Abstract Rescue excavations recovered a skeleton that resurrect the contemporary dramatic history of Austria in the 17th century as troops besieged Vienna in the second Osmanic-Habsburg war. Unique for Central Europe is the evidence of a completely preserved camel skeleton uncovered in a large refuse pit. The male individual of slender stature indicates a few but characteristic pathological changes revealing not a beast of burden but probably a valuable riding animal. Anatomical and morphometrical analyses suggest a hybrid confirmed by the ancient DNA analyses resulting in the presence of a dromedary in the maternal and a Bactrian camel in the paternal line. 
 
 Citation: Galik A, Mohandesan E, Forstenpointner G, Scholz UM, Ruiz E, Krenn M, et al. (2015) A Sunken Ship of the Desert at the River Danube in Tulln, Austria. PLoS ONE 10(4): e0121235. https://doi.org/10.1371/journal.pone.0121235 Academic Editor: Serge Muyldermans, Vrije Universiteit Brussel, BELGIUM Received: October 30, 2014; Accepted: January 28, 2015; Published: April 1, 2015 Copyright: © 2015 Galik et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited Data Availability: All relevant data are within the paper and its Supporting Information files. Funding: EM and RF were supported by the Austrian Science Foundation FWF project (P24706-B25) to P. Burger, who is recipient of an APART fellowship (11506) from the Austrian Academy of Sciences. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. Competing interests: The authors have declared that no competing interests exist. 
 
 Introduction Prior to the construction of a shopping center, excavations rescued the archaeological heritage of Tulln (Fig. 1) in Lower Austria in 2006 and 2007. The permission for the excavation was issued by the Federal Office for the Protection of Monuments in Vienna. Significant finds revealed large parts of the medieval and the early modern town including several building plots of an urban district [1]. An ensemble of buildings was specified as the historically mentioned tavern called “Auf der Rossmühle” near the large market place [2]. An abandoned cellar associated with this plot yielded the most extraordinary find of a complete skeleton originating from a large animal. The partly excavated skeleton was suspected to be a large horse or cattle, until the first author undoubtedly identified it as the remains of a camel (Fig. 1). The skeleton was found in a typical post-mortal posture with its neck bent backwards and drawn-up extremities [3]. PPT PowerPoint slide 
 
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 larger image TIFF original image Download: Fig 1. In situ view of the camel in the cellar. Inserted map indicates the geographical position of the town Tulln in Austria with a triangle. https://doi.org/10.1371/journal.pone.0121235.g001 Archaeological camel finds in Central Europe are not as unusual as one may expect covering a chronological span from Roman period till early modern age [4]. Yet, to this date only isolated bones or partly preserved camel skeletons have been found, e.g. at the Amphitheatre Viminacium in Serbia [5]. Its chronological position and the completeness of the skeleton highlights the uniqueness of the Tulln record. It is the first complete camel skeleton found in Central Europe and Central European territories under the control of the Ottoman Empire [6,7,8,9,10,11,12], apart from the complete skeleton of a dromedary recovered from the sediments of the Theodosius harbor on the European part of Istanbul [13]. In this study we combine archaeological as well as complementary morphologic and molecular genetic analyses to unravel the mystery of the sunken ship of the desert at the river Danube in Tulln. 
 
 Archaeological and Historical Context The backfill of the cellar yielded masses of domestic refuse like animal bones and ceramics (e.g. plates, pans and flagons), pieces of a tiled stove and enameled pipe bowls which date the filling in the early modern period, although the main part of the excavated material is not completely studied yet. Metal finds allowed a narrow and high resolution dating of the context. A coin—a so called “Rechenpfenning” depicts the countenance of King Louis XIV of France was dated from 1643 to 1715. A medicinal bottle made of lead contained the remedy “Theriacum” produced in the chemist’s shop “Apotheke zur Goldenen Krone” (approximately 1628/1665) in Vienna. These two exceptional finds place the filling containing the skeleton to the late 17th century. Tulln was affected by floods of the Danube and plagues such as the Black Death, which drastically reduced the urban development [14] and a reduction of the inhabited area occurred due to conflagration and demolition of houses in the early 17th century [2]. Two building plots got new owners at the end of the century around the 1690, certainly the time when the cellar was backfilled, offering enough space to bury such a big cadaver in the center of the town. In summer 1683, Ottoman troops tried to reach Vienna and combed the region south of the Danube. The surrounding of Tulln was besieged by the army division, but the town itself never was conquered [14]. On the contrary, written sources describe a peaceful surrender of two prisoners of war. The Ottoman commander released the imperial ambassador and his secretary in Tulln in August 1683 and obviously, as proven by written sources, the Ottoman troops were in contact with the inhabitants of Tulln [14]. 
 
 Discussion The identification of isolated archaeological faunal remains often entails difficulties and uncertain results. Although the Tulln specimen was preserved almost completely and subjected to intensive morphological and morphometrical examinations, the species identification turned out to be problematic due to its intermediate position between Bactrian camel and dromedary (Fig. 7). Today, the two domestic species exist in geographically different areas. The dromedary ranges from North Africa to Western Asia and Australia [29,30,31] while the Bactrian camel occurs in the Far East and Central Asia [32]. Different adaptations and ecotypes as well as different types of breeds cause variability of the habitus and size for various purposes, as riding animals, beasts of burden or for exploitation of meat and dairy products [29,33,34]. The sexual dimorphism also effects the discrimination of camel remains [35] as male dromedaries are about 10% heavier and 10 cm taller at shoulder height [36,37]. Depending on the animal’s age, castration certainly influences the skeleton and induces uncertainties in the identification of Bactrian camel versus dromedary bones. However, castrated camels are easier to handle and more effective as working animals and in meat production [34,38]. The cranial part of the pelvic symphysis certainly indicated a male Tulln specimen, but the smooth and weakly structured surface of the cranial part of the corpus pubis probably might hint to a castration. The two species are able to interbreed, which results in larger, more powerful and efficient hybrid offspring [33,39,40,41]. Crossbreeding probably took first place in Assyria at the beginning of the 1st millennium BC [42] and this technique continued over Antiquity towards the modern times [4,34]. Hybridization was improved as the Arabs went into Iran and Central Asia [29,33]. Obviously, such hybrids were of great importance in the Ottoman troops mainly for transportation but also as riding animals [43,44,45]. The first appearance of a camelry, soldiers fighting with bows on camelback, had been recorded during the invasion of Xerxes in Greece 481 BC [39]. Today, hybridization facilitates improved milk and wool yield in hybrid Tulu or Nar camels from Middle Eastern and Central Asian countries. Commonly, two hybridizing methods are recognized, Kurt-nar (dromedary female x Bactrian male) and Kez-nar (Bactrian female x dromedary male) followed by F 1 -backcrossing with either dromedary for increased milk productivity or Bactrian camel for a higher wool yield and cold resistance [46,47]. A remarkable and relished sport is common in western Turkey, where highly valued male Tulus fight against each other in strictly regulated “camel wrestling” competitions (47, 48). A lack of osteo-morphological characterization of hybrids [4,18,48] in literature and reference collections prevents a discrimination of hybrids and parental species. Nevertheless, the dromedary has more slender and the Bactrian camel broader long bones [17,40]. The long bones of the Bactrian camel are more robust while the dromedary has longer zygopodial and metapodial bones [32]. Most of the long bones of the Tulln specimen are rather long while their robustness addresses a Bactrian camel. Especially radius and femur have an intermediate position between both species (Fig. 7). Not so for the metacarpals, which are short and broad like in Bactrian camel (Fig. 7). The morphological features incorporate characteristics of both species, confirmed by the results of the nuclear DNA analyses as the Tulln-specimen turned out to be an F 1 hybrid Tulu with a dromedary in its maternal and evidences of a Bactrian camel in its paternal line. We compared the detected mitochondrial haplotpye with a modern sequence data set of dromedaries throughout their distribution range. The hybrid camel from Tulln shares a main mitochondrial haplotype, which is common in nearly 70% of the modern dromedaries [49]. The results confirm previously described traditions that people in Anatolia imported Bactrian males to interbreed with Arabian female dromedaries but Bactrian females were and are of less interest [41,46]. Pathological deformities on the scapulae and the upper arm bones require special attention because they occur symmetrically. Probably, the periostitis ossificans at the mandible is caused by pressure of the harness. The periostitically changed bone surface medial on both shoulder blades perhaps originates in high strains on the musculus serratus ventralis that could be related to frequent coming down and going up of the individual to let the camel rider on and up (Fig. 8). The lesions under the exposed tuberculum intermedium (Fig. 9) are not easily explainable but might be related to stress-born deformities of the overlaying tendons of origin of the musculus biceps brachii. However, if the Tulln-specimen was used as a beast of burden, a higher frequency of stress related osteological changes should be expected. In this case only a few deformities mainly at the acropodium (Fig. 5, Fig. 6) and some bent dorsal spines of the thoracic vertebrae are proven. Such an ensemble of pathologies rather indicates a camel that was used as a riding animal and kept under good conditions. It is impossible to reconstruct how the camel did arrive within the town walls of Tulln. Its appearance might be linked to an exchange of local people with the troops or the Ottoman army simply left it behind. Apparently, the citizens took it inside the town, where they probably kept and displayed it as an “exotic animal”. Further archeological research might answer this question. It seems quite conceivable that being not familiar with behavioral and feeding habits, the scarcity of food in wartimes, people did not keep it for long. As the camel died the citizens of Tulln did not exploit this alien animal, which the soldiers of the Ottoman troops certainly would have done. Exploitation of camel flesh especially in times of need was absolutely necessary. The dismemberment of the carcasses certainly is a reason for the scarce preservation of camel finds in general and is indicated by bones with butchering marks in particular. However, the citizens buried this camel in a typical post-mortal position, and together with rubbish in the remnants of a cellar that was leveled. The skeleton remained there for more than 300 years to raise questions in the future. Finally, it served as a perfect study example for a successful cooperation of the complementary fields of archaeological and genetic sciences. The camel specimen from Tulln is the first archaeozoologically and genetically confirmed evidence of a Tulu hybrid camel. 
 
 Acknowledgments We are grateful to H. Emmerig, Institut für Numismatik und Geldgeschichte, University of Vienna, who analyzed the coins. We thank the camel owners and collectors for providing the precious samples, Gräfliche Kamelheimat (AT), O. Abdelhadi and B. El-Zain (SD), and G. Konuspayeva (KZ). We are very grateful to J. Peters for access to the Paleogenetic Core Facility, ArchaeoBioCenter, LMU Munich (DE) and to C. Schlötterer, Institute of Population Genetics, Vetmeduni Vienna for general support. 
 
 Author Contributions Conceived and designed the experiments: AG EM GF UMS ER MK PB. Performed the experiments: AG EM GF UMS ER MK PB. Analyzed the data: AG EM GF UMS ER MK PB. Contributed reagents/materials/analysis tools: AG EM GF UMS ER MK PB. Wrote the paper: AG EM GF UMS ER MK PB. ||||| In 2006 construction began on a new shopping centre in Tulln. The works unearthed various archaeologically valuable objects that were salvaged during rescue excavations. Among these objects was also the complete skeleton of a large mammal. 
 
 Large mammal uncovered during excavations in Tulln 
 
 "The partly excavated skeleton was at first suspected to be a large horse or cattle," says archaeozoologist Alfred Galik from the Institute for Anatomy, Histology and Embryology at the University of Veterinary Medicine Vienna. "But one look at the cervical vertebrae, the lower jaw and the metacarpal bones immediately revealed that this was a camel." 
 
 Camel bones have been found in Europe dating back to the Roman period. Isolated bones or partly preserved skeletons are known from Mauerbach near Vienna as well as from Serbia and Belgium. But a complete camel skeleton is unique for Central Europe. 
 
 "Exotic animal" died in Tulln 
 
 In addition to horses, the Ottoman army also used camels for transportation and as riding animals. In cases of scarcity, the soldiers also ate the animal's flesh. But the skeleton found in Tulln was complete. "This means that the animal was not killed and then butchered. It may have been acquired as part of an exchange," says first author Galik. "The animal was certainly exotic for the people of Tulln. They probably didn't know what to feed it or whether one could eat it. Perhaps it died a natural death and was then buried without being used." 
 
 Camel was a hybrid 
 
 Extensive DNA analysis showed that the animal was a hybrid: its mother was a dromedary and its father a Bactrian camel. The genetic diagnosis confirmed what the scientists saw morphologically. Several of the physical features were that of a dromedary, others of a Bactrian camel. "Such crossbreeding was not unusual at the time. Hybrids were easier to handle, more enduring and larger than their parents. These animals were especially suited for military use," Galik explains. 
 
 The camel was male, around seven years old and most likely castrated. 
 
 Find dated to the 17th century 
 
 Besides animal bones, the excavations also unearthed ceramic plates and other items. A coin - a so-called "Rechenpfenning" - from the time of Louis XIV dates the find to the years between 1643 and 1715. A medicinal bottle containing Theriacum, a medieval remedy from the chemist's shop "Apotheke zur Goldenen Krone" in Vienna was also found at the site. This pharmacy existed between 1628 and 1665, which helped date the site with further precision. 
 
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 The article „A sunken ship of the desert at the river Danube in Tulln, Austria", by Alfred Galik, Elmira Mohandesan, Gerhard Forstenpointner, Ute Maria Scholz, Emily Ruiz, Martin Krenn and Pamela Burger was published in the scientific journal PLOS ONE. DOI 10.1371/journal.pone.0121235 http://dx. plos. org/ 10. 1371/ journal. pone. 0121235 
 
 About the University of Veterinary Medicine, Vienna 
 
 The University of Veterinary Medicine, Vienna in Austria is one of the leading academic and research institutions in the field of Veterinary Sciences in Europe. About 1,300 employees and 2,300 students work on the campus in the north of Vienna which also houses five university clinics and various research sites. Outside of Vienna the university operates Teaching and Research Farms. http://www. vetmeduni. ac. at 
 
 Scientific Contact (archaeozoology): 
 
 Dr. Alfred Galik 
 
 Institute of Anatomy, Histology and Embryology 
 
 University of Veterinary Medicine Vienna (Vetmeduni Vienna) 
 
 T +43 1 25077-2521 
 
 alfred.galik@vetmeduni.ac.at 
 
 Scientific Contact (genetics): 
 
 Elmira Mohandesan, PhD 
 
 Institute of Population Genetics 
 
 University of Veterinary Medicine Vienna (Vetmeduni Vienna) 
 
 T +43 681 815 033 12 
 
 elmira.mohandesan@vetmeduni.ac.at 
 
 Released by: 
 
 Susanna Kautschitsch 
 
 Science Communication / Public Relations 
 
 University of Veterinary Medicine Vienna (Vetmeduni Vienna) 
 
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– The study on the Austrian find calls it a "sunken ship of the desert" and an "exotic animal." What exactly was found in a cellar containing centuries-old trash during excavations for a planned shopping center in Tulln: the full skeleton of a 17th-century camel. That a complete skeleton was found is unique, but the presence of camel bones is less so: The Ottoman army used the creatures for transportation and, in some cases, food, and their bones have been found in the region. But the completeness of the skeleton shows "the animal was not killed and then butchered," says archaeozoologist and study author Alfred Galik in a press release. He suggests it may have been traded or left behind by the troops, possibly following their siege of Vienna in 1683, reports the BBC. The locals "did not exploit this alien animal" for food upon its death, as the Ottoman troops "certainly would have done," write the authors in the study, published in PLOS One. "Exploitation of camel flesh especially in times of need was absolutely necessary," and the regularity with which that happened is "a reason for the scarce preservation of camel finds in general." What researchers were able to glean from this find: The male camel was the hybrid offspring of a female dromedary and male Bactrian camel and probably castrated, which would have made him easier to handle. Crossbreeding, which was not unusual for the time, made for larger animals that "were especially suited for military use," Galik explains. To wit, the study points to bone defects that indicate the animal was harnessed and ridden, reports the BBC. (Oil workers recently made a big find in Siberia.)