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Elucidating anuran accumulations: massive taphocenosis of tree frog Hyla from the Chalcolithic of El Mirador cave (Sierra de Atapuerca, Spain)
Abstract
Anurans, such as frogs and toads, are occasionally very abundant in archaeo-paleontological sites and representing >80% of the remains. These accumulations have been linked mainly to human consumption in the European context, by the preferential selection of body parts and by the presence of burning, and to a lesser extent, cut marks (as in Chalain 3, Baume d’Ogens, Kutná Hora-Denemark). However, there are also records of accumulations attributed to natural catastrophic events (Bois-Roche). This research presents a taphocenosis of microvertebrates of reduced dimensions (7.8 × 2.7 × 2.4 cm) containing 2526 bones, from El Mirador cave of Atapuerca (Burgos, Spain) contemporaneous to an adjacent Chalcolithic burial (ca. 4500 BP). The taxonomical study has identified at least 12 different taxa, which include one urodele, two anurans, seven squamates and two rodents. Tree frogs, Hyla gr. arborea (Hyla molleri), constitutes 84% of the assemblage. Quantitative and qualitative taphonomic analyses are focused on Hyla bones and show the presence of breakage and digestion marks associated with animal predation, albeit in low percentages and in light or moderate degrees. Due to its characteristics, El Mirador accumulation has been identified as a pellet accumulation produced by a medium-large owl (category 2). The nocturnal raptor hunted its preys during a wet period in spring, when the reproduction of the tree frogs occurs, showing a possible evidence of opportunistic predation on a seasonal resource. These findings help to understand the importance of anurans in the trophic chains of Prehistoric times, not only among human groups but also among non-human predators.
... Rodent and to a lesser extent bird remains recorded in scats and pellets have probably been the most studied taxa since they correspond to the main dietary components of both terrestrial carnivorans and avian raptors. Studies of other groups such as amphibians and fish remain comparatively underdeveloped because of both their low density in archaeological and paleontological contexts and because of the dynamics of these disciplines (Bisbal Chiniesta et al., 2020;Butler and Schoeder, 1998;Guillaud et al., 2017;Mignino, 2021;Nicholson, 1993;Stoetzel et al., 2012). Considering mainly the incidence and intensity of gastric acids and bone fragmentation, is broadly agreed that a distinction can be made between three broad categories of predators can be distinguished: mammalian carnivores, diurnal and nocturnal raptors (Andrews, 1990;Bochenski et al., 2018;Guillaud et al., 2018;Montalvo and Fern andez, 2019, among others). ...
... In turn, VD of Sardinops sagax (Falabella et al., 1994) was correlated with fish %MAU. Regarding anurans, we evaluated anatomical proportions based on the following indices (Bisbal-Chiniesta et al., 2020): (1) representation of postcranial and cranial elements, considering long bones (femora þ humeri þ tibiofibulae þ radioulnae x 4), divided by cranial remains (most represented cranial bone þ most represented mandibular bone x 8); (2) representation of anterior and posterior long bones, considering humeri þ radioulnae, divided by femora þ tibiofibulae; (3) representation of proximal and distal elements, considering the number of distal elements (tibiofibulae þ radioulnae), divided by proximal limb elements (femora þ humeri). For birds, we calculated the following indices: ...
... To evaluate completeness and breakage rates of cranial and long bones, we adopted a combination of methodologies following those used in several taphonomic studies on anurans, rodents and birds (Andrews, 1990;Bochenski et al., 1993;Bisbal-Chiniesta et al., 2020;Montalvo et al., 2020). For mandibular rodent elements, we employed the following categories: (1) complete, (2) mandibles with ascending ramus missing, (3) mandibles with ascending ramus missing and inferior border broken, and (4) diastema or body fragments (Andrews, 1990). ...
Taguatagua 1 is a late Pleistocene open-air archaeological site located on an ancient lakeshore in central Chile and dated to around 12,600 cal BP. It presents clear evidence of human and megafauna (Equidae, Gomphotheriidae and Cervidae) interaction that includes burned, fractured and cut-marked bones, as well as tusk and bone-made instruments. Mixed with artefacts and megafauna bones, an outstanding small vertebrate record (Class Actinopterygii, Amphibia, Reptilia, Aves and Mammalia) has also been reported, which has remained largely unstudied. In this paper, we present the first comprehensive taphonomic study of a selected sample of this ensemble. Birds are the most common taxa, followed by anurans, fish and rodents. Surface modifications, bone breakage, skeletal frequencies and the sample's ecological/biological attributes indicate different taphonomic trajectories for an averaged sample. A portion of the ensemble entered the context via predators and natural deaths, but the presence of anthropogenic marks, especially in aquatic birds, and to a lesser extent in Myocastor coypus and Calyptocephalella sp., indicates cultural exploitation of these taxa. These results portray a more precise image of the late Pleistocene hunter-gatherer subsistence strategies in central Chile, which is consistent with the settlement's lacustrine context.
... The main proxies used to draw paleoenvironmental and paleoclimatic inferences for the Holocene include pollen (Carrión et al., 1999;Jalut et al., 2000;Davis et al., 2003;Fernández et al., 2007;Ruíz-Zapata et al., 2010;Valero-Garcés et al., 2010;Pérez-Obiol et al., 2011;López-Merino et al., 2012;Expósito & Burjachs, 2016), marine cores (Johnsen et al., 1972;Shackleton et al., 2000;Cacho et al., 2001;Kageyama et al., 2005), continental cores (Pons & Reille, 1988;Moreno et al., 2007;Bernárdez et al., 2008;Morellón et al., 2009;Sancho et al., 2011), charcoal (Badal et al., 2012;Kaal et al., 2011;López-Doriga et al., 2011;Euba et al., 2016), and sediments (Benito et al., 2010;Domínguez-Villar et al., 2012;Gómez-Paccard et al., 2013;Pérez-Lambán et al., 2014). Nevertheless, in recent decades, small vertebrates have been used as paleoenvironmental and paleoclimatic proxies in studies of Europe (Stoetzel et al., 2011;Ponomarev et al., 2013;López-García et al., 2014;Popova, 2015;Berto et al., 2016), and more specifically, in studies of the Iberian Peninsula (Laplana Conesa & Cuenca-Bescós, 1995;Guillém-Calatayud, 1999;Murelaga et al., 2007;Oms et al., 2009;Cuenca-Bescós et al., 2009;Bañuls-Cardona & López-García, 2009López-García et al., 2011;Sesé, 2011;Cuenca-Bescós & García-Pimienta, 2012;Bañuls-Cardona et al., 2013, 2017aBisbal-Chinesta et al., 2020;Domínguez-García et al., 2019;Fagoaga et al., 2021). ...
... The presence of this frog is highly conditioned by temperature, environmental humidity, and rainfall (Álvarez, 2014), so it is highly susceptible to being negatively affected by aridity crises. However, the high number of Hyla frogs (66 adult tree frog individuals) suggests that the level was formed when there was also an abundance of tree frogs in the environment (Bisbal-Chinesta et al., 2020). ...
... Another aspect to consider is the seasonal accumulation of small vertebrate remains. The pellets studied from MIR5 and MIR4 (Bisbal-Chinesta et al., 2020 are interpreted as having different seasonal ranges. Based on the timing of the reproduction period of the tree frog Hyla gr. ...
This chapter summarizes the climate and environmental information that can be inferred for the Middle to Late Holocene on the Iberian plateau based on the analyses of small vertebrates from El Mirador cave. Due to their size, these small vertebrates are more sensitive than larger fauna, making it possible to register small changes that would not be noticeable with other proxies. In this study, climate variation during the Holocene is reflected in changing vegetation and biodiversity, but this diversity is also under threat from human activities, which have altered its composition and structure, reducing the species richness, and, in short, degrading it. In agricultural and livestock-rearing areas, humans plan and directly control the vegetation cover. Early Neolithic human groups also modified the landscape through these activities, but the modifications have intensified as the Holocene has advanced. These changes are reflected in the modification of forests and undergrowth in favor of open spaces and the deterioration of biodiversity from the end of the Neolithic onward, with a consequential rise in synanthropic species. This trend would increase exponentially starting in the Bronze Age.KeywordsSmall vertebratesEnvironmentClimateHuman impactHolocene
... Besides physical marks of predations in specimens (gnawing, perforations), other markers encompass taxa-level digestion, breakage, and element representation patterns that, together, can indicate species-specific predation. Additional studies have expanded on these foundational studies while considering multiple lines of evidence, from archaeological and cultural contexts, to site taphonomy and paleoenvironmental conditions [19][20][21][22][23]. Biton et al. [22] and Bisbal-Chinesta et al. [23] hypothesized that the accumulation of anuran remains in an open-air site in Israel and a Spanish cave resulted from diverse processes such as natural death, human exploitation for consumption or ritual/symbolic use, accidental human encounter, and animal predation. ...
... Besides physical marks of predations in specimens (gnawing, perforations), other markers encompass taxa-level digestion, breakage, and element representation patterns that, together, can indicate species-specific predation. Additional studies have expanded on these foundational studies while considering multiple lines of evidence, from archaeological and cultural contexts, to site taphonomy and paleoenvironmental conditions [19][20][21][22][23]. Biton et al. [22] and Bisbal-Chinesta et al. [23] hypothesized that the accumulation of anuran remains in an open-air site in Israel and a Spanish cave resulted from diverse processes such as natural death, human exploitation for consumption or ritual/symbolic use, accidental human encounter, and animal predation. Blain and Villa [19], Blain [20], and Martin et al. [21] also explored the taphonomic processes responsible for anuran accumulation in archaeological deposits, but complement their interpretations with discussions on paleoclimatic and paleoenvironmental conditions. ...
... As taphonomic studies of anuran remains the world over show, one of the principal taphonomic agent for introducing these bones into archaeological assemblages are the natural predators of frogs/toads [17][18][19][20][21][22][23]61,62]. In Moquegua, identifying one of the probable taphonomic agents that produced the accumulations of frog and toad skeletons described here was largely serendipitous. ...
We examine the processes that resulted in the deposition of bones of at least three anuran genera on four archaeological sites associated with the Tiwanaku culture occupied between 700–1100 CE in the Moquegua Valley of far southern Peru. We review archaeological data and ethnographic accounts of Andean peoples using frogs and toads for food and in rain-quest rituals. Anuran bones are common in prehispanic cemeteries, but far less common in habitational and ceremonial sites. The quantity of anuran remains in certain cemeteries is anomalous in comparison to other archaeological sites in the valley and to Tiwanaku sites in other geographic settings. We conclude that anurans are most common where abandoned below-ground rock-covered tombs are likely to have been reused by nesting owls, and propose that most anuran remains in archaeological contexts in Moquegua are the result of predation. We consider environmental, cultural and taphonomic explanations and posit that the abundance of anuran remains relates to the 14th-century Miraflores ENSO event. This event generated increased rainfall in the desert, creating conditions favorable for frogs and toads, and predation by owls. We also advocate for the use of fine-screening to recover small-sized animal remains, such as anurans, that can be used to understand taphonomic processes and paleoenvironmental conditions.
... Nonetheless, the main factor accumulating microvertebrate assemblages in both, modern and fossil sites is predation (e.g. Andrews, 1990;Bisbal-Chinesta et al., 2020;Fernández and Pardiñas, 2018;García-Morato et al., 2019a;Lloveras et al., 2014;Marin-Monfort et al., 2019. Different factors such as the size ratio of the predator/prey relationship, diurnal or nocturnal hunting behaviour, hunting strategy and territory size may vary from one predator to another, and this will be reflected in the species represented in the small vertebrate assemblages (Andrews, 1990). ...
... Palaeoenvironmental analyses from El Mirador cave have been carried out through a multiproxy approach (zooarchaeological, microvertebrate and pollen studies), providing a comprehensive view of the evolution of the landscapes surrounding the cave (e.g. Bisbal-Chinesta, 2020;Bisbal-Chinesta et al., 2020;Expósito et al., 2017;Martín et al., 2016;Vergès et al., 2016). Both, small mammals and palynological data have provided evidence of the intensification of anthropization in the area. ...
The regional climatic context in which Chalcolithic (MIR5) and Bronze Age (MIR4) levels from El Mirador cave (Sierra de Atapuerca, Burgos, Spain) are framed is affected by the 4.2 ka cal. BP event, a global event defined as a cooling and aridification phase. Previous works based on palaeoenvironmental inferences indicate conflicting results regarding the possible impact of the event on vegetation and small mammals from MIR5. Pollen record illustrates a possible aridification episode that could match with the 4.2 ka cal. BP event, while the signal of this event is not clearly recorded in the small mammal assemblage, which indicates more humid environmental conditions than pollen record. Taphonomic analyses confirmed that the small mammal assemblages from MIR4 and MIR5 are the result of predation, supporting the involvement of European eagle owls (Bubo bubo) in its formation. This avian raptor shows a marked preference for hunting animals living in the more open and wetter parts of their hunting range. Likewise, spontaneous specialisation on abundant prey species could also be observed under certain environmental conditions. This characteristic behaviour of eagle owls may have provided the contradictory results observed between the small mammal assemblage and palynological evidence. Nonetheless, taphonomic analyses also provided information about climatic conditions and fluctuations along time. The low incidence of manganese coatings and carbonate crusts deposits in small mammal bone remains from MIR5 support the presence of arid conditions during the formation of this level, which agreed with the aridification phase probably related to the 4.2 ka Bond Event inferred by palynological data from MIR5. These results provided a more robust conclusion about the paleoenvironmental contexts during the formation of Chalcolithic and Bronze Age levels at El Mirador cave.
... Ortiz et al. (2012a) proposed that the assemblage was formed in a short lapse (i.e., low time averaging), which is suggested by the high concentration of bones in a very small area and the thinness of the fossiliferous level and because this assemblage was produced by strigiform birds. We hypothesize that the short lapse in which these deposits were produced corresponded to late spring, a time when many juveniles would be available prey for owls (Bisbal-Chinesta et al., 2020). We base this suggestion on the fact that at this time of the year, in temperate areas, where the fossil comes from, Rhinella larvae with early reproduction begin to metamorphose and it is very common to find large numbers of juveniles abandoning the streams (MLP, pers. ...
... Sánchez et al. (2008) reported that juvenile amphibians are eaten whole, whereas body parts of adults, such the skin and the head, are discarded. In addition, the biggest account of the hind parts of the frogs' body could be related to its greater meat supply (Bisbal-Chinesta et al., 2020). This could explain the pattern observed in the fossil sample. ...
The best known Neotropical fossil bufonid discovered to date is described herein. This material includes well-preserved cranial and postcranial disarticulated remains that were part of a microvertebrate fossil assemblage generated by the accumulation of predatory bird pellets. The remains were recovered from the Middle Unit of the Uquía Formation (late Pliocene) that crops out next to the locality of Humahuaca, Jujuy province, northwestern Argentina. In this contribution, we present a detailed osteological description of this taxon and compare the morphological characters with respect to other related members of the large genus Rhinella. Several distinctive character states present in these fossil remains allows us to refer to them as part of a new species of the Andean R. spinulosa Group. The new species described here together with the fossil marsupials and sigmodontines suggest that by the late Pliocene, the Uquía area, the Andean region of Bolivia, and southern Peru were part of the same biogeographic unit.
... Por otro lado, si bien las investigaciones tafonómicamente orientadas de huesos de anfibios en contextos europeos y africanos han sido desarrolladas (Pinto-Llona & Andrews, 1999;Blain et al., 2008;Stoetzel et al., 2012), los estudios actualísticos sobre acumulaciones por causas no antrópicas son aún escasos (Bisbal-Chinesta et al., 2020). ...
... Sobre estos, se analizó la acción digestiva y sus efectos, como astillamiento, redondeamiento, adelgazamiento, colapso y descamación. Las categorías empleadas para estudiar las modificaciones por digestión siguieron las pautas de Bisbal-Chinesta et al. (2020). Por ello, se clasificaron como leves (pequeños agujeros en articulaciones), moderadas (aumento de tamaño de orificios y exposición parcial de tejido esponjoso interno), y fuertes (pérdida parcial de región articular), y se incluyó la categoría extrema cuando existió pérdida total de región articular por corrosión. ...
Se presenta un análisis tafonómico comparativo de restos de anfibios recuperados en egagrópilas de aves rapaces Strigifomes provenientes del centro de Argentina, en la provincia de Córdoba: Tyto furcata, Megascops choliba y Athene cunicularia. El estudio se basó en el promedio de abundancia relativa de elementos anatómicos, índices de representatividad de partes esqueletales y se evalúo fracturación y digestión. Los resultados derivados del análisis de representatividad y abundancia anatómica revelaron promedios elevados de abundancia relativa en todos los casos. Los grados de fractura y digestión fueron evaluados como leves para T. furcata y M. choliba y moderados para A. cunicularia. En general, estas evaluaciones coinciden con las referencias previas. Sin embargo, se detectaron variaciones en los patrones de fracturación y digestión. Los datos presentados aportan información como análogos modernos para evaluar la presencia de esta clase de restos en contextos arqueológicos y paleontológicos.
A comparative taphonomic analysis of amphibian remains recovered from Strigiforms raptors pellets from central Argentina in the Córdoba province is presented: Tyto furcata, Megascops choliba and Athene cunicularia. The results derived from the analysis of the representativity and anatomical abundance revealed high percentages of relative abundance in all cases. The degrees of breakage and digestion were evaluated as light for T. furcata and M. choliba and moderate for A. cunicularia. In general, these evaluations are consistent with previous references. However, variations in breakage and digestion patterns were detected. Data presented provide information as modern analogs to assess the presence of this class of remains in archaeological and palaeontological contexts.
... Finally, we conducted a preliminary taphonomic study of the reptiles and amphibians of SU V to discern the agent(s) responsible for the fossil accumulation and to verify the degree of reliability of the information provided by the fossil assemblages. Our taphonomical approach is based primarily on Pinto Llona and Andrews (1999), Stoetzel et al., 2011, Lebreton et al. (2020), and Bisbal-Chinesta et al. (2020a. These authors have studied skeletal representation, as well as bone fragmentation and modification. ...
... Although taphonomical studies of micromammals are common, those addressing herpetofauna are scarce (Lyman, 1994;Bailon, 2011). In fact, papers exclusively focused on amphibian and/or reptilian taphonomy are extremely rare (Pinto Llona and Andrews, 1999;Stoetzel et al., 2011;Bisbal-Chinesta et al., 2020a); they have usually been incidental to other studies (i. e., Castillo et al., 2001). ...
The locality of El Salt (Alcoi, Spain) is a key site for understanding the extirpation of Neanderthals in the eastern part of Iberia. In this paper, we analyse an assemblage of amphibians and reptiles from Stratigraphic Unit V (45.2 ± 3.4 ka to 44.7 ± 3.4 ka), which corresponds to one of the last regional records of Neanderthals, to improve knowledge of the palaeoecology and palaeoclimate of this event. The assemblage comprises three anurans (Pelodytes sp., Alytes obstetricans, and Epidalea calamita), two lizards (Lacertidae indet. and Chalcides bedriagai), and five snakes (Colubridae indet., Coronella sp., Coronella sp./Zamenis sp., Natrix maura, and Vipera latastei). Palaeoclimatic reconstruction, based on the Mutual Ecogeographic Range method, indicates that climate was cooler and slightly wetter climate than the present day climate of the Alcoi area. Applying the Habitat Weighting Method, we infer that the area surrounding El Salt was dominated by open dry regions, alternating with rocky areas with scarce scrubs and forest patches that would have developed under mesomediterranean conditions. These results are not entirely consistent with those obtained with other proxies (charcoal and small mammal assemblages) from the same site, which suggest slightly warmer and drier conditions. We hypothesise that these divergences may be partly related to the current wide distribution of reptiles and amphibians across Iberia. A key finding is that the extirpation of the Neanderthals in Iberia coincided with aridification.
... Limb bones are scarce (S4 Table). If [59] indexes are considered, the lower level is characterized by a marked predominance of cranial bones (0.28), more anterior than posterior limb bones (1.16) and more distal than proximal ones (1.16). In fish, three individuals were identified. ...
We present the results of the excavations and analyses of the diverse and exceptional archaeological assemblage of Taguatagua 3, a new late Pleistocene site located in the ancient Tagua Tagua lake in Central Chile (34°S). The anthropogenic context is constrained in a coherently dated stratigraphic deposit which adds new information about the mobility, subsistence strategies, and settlement of the early hunter-gatherers of southern South America. The age model constructed, as well as radiocarbon dates obtained directly from a combustion structure, indicate that the human occupation occurred over a brief time span around 12,440–12,550 cal yr BP. Considering taphonomic, geoarchaeological, lithic, archaeobotanical, and zooarchaeological evidence, as well as the spatial distribution combined with ethnographic data, we interpret Taguatagua 3 as a logistic and temporary camp associated mainly with gomphothere hunting and butchering. Nevertheless, several other activities were carried out here as well, such as hide and/or bone preparation, small vertebrate and plant processing and consumption, and red ochre grinding. Botanical and eggshell remains suggest that the anthropic occupation occurred during the dry season. Considering the contemporaneous sites recorded in the basin, we conclude that the ancient Tagua Tagua lake was a key location along the region’s early hunter-gatherer mobility circuits. In this context, it acted as a recurrent hunting/scavenging place during the Late Pleistocene due to its abundant, diverse, and predictable resources.
... Unfortunately, considering the high amount of data published for the Late Pleistocene sites, the Holocene sequences bearing both small vertebrates and molluscs are still poorly investigated. Although several works have been published to cover this gap, they mainly cover Alpine and Mediterranean Europe (López-García et al., 2013;Berto et al., 2016;Bañuls-Cardona et al., 2017;Bisbal-Chinesta et al., 2020). ...
The Holocene has always been considered a crucial epoch where the major cultural steps of humankind took place. Understanding past Holocene climatic variability, shifts in vegetation, and faunal communities are among crucial challenges in predicting the upcoming changes of the natural environment.
In Central Europe, the Kraków-Częstochowa Upland is one of the best study areas for detecting faunal, environmental, and climate shifts. For that reason, we choose to study the small vertebrate and mollusc communities of three archaeological sites with Holocene horizons: Sąspowska Zachodnia Cave, Ciasna Cave, and Małe Rockshelter.
We analysed the faunal community shifts, and we compared our results with the major Holocene sites in the area with the correspondence and cluster analyses. The environment and climate were reconstructed through Habitat Weighting, Quantified Ecology, malacological spectra and Bioclimatic methods.
Environment and climate values showed a temporary replacement of the Late Glacial tundra and grasslands with boreal forest mixed with open wetlands during the Preboreal and then stable landscapes dominated by temperate forests. Models based on rodent data show that temperature and precipitation generally increased, starting from the Boreal, to reach values near, or sometimes overwhelm, the current climate values of the area during the Atlantic and Subboreal. The climate values obtained from herpetofaunal analyses show very slight changes between the Preboreal and the Atlantic (Ciasna Cave layers 3 and 4), except for the winter temperatures, whereas results obtained for Sąspowska Zachodnia Cave are probably biased by a very low number of recovered remains. Among rodents, a major negative oscillation detected in layers related to the edge of Early Holocene was related to the Preboreal Oscillation or long-lasting effect of the Younger Dryas.
... Each bone was examined under a light microscope at 10× to 40× magnification for surface modifications caused by abiotic and biotic processes, such as carnivores, raptors, and human predation [29]. Surface modification in anurans was inspected following literature [30][31][32]. Actualistic taphonomic studies directed toward raptors preying on anurans and squamates were also consulted [12,[33][34][35][36]. For the snake vertebrae alterations, we followed the five digestion categories as published and well-illustrated by Lebreton et al. [36] that were available to us for comparison. ...
During the Natufian period, more than 12,000 years ago, Eynan (Ain Mallaha) was an important human settlement in the Hula Valley, Israel. This study concentrates on the anuran and squamate assemblage from the ultimate stage of the Natufian period at the site, the Final Natufian. Over five thousand bones assigned to at least sixteen taxa were studied from a sampled segment of the excavated open-air site. Relative species abundance, spatial distribution, taphonomic observations and ecological considerations all pointed to the conclusion that the inhabitants of Eynan intensively exploited three large “colubrine” snakes species: the Large Whip Snake (Dolichophis jugularis), the Eastern Montpellier Snake (Malpolon insignitus) and an Eastern Four-lined Ratsnake (Elaphe cf. sauromates). These snakes were the most desired and were intensively gathered, while other snakes and lizards could have been opportunistically collected when encountered. We raise questions about whether the large “colubrines” exploitation should be interpreted as additional evidence of increasing diet breadth. We suggest challenging this line of reasoning and offer possible alternative motives.
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El Mirador cave has become a reference for the study of the economic, social organization and funerary behavior of first farming communities in the Iberian Peninsula. In this chapter we put into context the set of studies carried out to date and presented in this monographic volume. The Neolithic occupations of El Mirador cave are among the earliest in the northern Meseta, dating from the 6th millennium cal BCE. These groups developed a consolidated agricultural and livestock economy that underwent a progressive intensification that is reflected in the development of cultural landscapes since the Neolithic. The cave was used as a sheepfold and also as a domestic space. Agricultural and livestock products formed the basis of the diet and were processed and consumed using ceramic and lithic remains found in the site. Late Neolithic and Chalcolithic human groups used the cave as a burial space. Funerary practices developed during the Late Neolithic and Chalcolithic were diverse, with cannibalism being practiced and bodies being buried both individually and collectively. Genetic studies indicate that these practices were carried out by a large group, which went beyond the family sphere.KeywordsEarly NeolithicHuman migrationsChalcolithicFunerary practicesHusbandryAgricultureFarming
The archeobotanical research focusing on El Mirador cave (Sierra de Atapuerca, Spain) has contributed to the reconstruction of the landscape in which human communities lived during the Neolithic and Bronze Age in the surroundings of the Sierra de Atapuerca. The analysis of the palynological and anthracological record has revealed a landscape of mixed forest with evergreen and deciduous oaks and pinewood. The evolution of the vegetation reflects a progressive trend towards greater aridity. This decrease in rainfall occurred parallel to the progressive increase in anthropic pressure indicator plants, leading towards gradual deforestation. In addition to illustrating the gradual opening of the forest formations around the settlement, the analysis has evidenced many of the anthropogenic features related to human interference throughout the sequence, such as an increase in the representation of heliophilic trees and shrubs, the development of anthropo-zoogenic pasture areas, the proliferation of nitrophilic plants and the increase in herbaceous and shrub communities typical of degradation phases. This progression towards an increasingly modified landscape from the Neolithic to the Bronze Age implies a decisive change in the relationship between the environment and the human groups that inhabited it.KeywordsAnthracologyPalynologyPollenNon-pollen palynomorphsCultural landscapeAnthropogenic impactHolocene
The dental and oral pathologies of the Chalcolithic and Early Bronze Age herders from El Mirador cave indicate good general oral health in both populations as deduced from the low prevalence of pathologies such as dental caries, dental calculus, linear enamel hypoplasia, periodontal disease and abscesses. In both groups, older individuals exhibit the highest prevalence and severity of these age-progressive pathologies. The ratio of dental caries and calculus in both populations is associated with a diet based primarily on proteins of animal origin complemented with carbohydrate-rich plant-based foods. The main difference in the typology of dental caries between the groups illustrates a shift in the way carbohydrates were processed. The Early Bronze Age group consumed softer and less fibrous plant items than the individuals from the Chalcolithic group.KeywordsBucco-dental pathologiesHealth statusHerder groupsChalcolithicBronze Age
El Mirador cave (Sierra de Atapuerca, Burgos) is one of the most significant sites for understanding the evolution of agricultural and livestock farming groups on the Iberian Peninsula. Its long sequence spans from the Early Neolithic (second half of the 6th millennium cal BCE) to the Bronze Age (second half of the 2nd millennium cal BCE). During this time, the cave was used as a sheepfold and burial space, providing a great deal of information on the economic activities of these groups, their social organization and their symbolic and/or funerary manifestations. This chapter presents the site with a description of its location, a historiographical overview of the research, and details of the excavation process and of the different methods used. It also includes a description of the stratigraphic succession, as well as a list of the radiocarbon dates currently available.KeywordsHistoriographyExcavation techniquesSampling methodsStratigraphyChronology
Neanderthals have been claimed to have had a selective adaptation to rugged, wooded landscapes that would have partially compensate their high basal metabolic rate and locomotor energetic costs through reducing search time and increasing diet breadth. The archaeological site of the Navalmaíllo rockshelter (Pinilla del Valle, Madrid), located in a mountain environment in central Iberian Peninsula, has been interpreted as a repeated short-term occupation hunting camp by Neanderthal groups, mainly focused on primary access to large bovids and cervids. Navalmaíllo rockshelter has also furnished fish, toads, frogs, tortoises, lizards and snakes remains. This association of amphibians and reptiles suggests a much warmer climate (þ2.8 C) for layer F than at present, with similar temperatures during the summer but higher temperatures throughout the remaining months. Rainfall was slightly more abundant (þ180.6 mm) than today, with a more contrasted regime during the year, with rainier winters and drier summers. A two-month period of aridity is observed during summer, representing a similar duration to present-day climatic conditions. Reconstructed landscapes are mainly constituted by open areas with dry grassland and rocky or stony areas, evolving laterally to humid meadows, probably close to the aquatic and peri-aquatic areas, such as rivers or ponds with riverside vegetation. Woodland environments are also well represented around the site, with medium scrublands to forest formations. Fish complete this reconstruction by documenting the presence of a pre-mountain well-developed river system characterized by relatively cold, permanent, oxygen-rich, and running waters. Such reconstruction is in disagreement with previous pollen interpretation for Layer F that suggested a very open and cold environment. Our new interpretation suggests that the Neanderthal occupation of the Navalmaíllo rockshelter occurred during a somewhat temperate and humid period, probably within the later part of the Marine Isotope Stage 5, effectively favouring the presence of a high biodiversity around the site.
La Lechucita de las Vizcacheras (Athene cunicularia) es un ave Strigiforme muy conspicua en las planicies de Sudamérica. Se caracteriza por ser una especie estacionalmente oportunista que consume una amplia diversidad de vertebrados y artrópodos. En este trabajo se evalúa la variabilidad estacional en la dieta de este predador y se brinda una caracterización tafonómica de los restos ingeridos y de carcasas de anuros que ayudará a identificar al predador en contextos fósiles. Los trabajos de campo se llevaron a cabo en un sector periurbano de la ciudad de Olavarría, provincia de Buenos Aires. Se recolectaron semanalmente las egagrópilas y restos no ingeridos hallados en asociación a cuatro nidos de lechucita. Los resultados obtenidos muestran que, entre los vertebrados, las presas principales en invierno fueron los roedores y en verano los anuros. Esta especie de Strigiforme produce modificaciones leves a moderadas sobre los especímenes óseos de anfibios. Entre los efectos digestivos identificados predominan el pulido, el adelgazamiento y astillamiento.
A fossil gastric pellet from the Danian 'Banco Negro Inferior' of the Salamanca Formation at Punta Peli-gro Locality (Chubut, Argentina) and its 3D preserved fossil content is studied herein. The structure of the pellet and the condition of the enclosed bones suggest that it was produced by a bird of prey, although birds of any kind are as yet unknown from osteological remains in the Banco Negro ecosystem. The content of the pellet originated from a single anuran individual, representing a new species of the genus Calyptocephalella, is described herein as C. sabrosa sp. nov. The new find highlights the broad temporal, geographical and taxonomic diversity of this frog genus in Patagonia's geological past.
Most small terrestrial vertebrate accumulations in archaeological and palaeontological sites result from
predation but we are far from having an exhaustive knowledge of modern predators’ diet, ecological
niches and bone modification patterns especially in North African sites. The few neotaphonomic
referentials available result from taxon-specialized palaeontologists’ initiatives. A survey of the literature
on North Africa predators shows that their prey diversity is high and not only include rodent and shrews
but also amphibians, squamates, bats and insects. We performed here a pilot taphonomic study of a
Moroccan Tyto alba nest pellets including the whole taxa consumed (birds, rodents, shrews, amphibians,
insects). We analyzed bone representation, fragmentation and digestion for each taxa and then compared
the results. We observe differences between the taxa but on the whole find higher modification levels
for this assemblage compared to roost sites of barn owls from other sites. We used for the comparisons
homologous bones like the femur and the humerus and also found differences between the taxa. For
example, digestion was lower on amphibians and birds than on micromammals. We discuss here some
methodological issues as well as archaeological and palaeoenviromental ones by comparison with the
Pleisto-Holocene site of El Harhoura II (Morocco).
The human impact on the environment in the Holocene has usually been characterized on the basis of palaeobotanical records, but attempts to distinguish the anthropogenic impact from natural events in landscape evolution have been the subject of much debate in recent years. The aim of this paper is to analyse small-mammal diversity and the presence of synanthropic species, whose small size makes them more sensitive to any changes in their environment that may occur. This study has allowed us to characterize palaeoclimatic and palaeoenvironmental changes, recording small changes whether resulting from a human influence or otherwise. Our object of study is El Mirador cave, which has a sequence with a well-documented human occupation extending from 7200 to 3000 cal. BP. The study has led us to differentiate two phases. In one phase, we can see small changes in diversity related to climatic oscillations from ca. 7200 to 6800 cal. BP, while in the second phase, lasting from ca. 6800 to 3000 cal. BP, the changes in diversity and in the assemblage of synanthropic species are associated with human economic strategies. Moreover, we distinguish which kinds of economic activity (crop and livestock farming) have influenced these changes, because some small-mammal species are influenced, positively or negatively, by environmental changes based on crop farming and animal husbandry. All this information is contrasted with other archaeological proxies, such as the large-mammal and palaeobotanical assemblages from El Mirador cave. Furthermore, this integrative analysis has made it possible to identify the existence of altered environments more generally throughout the Iberian Peninsula from ca. 6000 cal. BP. It additionally confirms the theory of low human occupation intensity in the northern Meseta and in high mountainous areas during the early Neolithic.
La gran diversidad en pequeños mamíferos del Pleistoceno superior y Holoceno de la Cueva del Mirón, con un total de 26 especies pertenecientes a seis órdenes de mamíferos (Erinaceomorpha, Soricomorpha, Chiroptera, Carnivora, Rodentia y Lagomorpha) hace de la secuencia del Mirón un importante documento sobre la distribución de las asociaciones de pequeños mamíferos durante la transición Pleistoceno-Holoceno; lo que nos permite conocer la evolución del clima y del paisaje en la Región Cantábrica. Al final del Cuaternario se registra la última extinción de microfauna Pleistocena, como la del topillo Pliomys lenki, relacionado con el actual Dinaromys, el topillo nival de los Balcanes o de Martino. También se documentan extinciones locales, como la desaparición del territorio español de los mamíferos característicos de las estepas del Norte de Europa y Siberia, como Microtus oeconomus y M. gregalis. Estas extinciones se pueden explicar por los cambios climáticos ocurridos al final del Cuaternario, sin descartar la competencia con especies inmigrantes, de ecología similar. El final del Holoceno, la actividad humana, sobre todo la deforestación y la cultura cerealística, se refleja también en la disminución de la diversidad específica de los pequeños mamíferos.
Humeri of 14 tailed amphibian species are described: Onychodactylus fischeri (BOULENGER, 1886), Salamandrella keyserlingii DYBOWSKI, 1870, Salamandrella tridactyla (NIKOLSKII, 1905), Ichthyosaura alpestris (LAURENTI, 1768), Lissotriton lantzi (WOLTERSTORFF, 1914), Lissotriton montandoni (BOULENGER, 1880), Lissotriton vulgaris (LINNAEUS, 1758), Ommatotriton ophryticus (BERTHOLD, 1846), Pleurodeles waltl MICHAHELLES, 1830, Triturus cristatus (LAURENTI, 1768), Triturus dobrogicus (KIRITZESCU, 1903), Triturus karelini (STRAUCH, 1870), Mertensiella caucasica (WAGA, 1876), Salamandra salamandra (LINNAEUS, 1758). Morphological characteristics and indices (ratios) almost always permit the distinction of tailed amphibian genera. Specific identification is more difficult because interspecific differences are insignificant.
Des milliers de restes de pélodytes ponctués ont été découverts dans la grotte de Bois-Roche (Pléistocène supérieur, Charente). Les caractéristiques taphonomiques de cet assemblage nous conduisent à émettre l'hypothèse d'une accumulation produite par une mortalité catastrophique en masse par noyade. Cet exemple démontre que la prédation n'est pas le seul processus responsable des taphocoenoses d'anoure à forte densité. Abstract The Bois-Roche cave site (Upper Pleistocene, Charente, France) has yielded thousands of remains of « Parsley Frogs » (Pelodytes punctatus). The taphonomic characteristics of this assemblage led us to put forward the hypothesis of an accumulation produced by a catastrophic mass mortality. All parsley frogs were drowned. This example demonstrates that the predation is not the only process responsible of anuran taphocoenosis with a high density. Mise en évidence d'une accumulation de batraciens par mortalité catastrophique en masse
Reptiles and frequently amphibians can be located inside cavities. Its presence is
voluntary in some cases and they use the hypogeum environment as a refuge. However, these animals are
trapped ocassionally without possibility to go outside and they die because of inanition, hypothemia or
dehidration. This article generally describes which aspects are necessary to differentiate if it is a specimen
using the cavernicola environment as a refugeor or if it is an animal which is trapped and will die. It also show
both specimenes located commonly in cavities and more general recomendations about their manipulation,
transport, recognition of their state and when and where to release if possible.
Las excavaciones arqueológicas iniciadas en 1999 en la cueva de El Mirador de la sierra de Atapuerca han permitido documentar hasta el momento una sucesión estratigráfica holocena de unos 2,5 m de potencia, formada por niveles del Neolítico y de la Edad del Bronce. Los datos preliminares sobre las ocupaciones de la Edad del Bronce señalan el uso de la cueva como redil, zona de hábitat y espacio sepulcral. Durante su uso como redil se llevó a cabo la quema periódica del estiércol depositado en el interior del corral. Es la primera vez que dicha práctica se documenta en la Meseta y, en el ámbito europeo, en contexto geográfico continental. El uso como cueva sepulcral viene seña- lado por la presencia de una inhumación colectiva. Ésta presenta evidencias de un ritual relacionado con el tratamiento de los cadáveres previo a su enterramiento. Cabe subrayar también la presencia de una hacha de bronce de rebordes en contexto estratigráfico.
We performed a comparative osteological analysis of middle trunk vertebrae (represented by V6) of representative species of all living genera of Salamandridae (Amphibia, Caudata). The qualitative morphological characters used were adapted from the traditional palaeontological literature; using this data set we inferred a phylogenetic hypothesis for the family. The same morphological matrix was then re-analysed using a weighting scheme for the characters derived from the answers of a psychological test taken by an international group of graduate students unfamiliar with palaeoherpetology. We compared the phylogenetic results of both groups with the currently accepted evolutionary model for this family, which is based on mitochondrial and nuclear gene sequences. The ranking of the relative (and subjective) conspicuity of vertebral structural units (prezygapophyses, neurapophyses, etc.) collectively made by the inexpert group, presumably, directly derives from a general (human) capability to recognise shapes. The same perceptive pattern also seems to be involved in the character set developed by the specialists, and both matrices obtained similar results in the quality of their respective phylogenetic inferences. Defining characters is the most important step in systematics and, therefore, we stress the importance of developing new tools and approaches for exploring new quantitative and qualitative characters in palaeontological research.
This article undertakes a palaeoenvironmental and palaeoclimatic reconstruction of the Middle Bronze Age on the
basis of a study of the small mammals from level MIR 4 of El Mirador Cave. The chronology of the level under study is
3,720-3,140 cal. yr BP. The palaeoenvironmental analysis indicates that it represents the period of transition between the
Subboreal and the Subatlantic, during which prevailed a very humid habitat, dominated by woodland and wet meadows.
Otherwise, the palaeoclimatic analysis, carried out using the Mutual Climatic Range method (MCR), reveals that level MIR
4 falls within an interval in which the temperatures were very similar to present-day ones, while mean annual precipitation
was notably higher than at present, i.e. 485 mm above the present level for Burgos.
Several Neolithic to Iron Age sites of the Mediterranean region contain archaeological sediments, called fumiers, which are composed mainly of burnt animal dung and vegetal remains, and are commonly interpreted as the product of pastoral activities. Here we address three main topics about these sediments, which occur almost exclusively in the entrance areas of karstic caves and rock-shelters: their characteristics; methodological aspects of their excavation and study; and their archaeological interpretation. For such purposes, we briefly review the information available about Neolithic fumiers and present the first data from the sites of El Mirador (Burgos, Spain) and Riparo Gaban (Trento, Italy).
During a peak in the reproductive season, males called from the edge of a pond and occasionally engaged in interactions involving aggressive calling and non-physical displacement of other calling males. Some silent males were also present in the chorus. Amplexus occurred only after females approached calling males and made physical contact with them. Calling males were significantly larger than amplectant males, and amplectant males were significantly larger than a random sample of males. Significant positive size-assortative mating was found for the 45 amplectant pairs measured. The dominant frequency of the advertisement call was significantly correlated with male size. The large-male mating advantage observed may be a result of different strategies (calling versus non-calling) adopted by males of different sizes. A mechanistic explanation based on the ontogenetic changes in female sensitivity to auditory frequency may explain the size-assortative mating. -from Authors
This project aims to study fossil amphibian and reptile (herpetofaunal) remains from Quaternary sites in the British Isles. This neglected group of vertebrates hold great potential for Quaternary Science. Collectively, they cover a wide range of ecological tolerances, although individual species often have very specific tolerances. The biology and ecology of individual species are discussed (Chapter 2) to facilitate their use in Quaternary palaeoenvironmental reconstructions, and an account of previous work on fossil herpetofaunas is given (Chapter 3). Very little work on fossil herpetofaunas has been carried out in the British Isles, mainly due to a lack of the required osteological expertise. The preparation and study of a modern osteological collection (Chapter 4), for comparative purposes, has therefore constituted a large and essential part of the project. The resulting manual for the identification of fossil herpetofaunal remains, appropriately illustrated with SEMs and hand-drawn figures, is presented (Chapter 5). The difficulties encountered in identifying some taxa are discussed in detail, and points of caution are stressed where necessary.
Amphibian and reptile remains found in archaeological contexts are still poorly studied, especially in North
Africa. This article presents the first taphonomic analysis realised on amphibian and squamate remains
coming from a North-African quaternary site. The bones were not transported/sorted by water; they were
quickly buried and no significant perturbation occurred within deposits. The main origin of the amphibian
and squamate assemblages is predation, although some animals probably died in the cave from natural
causes. A comparison with taphonomic data from small mammals of the same cave gave more accurate
taphonomic and reliable palaeoecological interpretation of the site. This study also highlights the numerous
problems of interpretation linked to the lack of taphonomic referentials based on predator pellets/scats and
on weathering/burying experiments in North Africa.
Small vertebrates found in archaeological context can provide important information on the evolution of biodiversity and paleoenvironments of a precise geographical region and sometimes on a large interval of time. However, very few studies are specifically dedicated to small fossil vertebrates in north Africa, specially for the end of the Quaternary period. The Late Pleistocene to Middle Holocene succession of El Harhoura 2 cave, situated in the region of Te´mara (Morocco), has revealed an exceptional richness of small vertebrates’ remains, as well as in bones abundance than in species diversity (rodents, shrews, hedgehogs, amphibians, chelonians and squamates). We present here an annotated preliminary axonomic list of this material.
The skull and dentition were analysed and compared in a number of species of Old World scincid lizards, genus Chalcides (including Sphenops). The major differences are related to the decrease in absolute skull size, which entails an increase in the relative size of the otic capsule and the progressive obliteration of the supra- and post-temporal fossae. The greatest reduction of the post-temporal fossa is observed in the smallest species, resulting
in close proximity of parietal and supraoccipital bones and in a possible reduction of metakinesis. Another major difference concerns the relationships between pre- and post-frontal bones, widely separated in Chalcides sensu stricto and articulated in the taxa formerly attributed to the genus Sphenops. This close connection likely represents an internal reinforcement preventing deflection or fracture in these sand-swimming taxa by reducing the mobility of the fronto-parietal mesokinetic joint. The teeth are conical, pleurodont and ankylosed to the inner margin of the dentary, pre-maxilla and maxilla. They exhibit two cusps separated by a single groove that probably aid in grasping hard-bodied
prey. The pre-maxillae usually bear an odd number of teeth; the number of pre-maxillary, maxillary and dentary teeth does not seem to increase with skull size. In all cases, the number of dentary teeth exceeds that of maxillary ones.
The present paper is a study of fossil amphibians and squamate reptiles found in 34 Spanish localities, dated from the Upper Pliocene to the Middle Pleistocene. This work permits to supplement the knowledge of the settlements of amphibians and squamate reptiles in Spain during the Quaternary and to define the contribution of these faunas to the reconstitution of the landscape and the climate. During the Late Pliocene, several thermophilous groups of squamates will gradually disappear from the Iberian Peninsula following successive climatic pejorations: the Eastern Vipers and the genus Pseudopus, during the MN16, in Cova Bonica (Barcelona), then the Agamidae, at the end of the MN17 in Almenara-Casablanca-1 (Castellón) and Vallirana (Barcelona). Only the genus Dopasia will persist until the end of the Early Pleistocene (MmQ3) in the South of Spain in Barranco León (Guadix-Baza). At the same time, the observed variations within hygrophilic species suggest a peak of aridity at 2.0 My, and an increase in moisture
between 1.3 and 1.0 My in the localities of Barranco León and Fuente Nueva 3 (Guadix-Baza), of Almenara-Casablanca-3 (Castellón) and especially Cal Guardiola (Barcelona) with the presence in the north of Eurosiberian taxa like the common brown frog (Rana temporaria) or the smooth snake (Coronella austriaca) and the withdrawal towards the South of some Mediterranean taxa (Blanus cinereus
and Chalcides bedriagai). The stratigraphic sequence of Gran Dolina (Atapuerca, Burgos), thick of 18 m
and covering the period going from 1.0 to 0.3 My constitutes a privileged site for the study of climatic and environmental changes. Variations within herpetofauna highlight the alternation of hot-dry periods and colder-wet periods. The evolution of the forest indices within the sequence makes it possible to correlate the fresher phases with a probable more significant development of forest cover. The modern chorologic data of these faunas allow hypothesis for estimating mean annual temperatures and precipitations.
UM~N.-Diferencios interltdbitar en la alitnentacidn del Bliho Real (Bubo bubo) eti el valle medio del Ebro (NE de Esparia): efecro de la disponibilidad de conejo {Oryctolagus cuniculus). En el presente estudio se examinan las diferencias tr6ficas enve dos subpoblaciones muy pdximas de B~iho Real que difieren tan-to en la proporci6n de habitats como en la dispnibilidad de conejos en habitat favocable en torno a 10s nidos (Norte: 25-50% de regadlo y 10,4 conejosf100 km; Sur: 0-10% de regadio y 56 conejosl100 km). Se anali-zaron 3670 presas aparecidas en egagr6pilas & 14 parejas. En el sector segtentrional Mm sprerus es la especie m& capturdda, cediendo importancia a Otycrolagus cuniculus en la zona meridional. En t6rmioos de biomasa et conejo es la presa principal en ambas zonas, aunque con mucha menor importancia en la zona none, don-de Ratflu y MW contribuyen tarnbikn de forma considerable a la dieta del ave. Otros taxones presentan dife-rentes tasas de captura entre ambos sectores; 10s peces unicamente son capturados en 10s territories septen-trionales mientras reptiles e invertebrados son significativamente mAs atrapados en el sur. Asimismo, se examinan las diferencias en diversidad tr6fica y tamarIo de presa entre ambas subpoblaciones. llegando a la conclusidn de que el Btiho Real diversifica sus fuentes de energIa en la zona norte debido a la escaset de una
The middle Pleistocene site of Atapuerca (Spain) has many archaeological, palaeontological and human remains in stratigraphic position. Amphibian remains are abundant and easily identifiable in many levels of the site. We have investigated the taphonomy of the amphibian remains, with the question to be answered being their source. Did the amphibians live and die inside the cave, as occurs naturally nowadays, or were their remains brought into the cave by predator activity or by other agents such as sedimentary motion, water transport, etc. There is no reference source of taphonomic alterations on amphibian remains and we provide some basic background for this, using two approaches. One consisted of analyzing a collection of scats and pellets of known predators, and based on the alterations and breakage present in the bones digested by each, we produce a set of categories that will be an analytical tool to be applied to the fossil record. Secondly, we have carried out several laboratory experiments to measure the effects of weathering and water transport on amphibian remains. We subsequently analyzed the fossil amphibian remains from the middle Pleistocene site of Dolina (Atapuerca, Spain). The results showed that the taphonomic processes observed in the Atapuerca sequence of TD4–TD5 are the same, and the amphibian remains at these levels entered the cave deposits in the scats of a mammalian predator. This has been identified as a mustelid, which is an opportunistic predator which would feed on amphibians from the surrounding environment.
BLAIN H.-A., VILLA P. 2006. Amphibians and squamate reptiles from the early Upper Pleistocene of Bois Roche Cave (Charente, southwestern France). Acta zoologica craco-viensia, 49A(1-2): 1-32. Abstract. The layers, ascribed to the early Upper Pleistocene (Middle Palaeolithic) of Bois Roche Cave, contain an abundant bone assemblage, as well as a few lithic artifacts, in a hyena den. The amphibians and squamate reptiles fauna consist of: Pelobates fuscus (Pe-lobatidae), Pelodytes punctatus (Pelodytidae), Bufo bufo and Bufo calamita (Bufonidae), Hyla sp. (Hylidae), Rana temporaria and Rana cf. R. lessonae (Ranidae), a small lacertid (Lacertidae), Anguis fragilis (Anguidae), Natrix natrix and Coronella austriaca (Colubri-dae) and a "Vipera berus complex" species (Viperidae). The herpetofauna may be con-temporaneous of cold-temperate climatic conditions. P. fuscus, R. lessonae and "V. berus complex" underline more continental climatic conditions mitigated by the occurrence of species like P. punctatus and B. calamita with western Atlantic affinities. The palaeoenvi-ronment, suggested by herpetofauna, was a very open environment, with damp meadows and small grove areas of broad-leaved trees and conifers. The proportion and distribution of species in the stratigraphic sequence allow hypotheses concerning minor climatic variations to be advanced.
In the temperate climate zone in Europe the composition of the diet of predatory vertebrates shows evident variability between
the warm and cold season. However, the recently observed climate warming can mitigate the effect of snow cover and low temperatures
on the winter foraging ecology of raptors, thus affecting trophic webs in ecosystems. We analysed diet variability in the
tawny owl Strix aluco, between the warm and cold seasons of four unusually warm years (as compared to reference years of 1950–2000) in two habitats
(forest vs. farmland) in Central Poland. The most important prey group in the tawny owl’s diet were mammals, constituting
over 80% of prey items. There were distinct diet differences between the two seasons: insectivorous mammals, birds and amphibians
were caught more often during the warm season, and Muridae and Arvicolidae during the cold season. The proportion of insectivorous
mammals, voles and amphibians was significantly higher in forest than in farmland. Diet diversity, analysed with rarefaction
methods and expressed as the expected cumulative mammal species number for a given number of randomly sampled preyed mammals,
was independent of season and higher in forest than in the agricultural habitat. We conclude that even during unusually warm
years tawny owls change significantly their feeding habits between the warm and cold season. The effect of season, habitat
and weather factors on diet variability in raptors are discussed.
Despite being described in 1993, the skeleton of Rana pyrenaica Serra-Cobo, 1993 has never been investigated. The present publication thus represents the first description of the osteology of the spring-dwelling Pyrenean small brown frog. Its specific status is osteologically supported by some discrete differences. When compared with other Iberian brown frogs, it shows a mosaic of characters that are difficult to interpret. However, according to ecomorphology, R. pyrenaica shares a robust skeleton morphology and various morphological traits with the more terrestrial Rana temporaria (Linnaeus, 1758), but also shares some more aquatic life osteological characters with the aquatic Rana iberica Boulenger, 1879. As well as increasing what is known about R. pyrenaica, this study also aims to facilitate the correct diagnosis of the species in fossil assemblages and its osteological identification as the prey of other vertebrates in trophic studies.
El Mirador is a cave in the Sierra de Atapuerca (northern Iberian Peninsula) that contains 27 archaeological layers from the Lateglacial to the Late Holocene. A total of 4436 small-mammal remains have been analysed from these layers, and 19 taxa have been identified (three insectivores, seven chiropters and nine rodents). The palaeoenvironmental reconstruction based on a small-mammal analysis suggests that the entire sequence is dominated by a woodland landscape. Our climatic analysis characterises the climate in terms of an evolution from a cool and arid period in the Pleistocene layers (16,000 to 14,000 cal yr BP), probably related to Heinrich Event 1, to humid conditions and temperatures similar to nowadays in the Holocene layers in general. In MIR23 and MIR22 (7300 to 6800 cal yr BP) we detect an increase in temperature to levels above current values and an important decrease in rainfall. These climatic characteristics could be related to the end of the African Humid Period. Moreover, a short, slightly cooler event is registered in MIR11 (ca. 6300 to 5900 cal yr BP), coinciding with a change in the economic pattern in El Mirador. The results obtained on the basis of small-mammal studies are compared with multiproxy terrestrial data (pollen, charcoal, phytoliths, geochemistry, large mammals) and the Greenland ice-core record, as well as various other core records closer to the coasts of the Iberian Peninsula (MD95-2042, MD99-2331, MD95-2043 and SMP02-3). These comparisons show the agreement of the palaeoenvironmental and palaeoclimatic results with all the previous multiproxy and core-record data.
Herpetological samples from the late Pleistocene wurmian site Cueva de Cobrante (San Miguel de Aras, Cantabria, Spain) are studied. The herpetological remains are associated with different Upper Palaeolithic levels. The predominant species in all excavated levels is Rana tempo-raria, accompanied by scant remains of Alytes obstetricans and Bufo bufo. Sporadic isolated remains of Lacerta sp., newborn snakes and perhaps Bufo calamita have also been detected. Within the osteological anomalies, two remarkable Rana temporaria opisthocoelous vertebrae have been found. Probably most of the herpetological material comes from raptor pellets different from Tyto alba, plus a minor component derived from carnivore mammal predation. Based on its availability as prey, the sedimentation of these remains seems to have occurred between April and October, probably at a higher rate during the early summer period.
The archaeological site of El Mirador is located in the southern slope of the Sierra de Atapuerca. The work developed at the site is providing a substantial set of data from the Upper Palaeolithic and Early Neolithic to the Middle Bronze Age. Throughout at least about 4000 years of occupation, the cave was used for various activities, among which, burial, habitation and animal stalling. The practices related with this last use is, at the moment, the main origin of the archaeological deposits, which are mainly composed by burnt animal dung with vegetal residues, potsherds, lithics and faunal remains. In addition, it is characterized by high sedimentation rates that have enabled an individual and clear record of different episodes, providing high resolution chronological data. Due to these particularities, specific excavation methodology and interdisciplinary studies of the archaeological data have been developed in order to understand the genesis of this archaeological sequence and, at the same time, to provide information about the introduction and development of the production economy in the Submeseta Norte region.
Owls are important consumers of small vertebrates, and because they regurgitate pellets rich in bone, they may be important potential contributors of the concentrated remains of small vertebrates to the fossil record. Owls of three sizes, the large great horned owl ( Bubo virginianus ), the medium-sized barn owl ( Tyto alba ), and the small screech owl ( Otus asio ), were fed a common diet of mice. The bony contents of the pellets were analyzed to determine the amount of bone loss by digestion, bone completeness, and sites of bone breakage. For all three species, only about half the number of bones ingested were recovered in the pellets. Mandibles and femora were most abundant, and pelves and scapulae were the least abundant. Screech owls broke 80% of the cranial and limb elements, barn owls only 30%. Skulls fared poorly in great horned and screech owl pellets, while barn owls returned 80% of the skulls intact, with only the caudal portion of the cranium damaged; barn owls also returned articulated strings of vertebrae and complete paws. These results provide a baseline for the recognition of owls as agents of accumulation of small bones in the fossil record and suggest that the actions of ancient predators may be revealed by species-specific patterns of bone destruction of an assemblage of fossil prey species.
Cultural dental wear provides useful information about the use of anterior teeth for non-masticatory purposes. Non-alimentary tasks are usually related to economic and cultural activities. The presence of cultural dental wear has been checked in four different Homo species from the Sierra de Atapuerca sites (Sima del Elefante, Gran Dolina-TD6, Sima de los Huesos and El Mirador cave). The chronology of these sites ranges from more than one million to 4000 years ago.Evidence of dental wear has been documented in the four Homo species analysed, confirming that hominins began using their teeth as tools as far back as one million years ago. Each species exhibits specific typologies and frequencies of dental wear features. Also, dental features are located on different dental surfaces and tooth types, indicating a diversity of activities carried out using the anterior teeth as a tool.The use of the teeth as a tool was a widespread habit in the genus Homo. However, the diversity of dental wear patterns can be related to cultural and economic activities, broadening our knowledge of the behaviour of ancient hominins.
We present a middle-aged Chalcolithic male with a supernumerary distomolar in the mandible. The prevalence of supernumerary teeth in present-day populations is low, ranging from 0.1% to 3.4%; most supernumerary teeth are documented in the anterior and molar regions of the maxilla in present populations. However, the prevalence of supernumerary molars in past populations is still unknown. Moreover, a complete pathological study has been done of this individual.Maxilla, mandible and teeth have been analyzed searching for dental pathologies. A cone-beam computed tomography (CBCT) analysis of mandibular and maxillary fragments was performed to check the evidence of hyperdontia. Dental wear and maxillary alveolar bone have been analyzed with environmental scanning electronmicroscope (ESEM) to improve the diagnosis of an abscess and evidence of tooth picking.This individual shows a left distomolar in the mandible without any evidence of other supernumerary teeth. The fourth molar is not associated with any congenital disease. However, this individual suffered severe dental wear and a variety of oral pathologies such as, dental decay, abscesses, pulpitis, periodontal disease, toothpicking marks in an upper molar, arthritis of the temporomandibular joint and malocclusion associated with high masticatory loads. To our knowledge, this individual from El Mirador Cave (Sierra de Atapuerca, Spain) shows the most ancient case of a fourth molar documented. The poor oral health of this individual corresponds to the general dental health of Chalcolithic populations.
Manuskript eingelangt am 11. November 1985 Zusammenfassung Nahrungsreste aus einem alpinen Uhubrutrevier in den Hohen Tauern (Salzburg/Österreich) enthielten 7593 Beutetiere in 34 Vogel-, 29 Säugetier-, 2 Fischarten, sowie je einer Amphibien-und Reptilienart. Hervorzuheben ist der Nachweis von Birkenmäusen, Sicista betulina (n = 16), Haselmäu-sen, Muscardinus avellanarius (n = 19), Siebenschläfern, Glis glis (n = 5), Baumschläfern, Dryomys nitedula (n = 2) und Hamstern, Cricetus cricetus (n = 4). Grasfrösche, Rana temporaria (n = 3668) stellen mit 48% der Stückzahl und 30% der Biomasse die wichtigste Nahrungsresource. Zusammen mit Kleinnagern (Schermaus, Arvicola terrestris, Microtus-Arten) erreichen sie 62% der Biomasse und 89% der Individuenanteile. Die Mandibula erwies sich als repräsentatives Skelettelement zur quantitativen Erfassung der Kleinnager. Das geborgene Material enthält auch Hinweise auf ehemalige Besiedelung der Uhuhorste durch Bartgeier, Gypaetus barbatus (Eischalenreste, Nahrungsreste, Kotballen). Summary Prey remains from an alpine territory of the Eagle Owl, Bubo bubo, situated in the Hohe Tauern (Province of Salzburg/Austria) contained 7593 prey animals (34 species of birds, 29 species of mammals, 2 species of fish, 1 of amphibian, 1 of reptile). Of special interest is the occurence of Northern Birch Mouse, Sicista betulina (n = 16), Common Dormouse, Muscardinus avellanarius (n = 19), Fat Dormouse, Glis glis (n = 5), Forest Dormouse, Dryomys nitedula (n = 2) and Hamster, Cricetus cricetus (n = 4). The Brown Frog, Rana temporaria (n = 3668) is the main feeding source (48% of prèy animal and 30% of biomass). Together with small rodents (Vole Rat, Arvicola terrestris, Snow Vole, Microtus nivalis and Common Vole, Microtus arvalis) they represent 62% of the biomass and 84% of prey individuals. The mandibula proved to be a representative item of skeleton for the quantitative determination of small rodents. The material recovered proved the former breeding of the Bearded Vulture, Gypaetus barbatus, in the Eagle Owl's nest (egg shells, food remains, excrements in ball shape).
This study provides a morphological characterization of the inner anatomy of the root canals of permanent first and second molars in Chalcolithic and early Bronze Age human fossils using cone-beam computed tomography. The general evolutionary trend in present-day human dentition is related to morphological simplification. As little is known about when this trend appeared in Homo sapiens populations, the aim of this work is to test the presence of modern radicular morphology 4,400 years ago. Fifty-four permanent first and second maxillary and mandibular molars of 17 individuals were included in the study. All maxillary first and second molars showed three separate roots. Almost all the lower molars analyzed (100% of first molars and 75% of second molars) had two separate roots. More differences in the canal system configuration were documented in the maxillary mesiobuccal roots than in the palatal or distobuccal roots. The most variable tooth in root and canal configuration is the maxillary second molar. It should be pointed out that 12.5% of the teeth analyzed showed a C-shaped root configuration. Anat Rec, 2014. © 2014 Wiley Periodicals, Inc.
Because of their modes of life, Amphibians and Reptiles are good environmental and climatic indicators and can therefore teach us a lot about present and past environments. However, the remains of these animals are little studied in archaeological contexts. This study shows that development of the field of analysis of fossils bones of Amphibians and Reptiles is of major interest, but also highlights the necessity to make osteological collections for comparison, to undertake neotaphonomic studies and to put palaeoclimatic indications in place for these taxa. These remarks are especially pertinent for geographical regions that have a high potential with regard to Prehistory, as is the case in Morocco. In this study, we present information on the environment of Neolithic people from the region of Rabat-Temara, Morocco. Five thousand and eight hundred years ago, although global climate was similar to that of today, it was more humid, as shown by the presence of a toad species (Bufo bufo) that is absent today in this same region.
Mesolithics as « frogsfans » ? A new finding at the site of la Baume dʼOgens (Vaud, Switzerland).- Occupied several times during the Mesolithic, the site of la Baume dʼOgens (Vaud, Switzerland) has yielded numerous and well preserved faunal remains. In particular, these include several hundred amphibian bones, apparently belonging to a single species, the Grass Frog (Rana temporaria LINNÉ, 1758). Most of these bones are burnt and the relative abundance of the different skeletal parts shows a strong over representation of the hind quarters. These findings leave little doubt that the frogs were consumed by Mesolithic people.
Within the site of Kutná Hora-Denemark (Kutná Hora distr., central Bohemia – Eneolithic, Řivnáč Culture, 3000–2800 BC), 893 frog bones or bone fragments were present in five archaeological features. All identified specimens are of the Common Frog (Rana temporaria Linnaeus, 1758). Males predominate, according to humeral morphology, and only adult individuals are present. The most important findings come from feature 36, where an accumulation of 739 frog bones (MNI=123) were found, of which 10% (NISP) were burned. This and other contextual evidence indicates that they relate to activity of the Eneolithic people in the settlement. The predominance of hind leg bones and other circumstantial evidence suggest that the frogs were part of the human diet. Behavioural studies of this species suggest that the frogs were gathered from small ponds during the months of March or April. Similar findings are rare.
Aim To study the patterns of genetic variation and the historical events and processes that influenced the distribution and intraspecific diversity in Hyla meridionalis Boettger, 1874.
Location Hyla meridionalis is restricted to the western part of the Mediterranean region. In northern Africa it is present in Tunisia, Algeria and Morocco. In south‐western Europe it is found in the south of France, north‐western Italy and north‐eastern and south‐western Iberian Peninsula. There are also insular populations, as in the Canaries and Menorca.
Methods Sampling included 112 individuals from 36 populations covering the range of the species. We used sequences of mitochondrial DNA Cytochrome Oxidase I ( COI ) for the phylogeographical analysis (841 bp) and COI plus a fragment including part of tRNA lysine , ATP synthase subunits 6 and 8 and part of Cytochrome Oxidase III for phylogenetic analyses (2441 bp). Phylogenetic analyses were performed with paup *4.0b10 (maximum likelihood, maximum parsimony) and M r B ayes 3.0 (Bayesian analysis). Nested clade analysis was performed using tcs 1.18 and G eo D is 2.2. A dispersal‐vicariant analysis was performed with diva 1.0 to generate hypotheses about the geographical distribution of ancestors.
Results We found little genetic diversity within samples from Morocco, south‐western Europe and the Canary Islands, with three well‐differentiated clades. One is distributed in south‐western Iberia and the High Atlas, Anti‐Atlas and Massa River in Morocco. The second is restricted to the Medium Atlas Mountains. The third one is present in northern Morocco, north‐eastern Iberia, southern France and the Canaries. These three groups are also represented in the nested clade analysis. Sequences from Tunisian specimens are highly divergent from sequences of all other populations, suggesting that the split between the two lineages is ancient. diva analysis suggests that the ancestral distribution of the different lineages was restricted to Africa, and that an explanation of current distribution of the species requires three different dispersal events.
Main conclusions Our results support the idea of a very recent colonization of south‐western Europe and the Canary Islands from Morocco. South‐western Europe has been colonized at least twice: once from northern Morocco probably to the Mediterranean coast of France and once from the western coast of Morocco to southern Iberia. Human transport is a likely explanation for at least one of these events. Within Morocco, the pattern of diversity is consistent with a model of mountain refugia during hyperarid periods within the Pleistocene. Evaluation of the phylogenetic relationships of Tunisian haplotypes will require an approach involving the other related hylid taxa in the area.
Although still ongoing, the archaeological and palaeontological excavations in the Gran Dolina cave (Sierra de Atapuerca, Burgos) every year since 1976 have already revealed a long, culturally and palaeontologically rich sequence dated by biostratigraphy, ESR, U-series, and the palaeomagnetic record at between roughly 1 my and 250 ky. Gran Dolina has one of the longest and most continuous vertical profiles. The section, 19 m thick, divided into 11 stratigraphic levels, represents a time span of nearly 700,000 years. The Gran Dolina sequence was partially excavated during a preliminary evaluation of its archaeological and palaeontological richness. This excavation, known as the “Gran Dolina Sondeo South”, lasted from 1993 to 1999. Hominin remains were unearthed in 1994 from level 6 of Gran Dolina, and level 6 was dated to somewhat more than 780 ky based on palaeomagnetic and microfaunal evidence, making these the oldest known hominins in Europe. Given the importance of this discovery, every effort is being made to reconstruct the landscapes where these hominins once thrived. The Gran Dolina Sondeo South (TDS) has furnished a great number of microvertebrate remains, and of these around 40,000 bones are attributed to amphibians and squamates. The faunal list of the squamates and amphibians of the TDS comprises at least 22 species, a diversity similar to what exists nowadays in Spain: Triturus sp., Alytes obstetricans, Pelobates cultripes, Pelodytes punctatus, Bufo bufo, Bufo calamita, Hyla arborea, Rana temporaria, Rana cf. dalmatina, Rana cf. iberica, Pelophylax (ridibunda) sp., Blanus cinereus, Lacerta s.l., small-sized indeterminate lacertids, Anguis fragilis, Natrix natrix, Natrix maura, Coronella austriaca, Coronella cf. girondica, Rhinechis scalaris, Vipera cf. aspis and Vipera cf. latasti. These ectotherm vertebrates show variations in their taxonomic diversity throughout the sequence. Although they do not differ specifically from the extant herpetofauna of the Iberian Peninsula, they do so in the composition of their taxonomic assemblage. When the squamates and amphibians of TDS are grouped by habitat categories (the habitat weightings method), and studied in terms of their vertical evolution throughout the sequence, it is possible to follow environmental changes across the Early–Middle Pleistocene transition. Results from the squamate and amphibian studies indicate that the landscape had open habitats in the vicinity of the Atapuerca caves throughout the sequence, with wet points in the surrounding area. Open-dry microhabitats were more developed during warm periods, whereas during cold periods the environment was slightly more humid and wooded. These results are compared with studies in small mammals, pollen analysis and geomorphology, giving a scenario for the palaeoclimatic and palaeoenvironmental changes that occurred during the Early to Middle Pleistocene boundary in Atapuerca, and hence a scenario for the hominins that once lived in the Sierra de Atapuerca.
The relationship between local and global climatic variations and the origin and dispersal of Homo sapiens in Africa is complex, and North Africa may have played a major role in these events. In Morocco, very few studies are specifically dedicated to small fossil vertebrates, and neither taphonomic nor palaeoecological studies have been undertaken on these taxa, particularly in archaeological contexts. The late Pleistocene to middle Holocene succession of El Harhoura 2 cave, situated in the region of Témara, yields an exceptionally rich small vertebrate assemblage. We present the results of a first systematic, taphonomic, and palaeoecological study of the small mammals from Levels 1 to 8 of El Harhoura 2. The absence of bone sorting and polishing, as well as the presence of significant traces of digestion indicate that the small mammal bones were accumulated in the cave by predators and that no water transport occurred. Other traces observed on the surface of bones consist mainly of root marks and black traces (micro-organisms or more probably manganese) which affected the majority of the material. The percentage of fragmentation is very high in all stratigraphic levels, and the post-depositional breakage (geologic and anthropogenic phenomena) obscure the original breakage patterns of bones by predators. According to the ecology of the different species present in the levels of El Harhoura 2, and by taking into account possible biases highlighted by the taphonomic analysis, we reconstruct the palaeoenvironmental evolution in the region. For quantitative reconstructions we used two indices: the Taxonomic Habitat Index (THI) and the Gerbillinae/Murinae ratio. Late Pleistocene accumulations were characterised by a succession of humid (Levels 3, 4a, 6, and 8) and arid (Levels 2?, 5, and 7) periods, with more or less open landscapes, ending in an ultimate humid and wooded period during the middle Holocene (Level 1). We discuss particular limits of our results and interpretations, due to an important lack of taxonomic, ecological, and taphonomic knowledge in North Africa.
Este trabajo trata de la alimentación de la lechuza común (T. alba) en quince localidades de Alicante (S. E. Iberia). El número total de presas vertebradas estudiadas es de 6.570. Las aves paseriformes presentan más importancia por término medio en su dieta que en otros lugares de España. Por el contrario el topillo P. duodecimcostatus, presa común en otros lugares de Iberia, casi no tiene importancia en las localidades estudiadas. Además se han analizado algunas regurgitaciones de búho real (B. bubo), siendo el conejo y el erizo las presas básicas. This paper deals with the feeding habits of the Barn owl (T. alba) in 15 localities of Alicante (S. E. Spain). The amount of the studied vertebrate preys goes up to 6.570. Passerine birds are, on the average, more important as a prey than elsewhere in Spain. Otherwise, the vole P. duodecimcostatus, common prey elsewhere in Iberia, is very rarely concerned here. Some pellets of Eagle owl (B. bubo) have been also analyzed, being rabbit and hedgehog the ground preys.