The latest steppe mammoths (Mammuthus trogontherii (Pohlig)) and associated fauna on the Late Middle...

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The latest steppe mammoths (Mammuthus trogontherii (Pohlig)) and associated fauna on the Late Middle Pleistocene steppe at Nosak, Kostolac Basin, Northeastern Serbia Vesna Dimitrijevi c a, * , Nemanja Mrdji c b , Miomir Kora c b , Seimi Chu e , Dejan Kosti c c , Mladen Jovi ci c b , Bonnie A.B. Blackwell d, e a Laboratory for Bioarchaeology, Department of Archaeology, Faculty of Philosophy, University of Belgrade, Serbia b Institute of Archaeology, Belgrade, Serbia c Military Medical Academy, Belgrade, Serbia d Department of Chemistry, Williams College, Williamstown, MA 01267-2692, USA e RFK Science Research Institute, Glenwood Landing, NY 11547-0866, USA article info Article history: Available online xxx Keywords: Late Middle Pleistocene Mammuthus trogontherii Loess Nosak Kostolac Serbia abstract In 2012, bones and tusks of mammoths and remains of other large mammals scattered in a line 130 m long have been discovered in the loess deposits at the Nosak mound in the Kostolac Basin (Northeastern Serbia). Preliminary analyses indicate that remains come from Mammuthus trogontherii, Equus ferus mosbachensis and Cervidae indet. Steppe mammoth remains originate from at least four individuals, three adults, and one juvenile. They include the largest steppe mammoth tusks ever reported. The mammalian remains were mostly found disarticulated, except for few articulated skeleton elements. Individual bones and teeth were encased within thick carbonate concretions. From their composition, morphology, and manner of formation, these carbonate concretions resemble the characteristic loess dollsthat form in loess. Bones were exposed to subaerial weathering for some time before burial. Some damage on the bones indicates biogenic agents of their fragmentation and dispersal. Animals were probably attracted to the spot by a water source in the vicinity, such as a spring or intermittent stream. The bones and teeth accumulated at a single level within the loess likely deposited in MIS 6, just above the upper of two paleosols formed during MIS 7. A sample from a mammoth molar plate has been ESR dated at 192 ± 5 ka, correlating with the very early MIS 6, near the MIS 6/7 boundary. This makes the Nosak mammoth the most recent occurrence of steppe mammoths in Europe, and the sole nd dated to MIS 6. © 2015 Elsevier Ltd and INQUA. All rights reserved. 1. Introduction In the Kostolac Basin, coal mining began at the end of the 19th Century. Three large mine pits, Kostolac, Cirikovac, and Drmno, have been successively opened, which has drastically changed the landscape in the area. Open pits have exposed geological forma- tions with thickness of over 50 m, deposited over 7 million years, from the Late Miocene until the present day (Stevanovi c, 1977; Vu ceti c, 2010; Stojanovi c et al., 2012). Open-pit coal mining requires removing the overburden covering the Neogene coal, which is overlain by Pleistocene alluvial sediment and a sequence of loessepaleosols. These deposits formed in the course of the Pannonian Basin uplift, followed by complex environmental changes. Although changing, this envi- ronment constantly provided the conditions for living to commu- nities that boasted the biggest land mammals, such as proboscideans and rhinoceroses, and large ungulate herds, like horses and bisons, with the carnivores preyed upon them. Conse- quently, nding these animals' remains when the deposits which house them were excavated in such enormous quantity was not surprising. Their recognition and recovery, however, is extremely difcult, given that large backhoes relentlessly cut through the sediment, leaving behind poorly accessible cliffs, 6e7 m high. Eventually, fossil remains, if the backhoe blades and wheels have * Corresponding author. E-mail addresses: [email protected] (V. Dimitrijevi c), nemanjamrdjic@gmail. com (N. Mrdji c), [email protected] (M. Kora c), [email protected] (S. Chu), [email protected] (D. Kosti c), [email protected] (M. Jovi ci c), [email protected] (B.A.B. Blackwell). Contents lists available at ScienceDirect Quaternary International journal homepage: www.elsevier.com/locate/quaint http://dx.doi.org/10.1016/j.quaint.2015.06.025 1040-6182/© 2015 Elsevier Ltd and INQUA. All rights reserved. Quaternary International xxx (2015) 1e14 Please cite this article in press as: Dimitrijevi c, V., et al., The latest steppe mammoths (Mammuthus trogontherii (Pohlig)) and associated fauna on the Late Middle Pleistocene steppe at Nosak, Kostolac Basin, Northeastern Serbia, Quaternary International (2015), http://dx.doi.org/10.1016/ j.quaint.2015.06.025

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Quaternary International

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The latest steppe mammoths (Mammuthus trogontherii (Pohlig)) andassociated fauna on the Late Middle Pleistocene steppe at Nosak,Kostolac Basin, Northeastern Serbia

Vesna Dimitrijevi�c a, *, Nemanja Mrdji�c b, Miomir Kora�c b, Seimi Chu e, Dejan Kosti�c c,Mladen Jovi�ci�c b, Bonnie A.B. Blackwell d, e

a Laboratory for Bioarchaeology, Department of Archaeology, Faculty of Philosophy, University of Belgrade, Serbiab Institute of Archaeology, Belgrade, Serbiac Military Medical Academy, Belgrade, Serbiad Department of Chemistry, Williams College, Williamstown, MA 01267-2692, USAe RFK Science Research Institute, Glenwood Landing, NY 11547-0866, USA

a r t i c l e i n f o

Article history:Available online xxx

Keywords:Late Middle PleistoceneMammuthus trogontheriiLoessNosakKostolacSerbia

* Corresponding author.E-mail addresses: [email protected] (V. Dimitrij

com (N. Mrdji�c), [email protected] (M. Kora(S. Chu), [email protected] (D. Kosti�c),(M. Jovi�ci�c), [email protected] (B.A.B

http://dx.doi.org/10.1016/j.quaint.2015.06.0251040-6182/© 2015 Elsevier Ltd and INQUA. All rights

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a b s t r a c t

In 2012, bones and tusks of mammoths and remains of other large mammals scattered in a line 130 mlong have been discovered in the loess deposits at the Nosak mound in the Kostolac Basin (NortheasternSerbia). Preliminary analyses indicate that remains come from Mammuthus trogontherii, Equus ferusmosbachensis and Cervidae indet. Steppe mammoth remains originate from at least four individuals,three adults, and one juvenile. They include the largest steppe mammoth tusks ever reported. Themammalian remains were mostly found disarticulated, except for few articulated skeleton elements.Individual bones and teeth were encased within thick carbonate concretions. From their composition,morphology, and manner of formation, these carbonate concretions resemble the characteristic “loessdolls” that form in loess. Bones were exposed to subaerial weathering for some time before burial. Somedamage on the bones indicates biogenic agents of their fragmentation and dispersal. Animals wereprobably attracted to the spot by a water source in the vicinity, such as a spring or intermittent stream.The bones and teeth accumulated at a single level within the loess likely deposited in MIS 6, just abovethe upper of two paleosols formed during MIS 7. A sample from a mammoth molar plate has been ESRdated at 192 ± 5 ka, correlating with the very early MIS 6, near the MIS 6/7 boundary. This makes theNosak mammoth the most recent occurrence of steppe mammoths in Europe, and the sole find dated toMIS 6.

© 2015 Elsevier Ltd and INQUA. All rights reserved.

1. Introduction

In the Kostolac Basin, coal mining began at the end of the 19thCentury. Three large mine pits, Kostolac, �Cirikovac, and Drmno,have been successively opened, which has drastically changed thelandscape in the area. Open pits have exposed geological forma-tions with thickness of over 50 m, deposited over 7 million years,from the Late Miocene until the present day (Stevanovi�c, 1977;Vu�ceti�c, 2010; Stojanovi�c et al., 2012).

evi�c), nemanjamrdjic@gmail.�c), [email protected][email protected]

. Blackwell).

reserved.

�c, V., et al., The latest steppem, Kostolac Basin, Northeaster

Open-pit coal mining requires removing the overburdencovering the Neogene coal, which is overlain by Pleistocene alluvialsediment and a sequence of loessepaleosols. These depositsformed in the course of the Pannonian Basin uplift, followed bycomplex environmental changes. Although changing, this envi-ronment constantly provided the conditions for living to commu-nities that boasted the biggest land mammals, such asproboscideans and rhinoceroses, and large ungulate herds, likehorses and bisons, with the carnivores preyed upon them. Conse-quently, finding these animals' remains when the deposits whichhouse them were excavated in such enormous quantity was notsurprising. Their recognition and recovery, however, is extremelydifficult, given that large backhoes relentlessly cut through thesediment, leaving behind poorly accessible cliffs, 6e7 m high.Eventually, fossil remains, if the backhoe blades and wheels have

ammoths (Mammuthus trogontherii (Pohlig)) and associated fauna onn Serbia, Quaternary International (2015), http://dx.doi.org/10.1016/

V. Dimitrijevi�c et al. / Quaternary International xxx (2015) 1e142

not smashed them totally, remain exposed, only until the next passby the backhoe. Thus, although large volumes of sediment havebeen excavated, no records of fossil skeletons or significant isolatedfinds of bones or teeth have occurred until 2009.

Expansion of the mines in the Kostolac Basin threatens Vim-inacium, a large Roman city and legionary fort (Fig. 1b). Therefore,since 1973, and especially since the youngest pit mine Drmno hasbeen opened, an archaeological team has been involved in rescueexcavations and protecting cultural heritage. Thus, in 2009, ar-chaeologists were on the spot when an almost complete steppemammoth (Mammuthus trogontherii (Pohlig)) skeleton wasdiscovered (Kora�c et al., 2010; Lister et al., 2012). They were able toexcavate, and, in cooperation with paleontologists, save most ofthat skeleton. The same is true for the remains at the Nosak site. In2012, 1900 m to the north and 20 m above the previous find,mammoth bones and tusks were seen sticking out of a profile madeby a large backhoe excavating the overburden at the Drmno coalmine. This new discovery of mammoths and other mammals' re-mains, as well as their preliminary description and dating, form thesubject of this paper.

2. The regional setting and the Nosak site

The site is located approximately 90 km east of Belgrade, and5 km north of Po�zarevac, within a low relief area known as theKostolac Basin (Fig. 1a, b). The basin is bordered by the DanubeRiver to the north, the Velika Morava River to the west, while to the

Fig. 1. Geography of the Nosak site. a. Satellite view of the South Europe region showing the2012, showing Drmno open coal pit, the location of Roman city and legionary fort Viminacium2009 (1) and the Nosak site (2). Google Earth March 28, 2012.

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east the basin stretches to the Po�zareva�cka greda, the foothills ofthe Goluba�cke planine (mountains). The basin hosts Late Miocenelignite. Nosak refers to a mound at the edge of loess plateau wherethe loessepaleosol sequence is ~26 m thick, thicker than in thesurrounding area. For example, where the mammoth skeleton wasdiscovered in 2009 the loessepaleosol sequence was about 20 mthick.

At the mammoth skeleton site, the Neogene and Pleistocenesuccession has been studied (Fig. 2a; Lister et al., 2012). TheNeogene sequence contains coal, gray and yellow fine grained siltysands, and orange and brown medium to coarse grained, cross-bedded sands (Fig. 2a, 1e3). The overlying Pleistocene sequencecontains alluvial sediment at the base, capped by loess with inter-calated paleosols (Fig. 2a, 4e5). The skeleton occurred in the lowerPleistocene fluvial sediment (Fig. 2b), while the Nosak remains layin the upper Pleistocene loess deposits (Fig. 2c).

At the Nosak site, in a profile orthogonal to the layer with thePleistocene mammal fossils, the loessepalaeosol sequence wassampled at 10 cm intervals for low-field magnetic susceptibility,sedimentological and geochemical analyses, while samples for TL/OSL luminescence dating were removed as blocks from the fossil-iferous layer (Markovi�c et al., 2014). The loess and palaeosol strat-igraphic units, designated as L and S respectively, were numberedin order of increasing age, following an initial stratigraphic labelingscheme first applied in the Chinese loess stratigraphic model(Kukla, 1987; Kukla and An, 1989). The mammalian remains accu-mulated just above the double pedocomplex S2, which had most

location of the Kostolac Basin. Google Earth February 10, 2015. b. The Kostolac Basin in(outlined in a rectangle), the site where a steppe mammoth skeletonwas recovered in

ammoths (Mammuthus trogontherii (Pohlig)) and associated fauna onn Serbia, Quaternary International (2015), http://dx.doi.org/10.1016/

Fig. 2. Steppe mammoth remains in the Kostolac Basin. a. A stratigraphic column in the Drmno open pit coal mine. 1e3: Neogene, Late Upper Miocene. 1. Coal and coal-rich clays. 2.Gray silty sand with intercalated gray yellow sand. 3. Cross bedded orange and brown sand. 4e5: Quaternary, Pleistocene. 4. Alluvial sediment. 4a: Riverbed sediment: gravel, yellowsand, gray sandy gravel, and clay, with mammoth remains at 58 m AMSL; 4b: Wetland sediments: silts, silty sands with mollusc and mammal remains. 5. Loess. 5a: Marshy loess:gray-brownish clayey silts; 5b: Land loess: gray yellow silt. 6: Holocene. Anthropogenic deposits (archaeological layers and humus) (adapted from Lister et al., 2012. Approximateposition of new steppe mammoth remains has been added). b. The location of the Nosak site in loess deposits. A view from the site where the steppe mammoth skeleton wasdiscovered in 2009 (Photo: N. MrCi�c, June 2012). c. The location where the steppe mammoth skeleton was discovered in 2009: A view from the Nosak site. (For interpretation of thereferences to colour in this figure legend, the reader is referred to the web version of this article.)

V. Dimitrijevi�c et al. / Quaternary International xxx (2015) 1e14 3

likely formed duringMarine Isotope Stage 7 (MIS 7), when the loesshorizon L2, likely corresponding to MIS 6, began to be deposited. Inthe profile, L2 was 600 cm thick, and intercalated with severalweakly developed paleosols. Above lie the last interglacial pedo-complex, S1, and the last glacial loess, L1, with the modern soil atthe top. TL/OSL luminescence dates estimated theminimum ages ofthe palaeontological layer to be >150 ka (Markovi�c et al., 2014).

3. Material and methods

3.1. Field methodology

In excavating the mammalian remains at the Nosak site, stan-dard archaeological field methodology was followed: Each spec-imen was uncovered, photographed, and its spatial positionrecorded using total station. Fossil remains were left in situ until thesediment removal was completed in order to establish whetherparticular skeletal elements were articulated and how the bones

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had accumulated. Unfortunately, the taphonomy of the accumula-tion precluded definitive answers about the bone or taxonomicidentification, the number of animals present, and their relation-ships in the field.

At the Nosak site, all the faunal remains were covered with acoating of calcium carbonate approximately 1 cm thick. In fact,most bones and teeth were completely encased in calcium car-bonate concretions (Fig. 3a, b). The carbonate crust has the samecomposition and most likely originates from the same processes asloess dolls. Namely, a well known feature in loessepalaeosol se-quences, carbonate concretions, or loess dolls, form after sedi-mentary leaching (Barta, 2011a, 2011b). Concretions mostly occurin interesting globular shapes, the reason why they are called“dolls”.

For many specimens, the carbonate concretions completelyconcealed their morphology and did not allow the skeletal elementinside to be identified. Therefore, even the number of specimens forthe mammal teeth and bones at Nosak could not be counted. Only

ammoths (Mammuthus trogontherii (Pohlig)) and associated fauna onn Serbia, Quaternary International (2015), http://dx.doi.org/10.1016/

Fig. 3. Bones coated in carbonate and loess dolls. a. What appears to be a mammalianlong bone coated in carbonate from the Trench 97. b. A cross section of horse meta-tarsal from the Trench 101 coated in carbonate. c. A globular concretion with a ho-mogenous silty core. d. A globular concretion with a bone fragment at its core.

Table 1U in the Nosak Mammoth.

Sample U concentration (ppm)

Enamel Inner dentine Outer dentine

AT27en1 5.95 70.07 59.67AT27en2 4.20 70.72 70.72AT27en3 2.98 59.86 30.28AT27en4 10.18 63.71 16.28AT27en5 5.09 60.18 17.71AT27en6 4.10 59.73 20.72AT27en7 8.33 270.08 52.48AT27en8 6.45 73.29 53.59

AT27 mean 5.91 90.96 40.18(n ¼ 8) ± 2.23 67.89 19.99

Uncertainty ± 0.02 0.02 0.02

Detection ~ 0.01 0.01 0.01limitsa e 0.02 0.02 0.02

Water conc'ns 0.02 0.05 0.05(wt%) ± 0.02 0.02 0.02

a Typical detection limits depend on the sample's mass and tissue type.

V. Dimitrijevi�c et al. / Quaternary International xxx (2015) 1e144

the largest mammoth bones and tusks were easily recognizable,simply because no other animal matches a mammoth in size. In thefew identifiable specimens, dimensions were measured using atape measure with a precision of 1 cm.

Beside mammoth cranial and postcranial elements and othermammalian bones and teeth encased in large concretions, mostlyless than 10e100 cm in size, numerous small globular concretions,1e10 cm in size, were collected during the excavation. Somecontain bone fragments, while others lack organic remains (Fig. 3c,d).

Excavated bones and large carbonate concretions that mightpreserve mammalian remains inside them were hand-collectedand stored at Scientific Center at Viminacium. A few impressivespecimens of mammoth skulls and tusks were removed in blockscut from the sediment and displayed at the Paleontological Gallerybuilt in 2014 near the Scientific Center.

Under the carbonate cover which had been broken in somespecimens, the bones appeared white and brittle. The carbonatecover was mostly firmly attached to the bone surface. Attempts todetach the crust mechanically, as a rule, damaged the bone.

3.2. X-ray and CT scanning

In order to establish which specimens were worth preparing, X-ray and CT scans were performed. Both analyses permitted mea-surements for bones and paleontological observations about theirmorphology. Digital X-ray radiography (Shimadzu RAD Speed Pro,Shimadzu Europa GmbH, Duisburg, Germany) was used to X-raythe specimens at a tube voltage of 75 kV, a tube current of 56 mA,with a film-focus distance of 110 mm. The CT examination wasperformed using 64 section MDCT scanner (Aquilion, ToshibaMedical Systems, Otawara, Japan), using a tube voltage of 135 kVp, a

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tube current of 200e300 mA, a slice thickness of 0.5 mm, a gantryrotation 0.75 s, and a bone reconstruction algorithmwith awindowwidth/window level of 2000/500. For the multiplanar re-constructions, the raw data files from bone and teeth CT's weretransferred to a Vitrea workstation (Vital Images, USA).

3.3. ESR dating methodology

An enamel plate for ESR dating was removed from amandibularmammoth tooth, Specimen M2eK1, found in Trench 99, from thesediment 7e14 cm below a mandible, and 19e20 cm below thesediment surface respectively. Using the standard ESR proceduresfor a Class 10,000 clean lab, eight subsamples were ESR preparedand dated independently from the enamel plate (Table 1). To pre-vent contamination, all glass and plastic ware were rinsed with 6 MHCl (aq) once and >15 times with doubly distilled deionized waterto remove the Cl� ions (Blackwell, 1989).

After drawing and photographing the sample in order to retain avisual record before its destruction, the dentine and enamelthicknesses were measured in 20 places with a hand-held digitalCD-4C caliper. After subsampling the fragment with a hand-helddiamond-tipped Dremel drill, all the dentine was drilled off theenamel surface and saved for NAA. After the thicknesses for eachenamel subsampleweremeasured in 20e30 spots using aMitutoyoID-C112E micrometer, 20 mm was shaved off both surfaces toremove the enamel dosed by a radiation. After powdering theenamel to 150e74 mm (200e400 mesh) with an agate mortar andpestle, 20.0 ± 0.1 mg of enamel was weighed into each of 14e16aliquots, and any extra saved for NAA. All aliquots, except one (the0 Gy dose), were irradiated using a 60Co g source with doses from0 to 20 kGy at ~16.0e17.0 krad/s, and were annealed at 90 �C for 3.0days to remove any unstable interference created by the irradiation.

All aliquots' ESR spectra were analyzed in a JEOL RE1X spec-trometer at 2 mW power under 100 kHz field with a modulationamplitude of 0.1 mT in the X-band. The spectra were scanned at336.0 mT using an 8.0 min sweep time and 0.1 s time constant withreceiver gains set to maximize peak heights. With EWWIN v. 4.5,derivative spectra for the HAP peak at g ¼ 2.0018 were collected,stored, and measured without deconvolution, because the twoHAP signal components follow identical growth (Skinner et al.,2001).

ammoths (Mammuthus trogontherii (Pohlig)) and associated fauna onn Serbia, Quaternary International (2015), http://dx.doi.org/10.1016/

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At the field site in August 2012, two sediment samples werecollected from the same sedimentary unit that had held themammoth tooth. The two bulk sediment samples were powderedto 50e200 mesh, and analyzed by NAA for U, Th, and K to deter-mine the sedimentary dose rate, Dext(t) (Table 2). Samples of eachenamel subsample and its associated dentine and cementumsamples were powdered to 50e200 mesh and analyzed mainly forU. Then, ~0.7 g of sediment and >1.0 g of powdered tooth tissuewere analyzed byNAA. For U, a 60 s delayed neutron count followeda 60 s irradiation and a 10 s cooling delay. One enamel and onedentine per tooth, and all sediment samples, were counted for K for20.0 min in a g counter after a 60 s irradiation and 24e30 h delay,and for Th, after a 1.0 h irradiation, and a 7e10 day delay (Blackwell,1989). Results were standardized to NIST 1633B. The sedimentsamples' water loss could not be analyzed, because the sedimenthad been exposed before collection.

Table 2Sedimentary radioactivity for the Nosak Mammoth.

By plotting the ESR peak heights against the accumulateddose, A S and its errors were calculated with Vfit assuming asaturating exponential fit weighted by 1/I2. Rosy v. 1.4.2, whichcorrects for b and g, but not a, attenuation due to the waterconcentration, tissue density, and thickness from sediment(Brennan et al., 1997), was used to calculate the dose rates, ages,and their errors for all subsamples. The a/g efficiency factor wasset to 0.15 ± 0.02, and the initial (234U/238U)0 ratio, to 1.20 ± 0.02(Blackwell, 2006). Since Rn loss is hard to measure accurately, itwas first assumed to be 0 ± 0 vol% and then modeled by recal-culating the ages with different loss parameters. Volumetricallyaveraged DextðtÞwere calculated by averaging the dose rates fromall adjacent sediment and bone, weighted by their volume withinthe 30 cm g sphere as estimated from field photos. Ramped boxmodels were used to assess the time-averaged cosmic and sedi-mentary dose rates. Weighted mean ages were calculated byweighting the subsample's age by the inverse of its error(Blackwell, 2006).

4. Results

4.1. Excavation and spatial distribution of the mammal remains

As noted above, large backhoes remove the overburden abovethe Miocene coal in the open-pit mine at Drmno by making cutsapproximately 6e7 m tall (Fig. 4a). These cuts are steep and fur-rowed by traces of the large buckets on the backhoe (Fig. 4b). The

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archaeological field survey crew discovered broken mammothtusks and bones in one such cuts 19 m below the land surface(Fig. 4c). In July 2012, paleontological excavations started intrenches approximately 10 m wide positioned in a line 150 m long(Figs. 5a, b and 6). The excavated area covered 900 m2.

Given the carbonate coatings, many remains have not beenidentified yet. Trench 97 (Fig. 7) yielded a fragment of amammoth skull with the right tusk in the alveolus, resting on itsdorsal side. Several meters to the northeast, a complete left tusk,most likely from the same skull, was found. Near the skull, a leftscapula was lying, complete but with ragged proximal andlateral rims and a longitudinally broken scapular spine. Belowthe right tusk tip sat a fused radius and ulna, and two frag-mented thoracic vertebrae. Another thoracic vertebra lay sepa-rated from this group but together with a few unidentifiablebones in concretions. Because all these bones lay close to oneanother on a single horizon, and all came from a large adult withno repeated elements, they (Fig. 7) probably originated from oneanimal.

To the east, Trenches 102and76yieldedmanyconcretions,mostly10e25 cm long. One concretion housed a cervid tooth. Separate fromthe pile of concretions, three articulated bones of a largemammal thesize of a horse were found. Trench 109 contained a fragmentedmammoth skull with the right tusk in the alveolus, resting on itsdorsal side. Severalmeters to thenortheast, a complete left tusk,mostlikely fromthesameskull,was found.Near theskull, a left scapulawaslying, complete but with ragged proximal and lateral rims and alongitudinally broken scapular spine. Below the right tusk tip sat afused radius and ulna, and two fragmented thoracic vertebrae.Another thoracic vertebra lay separated from this group but togetherwith a few unidentifiable bones in concretions. Because all thesebones layclose tooneanotherona singlehorizon, andall came fromalarge adult with no repeated elements, they (Fig. 7) probably origi-nated from one animal.

To the east, Trenches 102 and 76 yielded many concretions,mostly 10e25 cm long. One concretion housed a cervid tooth.Separate from the pile of concretions, three articulated bones of alargemammal the size of a horse were found. Trench 109 containeda fragmented mammoth tusk that laid near a cluster of fragmentsthat may come from a skull, but are too fragmented to be identified(Fig. 9).

The largest concentration of remains occurred scatteredNNEeWSW over some 10 m in Trench 101 (Fig. 10). Besidesmammoth ribs and long bones, there were remains of othermammals. A few long, curved concretions with circular cross-sections resemble deer antlers. In another pile were a tibia,metatarsal, thoracic vertebra, and third phalanx from a horse. Onyet another lay ribs, curved elements that are probably antlers,and a long bone from an unidentified mammal. A horse femur(Fig. 11) was found separately, laying on an unidentifiable bone (?vertebra).

Trench 108 produced no mammal remains. In this area, theuniformity of loess deposit was disturbed by the appearance of finegravel 2e5 mm in diameter extending NNEeWSW, parallel to theconcretions in Trench 101. They suggest a nearby spring or anintermittent stream.

In Trench 103, three mammoth ribs were found with a fewmoreunidentifiable bones encased in concretions. In Trench 110, con-cretions and several unidentified bones of different sizescompletely covered by carbonate were found. They were scatteredand cannot be mutually related.

Trench 99 produced another mammoth skull (Fig. 8), amandible, and a tusk fragment. These cranial elements mostprobably originated from a single animal. Lying on its caudal side,the facial portion of the skull is preserved, alongwith the zygomatic

ammoths (Mammuthus trogontherii (Pohlig)) and associated fauna onn Serbia, Quaternary International (2015), http://dx.doi.org/10.1016/

Fig. 4. The discovery of the mammoth and other mammalian remains in loess depositsat the Nosak site. Photos: N. MrCi�c, May 2012. a. A large backhoe at work at the Drmnocoal mine observed from the surface. b. The location for the damaged bones in theprofile at the Nosak site. c. A mammoth tusk transversely cut by the large backhoe.

Fig. 5. Excavations at the Nosak site. a. The test trench on the edge of the profile(Drone aerial image, June 2012). b. The final excavation phase (Photo: M. Jovi�ci�c,September 2012).

V. Dimitrijevi�c et al. / Quaternary International xxx (2015) 1e146

arches and orbits. The skull had been damaged prior to the sedi-ment deposition. In contrast, the mandible was complete, lying onits ventral side with teeth in both left and right branch, but it wastransversally bisected and badly fragmented when the machinesleveled the surface for the excavations.

At the western end, Trench S74 was opened to ascertain that nomore mammal remains lay to the west. Since the trenches werewidened to the north to encompass all remains and concretions,

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likely all mammal remains accumulated in the areawere excavated,except for few specimens destroyed by the overburden removal.

4.2. X-ray and CT scanning

Three specimens were both X-rayed and CT scanned. The firstone (Fig. 12) was encased in a curved concretion with a length of26 cm along its outer curvature. On both the X-ray and CT scans, notonly is the outline of the encased specimen's visible, but also itsinternal structure, positively identifying it as a young mammothtusk. Namely, elephantid tusks have cone-shaped internal lamellaethat develop as the tusk grows (Rountrey et al., 2012). At its brokenend the tusk has a diameter of 4 cm, which is too small for massivetusks of adults, thus, suggesting a juvenile.

The second specimen (Fig. 13) was a bone encased in a straightconcretion 31.5 cm long. Broken during the excavation, it showedthe cross-section typical of a mammalian long bone. The bone'soutline in cross-section and its arrangement of compact andcancellous tissue point to an ungulate metapodial. Measuring thebone's length, articular and diaphysis diameters on the X-rayidentified it as a horse metatarsal. CT scans permitted more precisemeasurements with a precision of 1 mm. CT scans also show theoutline of the bone more clearly and allowed it to be rotated asneeded for measuring defined points. Following dimensions weremeasured: length (307mm), medio-lateral breadth of the proximalend (64 mm), medio-lateral breadth of the diaphysis in the middle(43 mm), medio-lateral breadth of the distal end (66 mm), andmedio-lateral breadth of the articular end (57 mm).

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Fig. 6. The excavation area at the Nosak site, August, 2014 (Drawing: N. Mrdji�c).

Fig. 7. Trench 97, with remains of a mammoth. The left tusk sits in the forefront. A splintered long bone with a crenulated edge occurs on the right, while further toward the edge ofTrench is a scapula. A skull with the right tusk in the alveolus, several vertebrae, a radius, and an ulna are also visible. A thoracic vertebra is to the left of the right tusk (Photo: N.MrCi�c, August 2012).

V. Dimitrijevi�c et al. / Quaternary International xxx (2015) 1e14 7

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Fig. 8. The skull of a mammoth in Trench 99. The skull is lying on its caudal side withthe tusk alveoli in the front view (Photo: M. Jovi�ci�c, August 2012).

Fig. 9. The tusk of a mammoth in Trench 109 (Photo: N. Mrdji�c, August 2012).

V. Dimitrijevi�c et al. / Quaternary International xxx (2015) 1e148

The third specimen was encased in a leaf-shaped concretion22 cm long. Although the concretion was broken, exposing thespecimen's cross-section neither observing the break, nor X-rayand CT analyses identify it. In cross-section, the specimen is whiteand homogenous, unlike any organic tissue. The specimen ends in awell rounded, blunt point, while its body is ornamented with anetwork of tendrils. The outline visible on the X-ray and CT scansdoes not match to any skeletal element.

4.3. Taxonomic identification

Preliminary analyses indicate that remains come from mam-moths, a horse, and a cervid. Generic attribution of the probosci-dean remains to Mammuthus is based on the morphology of thetusks' alveoli, the shape and size of tusks, and the molars' features.

On the skull from Trench 97, the alveoli are mutually paralleland close to each other. In the sole proboscidean genus that alsoexisted in the area in the same time, Palaeoloxodon, they aredivergent at their endings and mutually quite apart (Osborn,1942). In the Nosak skull, the tusk's torsion also matches thatseen in Mammuthus. Namely, the right tusk from the same skulldeparts from the alveolus forward and downward, then spiralsupwards and inwards. In Palaeoloxodon, however, the tusks

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extend straight downward and then swing slightly upward withina single plane. The tusk shape in M. trogontherii andM. primigenius is similar, but usually more curved in the latter.Moreover, the size of the Nosak tusks indicates M. trogontherii. Infact, its very large size indicates a very large, male, steppemammoth. Including the carbonate cover, the diameter of thetusk at the base measured 28 cm. Since the thickness of the car-bonate crust measured 14 mm at one spot, the tusk diameter atthe base is approximated to about 25 cm. This exceeds the tuskdiameter for a very large male steppe mammoth from Steinheimin Germany, which measured 217 mm (Dietrich, 1912), or a tuskfragment from a Peloponnese steppe mammoth with maximumdiameter of 203 mm (Athanassiou, 2012).

Sexual dimorphism in mammoths is reflected in the consider-ably larger size of the males. In that respect, one of the mostindicative features is the tusk diameter at the base. Unfortunately,the variation range for M. trogontherii males and females has notbeen established, due to the paucity of fossils, unlike it has been forM. primigenius tusks for the fossil population from Siberia, forwhich Vereshchagin and Tikhonov (1986) gave a range of47e90 mm in females and 89e180 mm in males.

The length of the two tusks from Trench 97 also point to a steppemammothmale of exceptional size. Along the outer curve, the righttusk measured 3.26 m from the alveolus to the distal tip, whileprobably only a small part of the tusk tip has been lost. The alveolusis approximately 46 cm long. Thus, the total tusk length withalveolus approximates 3.72 m. The left tusk is 4.2 m long. Since itwas detached from the alveolus, it was not possible to concludehow long the unexposed portion of the tusk was. Complete steppemammoth tusks for comparison with the Nosak tusks are rare.From England in the Early Middle Pleistocene, the West Runtonskeleton, had a complete right tusk at 280 cm long, counting theexposed part of 211 cm and 69 cm of the tusk buried in the skull(Lister and Stuart, 2010).

The preservation of the Nosak remains largely precluded theidentification of the dental features that are the most importantfor establishing taxonomic identity. Cheek teeth were found onlyin a mandible from Trench 99. This mandible was transverselybisected, and thus, badly damaged while leveling the plateau inpreparation for the paleontological excavation. It broke intopieces, which are brittle and prone to more degradation. How toreconstruct the mandible and its teeth remains a conservationproblem not yet solved. Hence, only a tooth fragment wasavailable for study (Fig. 14). The enamel in this tooth fragment isapproximately 3 mm thick. While it was not possible to measurethe average occlusal thickness across the tooth whole length, thisvalue roughly compares with M. trogontherii, and is too thick forM. primigenius. Maglio (1973) notes that it averages 1.5e3 mm inM. trogontherii, but only 1e2 mm in M. primigenius. InM. trogontherii found in Kostolac in 2009, the enamel thicknessaveraged 2.9e3.1 mm for the lower third molars (Lister et al.,2012). Number of plates in the Nosak molar fragment is 5. Itslamellar frequency counted as the average number of plates on10 cm from the base and top of the crown, between the buccaland lingual sides (after Maglio, 1973) could only be assumed to 6plates on buccal, and 7 plates on lingual side, both at the baseand on the occlusal surface on the crown, which gives an esti-mated average of 6.5. In 2009 find, third molars lamellar fre-quency measured 6.8 (Lister et al., 2012). Lamellar frequency inM. trogontherii is 5e8, but 7e12 in M1eM3 for M. primigenius(Maglio, 1973).

A fragmented skull, complete left and right tusks, at least fivevertebrae, several ribs, a scapula, a fused radius and ulna in Trench97 originated from an adult mammoth (Fig. 7). The remains fromanother adult included a fragmented skull, a mandible, and one

ammoths (Mammuthus trogontherii (Pohlig)) and associated fauna onn Serbia, Quaternary International (2015), http://dx.doi.org/10.1016/

Fig. 10. Scattered mammalian remains in Trench 101 (Photo: M. Jovi�ci�c, August 2012).

V. Dimitrijevi�c et al. / Quaternary International xxx (2015) 1e14 9

tusk in Trench 99. The third mammoth is represented by a frag-mented tusk from Trench 109. The tusk from Trench 109 could notbelong to the mammoth from Trench 97, since the latter's tuskswere almost complete. The tusk from Trench 109 could not belongto the mammoth from Trench 99, because the latter's diameter wasmuch larger. As noted above, the X-rays and CT scans also showedthe presence of juvenile. Thus, the presence of the fourth individualhas been established.

Fig. 11. A horse femur in Trench 101 (Photo: N. MrCi�c, August 2012).

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The equid remains included a thoracic vertebra, femur, tibia,metatarsus, and third phalanx. Although coated in carbonate, thefemur was identified as an equid (Fig. 11) by its third trochanter,and the third phalanx, by its arched outline. The best opportunity

Fig. 12. The juvenile mammoth tusk fragment from Trench 109. a. The specimencoated in carbonate (Photo: N. Mrdji�c) b. A radiograph of the specimen (MilitaryMedical Academy, Belgrade). c. A CT scan of the specimen (Military Medical Academy,Belgrade).

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Fig. 13. A horse metatarsal from Trench 109. a. The metatarsal coated in carbonate(Photo: N. Mrdji�c) b. A radiograph of the metatarsal (Military Medical Academy, Bel-grade). c. A CT scan of the metatarsal (Military Medical Academy, Belgrade).

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for taxonomic identification of the Nosak horse remains came fromthe X-rays and CT scans of metatarsal. The metatarsal's size in-dicates a large Middle Pleistocene horse related to Equus ferusmosbachensis von Reichenau (Eisenmann and Beckouche, 1986;Eisenmann, 1991).

The presence of a cervid is supposed only from a few long,curved antler-shaped carbonate concretions, and a tooth from aconcretion found in 102. Although most of the tooth was coveredwith carbonate, the observable portion showed the dental rugositycharacteristic in cervids (Cervidae indet.)

Fig. 14. A fragment of a mammoth molar from a mandible from Trench 99. a. Theocclusal surface. b. The buccal side.

4.4. Dates

Limited to just three taxa, Mammuthus trogontherii (Pohlig),Equus ferus mosbachensis von Reichenau, and Cervidae indet., thepaleontological association broadly correlates the Nosak site to theMiddle Pleistocene. However, the potential for dating is given bythe paleontological layer's stratigraphic position within a sequenceof loess accumulated during a glacial period, intercalated withpaleosols formed under interglacial conditions.

In the central Europe, including the southern rim of the Pan-nonian Plain and Kostolac Basin, loess deposition occurred duringthe Middle and Late Pleistocene (Raki�c, 1978; Buggle et al., 2009).The thickness of the loess cover and the completeness of sequencedepend on many regional and local factors. At the Nosak site, thefollowing succession was observed from bottom to top: a doublepedocomplex S2, a loess horizon L2, a paleosol S1, a loess horizonL1, and themodern soil S0. Mammal remains were found just abovethe upper of the double paleosols S2, or at the very bottom of thelower loess horizon L2. The double pedocomplex likely formedduring MIS 7, while the loess probably was deposited in MIS 6.Preliminary TL/OSL thermoluminescence dating for two samples

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from the paleontological layer gave a minimum age of >150 ka(Markovi�c et al., 2014).

To narrow the age range for the Nosak mammoths, onemammoth tooth fragment was ESR dated.

Eight subsamples were ESR dated independently from oneenamel plate (Table 1). Two bulk sediment samples were analyzedby NAA to determine the sedimentary dose rate, Dext(t) (Table 2).Their U concentrations agreewith expectations for loess containingcarbonate concretions, but the high Th concentrations suggest clayin the sediment, possibly as clay coatings on the grains. The high Kconcentrations hint that evaporation was high, and thus, that theclimate was fairly dry. The time-averaged cosmic and sedimentarydose rates together averaged 982 ± 63 mGy/y.

Except for AT27en4, the accumulated doses, A S0s were

consistent, and the ages agreed well among the subsamples

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(Table 3). At 10.18 ± 0.02 ppm, AT27en4 also had roughly doublethe U in its enamel of that found in any other subsample. Mostlikely, most of this U was absorbed very late in its history. None-theless, removing AT27en4 from the mean age (Table 3) does notchange the ages significantly, given the 1 s error. Therefore,AT27en4 is not an outlier to the distribution and is included in themean age of 192 ± 5 ka. Ranging from 2.98 to 10.18 ± 0.02 ppm inthe enamel, and from 16.28 to 270.08 ± 0.02 ppm in the dentine(Table 3), the variability in the U concentrations suggested that anisochron might be possible (e.g., Blackwell and Schwarcz, 1993).Such high U concentrations rarely occur in teeth that have expe-rienced early or linear U uptake (EU or LU models), making Uuptake more likely to have followed recent U uptake (an RUmodel). These values also hint that the tooth has spent some of itshistory exposed to saline or hypersaline water (Blackwell et al.,2002).

Table 3ESR ages for the Nosak Mammoth.

The isochron analysis for AT27 (Fig. 16) yielded a family ofisochrons with the external component of the accumulated dose,A ext at ~174e183 Gy. With an age of 181 ka, the RU isochron gavea time-averaged external dose rate, Dext(t) ¼ 1010 ± 245 mGy/y.Since the time- and volumetrically averaged external dose rate,

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Dext(t) ¼ 982 ± 63 mGy/y from the standard ESR analysis, the toothlikely experienced recent U uptake. Most of the eight subsamplesclustered on the modelled RU isochron with t ¼ 180 ka and p ¼ 10,but one sample, AT27en4, showed very recent U uptake, sug-gesting a somewhat older age with p ¼ 20. Although all the agesare preliminary pending the coupled ESR-230Th/234U analyses, theage for the tooth appears likely to be 192 ± 5 ka. This age corre-lates with the very early phases of MIS 6, near the MIS 6/7boundary.

5. Discussion

5.1. Taphonomy of the mammalian remains

At Nosak, the mammalian remains probably accumulated in asingle geologic event, making the site taphonomically intriguing.

The bones and teeth accumulated on a single horizon level,reflecting presumably a contemporary land surface. In instanceswhere several bones occurred together they were tightly packed,i.e. little, if any, sediment was found between them, suggesting theywere likely not deposited in separate episodes. Also, given that all

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Fig. 15. Mammoth bones in Trench 97. Photos: V. Dimitrijevi�c, August 2012. a. Fusedradius and ulna lying on fragmented vertebrae. b. The distal end of a radius with anincised semi-circular groove. c. A close up of the uncoated bone surface.

Fig. 16. The isochron analysis for AT27. The isochron analysis gave a family of isochronswith A ext ¼ 174e183 ± 11e15 Gy. The EU and LU isochrons gave Dext(t) values farexceeding that from the sedimentary NAA analyses. With an age of 181 ka, however,the RU isochron gave Dext(t) ¼ 1010 ± 245 mGy/y, which agrees well the volumetricallyaveraged external dose rate, DextðtÞ ¼ 982 ± 63 mGy/y, determined from the NAA an-alyses, the best U uptake model appears to have p y 10.

V. Dimitrijevi�c et al. / Quaternary International xxx (2015) 1e1412

the bones occurred in a sedimentologically homogeneous, siltmatrix, no signs of different depositional conditions were evidentduring the bone accumulation.

Nonetheless, the bones had been exposed to subaerial weath-ering for some time before burial. On the distal radius found inTrench 97, a small area of the bone surface (Fig. 15c) shows denselongitudinal cracks, filled with black mineral oxides. Bone crackingprobably occurred due to the atmospheric exposure, while themineral oxides deposition postdated the cracking. Formation of thecarbonate concretions occurred last when bones were alreadyburied under the loess.

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The Nosak bones were mostly found disarticulated, although afew skeleton elements were articulated: Three articulated bones ofa large mammal about the size of a horse occurred in Trench 102,and hind leg bones from a horse, in Trench 101. Moreover, althoughnot articulated, the mammoth cranial and postcranial skeletal el-ements in Trench 97, as well as the mammoth skull and mandiblefrom Trench 99 were found close to each other, only a few metersapart. The question remains: What agent(s) disarticulated, scat-tered, and broke the bones?

Some damage on the bones indicates biogenic agents. On ablade of the mammoth scapula found in Trench 97, a deep incisedgroove runs parallel to its cranial edge, and a large semi-circulargroove has been incised along its caudal edge. The splintered par-tial shaft of the mammoth long bone is found in the same trench. Itslongitudinal breaks are deeply serrated. Also, a semi-circulargroove has been incised close to the distal end of the mammothradius, and two more pits occur nearby (Fig. 15a, b). The grossmorphology of these marks resembles predator damage, but post-depositional processes have obscured these taphonomical traces,making their causes not easily discernable. Thus, traces frompredators' teeth or claws are not recognizable. Nonetheless, if themammalian remains had been exposed for any time sometime, onecan reasonably assume that they would attract local carnivores thatcould have dispersed at least some remains. Trampling by animals,and particularly mammoths, also might have happened, supposingthat four mammoths did not die all in the same time. A watersource in the vicinity, such as a spring or intermittent stream, assuggested by the gravel found in Trench 108, would keep drawingthe mammoths and other animals repeatedly to the site, causingthem to trample the bones of animals that died in preceding sea-sons, as is known for elephants' bone deposits studied in SouthAfrica (Haynes, 1988). A gravitation force could also add fossils'scatter, but the sedimentological data excludes any significanttransport by water.

5.2. The Nosak mammoths and M. trogontherii/M. primigeniusreplacement

After evaluating of data on the late occurrence of M. trogontheriiand the earliest appearance of M. primigenius in Europe, Lister andcolleagues (e.g., Lister, 2001; Lister and Sher, 2001; Lister et al.,2005) disputed the previously broadly accepted hypothesis onthe gradual transformation of M. trogontherii into M. primigenius.

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Instead, they proposed thatM. trogontheriiwas replaced in the LateMiddle Pleistocene by immigrants from Siberia, whereM. primigenius evolved some 200 ka earlier. Nosak finds mightcontribute to the challenge of dating this event of replacement.

The loessepaleosol stratigraphy (Markovi�c et al., 2014) and theESR enamel dates indicate that the age of Nosak mammoths isrelated to the latest occurrence of steppe mammoths in Europe.Namely, the steppe mammoth is last recorded in MIS 7, and mostprobably replaced by wooly mammoth in the next stage, MIS 6(Lister et al., 2005). For the occurrences of the latest steppe mam-moths in Europe Lister et al. (2005) cite Ilford and Brundon, En-gland, and Ehringsdorf, Germany. Ilford was correlated with MIS 7based on amino acid racemization (AAR) in shells (Bowen et al.,1989). Ehringsdorf was dated to MIS 7 by 230Th/234U analyses ontravertines (Blackwell and Schwarcz, 1986). Yet another importantsite related to this question is Lower Channel at Marsworth, inEngland, dated to either late MIS 7, or early MIS 6 (about190e150 ka), wheremolars with enamel plate counts characteristicfor both M. trogontherii and M. primigenius occurred in a singlestratigraphic level (Lister and Sher, 2001). Apparently, this repre-sents the only site yet discovered where both species coexisted.

The list of sites with finds of molars with morphology indis-putably assigned to M. primigenius in MIS 6 is much longer, andincludes sites in England, Belgium, and Italy. In Italy, in the sametime, Palaeoloxodon antiquus is abundant. The straight-tuskedelephant is also abundant throughout the Middle Pleistocene onthe Apennine Peninsula and in Greece, where M. trogontherii ispoorly represented (Palombo and Ferretti, 2005; Tsoukala et al.,2011). This is contrary to the situation in the Balkan region,where steppe mammoth remains are more abundant and thestraight-tusked elephant rarely has been found (Raki�c, 1977;Dimitrijevi�c and Pavlovi�c, 1987; Pavlovi�c and Dimitrijevi�c, 1987).

6. Conclusions

With an in situ assemblage of mammalian remains from aPleistocene steppe, Nosak is a valuable paleontological and tapho-nomically intriguing site. In fact, Nosak is the only Pleistocene siteof this kind in the region. Throughout the central Balkans, includingthe south Pannonian Basin, as well as throughout the mountainousarea between the Carpatho-Balkanids on the east to the Dinarideson the west, large mammal remains have mostly been found incaves or as transported material in alluvial deposits (Dimitrijevi�c,1997, 1998). Therefore, most of mammoth finds have beenreduced to individual, disarticulated bones and loose teeth, whichprovide little data compared to faunal assemblage found in situ.

Steppe mammoth remains, from at least four individuals, threeadults, and one juvenile, dominate this assemblage. One adult malewas very large, judging on its tusks. Along the right tusk's outercurve it measured 3.26 m from the alveolus to the distal tip, whileits left tusk, found separated from the skull, measured 4.2 m. Theleft tusk measured ~25 cm in diameter at its base. These are thelargest steppe mammoth tusks ever reported.

The bones and teeth accumulated on a single horizon within aloess unit, L2, just above the double pedocomplex, S2. A samplefrom a mammoth molar plate has been ESR dated at 192 ± 5 ka,correlating with the very early MIS 6, near the MIS 6/7 boundary.This makes the Nosak mammoth the most recent occurrence ofsteppe mammoths in Europe, and the sole find dated to MIS 6.

Finally, the Nosak site illustrates a rare phenomenon in whichthe mammalian remains were enveloped and preserved in car-bonate concretions. This unusual fossilization process hamperedthe identification of mammal remains and reduced their presen-tation in this paper to a preliminary report. However, it inspired theintroduction of X-ray and CT-scanning that showed promising first

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results for studying the encrusted bones and teeth, paving the wayfor further research.

Acknowledgements

We thank the Kostolac power plant for financing, and theViminacium archaeological team for performing, the excavations atthe Nosak site. The Ministry of Education and Science, the Republicof Serbia funded the projects “Bioarchaeology of Ancient Europe:Humans, animals and plants in the prehistory of Serbia” (III47001)and ”IRS e Viminacium, Roman city and military legion camp e

research of material and non material culture of inhabitants byusing the modern technologies of remote detection, geophysics,GIS, digitalisation and 3D visualization” (III 47018). The US NationalScience Foundation (ILI 9151111 to Anne R. Skinner), WilliamsCollege, RFK Science Research Institute, and McMaster NuclearReactor funded the ESR dating work. Jon Florentin, Israt Ahmed,and 2012e2014 RFKSRI lab crews assisted with the analyses. AlicePidruczny, McMaster University Nuclear Reactor, did the NAA. Wecordially thank the reviewers, Athanassios Athanassiou, Ministry ofculture, Athens, and Dick Mol, Het Natuurhistorisch, Rotterdam, fortheir helpful comments.

References

Athanassiou, A., 2012. A skeleton of Mammuthus trogontherii (Proboscidea, Ele-phantidae) from NW Peloponnese, Greece. Quaternary International 255, 9e28.

Barta, G., 2011a. The structure and origin of loess dolls: a case study from the loess-paleosoil sequence of Sütt}o, Hungary. Journal of Environmental Geography 4(1e4), 1e10.

Barta, G., 2011b. Secondary carbonates in loess-paleosoil sequences: a general re-view. Central European Journal of Geosciences 3 (2), 129e146.

Blackwell, B.A., 1989. Laboratory Procedures for ESR Dating of Tooth Enamel.McMaster University Department of Geology Technical Memo 89.2.

Blackwell, B.A.B., 2006. Electron spin resonance (ESR) dating in karst environments.Acta Carsologica 35, 123e147.

Blackwell, B.A., Schwarcz, H.P., 1986. U-series analyses of the lower travertine atEhringsdorf, DDR. Quaternary Research 25.2, 215e222.

Blackwell, B.A., Schwarcz, H.P., 1993. ESR isochron dating for teeth: a briefdemonstration in solving the external dose calculation problem. Applied Ra-diation & Isotopes 44, 243e252.

Blackwell, B.A.B., Skinner, A.R., Brassard, P., Blickstein, J.I.B., 2002. U uptake in toothenamel: lessons from isochron analyses and laboratory simulation experiments.In: Whitehead, N.E., Ikeya, M. (Eds.), Proceedings of the International Sympo-sium on New Prospects in ESR Dosimetry and Dating, Advances in ESR Appli-cations, vol. 18. Society of ESR Applied Metrology, Osaka, pp. 97e118.

Bowen, D.Q., Hughes, S., Sykes, G.A., Miller, G.H., 1989. Land-sea correlations in thePleistocene based on isoleucine epimerization in non-marine molluscs. Nature340, 49e51.

Brennan, B.J., Rink, W.J., McGuirl, E.L., Schwarcz, H.P., 1997. b doses in tooth enamelby “one-group” theory and the ROSY ESR dating software. Radiations Mea-surements 27, 307e314.

Buggle, B., Hambach, U., Glaser, B., Gerasimenko, N., Markovi�c, S., Glaser, I., Z€oller, L.,2009. Stratigraphy, and spatial and temporal paleoclimatic trends in South-eastern/Eastern European loessepaleosol sequences. Quaternary International196 (1), 86e106.

Dietrich, W.O., 1912. Elephas primigenius Fraasi, eine schwabische Mammutrasse.Jahreshefte des Vereins für Vaterl€andische Naturkunde in Württemberg 68,42e106.

Dimitrijevi�c, V., 1997. Upper Pleistocene mammals from cave deposits of Serbia.Geolo�ski anali Balkanskog poluostrva 61 (2), 179e370.

Dimitrijevi�c, V., 1998. Fossil remains of vertebrates in Serbian Cave deposits. In:Djurovi�c, P. (Ed.), Speleological Atlas of Serbia, Geographical Institute of theSerbian Academy of Sciences and Arts “Jovan Cviji�c”, Special Issues, vol. 52,pp. 51e57. Belgrade.

Dimitrijevi�c, V., Pavlovi�c, M., 1987. Kvartar. In: AnCelkovi�c, M. (Ed.), Geologija �sireokoline Beograda, I, Geologija i geodinamika. Zavod za regionalnu geologiju ipaleontologiju Rudarsko-geolo�skog fakulteta Univerziteta u Beogradu, Beograd,pp. 247e251 (in Serbian).

Eisenmann, V., 1991. Les chevaux quaternaires europ�eens (Mammalia, Peri-ssodactyla). Taille, typologie, biostratigraphie et taxonomie. Geobios 24,747e759.

Eisenmann, V., Beckouche, S., 1986. Identification and discrimination of metapodialsfrom Pleistocene and modern equus, wild and domestic. In: Meadow, R.H.,Uerpmann, H.P. (Eds.), Equids in the Ancient World, Beihefte zum TübingerAtlas des Vorderen Orients, A, 19, 1, pp. 117e163. Wiesbaden.

ammoths (Mammuthus trogontherii (Pohlig)) and associated fauna onn Serbia, Quaternary International (2015), http://dx.doi.org/10.1016/

V. Dimitrijevi�c et al. / Quaternary International xxx (2015) 1e1414

Haynes, G., 1988. Longitudinal studies of African elephant death and bone deposits.Journal of Archaeological Science 15, 131e157.

Kora�c, M., Markovi�c, Z., Milivojevi�c, M., Mol, D., 2010. The Kostolac mammoth,Serbia: an almost entirely complete skeleton of the steppe mammoth, Mam-muthus trogontherii. Quaternaire, Hors S�erie 3, 79e80.

Kukla, G., 1987. Loess stratigraphy in central China. Quaternary Science Reviews 6,191e219.

Kukla, G.J., An, Z., 1989. Loess stratigraphy in Central China. Palaeogeography,Palaeoclimatology, Palaeoecology 72, 203e225.

Lister, A.M., 2001. “Gradual” evolution and molar scaling in the evolution of themammoth. In: Cavarretta, G., Gioia, P., Mussi, M., Palombo, M.R. (Eds.), TheWorld of Elephants, Proceedings of the First International Congress, Rome,pp. 648e651.

Lister, A.M., Sher, A.V., 2001. The origin and evolution of the woolly mammoth.Science 294, 1094e1097.

Lister, A., Sher, A., van Essen, H., Wei, G.B., 2005. The pattern and process ofmammoth evolution in Eurasia. Quaternary International 126e128, 49e64.

Lister, A.M., Stuart, A.J., 2010. The West Runton mammoth (Mammuthus trogon-therii) and its evolutionary significance. Quaternary International 228 (1),180e209.

Lister, A., Dimitrijevi�c, V., Markovi�c, Z., Knezevi�c, S., Mol, D., 2012. A skeleton of‘steppe’ mammoth (Mammuthus trogontherii (Pohlig)) from Drmno, near Kos-tolac, Serbia. Quaternary International 276e277, 129e144.

Maglio, V.J., 1973. Origin and evolution of the Elephantidae. Transactions of theAmerican Philosophical Society 63, 1e149.

Markovi�c, S.B., Kora�c, M., MrCi�c, N., Buylaert, J.P., Thiel, C., McLaren, S.J., Stevens, T.,Tomi�c, N., Peti�c, N., Jovanovi�c, M., Vasiljevi�c, D.A., Sümegi, P., Gavrilov, M.B.,Obreht, I., 2014. Palaeoenvironment and geoconservation of mammoths fromthe Nosak loess-palaeosol sequence (Drmno, Northeastern Serbia): Initial re-sults and perspectives. Quaternary International 334e335, 30e39.

Osborn, H.F., 1942. Proboscidea, vol. 2. American Museum of Natural History, NewYork.

Please cite this article in press as: Dimitrijevi�c, V., et al., The latest steppemthe Late Middle Pleistocene steppe at Nosak, Kostolac Basin, Northeasterj.quaint.2015.06.025

Palombo, M.R., Ferretti, M.P., 2005. Elephant fossil record from Italy: knowledge,problems, and perspectives. Quaternary International 126, 107e136.

Pavlovi�c, M., Dimitrijevi�c, V., 1987. Pleistocenski sisari. In: AnCelkovi�c, M. (Ed.),Geologija �sire okoline Beograda, II, Fosilna fauna i flora. Zavod za regionalnugeologiju i paleontologiju Rudarsko-geolo�skog fakulteta Univerziteta u Beog-radu, Beograd, p. 295, 302e303.

Raki�c, M., 1977. Geneza i stratigrafija kvartarnih sedimenata u slivu Ju�zne i ZapadneMorave. Rasprave Zavoda za geolo�ska i geografska istra�zivanja, 18, Beograd.

Raki�c, M., 1978. Tuma�c Za OGK SFRJ, list Bela Crkva 1:100,000. Savezni geolo�skizavod, Beograd.

Rountrey, A.N., Fisher, D.C., Tikhonov, A.N., Kosintsev, P.A., Lazarev, P.A.,Boeskorov, G., Buigues, B., 2012. Early tooth development, gestation, and seasonof birth in mammoths. Quaternary International 255, 196e205.

Skinner, A.R., Chasteen, N.D., Shao, J., Blackwell, B.A., 2001. Q-band studies of theESR signal in tooth enamel. Quaternary Science Reviews 20 (5), 1027e1030.

Stevanovi�c, P., 1977. Oblast zapadno do Karpatobalkanida. In: Petkovi�c, K. (Ed.),Geologija Srbije, II-3 (Stratigrafija, kenozoik). Zavod za regionalnu geologiju ipaleontologiju Rudarsko-geolo�skog fakulteta. Univerzitet u Beogradu, Beograd,pp. 72e74.

Stojanovi�c, K., �Zivoti�c, D., �Sajnovi�c, A., Cvetkovi�c, O., Nytoft, H.P., Scheeder, G., 2012.Drmno lignite field (Kostolac Basin, Serbia): origin and palaeoenvironmentalimplications from petrological and organic geochemical studies. Journal of theSerbian Chemical Society 77, 1109e1127.

Tsoukala, E., Mol, D., Pappa, S., Vlachos, E., van Logchem, W., Vaxevanopoulos, M.,Reumer, J., 2011. Elephas antiquus in Greece: new finds and a reappraisal ofolder material (Mammalia, Proboscidea, Elephantidae). Quaternary Interna-tional 245, 339e349.

Vereshchagin, N.K., Tikhonov, A., 1986. A study on mammoth tusks. In: Proceedingsof the Zoological Institute, USSR Academy of Sciences, vol. 149, pp. 3e14 (inRussian).

Vu�ceti�c, M., 2010. Jame Kostola�ckog majdana: iz istorije srpskih ugljenokopa.Elektroprivreda Srbije, Beograd, Novi Sad.

ammoths (Mammuthus trogontherii (Pohlig)) and associated fauna onn Serbia, Quaternary International (2015), http://dx.doi.org/10.1016/