pleistocene of Valea Grăunceanului (Southern Romania) and Rucervus gigans sp. nov. from the late Early Pleistocene of Apollonia-1 (Greece). The described cervid species represent two different evolutionary radiations of Rucervus that are grouped into the extinct subgenus Arvernoceros that represents the northern evolutionary radiation and the nominotypical subgenus that is regarded as the southern evolutionary radiation and represented today by only one species Rucervus duvaucelii. The evolutionary radiation and dispersals of Rucervus are regarded in the paleobiogeographic context of faunal exchanges between southeastern Europe, Caucasus, and Near East during the Early Pleistocene and the westward dispersal of early hominins in Eurasia. OPEN ACCESS: http://www.mdpi.com/2571-550X/1/2/17/htm

The article describes the early Middle Age horse skeletal remains yielded by the Saltovo-Mayaki archaeological sites from Eastern Ukraine. According to the obtained results, the Saltovo-Mayaki domestic breed represents an improved riding horse with the medium height at the withers, thin and semi-thin metacarpals, elongated phalanxes, moderately short muzzle, relatively large cheek teeth and long protocone in upper cheek teeth. A large number of juveniles and few seniles in the studied sample suggest that Saltovo-Mayaki Culture bearers practiced hippophagy. This conclusion is supported by the recorded cutting tool marks on horse bones.

Монография посвящена анализу археологического материала кизил- кобинской культуры поселений Инкерманской долины. В основе ана- лиза — материалы недавних раскопок поселения Уч-Баш, а также ар- хеологические коллекции предыдущих раскопок. Археологический источник анализируется в контексте реконструкций окружающей среды, что необходимо для воссоздания палеоэкономики жителей Инкерманской долины. Предпринята попытка синтеза разных на- правлений научных исследований как гуманитарных, так и естест- венных наук для реконструкции социально-экономической модели населения времени перехода от бронзового к железному веку. Для историков, археологов, преподавателей средней и высшей школы, студентов, а также всех интересующихся древней истори- ей юга современной Украины.

The article presents a description of horse remains (teeth and limb bones) from two Belozerka Culture sites (Late Bronze and Early Iron Age of Southeast Europe) from South Moldova.

The ecological and evolutionary interaction during the last 3 million years between communities of large-sized herbivores (67 species, proboscideans and rhinocerotids excluded) and carnivores (30 species with body mass above 10 kg) from western Eurasia is analyzed. This frame can bring information about the ecologic context of early human dispersal. The ecologic and systematic structures of herbivore and carnivore communities during the Late Pliocene and Early Pleistocene show little evidence on mutual evolutionary influence triggered by climate change. Herbivores were highly sensitive to climate fluctuations that caused a high species turnover due to extinctions of local stenobiont forms and dispersals of new large-sized oriental species. The body size structure of the whole herbivore guild sifted significantly toward the large-sized body class. The dispersal of first hominines into the European subcontinent (1.1-1.4 Ma) coincided with this restructuration of the herbivore guild. Unlike herbivores, carnivores were long-living ubiquitous species with a large distribution range and show a lesser dependence on climate shifts caused by the 41 kyr climatic cycles. This archaic predator assemblage lasted in Europe until ca. 1.2-1.0 Ma. The high-amplitude 100 ka cycles caused a significant increase of carnivore species turnover marked by the extinction of specialized felid predators, however, the main ecological parameters of the carnivoran guild (mean body mass and species richness) changed insignificantly. Early Pleistocene predators faced to the "large body size-cursorial ability" evolutionary dilemma that impeded the adaptation of solitary carnivores to megaherbivores in the open landscape conditions. Such dilemma was solved independently in the lineages of Felidae (Panthera leo), Hyaenidae (Crocuta crocuta), and Canidae (Canis lupus, Lycaon lycaonoides) by acquiring ethological characteristics such as social behavior and cooperative hunting that in combination with large body mass (lions) or high cursorial abilities (spotted hyena and dogs) to take a larger prey. Such a scenario will be put in parallel with the arrival and permanent settlement of human groups in Western Eurasia. Factors as ecological flexibility (especially in diet) and sophisticated social behaviour of early hominines were particularly advantageous in the specific faunal ecosystems of Western

Paleobiogeography is proposed here as a helpful source of information on limiting environmental factors and the shape and range of the archaic hominin ecological niche. The cluster analysis of Early Pleistocene ircum-Mediterranean mammal faunas confirms a single way of early hominin dispersals into the Eurasian mainland via Near East. Despite of the assumed belonging of early hominins to the carnivore guilds as a commensal scavenger, the biogeography of early hominin dispersal and area of distribution dynamics were different from those of true ubiquitous carnivores, which dispersed over entire continents. The hominin area of distribution on the Eurasian mainland during the Early and beginning of Middle Pleistocene was limited by geographic and climatic factors. The hominin dispersal toward Sundaland (ca.

The article proposes an interpretation of hunting strategy of Late Paleolithic hunters from Moldova based on demographic structure of reindeer remains (sex ratio and proportion of juvenile remains) and reindeer paleobiology and ecology. The obtained results demonstrate a flexible strategy of game procurement of Paleolithic hunters ensuring the optimal energy investment/ food gain ratio. The hunting strategy was influenced by prey ecology, seasonal biological cycle, paleogeographic conditions, prey availability, cultural traditions, and available human resources.

A multivariate cluster analysis of western Eurasian regional herbivorous mammalian faunas is applied in order to reveal the paleobiogeographic context of early human dispersal in the area under study. During the Early Pleistocene, the north Mediterranean area and Caucasian Land acted as refugia for warm-loving Pliocene faunal holdovers. The Italian Peninsula was biogeographically partially isolated during most of the Early Pleistocene due to the forested Dinaric Alps zoogeographic filter, which possibly caused the late arrival of hominines on the Italian Peninsula. The multivariate analysis confirms a firm paleobiogeographic border between the Iberian Peninsula and northwestern Africa. The Pannonian – western European path is proposed here as the most plausible dispersal route for early hominines. The article gives a brief discussion of paleobiogeographic significance of the Alpine-Himalayan Mountain Belt and the Movius Line in western Eurasia.

The articler describes an archaeozoological assemblage from Late Eneolithic settlement of Gordineşti II-Stînca goală. The studied osteological material is characterized by low number of remains of wild animals, the predominant position of cattle and small cattle (with apparent importance of Capra hircus among small cattle), sparce remains of horse and apparent absence of pig in this sample. Systematical structure of the archaeozoological material from Gordinești II-Stînca goală and specific post-mortem modifications of bone remains are similar to the animal remains complexes from the described earlier sites of the Cucuteni-Tripolie Culture Horodca X and Trinca-La Șanț.

The seasonal Palaeolithic sites from the middle Dniester valley were reported as kill sites by analogy with the modern "caribou kill sites". These sites yielded abundant remains of the extinct reindeer subspecies Rangifer tarandus constantini that was present in the region under study during the coldest phase of the last glaciation. The excavation of the site of CosǍuţi (Moldova) unearthed numerous specimens of lethal weapons. The close range weapons are represented by pikes made of longitudinally split diaphysis of long bones, pikes with a dihedral tip and groove "to drain the blood" made of shed reindeer antlers. The group of medium-range weapons includes spearheads and darts made of bone, horn and, less frequently, ivory. Usually, the weapons of this type are double-ended, fusiform, 10-20 cm long, round in cross-section. The group of throwing weapons includes a harpoon made of reindeer antler with notched and serrated edges and a hole in its diamond-shaped basal part. The ability to hit targets at long distances is evidenced by the finds of bone arrowheads whose form and mass meet all the requirements associated with archery. They are divided into two types: the rod-like arrowheads with a displaced center of gravity and the leaf-shaped arrowheads with a partial stem.

The article describes fossil reindeers from main Palaeolithic sites of Moldova: Duruitoarea Veche, Brynzeni 1, Rascov 7, and Cosauti. Biometrical and statistical analyses of osteological remains revealed presence of several diff erent forms of reindeer: a larger ancient woodland form Rangifer tarandus ssp. from Duruitoarea Veche and Brynzeni 1, a smaller younger form with comparatively small teeth Rangifer tarandus cf. guettardi from Rascov 7, which superfi cially resembles Upper Palaeolithic reindeer from Western Europe, and Rangifer tarandus constantini with very large cheek teeth similar to Upper Palaeolithic reindeer from Baikal Area and Russian Plain. The author discusses the ecology, systematics and evolution of the Upper Palaeolithic reindeer, as well as the demographic composition of osteological remains from the archaeological sites included in the study.

The article presents a description and analysis of archaeozoological material from Early Medieval site Upper Saltiv (Eastern Ukraine).

Hominids of the genus Homo are believed to follow an unusual evolutionary direction of scavenger joining the guild of carnivores (Dunbar 1983; Walker 1984; Blumenschine 1987; Bunn, Kroll 1986; Bunn, Ezzo 1993; Marean 1989; Lewis 1997; Brantingham 1998; Stiner 2002; Brugal, Fosse 2004). Arribas and Palmquist (1999) proposed a model of Homo erectus dispersal in Eurasia facilitated by the commensal relationship with sabertooth cats Megantereon whitei and Homotherium latidens. However, the general ecology of scavenging and ecological constraints for a scavenger were never discussed, as well as biological and physiological constraints that can make the scavenger niche unavailable for hominids. The common meaning of the guild of scavengers implies the ecological group that includes large-sized terrestrial vertebrate animals that feed in a significant proportion on carrion and kill remains left by specialized predators. The specialized scavengers, mostly mammals (hyenas) and birds (vultures, condors, carrion-crows), represent the core of the group of scavengers. However, many animals lacking the specializations for scavenging may occasionally consume carrion if such an opportunity occurs. The availability and stability of food resource for large scavengers are defined by two basic factors: the high productivity of ecosystem and open landscape that makes the carrion visible and easily available for large animals. It means that the specialized large-sized scavengers evolve in open landscapes of tropical and subtropical latitudes, where the high ecological productivity can assure a sufficient animal biomass, first of all the biomass of herbivores, and the food object (carrion) may be easily identified from the large distance (Van Valkenburgh 1989). Indeed, the emergence of the genus Homo is bound up with the forested area decline and the extension of dry savanna in Africa during the Late Pliocene (Bobe, Behrensmeyer 2004; deMenocal 2004). However, we have first to discuss the specific physiological peculiarities of primates (and hominids in particular) before estimating the chances of archaic humans to enter the guild of scavengers. A comparatively weak sense of smell is a specific character of primates (including modern humans) that distinguish them from carnivores. A weak sense of smell in ancient humans must be a serious mal-adaptation in the competition with scavenging carnivores, which certainly are better equipped for the localization of carrion. The sharp eye for carrion identification is useful for a soaring vulture, but is of little help for a terrestrial scavenger. Therefore, the archaic Homo was inferior to specialized scavengers in localization and reaching the carrion because of their weak sense of smell and lack of effective adaptations for fast locomotion. This human mal-adaptation for scavenging niche can not be compensated by stone industry. The second physiological constraint for archaic Homo to join the carnivore guild is the human mechanism of thermoregulation. Modern humans are characterized by a very high density of sweat glands that distinguish them from the rest of primates (Sarmiento 1998). The sweat production is an important physiological adaptation that eliminates the excessive heat and prevents the body overheating (Lupi 2008). Apparently, such mechanism of thermoregulation was very effective and important in the conditions of hot dry savannas. However, this physiological adaptation of humans distinguishes them from carnivores, which lack the sweat glands. The evaporation of moisture from mouth and tongue surfaces ensures the elimination of excessive heat in carnivores. This specific way of thermoregulation in carnivores has an important adaptive significance, since the sweat must disclose the ambushing predator and significantly diminish the success of hunting. The example of modern human is quite appropriate here, since, according to the viewpoint of Brantingham (1998) and Stiner (2002), modern Homo sapiens is the result of two-million years evolution in the composition of the carnivore guild. What about ancient humans, they could be easily disclosed first of all by ambush predators. Comparatively small-bodied (30-60 kg according to Kappelman 1996) ancient humans could be an easy target for many predators of that epoch, and first of all for sabetrooth felids (Tab. 1). The available data on functional morphology of sabertooth cats suggest that those animals were ambushing predators. No doubts, the small-bodied Megantereon with short and powerful limbs was a forest dweller perfectly adapted for tree-climbing (Savage 1977; Marean 1989; Lewis 1997; Turner, Anton 1997; Christiansen, Adolfssen 2007). The data on functional morphology of Homotherium are quite controversial. Comparatively short semiplantigrade hindlimbs and shortened inflexible lumbar portion of backbone suggest that Homotherium was poorly adapted for fast running and was not able to pursuit the prey in open landscapes (Ballesio 1963; Marean 1989; Anton et al. 2005). Anton et al. (2005) supposed that H. latidens might be an open-landscape predator who compensated its low running capability by social hunting behavior. Rawn-Schatsinger (1992) and Therrien (2005) assumed that Homotherium was a cursorial predator because of its very long forelimbs. The postcranial and dental specializations of H. latidens suggest that this species was rather an ambush predator, which required a wooded environment as a necessary condition for a sudden attack and fast killing by shear-bite. The specialized dentition of sabertooth cats implies some constraints, however it perfectly adapted for fast killing of prey with minimal energy consumption (Akersten 1985; Turner, Anton 1997; Salesa et al. 2005; Christiansen 2007). Such hunting strategy must be efficient only if the prey is suddenly attacked from ambush. The long, sharp, but rather fragile upper canines are ill-adapted for a frank pursuing and fight with prey, being exposed to a high risk of damage. Most probably, Homotherium inhabited riparian woodlands and reeds. The social behaviour of Homotherium seems to be improbable, since the pack hunting requires an open environment that enables the visual interaction between members of predator group. The large body size of Homotherium (ca 160 kg) rules out the cursorial abilities. Andersson and Werdelin (2003) have noted that the adaptation to cursoriality and large body mass are two mutually exclusive directions of evolution in Tertiary carnivores. Thus, body mass of a cursorial predator rarely reaches 100 kg, and larger mammal predators are not able to develop the adaptations to cursoriality. The majority of specialized cursorial carnivores, like extant canids, hyaenids and cheetah Acinonyx jubatus are far below the 100 kg limit (Andersson, Werdelin 2003). A group of ancient humans could be a particularly easy target for the ambush sabertooth predator in riparian reeds, woodland, or closed forest, since the effectiveness of cooperative defense of humans should decrease in the conditions of limited visibility. There are no evidences that hominids were immune to the sabertooth predation or predation of other ambush carnivores. On the contrary, modern humans and even significantly larger gorillas quite often fall a victim of leopards and lions (Fay et al. 1995; Treves, Naughton-Treves 1999). Lee-Thorp et al. (2000) found that Megantereon and Panthera pardus hunted early Homo and Australopithecus robustus. The numerous remains of comparatively larger Homo erectus from Zhoukoudian (middle Pleistocene of China) show numerous tooth marks and other specific damages produced by Pachycrocuta brevirostris (Boaz et al. 2000, 2004). Therefore, Blumenschine’s (1987) suggestion that a free ecological niche in the riparian forests could be available for early Homo sounds improbable. Firstly, the ecological niche of carrion destroyers is not unfilled in the modern riparian forest ecosystems. This link of the ecological food chain is occupied by small-sized animals, mostly insects and other invertebrates, and there is a fossil evidence for that from Laetoli (Northern Tanzania). Kaiser (2000) reports the specific damage of fossil bones from Laetoli produced by, most probably, termites. Secondly, the main obstacle for large bodied scavengers in the forested ecosystems is the difficulty in carrion identification that results increased energy expenses and makes the subsistence on carrion too costly. For this reason there are no specialized large scavengers in forest ecosystems (Van Valkenburgh 1989) and this niche in the food chain is occupied by smaller animals, mostly insects. The situation must be even more difficult for early Homo, since they apparently relied upon their vision, which is less helpful for carrion search in the forest conditions than the sense of smell. Large-sized specialized scavengers evolved mainly in the ecosystems of tropical and subtropical ecosystems with high ecological productivity. Specialized scavengers are almost missing ecosystems with low productivity (probably, wolverine Gulo gulo, which shows some specific adaptations, may be the exception, but its body mass is fairly small, 10.9-18.1 kg [Pasitschniak-Arts, Lariviere 1995]), since the food resource for them is scanty and unstable. The ecological space of scavengers in the low-productivity ecosystems is occupied by facultative scavengers, such as jackals, opossums (McManus 1974), wolverines (Pasitschniak-Arts, Lariviere 1995), bears (Pasitschniak-Arts 1993), wild boars (Соколов 1959), and even leopards (Гептнер, Слудский 1972). However, the clearly defined guild of scavengers is not evolved in higher latitudes. Therefore, the proposed by Arribas and Palmquist (1999) model of hominid dispersal into the temperate climate zone of Europe because of the high quantity of carcasses produced by sabertooths is not reliable. Possibly, the productivity of Early Pleistocene European ecosystems was higher than the productivity of modern temperate zone ecosystems; however it was still lower than the productivity of Early Pleistocene African ecosystems. The ecosystem capacity for scavengers in Early Pleistocene of Europe was quite limited. The arrival of Pachycrocuta brevirostris in Europe was followed by disappearance of two smaller hyaenid species, Chasmaporthetes lunensis and Hyaena brunnea. The latter species appears again in Europe for a short period in Middle Pleistocene, but it never occurs with P. brevirostris in the same faunas (Turner 1992). Early man hardly could have more success in the competition with the lion-sized P. brevirostris than specialized and comparatively large C. lunensis and H. brunnea. I am not sure that Homo should be regarded as a true carnivore species in the strict sense of this word. Hominids’ tendency to harvest prime-adult hoofed animals reported by Stiner (1992) suggests rather an inadaptive for predator behaviour than resource partitioning with other carnivores. The human emphasis on prime-adult prey destroys the reproductive core of the prey species and causes the exhaustion of the ecological resource. It means that hominids normally are unable to achieve the “predator-prey” ecological equilibrium, unlike true carnivores. Hominids of the genus Homo must be regarded as opportunistic, omnivorous and ecologically versatile creatures, which opportunistically, of course, can consume carrion. I join the opinion of Ungar et al. (2006) that the capability to adapt in changeable and unstable ecological conditions and to switch to new food resources is the fundamental of Homo evolutionary success. The almost simultaneous arrival in Europe of two African species, Homo erectus and Megantereon whitei (Martinez Navarro, Palmqvist 1995, 1996; Arribas, Palmqvist 1999), apparently, represents two independent events caused by climate change. According to Turner (1988), the extinction of sabertooth predators in Africa was caused by extension of savanna and appearance of new species of fleet-footed hoofed mammals that were better adapted to escape in the conditions of open landscape. Indeed, the sabertooth cats with low running abilities and fragile upper canines were incapable to fight with a large prey in an open attack and were ill-adapted for a life in savanna. African carnivore guild got the modern structure ca. 1.5 Ma when the sabertooth cats became extinct because of the progressively increasing deforestization and climate aridization (Turner 1990, 1992). The carnivore guild in Europe maintained its archaic structure due to the presence of Megantereon that became extinct only shortly before Middle Pleistocene, and Homotherium that probably survived in Western Europe until Late Pleistocene (Reumer et al. 2003). The survival of sabertooth cats in Europe may be explained by the climatic-geographical peculiarities of this continent. Gulf Stream influence mitigated the climate deterioration and aridization in Europe during Early Pleistocene, particularly in its North-Western part (Brugal, Croitor 2007; Croitor, Brugal 2007). Relic forests in Western Europe served as a refugia for many Villavranchian species, mostly cervids (Croitor, Brugal 1997). Possibly, sabertooth cats were among those forest-dwelling species that survived in mild climate of Western Europe refugia. Megantereon became extinct soon during the End-Villafranchian climate shift, since this arboreal species strongly depended of dense forests that vanished by the end of Villafranchian. Terrestrial Homotherium could survive in Europe during the major part of Pleistocene in reeds near water bodies. Unlike the declining sabertooth felids, Homo erectus was a successful species adapted to open dry savanna and significantly increased its area of distribution during Early Pleistocene.

The paper presents a morphological description of the reindeer fossils from the Upper Paleolithic site of Raşcov-8. The studied reindeer has relatively enlarged cheek teeth and small body size and is referred to the fossil subspecies Rangifer tarandus constantini FLEROV 1934. The studied material from Raşcov-8 represents the earliest occurrence of this subspecies in Moldova. R. tarandus constantini substituted R. tarandus cf. guettardi on the territory of modern Moldova ca. 19,000 – 20,000 years ago. Evolution, paleobiology, and systematics of the fossil reindeer are discussed.