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Variation of antler shape in fallow deer species: A, Dama dama; B, Dama mesopotamica; C, Dama eurygonos, D. Dama vallonnetensis.
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Reliable, informative and up-to-date information regarding the location, mobility and behavioral patterns of animals enhances our ability to preserve biodiversity, manage invasive species and conduct research. The basis of which is an accurate count of the animals present in a specified region. In literature, previous studies have presented automat...
Citations
... Despite their abundance and frequency in the fossil record, the systematics of this group of cervids is highly debated (Di Stefano and Petronio, 1998;Azzaroli, 2001). Different classifications have been proposed over the years, with specimens being ascribed to both fossil (Pseudodama, Metacervoceros, "Euraxis", Praeelaphus) (Azzaroli, 1992;Di Stefano and Petronio, 1998;Croitor, 2006Croitor, , 2014Croitor, , 2018 and extant deer genera (Rusa, Axis, Dama, Cervus) (De Vos and Reumer, 1995;Kahlke, 1997;Pfeiffer, 1999;van der Made, 2001van der Made, , 2015van der Made et al., 2017van der Made et al., , 2023Di Stefano and Petronio, 2002;Petronio et al., 2007) and either lumped into a single genus (e.g., Pseudodama or Dama), or split into different genera based on cranial (especially antler) or postcranial morphology (SI1). Most studies have thus traditionally focused on their taxonomy and systematics, with fewer research aiming at investigating the palaeobiology of these cervids and their evolution in relation to the Early Pleistocene major environmental and climatic changes following the onset of the Quaternary glaciations. ...
... The neuroanatomy of the Pirro Nord cranium compared with that of other extinct and extant cervids suggests that this group gradually developed "Dama-like" features at least in terms of brain morphology, with specimen from Pirro Nord displaying the closest affinity to modern fallow deer. If this is evidence that Late Villafranchian and Epivillafranchian forms may belong to the fallow deer evolutionary lineage as proposed by Croitor (2018), or if instead it is evidence of evolutionary convergence between Dama and an extinct genus (e.g., Pseudodama), needs to be tested with a larger set of both Early Pleistocene fossils with well-defined craniodental features, and modern and fossil specimens of Dama. ...
Medium-sized deer are frequently found in Early Pleistocene fossiliferous deposits of Southern Europe, and hence they represent a reference group for the Villafranchian fauna. Their evolutionary history is still highly debated, as multiple classifications and systematic revisions have been proposed with specimens being ascribed to several genera such as Pseudodama, Metacervoceros, Euraxis, Praeelaphus, Axis, Dama, or Cervus. Whereas most studies focus on the taxonomy of the group, a few have addressed how these cervids evolved in relation to the major Early Pleistocene climatic events such as the onset of the Quaternary glaciations. A remarkably rich collection of Dama-like deer has been unearthed from the Early Pleistocene site of Pirro Nord (Apricena, south-eastern Italy). Here we analyse palaeoecological and palaeoneurological data of the Pirro Nord sample to investigate both habitat occupation and the evolutionary history of this group during the Early Pleistocene. To do so, we integrate dietary proxies (dental wear patterns and stable isotope signal) with morphological data from a virtual endocast of a well-preserved male specimen, a largely unexplored research path in cervid palaeobiology studies. Moreover, palaeoneurological data may also provide clues to solve the systematic issue of this group. Dental mesowear results point to a long-term mixed diet for the Pirro Nord deer, while a leaning towards a grazing behaviour is suggested by dental microwear patterns. The range of the stable isotope δ13C ratios suggests that it foraged on abrasive water-stressed C3 vegetation in warm woodland and semi-open habitats. Palaeoneurological data seems to confirm a closer affinity in the endocranial morphology of this taxon to the extant fallow deer Dama compared to other Early Pleistocene forms. Our research represents a novel approach to the study of Early Pleistocene fossil deer palaeoecology that can be extended also to other groups to investigate their evolutionary history in relation to climate changes.
... Antlers are an exoskeletal appendage characteristic of the Cervidae (deer) family with a yearly cycle of growth, fall and regrowth [38][39][40]. Shape and size are highly variable between species; thus, their morpho-structural properties are very diverse [41][42][43]. Such properties undoubtedly restrict their potential technical exploitation. ...
Antler is one of the primary animal raw materials exploited for technical purposes by the hunter-gatherer groups of the Eurasian Upper Palaeolithic (UP) all over the ecological range of deers, and beyond. It was exhaustively employed to produce one of the most critical tools for the survival of the UP societies: hunting weapons. However, antler implements can be made from diverse deer taxa, with different ecological requirements and ethological behaviours. Identifying the antler's origin at a taxonomic level is thus essential in improving our knowledge of humans' functional, practical and symbolic choices, as well as the human-animal interface during Prehistoric times. Nevertheless, palaeogenetics analyses have focused mainly on bone and teeth, with genetic studies of antler generally focused on modern deer conservation. Here we present the results of the first whole mitochondrial genome ancient DNA (aDNA) analysis by means of in-solution hybridisation capture of antlers from pre-Holocene archaeological contexts. We analysed a set of 50 Palaeolithic and Neolithic (c. 34-8ka) antler and osseous objects from SouthWestern Europe, Central Europe, SouthWestern Asia and the Caucasus. We successfully obtained aDNA, allowing us to identify the exploited taxa and demonstrate the archaeological relevance of those finds. Moreover, as most of the antlers were sampled using a minimally-invasive method, further analyses (morphometric, technical, genetic, radiometric and more) remain possible on these objects. Significance statement Antlers from the Cervidae family are one of the most exploited raw materials from the Palaeolithic period. Nevertheless, antler implements can be made from diverse deer taxa with different ecological requirements and ethological behaviours. It is thus essential to know their taxonomic origin to evaluate the synergy between the hunter-gatherers, their prey and their environment. Here, we successfully conducted a whole mitochondrial genome aDNA analysis of antlers by means of in-solution hybridisation capture from pre-Holocene archaeological contexts. We demonstrate that implements made from the antlers of undefined taxa can be attributed to a species, enabling deeper archaeological inferences. Although other methods, like palaeoproteomic analyses, can identify at a family taxonomic level, only aDNA allows us to identify the exploited species and perform further phylogenetic analyses.
... Reduced body surface to volume ratios, leading to a stout stature, have been previously proposed as a common evolutionary trajectory for species moving into colder climates, a process known as "Allen's rule" [12]. Another important morphological characteristic of these reindeer is an increased volume of the nasal cavity [13,14], which reduces the heat and water loss from the respiratory tract. It has been argued that these features contribute to weathering the extreme harsh and cold [15] conditions in the High Arctic. ...
Background
Reindeer (Rangifer tarandus) are iconic mammals that inhabit the Arctic and sub-Arctic regions. In these areas, reindeer not only play a vital ecological role, but they also hold cultural and economic significance for indigenous communities. In order to thrive in the harsh conditions of the northernmost areas of the world, reindeer have developed an array of phenotypic adaptations, especially in the ecotypes living in the High Arctic. Therefore, a thorough understanding of population structure, history, and genetic diversity of reindeer is useful for their sustainable management and to guide long-term conservation efforts.
Results
We conducted whole-genome sequencing of a male R. t. tarandus specimen from Norway's Hardangervidda region, generating a highly continuous and complete genome assembly that can be used as a reference genome for genetic analyses focusing on the Fennoscandian reindeer. We also sequenced reindeer ecotypes from across the globe and generated de novo sequences from two ancient samples. Our analysis suggests an independent evolution of small-sized phenotypes in specific high-arctic ecotypes, such as the Svalbard reindeer (R. t. platyrhynchus) and Peary caribou (R. t. pearyi). We describe how the demographic bottleneck that affected the reindeer in the Svalbard archipelago resulted in reduced genetic variability compared to mainland Norway reindeer. Our data suggests that these two distinct ecotypes were likely independent populations before the last glaciation. Finally, we also observe an enriched number of genes associated with cilium motility and cilium assembly presenting missense variants between these two ecotypes, potentially linked to adaptations in the extreme arctic environment. For instance, some of these genes play a role in respiratory cilia movement, potentially improving respiratory function in cold environments.
Conclusions
Our findings provide new insights into the genetic basis of small body size adaptations in reindeer ecotypes and highlight the impact of environmental constraints on their populations. Our high-quality reference genome and associated resources will aid in addressing epidemiological, conservation, and management challenges faced by reindeer populations in a rapidly changing world.
... Few remains attest to the presence of the fallow deer, Dama sp., at Atella. The taxonomy of Dama-like deer is far from unanimously agreed upon (e.g., Di Stefano and Petronio, 2002;Breda and Lister, 2013;van der Made et al., 2017;Croitor, 2018), and thus it is somewhat contentious to attempt to place occurrences of the genus in a chronological context. Differences in Middle Pleistocene fallow deer are mainly documented on antlers, and when these are not available, specific attributions are often based on chronological grounds. ...
... They placed a parallel lineage in Axis, which acquired a fourth point (here: A, B, C, D), and then gave rise to A. farnetensis, which survived till about 500 ka. Croitor (2006Croitor ( , 2018 believed that the Dama e. eurygonos-D. e. farnetensis lineage gave rise to three lineages, leading to D. vallonnetensis and the two living species (Fig. 1), which implies the loss of the middle tine (point B) in D. vallonnetensis, while the palmate species retained it. ...
... Lister (1986) considered the first appearance of Dama dama (including D. d. clactoniana) to be in the early Cromerian and perhaps Beestonian. Croitor (2006Croitor ( , 2018 considered the antlers from Soleilhac and Pakefield (on which the species D. roberti is based) as juveniles of Dama clactoniana, which implies a first appearance not later than MIS17 (712-676 ka). In the scenario that D. clactoniana evolved from D. roberti (Breda & Lister 2013), the transition must be after Soleilhac (MIS15, 563-621 ka) and older records are discarded. ...
We describe fossils of a new species of fallow deer, Dama celiae. It is the end member of the lineage Dama farnetensis–D. vallonnetensis–D. roberti–D. celiae, which reduced the number of points of the antler from four to two, while the parallel lineage leading to the living fallow deer evolved more complex and palmate antlers. The fossils are from localities Pedro Jaro I and Orcasitas in the + 25–30-m terrace of the Manzanares river, which is correlated to MIS9 (337–300 ka) and which also yielded fossils of Megaloceros matritensis, a recently named species, end member of a lineage that survived longer than previously believed. A younger terrace of the Manzanares yielded remains of Haploidoceros, a rare deer known from two older localities in southern France and one younger locality in Spain. So many rare deer species in this valley indicates either endemism and a very special environment or that the record of fossil deer is much less known than generally assumed. Until recently, the European Middle Pleistocene record of deer had only one middle-sized species at a time. Now, it appears that there were up to three contemporaneous species of the size of a fallow deer. Acheulean lithic assemblages have been documented from the same sites as Dama celiae. This species was contemporaneous to Neanderthals with Acheulean culture. Cut marks suggest that it was consumed by them and probably was hunted
... Red deer have a fossil history of over 0.9 million years, since the early Middle Pleistocene (Lister et al., 2010;Croitor, 2018). The maternal divergence between elaphoids and wapitoids has been estimated differently based on mitochondrial DNA (mtDNA), from 7 to 3 Ma (Ludt et al., 2004;Lorenzini and Garofalo, 2015), 2.49 Ma (Mackiewicz et al., 2022), 1.4e1.3 ...
Red deer (Cervus elaphus/hanglu/canadensis) are a group of closely-related species that survived through the Last Glacial period in Eurasia and expanded into North America. The evolutionary modes of red deer in Europe through glacial and interglacial cycles have been established relatively well based on both modern and ancient genomes, while the evolutionary and migratory history of red deer in Asia are not so clear. Northern China, as the southern border of their current distribution, is a significant area for exploring the evolution of wapitoids (Cervus canadensis). Here, we constructed six mitochondrial genomes (lengths of 13,282–16,306 bp) from Late Pleistocene fossil cervid materials in Northeastern China, and four modern complete mitochondrial DNA sequences (16,798–16943 bp) from Northern China. Phylogenetic analyses suggest four clades in wapitoids, with our new individuals clustered into three out of the four clades. BEAST analysis indicates that the divergence between elaphoids (including Cervus elaphus and Cervus hanglu) and the complex of ‘wapitoids + sika deer (Cervus nippon)’ occurred at approximately 1.37 million years ago (Ma, 95%Cl: 1.64–1.15 Ma). Bayesian skyline plots (BSPs) show that the effective maternal population size of wapitoids had a slight and stable expansion between Marine isotope stages (MIS) 7–3, and then shrank from the beginning of the Last Glacial Maximum (LGM). Combined with the divergence dates, population dynamics, and fossil findings in Northern China, we infer that after wapitoids migrated eastward from Central Asia to Northeastern Asia, they had multiple migrations southward into Northern China. The current distribution of wapitoids in Northern China was likely shaped by these migrations as well as habitat fragmentation. Meanwhile, the radiocarbon dates of ancient individuals indicate that the Northeast China Plain, located at the same latitude as European refugia, may have provided a refugium for megafaunal species such as red deer to survive the LGM.
... from China to Britain and as far south as the Caucasus (Valli 2010;Croitor 2018). In body mass, they were broadly similar to modern roe (ca. ...
This comprehensive species-specific chapter covers all aspects of the mammalian biology, including paleontology, physiology, genetics, reproduction and development, ecology, habitat, diet, mortality, and behavior. The economic significance and management of mammals and future challenges for research and conservation are addressed as well. The chapter includes a distribution map, a photograph of the animal, and a list of key literature.
... Comparisons and discussion -The most common largesized deer occurring in the Villafranchian is Eucladoceros, characterized by spectacular comb-like antlers (Azzaroli & Mazza, 1992;Croitor, 2018). Nevertheless, other groups of cervids of comparable size can be found in the same time interval, although their record is poorer. ...
... Nevertheless, other groups of cervids of comparable size can be found in the same time interval, although their record is poorer. These include the enigmatic Arvernoceros, whose systematic status and chronological occurrence are debated (Heintz, 1970;Petronio & Pandolfi, 2011;Croitor, 2018). The genus Praemegaceros seems to replace Eucladoceros, at least in southwestern Europe, in the latest Villafranchian (Abbazzi, 2004). ...
The region of Umbria (Central Italy) represents one of the most interesting areas for the study of Mediterranean Plio-Pleistocene mammal faunas due to the occurrence of numerous paleontological deposits. Most of the fossils have been discovered within the Tiberino Basin, which extends across the centre of the territory for ca. 1800 km 2 , and the small Tavernelle-Pietrafitta Basin, south of Lake Trasimeno. Herein, we provide an additional report on an assemblage within continental deposits from the locality of Vigna Nuova (Piegaro, Perugia) in the Valdichiana Basin, which are observed to crop out in only a few areas of western Umbria. Taphonomic and geological evidence, coupled with rare excavation notes, have allowed the sample to be divided into two assemblages. The younger was collected from the conglomerates (here called "upper layer") which are still visible on the outcrop and is composed only of remains of cf. Leptobos. The data obtained from this fragmentary sample together with some sedimentological-stratigraphic considerations suggest that it can be correlated with other well-known Valdichiana assemblages (e.g., Selvella, Farneta), dated at ca. 1.5 Ma. The older sub-sample comes from a putative paleosol (here called "lower layer") no longer visible in the field. It includes cf. Megantereon, Canis sp., Mustelidae indet., Proboscidea indet., Equus cf. senezensis, cf. Leptobos, Croizetoceros ramosus, Pseudodama sp., Cervidae indet. (large size), and Sus cf. strozzii. This assemblage can be referred to the Coste San Giacomo Faunal Unit (late Middle Villafranchian; ca. 2.2-2.1 Ma). It represents the first well-preserved sample of this age in Umbria and one of the few recorded in Italy, opening new interesting research perspectives on the distribution of mammals within the Peninsula and also on the stratigraphy and evolution of the Valdichiana Basin. "Aggiungeva che per scavare le poche ossa che mi avrebbe subito inviate aveva dovuto faticare enormemente e lottare per due giorni contro ogni sorta di difficoltà. Ricevuto il graditissimo invio mi affrettai a consolidare e restaurare con ogni cura quelle primizie ed oggi sono lieto di poter annunziare, in così solenne adunanza, la interessante scoperta…" ("He added that in order to excavate the few bones that he would immediately send me he had had to work enormously and struggle for two days against all sorts of difficulties. Having received the very welcome shipment,
... Panolia from the Upper Siwaliks (PUPC 66/17, 66/13, and 68/291) approaches this cervid form to the large-sized cervid Praemegaceros obscurus (Azzaroli, 1953) from the Early Pleistocene of Western Eurasia (Croitor, 2018b). The accessory prong in the area of basal ramification in Panolia is a homology of the so-called "outer" (according to Azzaroli and Mazza, 1993) or "dorsal" (according to Croitor, 2006b) tine characteristic of P. obscurus. ...
... The above enlisted features of the antler morphology suggest a close phylogenetic relationship between Panolia and P. obscurus (Croitor, 2018b). One can assume that, as R. (Suraprasit et al., 2016). ...
... Possibly, this is also the case of the deer from Sardhok that shows certain morphological affinity with "Eucladoceros sp." from Kuruksai (Tajikistan). Metacervocerus punjabiensis belongs to a very successful lineage with simple three-pointed antlers that dispersed over the vast Eurasian area and survived until the end-Villafranchian climate crush (Croitor, 2018b). Rucervus is related to European Arvernoceros that was recently included as a subgenus in Rucervus (Croitor, 2018a). ...
This article presents a description of new antler remains from five fossiliferous sites (Sardhok, Panjan Sher Shahana, Puran, Jari Kas, and Potha) of the Upper Siwaliks in Pakistan. The systematic study of the antler material revealed the presence of six cervid forms: Metacervocerus punjabiensis, Rucervus sp., Panolia sp., Hyelaphus sp., Praesinomegaceros bakri, and a poorly represented large cervid that shows a certain affinity with “Eucladoceros sp.” from the Early Pleistocene of Kuruksai (Tajikistan). The remains of Panolia represent the earliest known paleontological record of this cervid lineage. Unlike Metacervocerus and Rucervus that have had phylogenetically closely related counterparts in east and north of the Alpine-Himalayan mountain belt, the evolution of Panolia took place in the Indian subcontinent. The entry of Panolia lineage into the Indian subcontinent marks its phylogenetic split from the main Cervus/Rusa evolutionary branch. The earliest dispersal events of cervids into the Indian subcontinent was preceded by the late Miocene evolutionary radiation and ecological diversification of the subfamily Cervinae in Southeastern Asia. Praesinomegaceros and Metacervocerus most probably entered the Indian subcontinent via Central Asia. Possibly, this is also the case of “Eucladoceros sp.” from Kuruksai (Tajikistan). The dispersal ways of Panolia and Rucervus remain unclear. The dispersals of small-sized cervids (Muntiacus and Hyelaphus) into the Indian subcontinent was triggered by the establishment of the 100-ky glaciation cycle during the Middle Pleistocene: the sea level dropped during glacial peaks and opened to them the dispersal route from Sundaland in the South.
... Khomenko (1913) established the Pliocervinae (Pliocervini proposed later) including three genera with one species for each genus: Cervavitus tarakliensis, Cervocerus novorossiae, and Damacerus bessarabiae. However, the validity and systematic attribution of this group raised much dispute afterwards (Zdansky 1925;Teilhard de Chardin and Trassaert 1937;Simpson 1945;Vislobokova 1990;Di Stefano and Petronio 2002;Croitor 2018Croitor , 2021. Several species or subspecies of Cervavitus were established in China, such as C. novorossidae, Cervavitus demissus, Cervavitus novorossiae shanxius, Cervavitus novorossiae minor, Cervavitus huadeensis, Cervavitus fenqii and C. ultimus (Zdansky, 1925;Teilhard de Chardin and Trassaert 1937;Ji and Li 1979;Qiu 1979;Han 1987;Dong and Hu 1994). ...
Bailong Cave (Bailongdong), dating back to the Middle Pleistocene, is a very important Palaeolithic site in southern China with some hominin remains and substantial mammalian fossils unearthed. Cervid fossils are one of the most common elements in this site, and antler fossils from Bailong Cave were preserved in a better condition than any other Pleistocene site in southern China. Here, we describe the materials of Elaphodus cephalophus, Muntiacus reevesi, Muntiacus muntjak, Cervavitus ultimus and Cervus grayi, while the remains of Rusa unicolor have been described separately in previous paper. The antler fossil of E. cephalophus represents the earliest and undisputed record of this species known currently. The fossils of C. ultimus represent the latest occurrence of this species according to the current knowledge. Overviewing the geological distribution of Quaternary cervids based on 19 sites in southern China, we can find that there was a significant turnover on deer fauna around Mid-Pleistocene Transition in southern China.
