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Canis (Xenocyon) lycaonoides from Dmanisi. (a–c) D6327a, left corpus with p1-p3 in buccal (a), lingual (b) and occlusal (c) views. (d)–(f), D6327b, left lower m1 in buccal (d), lingual (e) and occlusal (f) views. QR code and Augmented Reality (AR) marker showing 3D comparison between the lower first molar morphologies of Canis (Xenocyon) from Dmanisi (red), Canis (Xenocyon) lycaonoides from Venta Micena (green) and Canis (Xenocyon) falconeri from Upper Valdarno (gray). Instructions: Scan the QR code on the left; open the link; allow the browser to access the camera of your device; point the camera toward the marker (on the right); and wait for the model to load (up to 10 s). It is possible to turn the device around the marker (or to move the marker) to see different parts of the model. Best visualization performances can be achieved by printing the markers, rather than pointing at them on screens. For common issues refer to Supplementary Information and Bartolini-Lucenti et al.³². Photos of the fossil specimens elaborated in Photoshop CC2019 (https://www.adobe.com/). Line drawing of C. (Xenocyon) and figure composition made by S. Bartolini-Lucenti in Inkscape ver. 0.92 (https://inkscape.org/). AR content made in Visual Studio Code ver. 1.50.0 (https://code.visualstudio.com/) and GitHub Desktop ver. 2.6.6 (https://desktop.github.com/).
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The renowned site of Dmanisi in Georgia, southern Caucasus (ca. 1.8 Ma) yielded the earliest direct evidence of hominin presence out of Africa. In this paper, we report on the first record of a large-sized canid from this site, namely dentognathic remains, referable to a young adult individual that displays hypercarnivorous features (e.g., the redu...
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Citations
... Apart from this large hyaenid, carnivores are well represented in BLD and FN3, including two mustelids (Martellictis ardea and Meles meles), three canids (Vulpes alopecoides, C. mosbachensis, and Lycaon lycaonoides), one ursid (Ursus etruscus), and three felids (Lynx pardinus, Megantereon whitei and Homotherium latidens) Palmqvist et al., 2023a;Rodríguez-Gómez et al., 2016a, 2016bRos-Montoya et al., 2021). This rich carnivore guild resembles those found in other late Early Pleistocene archaeological and paleontological sites of Eurasia, such as Dmanisi in Georgia, dated to ~ 1.8 Ma (Medin et al., 2019;Bartolini-Lucenti et al., 2021, Pirro Nord in Italy, dated to 1.6-1.3 Ma (Petrucci et al., 2013), and Vallonnet in France, dated to ~ 1.1 Ma (Moullè et al., 2006), or at the early Acheulian site of 'Ubeidiya in Israel, dated to 1.6-1.2 ...
The late Early Pleistocene archaeological site of Fuente Nueva-3 (FN3), which lies in the Guadix-Baza Depression (SE Spain) and is dated to ~ 1.4 Ma, contributes some of the oldest evidence of hominin presence in Western Europe, including a huge tool assemblage of Oldowan tradition, manuports (i.e., unmodified stones used as percussion tools) and abundant fossils of large mammals, some of which preserve anthropogenic marks related to defleshing, butchering and marrow processing. In addition, there are bones with tooth marks produced by scavenging carnivores. The fertile layers of the FN3 section have been grouped in a lower archaeological level (LAL) and an upper archaeological level (UAL). Both levels preserve abundant skeletal remains and lithic tools. However, the LAL shows a high density of manuports, which suggests that hominin activity was more intense at this level, while the UAL preserves many remains of megaherbivores, particularly proboscideans (Mammuthus meridionalis), and almost all coprolites unearthed from the site, which points to a greater involvement of the giant hyenas (Pachycrocuta brevirostris). In this paper, we (i) test for statistical differences in the composition of the faunal assemblages preserved in the UAL and LAL; and (ii) analyze particle size in the fertile layers of both archaeological levels. Our results show that megaherbivores are comparatively overrepresented in the UAL, specially by young elephants, while other medium-to-large and large-sized ungulates, particularly equids, are more abundant in the LAL, showing a predominance of adult individuals. Concerning the sedimentology, layers 2-3 of the LAL show a predominance of silts and clays, with sands representing a minor fraction. In contrast, layer 5 of the UAL is composed of two-thirds of fine and very fine sands, with the remaining third consisting of silts and clays. These data and the paleoenvironmental reconstruction of layer 5 suggest that it may be interpreted as a paleo-quicksand in which megaherbivores were trapped due to the elevated weight per unit area supported by their feet, and their half-sunken carcasses attracted the scavengers, particularly the hyenas, which fed on these carcasses and defecated in their surroundings. El yacimiento del Pleistoceno inferior de Fuente Nueva-3 (depresión de Guadix-Baza, sureste deEspaña): ¿Una letrina de hienas desarrollada en una trampa de arenas movedizas para la megafauna? Resumen El yacimiento arqueológico de Fuente Nueva-3 (FN3), de finales del Pleistoceno temprano, se sitúa en la Depresión de Guadix-Baza (SE de España) y ha sido datado en ~1,4 Ma. Aporta algunas de las evidencias más antiguas de la presencia de homininos en Europa occidental, incluyendo un amplio conjunto de herramientas de tradición olduvayense, manuports Extended author information available on the last page of the article Journal of Iberian Geology (es decir, piedras no modificadas utilizadas como herramientas de percusión) y abundantes fósiles de grandes mamíferos, algunos de los cuales conservan marcas antropogénicas relacionadas con el desollamiento, la carnicería y el procesamiento del tuétano. Además, hay huesos con marcas de dientes producidas por carnívoros carroñeros. Las capas fértiles de la sec-ción de FN3 se agrupan en un nivel arqueológico inferior (LAL) y un nivel arqueológico superior (UAL). Ambos niveles conservan abundantes restos óseos y herramientas líticas. Sin embargo, el LAL presenta una alta densidad de manuports, lo que sugiere que la actividad de los homininos fue más intensa en este nivel, mientras que el UAL conserva muchos restos de megaherbívoros, en particular proboscídeos (Mammuthus meridionalis), y casi todos los coprolitos exhumados del yacimiento, lo que apunta a una mayor participación de las hienas gigantes (Pachycrocuta brevirostris). En este tra-bajo (i) comprobamos si existen diferencias estadísticas en la composición de los conjuntos faunísticosconservados en el UAL y el LAL; y (ii) analizamos el tamaño de las partículas en las capas fértiles de ambos niveles arqueológicos. Nuestros resultados muestran que los megaherbívoros están comparativamente sobrerrepresentados en el UAL, especialmente por elefantes jóvenes, mientras que otros ungulados de tamaño medio-grande y grande, particularmente los équidos, son más abundantes en el LAL, mostrando un predominio de individuos adultos. En cuanto a la sedimentología, las capas 2-3 del LAL muestran un predominio de limos y arcillas, representando las arenas una fracción reducida. Por el contrario, la capa 5 de la UAL está compuesta en dos tercios por arenas finas y muy finas, y el tercio restante por limos y arcillas. Estos datos y la reconstrucción paleoambiental de la capa 5 sugieren que se puede interpretar como una paleoarena movediza en la que los megaherbívoros quedaban atrapados debido al elevado peso por unidad de superficie que soportaban sus extremidades, atrayendo sus cadáveres a medio hundir a los carroñeros, en particular a las hienas, que se alimentaron de estos cadáveres y defecaron en sus alrededores. Palabras clave Orce · Pleistoceno inferior · Tafonomía · Paleoecología · Proboscidios · Coprolitos
... The sparse and scattered fossil records of large hypercarnivorous canids across the Old World add complexity to our understanding of the evolutionary history of these wild dogs. These hypercarnivorous dogs are welldocumented during the Early Pleistocene in Europe, Asia, and Africa, though different taxa are attributed to them depending on the author [30,[124][125][126][127][128][129][130][131][132][133][134][135][136][137][138][139]. ...
... The large-sized hypercarnivorous canid Xenocyon lycaonoides was one of the most characteristic taxa of the second half of the Early Pleistocene ( [141], FAD ca. 1.76 Ma [136]) persisting until the Middle Pleistocene age (LAD ca. 450-400 ka [139]). ...
The Vallparadís Section encompasses various geological layers that span a significant chronological range, extending from the latest Early Pleistocene to the early Middle Pleistocene, covering a timeframe from approximately 1.2 to 0.6 Ma. This period holds particular importance as it coincides with a significant climatic transition known as the Early-Middle Pleistocene Transition, a pivotal phase in Quaternary climatic history.
This transition, marked by the shift from a 41,000-year obliquity-driven climatic cycle to a 100,000- year precession-forced cyclicity, had profound effects on the Calabrian carnivorous mammal com- munities. Notably, the once diverse carnivore guild began to decline across Europe during this pe- riod, with their last documented occurrences coinciding with those found within the Vallparadís Section (e.g. Megantereon or Xenocyon). Concurrently, this period witnessed the initial dispersals of African carnivorans into the European landscape (e.g. steppe lions), marking a significant shift in the composition and dynamics of the region's carnivorous fauna.
... Studies of pathologies in extinct animals provide fascinating insight into diseases that affected prehistoric animals (e.g., Anné et al., 2015;Chinsamy & Tumarkin-Deratzian, 2009;de Souza Barbosa et al., 2013;Ekhtiari et al., 2020;Fernández-Monescillo et al., 2019;González et al., 2017;Haridy et al., 2019;Heckert et al., 2021;Lucas & Schoch, 1987;Moncunill-Solé et al., 2019;Moodie, 1923;Rabe et al., 2022) and in some cases permit deductions regarding how the particular trauma may have affected the behavior of the animal (e.g., Iurino et al., 2013;Rabe et al., 2022;Redelstorff et al., 2015). Several studies have described pathologies in extinct carnivores (e.g., Domingo et al., 2012;Iurino et al., 2013Luna et al., 2023;Slabá et al., 2018;Wang et al., 2023), and some have specifically focused on canids (e.g., Bartolini-Lucenti et al., 2021;Rothschild et al., 2001;Tong et al., 2020;Wang & Rothschild, 1992;Werdelin & Lewis, 2013), although it is evident that the majority of these studies have focused on dental or oral pathologies. Furthermore, most of these studies combined anatomical descriptions with computer tomography (CT) scanning but rarely has the osteohistology of the pathology been investigated (e.g., Woolley et al., 2019). ...
Langebaanweg is a Mio-Pliocene locality located on the West Coast of South Africa. It is renowned for its rich diversity of both terrestrial and marine vertebrate fossils. Several carnivorans have been identified from this site, amongst which is the recently described jackal-like canid, Eucyon khoikhoi. One of the skeletons assigned to E. khoikhoi exhibits anatomical deformities on several bones of the skeleton. Here, we use multiple methodologies (anatomical descriptions, CT scanning and histology) to investigate the bony overgrowths or exostoses evident in the radius, and we compare these findings with those of a radius from a healthy individual of the same species from Langebaanweg. Our results show that anatomical observations are important for first level observation of the pathology, but that micro-CT scanning permits a more precise assessment of how the pathology affected the internal organization of the bone, both periosteally and endosteally. This methodology permitted us to diagnose the tumors as benign rather than cancerous. Our observations of calcified cartilage in the histological thin sections in the region of the exostosis allowed us to further diagnose the exostosis as an osteochondroma. This study has demonstrated the usefulness of applying multiple techniques to characterize and diagnose pathological bony growths in a fossil canivoran. We have also demonstrated the usefulness of histological studies in permitting a more refined diagnosis of the exostosis as an osteochondroma.
... Apart from this large hyaenid, carnivores are well represented in BLD and FN3, including two mustelids (Martellictis ardea and Meles meles), three canids (Vulpes alopecoides, C. mosbachensis, and Lycaon lycaonoides), one ursid (Ursus etruscus), and three felids (Lynx pardinus, Megantereon whitei and Homotherium latidens) Palmqvist et al., 2023a;Rodríguez-Gómez et al., 2016a, 2016bRos-Montoya et al., 2021). This rich carnivore guild resembles those found in other late Early Pleistocene archaeological and paleontological sites of Eurasia, such as Dmanisi in Georgia, dated to ~ 1.8 Ma (Medin et al., 2019;Bartolini-Lucenti et al., 2021, Pirro Nord in Italy, dated to 1.6-1.3 Ma (Petrucci et al., 2013), and Vallonnet in France, dated to ~ 1.1 Ma (Moullè et al., 2006), or at the early Acheulian site of 'Ubeidiya in Israel, dated to 1.6-1.2 ...
The late Early Pleistocene archaeological site of Fuente Nueva-3 (FN3), which lies in the Guadix-Baza Depression (SE Spain) and is dated to ~ 1.4 Ma, contributes some of the oldest evidence of hominin presence in Western Europe, including a huge tool assemblage of Oldowan tradition, manuports (i.e., unmodified stones used as percussion tools) and abundant fossils of large mammals, some of which preserve anthropogenic marks related to defleshing, butchering and marrow processing. In addition, there are bones with tooth marks produced by scavenging carnivores. The fertile layers of the FN3 section have been grouped in a lower archaeological level (LAL) and an upper archaeological level (UAL). Both levels preserve abundant skeletal remains and lithic tools. However, the LAL shows a high density of manuports, which suggests that hominin activity was more intense at this level, while the UAL preserves many remains of megaherbivores, particularly proboscideans ( Mammuthus meridionalis ), and almost all coprolites unearthed from the site, which points to a greater involvement of the giant hyenas ( Pachycrocuta brevirostris ). In this paper, we (i) test for statistical differences in the composition of the faunal assemblages preserved in the UAL and LAL; and (ii) analyze particle size in the fertile layers of both archaeological levels. Our results show that megaherbivores are comparatively overrepresented in the UAL, specially by young elephants, while other medium-to-large and large-sized ungulates, particularly equids, are more abundant in the LAL, showing a predominance of adult individuals. Concerning the sedimentology, layers 2–3 of the LAL show a predominance of silts and clays, with sands representing a minor fraction. In contrast, layer 5 of the UAL is composed of two-thirds of fine and very fine sands, with the remaining third consisting of silts and clays. These data and the paleoenvironmental reconstruction of layer 5 suggest that it may be interpreted as a paleo-quicksand in which megaherbivores were trapped due to the elevated weight per unit area supported by their feet, and their half-sunken carcasses attracted the scavengers, particularly the hyenas, which fed on these carcasses and defecated in their surroundings.
... The relationship between carnivores and hominids has shaped our lineage's evolutionary pathway and behavior since its inception (see [1][2][3][4][5] among others). Over the millennia, landscape sharing and resource competition have resulted in fatal encounters for hominins [6][7][8][9][10] but also in increased occasional and systematic large carnivore exploitation [11][12][13][14][15][16][17][18] . ...
During the Upper Paleolithic, lions become an important theme in Paleolithic art and are more frequent in anthropogenic faunal assemblages. However, the relationship between hominins and lions in earlier periods is poorly known and primarily interpreted as interspecies competition. Here we present new evidence for Neanderthal-cave lion interactions during the Middle Paleolithic. We report new evidence of hunting lesions on the 48,000 old cave lion skeleton found at Siegsdorf (Germany) that attest to the earliest direct instance of a large predator kill in human history. A comparative analysis of a partial puncture to a rib suggests that the fatal stab was delivered with a wooden thrusting spear. We also present the discovery of distal lion phalanges at least 190,000
old from Einhornhöhle (Germany), representing the earliest example of the use of cave lion skin by Neanderthals in Central Europe. Our study provides novel evidence on a new dimension of Neanderthal behavioral complexity.
... The Early Pleistocene dispersal of C. simensis' ancestors from Eurasia to Africa appears to have been coeval with that of the ancestors of the hypercarnivorous Lycaon pictus (African hunting dog), which arrived to Africa later than 1.8 Ma 22,23 . At the same time, 2.0-1. ...
In 2017, a hemimandible (MW5-B208), corresponding to the Ethiopian wolf (Canis simensis), was found in a stratigraphically-controlled and radio-isotopically-dated sequence of the Melka Wakena paleoanthropological site-complex, on the Southeastern Ethiopian Highlands ,~2300 m above sea level. The specimen is the first and unique Pleistocene fossil of this species. Our data provide an unambiguous minimum age of 1.6-1.4 Ma for the species' presence in Africa and constitutes the first empirical evidence that supports molecular interpretations. Currently, C. simensis is one of the most endangered carnivore species of Africa. Bioclimate niche modeling applied to the time frame indicated by the fossil suggests that the lineage of the Ethiopian wolf faced severe survival challenges in the past, with consecutive drastic geographic range contractions during warmer periods. These models help to describe future scenarios for the survival of the species. Projections ranging from most pessimistic to most optimistic future climatic scenarios indicate significant reduction of the already-deteriorating territories suitable for the Ethiopian Wolf, increasing the threat to the specie's future survival. Additionally, the recovery of the Melka Wakena fossil underscores the importance of work outside the East African Rift System in research of early human origins and associated biodiversity on the African continent.
... The situation changes with the record of the early forms of the extant dhole, Canis alpinus, around the beginning of the Middle Pleistocene, as it would later replace C. lycaonoides throughout Eurasia. Taxonomy of the African Canis-grade taxa is probably one of the most cogent topics in Quaternary Canidae paleontology, due to the scantiness of the record, unreliability of time correlations, etc. Ecologically, we can identify several jackal-like mesocarnivorous species (among which are the extant side-striped jackal, Canis adustus, and black-backed jackal, Canis mesomelas, already present in the Early Pleistocene) but also large hypercarnivorous forms, like Canis africanus (similar to European Canis falconeri) and others more derived (Bartolini-Lucenti et al., 2021). ...
This chapter introduces the beginning of Pleistocene vertebrate records, starting from an overview of the Plio-Pleistocene transition. Understanding the Early Pleistocene vertebrate record is impossible without the knowledge of the events characterizing the setting of Early Pleistocene land vertebrate communities. A number of taxa are selected to describe evolutionary and dispersal patterns within the Early Pleistocene. These are the dispersals and evolutionary history of Canidae, elephants, horses, Rhinocerotidae, Hippopotamidae and Cervidae. The last section offers an overview of the Early Pleistocene herpetofauna.
... Wild dogs have become endangered in East Africa from human hunting and interference competition by larger carnivores, and jackals have been in decline in for many years (e.g., Durant et al., 2011). There is only one individual (NISP ¼ 2) of Canis (Xenocyon) from Dmanisi (Bartolini-Lucenti et al., 2021). Lions live in the largest social groups of any extant felids and are able to suppress wild dog numbers in the Serengeti (Swanson et al., 2014). ...
There are many hypotheses regarding influences on the early hominin biogeographic spread into Eurasia; among them is increased meat-eating. Dmanisi in Georgia is one of the rare Early Pleistocene sites in Eurasia, and here we present primary information and analysis of the medium and large mammal taphonomy, contributing information about site formation and the hominins' interaction with the fauna. Nearly 85% of the specimens come from the B1 stratum. Relative abundances of mammal families demonstrate some bias toward carnivores, especially Canis borjgali, and diverse Felidae species. Bones display little weathering. Post-depositional surface modifications and matrix obscure many bone surfaces, but carnivore tooth marking is the most common bone surface modification from the nutritive taphonomic phase. Tooth pits are large, in the size range of those made by modern Crocuta crocuta and Panthera leo. Breakage variables indicate most breaks occurred while the bones were still fresh, many by carnivore consumption. Fairly even limb bone representation of herbivores suggests carcasses were introduced to the site nearly whole. Hominin tool marks are present in low frequencies, but they suggest a variety of behaviors. These marks are found on Equus, Palaeotragus, Bison, large cervids, Pseudodama, Canis, and Mammuthus. Some were made by filleting proximal limb segments, and so are likely indicative of early access to carcasses, while other marks suggest scavenging. The Homo taphonomic variables resemble the rest of the taphonomic signatures from the site with little weathering, a slightly higher percentage of their bones are whole, but only a few have probable carnivore damage. The assemblage characteristics are compared to modern actualistic and experimental assemblages, and it is concluded that Dmanisi presents a palimpsest of hyena denning, felid activity, hominin meat-eating and likely natural deaths.
... The recent taxonomic attributions for Panthera spelaea (distinguished into the Middle Pleistocene P. s. fossilis and the Late Pleistocene P. s. spelaea), 'Hyaena' prisca (distinguished from Pliocrocuta perrieri), Canis (Xenocyon) lycaonoides (instead of Lycaon lycaonoides), and Megantereon adroveri (for the late Villafranchian-Epivillafranchian Megantereon distinguished from the middle Villafranchian M. cultridens), of Marciszak et al. (2021), Iannucci et al. (2021), Bartolini-Lucenti et al. (2021) and Lavrov et al. (2022), respectively, are followed here. Finally, for sake of simplification we refer to all species of the Canis mosbachensis lineage (C. ...
This study investigates the community structure, dynamics and evolution of the guilds of large carnivorans during the Pleistocene of Europe. Emphasis is given to important renewals, the composition of the guilds in terms of dietary preferences and foraging strategies, and to intraguild competition for access to food resources. For this purpose, cluster, principal component, and guild structure analyses are performed combining four ecological/behavioural parameters —body mass, diet, prey acquisition strategy, sociality— of large carnivorans that practice hunting and/or scavenging on large prey. Results show only minimal niche overlap, indicating that large predators may have
reduced/avoided competition by almost exclusively occupying different niches, i.e. they did not compete for the same resources and/or employed different foraging strategy. Such niche partitioning and competition avoidance may have reduced the occurrences
of potential trophic conflict and could explain the cooccurrence of a high diversity of large predators within the same broad feeding guild. Furthermore, the major predator guild remodeling took place close to the Early/Middle Pleistocene transition, when previously dominant carnivorans (e.g. Pachycrocuta, Megantereon, Acinonyx, Panthera gombaszoegensis) went extinct and new immigrants arrived (e.g. Panthera spelaea, Panthera pardus, Crocuta crocuta), forming the Galerian to Late Pleistocene guilds. Finally, the inclusion of the meat-eating Homo in the carnivore guild is discussed, including its possible
impact to the demise of carnivoran diversity and accordingly of the several large carnivoran niches towards the end of the Pleistocene.
... However, the fossil record of L. lycaonoides is scarce in Guadix-Baza (a few specimens from VM, see details in Palmqvist et al., 2022b) and VM-7000 is the only complete skull. This limitation on sample size precludes the hypothetico-deductive method and forces to use the 'abductive research method, ' which means that the best explanation at hand for interpreting the anomalies of VM-7000 is to consider that they evidence FA resulting from developmental instabilities caused by inbreeding depression Bartolini-Lucenti et al., 2021). ...
... In the case of VM, this would result in an effective population size for L. lycaonoides of around one hundred individuals, which would have promoted further inbreeding. However, the dog from VM was able to reach adulthood, as shown by its moderately worn permanent dentition, despite severe developmental handicaps, which suggests that cooperative behavior from other members of the hunting pack may have helped it to survive Bartolini-Lucenti et al., 2021). Similarly, cooperation among hominins has been reported in Dmanisi. ...
The chronology and environmental context of the first hominin dispersal in Europe have been subject to debate and controversy. The oldest settlements in Eurasia (e.g., Dmanisi, ∼1.8 Ma) suggest a scenario in which the Caucasus and southern Asia were occupied ∼0.4 Ma before the first peopling of Europe. Barranco León (BL) and Fuente Nueva 3 (FN3), two Early Pleistocene archeological localities dated to ∼1.4 Ma in Orce (Guadix-Baza Depression, SE Spain), provide the oldest evidence of hominin presence in Western Europe. At these sites, huge assemblages of large mammals with evidence of butchery and marrow processing have been unearthed associated to abundant Oldowan tools and a deciduous tooth of Homo sp. in the case of BL. Here, we: (i) review the Early Pleistocene archeological sites of Europe; (ii) discuss on the subsistence strategies of these hominins, including new estimates of resource abundance for the populations of Atapuerca and Orce; (iii) use cartographic data of the sedimentary deposits for reconstructing the landscape habitable in Guadix-Baza; and (iv) calculate the size of the hominin population using an estimate of population density based on resource abundance. Our results indicate that Guadix-Baza could be home for a small hominin population of 350–280 individuals. This basin is surrounded by the highest mountainous reliefs of the Alpine-Betic orogen and shows a limited number of connecting corridors with the surrounding areas, which could have limited gene flow with other hominin populations. Isolation would eventually lead to bottlenecks, genetic drift and inbreeding depression, conditions documented in the wild dog population of the basin, which probably compromised the viability of the hominin population in the medium to long term. This explains the discontinuous nature of the archeological record in Guadix-Baza, a situation that can also be extrapolated to the scarcity of hominin settlements for these ancient chronologies in Europe.
