Jinhua Wang

Pan Nie

Xiaokang Li

Huakun Zuo

Benoît Fauqué

Zengwei Zhu
email: zengwei.zhu@hust.edu.cn

Kamran Behnia

Critical point for Bose-Einstein condensation of excitons in graphite

An exciton is an electron-hole pair bound by attractive Coulomb interaction. Short-lived excitons have been detected by a variety of experimental probes in numerous contexts. An excitonic insulator, a collective state of such excitons, has been more elusive. Here, thanks to Nernst measurements in pulsed magnetic fields, we show that in graphite there is a critical temperature (T = 9.2 K) and a critical magnetic field (B = 47 T) for Bose-Einstein condensation of excitons. At this critical field, hole and electron Landau sub-bands simultaneously cross the Fermi level and allow exciton formation. By quantifying the effective mass and the spatial separation of the excitons in the basal plane, we show that the degeneracy temperature of the excitonic fluid corresponds to this critical temperature. This identification would explain why the field-induced transition observed in graphite is not a universal feature of three-dimensional electron systems pushed beyond the quantum limit. Excitonic insulator | Bose-Einstein condensation | critical point | highmagnetic-field-induced transition

A macroscopic number of non-interacting bosons condense to a single-particle state below their degeneracy temperature (1). This phenomenon, known as the Bose-Einstein condensation (BEC) of bosons was unambiguously detected in ultra-cold atomic gases (2) seven decades after its prediction (3). The critical temperature for this phase transition depends on the mass m * and density n of bosonic particles (4):

kBT BEC = 3.31 2 m * n 2/3 [1]
In all known cases of BEC, the particles are composite bosons made of "elementary" fermions. This is the case of 4 He, which becomes superfluid below 2.17 K, and dilute cold atoms, which display BEC features below 0.17 µK (2). The difference in critical temperature reflects what is expected by Eq. 1. Denser fluids and lighter bosons have higher T BEC . The possible occurrence of BEC for excitons [bosonic pairs of electrons and holes (5)] has become a dynamic field of research in the past couple of decades (6,[START_REF] Eisenstein | Bose-Einstein condensation of excitons in bilayer electron systems[END_REF]. Individual excitons have been observed in semiconducting heterostructures stimulated by light creating electrons and holes in equal numbers. However, such excitons are ephemeral entities. The emergence of a collective state of spontaneously created excitons was postulated in the context of semimetal-to-semiconductor transition [START_REF] Jérome | Excitonic insulator[END_REF] and was dubbed an excitonic insulator (EI).

Condensation of excitons into a collective and thermodynamically stable state [START_REF] Kogar | Signatures of exciton condensation in a transition metal dichalcogenide[END_REF][START_REF] Lu | Zero-Gap semiconductor to excitonic insulator transition in Ta 2 NiSe 5[END_REF][START_REF] Butov | Cold exciton gases in coupled quantum well structures[END_REF][START_REF] Liu | Quantum Hall drag of exciton condensate in graphene[END_REF][START_REF] Li | Excitonic superfluid phase in double bilayer graphene[END_REF][START_REF] Burg | Strongly enhanced tunneling at total charge neutrality in double-bilayer graphene-WSe 2 heterostructures[END_REF][START_REF] Wang | Evidence of high-temperature exciton condensation in two-dimensional atomic double layers[END_REF][START_REF] Karni | Infrared interlayer exciton emission in MoS 2 /WSe 2 heterostructures[END_REF] would require three conditions: 1) a sufficiently large binding energy; 2) a lifetime exceeding the thermalization time and 3) a concentration high enough to allow a detectable degeneracy temperature. An independent issue is the identification of such a state in distinction from other collective electronic states of quantum matter. In the case of bulk 1T-TiSe2, scrutinizing the plasmon dispersion has revealed a signature of EI unexpected in the alternative Peierlsdriven charge density wave (CDW) [START_REF] Kogar | Signatures of exciton condensation in a transition metal dichalcogenide[END_REF]. Other indirect signatures of BEC transition have been reported in two-dimensional systems, such as quantum wells [START_REF] Butov | Cold exciton gases in coupled quantum well structures[END_REF], graphene [START_REF] Liu | Quantum Hall drag of exciton condensate in graphene[END_REF][START_REF] Li | Excitonic superfluid phase in double bilayer graphene[END_REF][START_REF] Burg | Strongly enhanced tunneling at total charge neutrality in double-bilayer graphene-WSe 2 heterostructures[END_REF] and transition metal dichalcogenides heterostructures [START_REF] Wang | Evidence of high-temperature exciton condensation in two-dimensional atomic double layers[END_REF][START_REF] Karni | Infrared interlayer exciton emission in MoS 2 /WSe 2 heterostructures[END_REF].

Here, we present the case of graphite subject to strong magnetic field where the existence of a thermodynamic phase transition is established [START_REF] Tanuma | Physics in High Magnetic Fields[END_REF][START_REF] Yaguchi | A high-magnetic-field-induced density-wave state in graphite[END_REF][START_REF] Fauqué | Basic Physics of Functionalized Graphite[END_REF]. We will show that a magnetic field of 47 T provides all necessary conditions for the formation of a BEC of excitons. At this field, the gap between the two penultimate Landau subbands vanishes. One of these subbands is electron-like and the other is hole-like. The combination of vanishing gap and the large density of states (DOS) at the bottom and top of the bands permits the formation of excitons. We will show that the mass and the the density of these excitons is such that they should become degenerate below a critical temperature of the order of 9.2 K. As the temperature is lowered, this collective state survives in a narrow field window, which widens with cooling at the lower end but not at the upper end of this window. The BEC scenario provides an explanation for this contrast. Increasing the field destroys the thermodynamic stability of electron-hole pairs. Decreasing it, on the other hand, leads to a reduction of the degeneracy, gradually pulling down the critical temperature. Questions which remained unanswered in the CDW scenario for this phase transition [START_REF] Yoshioka | Electronic phase transition of graphite in a strong magnetic field[END_REF][START_REF] Arnold | Charge density waves in graphite: Towards the magnetic ultraquantum limit[END_REF] find answers by this identification.

Graphite is a semi-metal with an equal density of electrons and holes (n = p = 3 × 10 18 cm -3 [START_REF] Brandt | Modern Problems in Condensed Matter Sciences[END_REF]). Above the quantum

Significance Statement

Bose-Einstein condensation of excitons (pairs of electrons and holes bound by Coulomb attraction) has become a dynamic field of research in the past couple of decades. Whilst individual excitons have been observed in many systems, a collective state of condensed excitons has proved to be more elusive. In this paper, by quantifying the mass and the density of the electron-hole pairs, we identify a critical temperature of 9.2 K and a critical magnetic field of 47 T as the cradle of the Bose-Einstein condensation of excitons in graphite. Our identification radically revises the nature of the field-induced phase transition in graphite and its boundaries. 1. A critical point in (field, temperature) plane (A) As magnetic field is increased, the gap ∆ between the electron spin-up and the hole spin-down subbands evolves from negative to positive. Both subbands are evacuated at 47 T and ∆ = 0.

T > 9.2 K E F T = 9.2 K T < 9.2 K Λ d ∆ < 0 A d < Λ d ~Λ d > Λ ∆ = 0 ∆ > 0 ∆ Fig.
Two other subbands with the same level index and opposite spin polarities remain occupied. (B) As the temperature is lowered, the thermal de Broglie wavelength Λ becomes longer. At 9.2 K, it becomes comparable to the interexciton distance d and BEC of exciton occurs. limit of 7.4 T, electrons and holes are both confined to their lowest Landau levels [START_REF] Zhu | Nernst effect and dimensionality in the quantum limit[END_REF][START_REF] Schneider | Using the de Haas-van Alphen effect to map out the closed three-dimensional Fermi surface of natural graphite[END_REF], which are each split to two spin-polarized subbands. A rich phase diagram consisting of distinct field-induced phases emerges above ∼20 T, which was documented by previous studies [START_REF] Fauqué | Two phase transitions induced by a magnetic field in graphite[END_REF][START_REF] Zhu | Graphite in 90 T: Evidence for strong-coupling excitonic pairing[END_REF][START_REF] Zhu | Magnetic field tuning of an excitonic insulator between the weak and strong coupling regimes in quantum limit graphite[END_REF][START_REF] Leboeuf | Thermodynamic signatures of the field-induced states of graphite[END_REF]. Our focus, here, is the identification of the peak transition temperature as the cradle of the excitonic instability. Theoretical calculations by Takada and Goto [START_REF] Arnold | Charge density waves in graphite: Towards the magnetic ultraquantum limit[END_REF][START_REF] Takada | Exchange and correlation effects in the three-dimensional electron gas in strong magnetic fields and application to graphite[END_REF], based on the SWM model [START_REF] Slonczewski | Band Structure of Graphite[END_REF][START_REF] Mcclure | Band structure of Graphite and de Haas-van Alphen effect[END_REF] of band structure and including self-energy corrections, predicted that the electron spin-up and the hole spin-down subbands simultaneously cross the Fermi level at ∼ 53 T. The first result of the present study is to confirm such a simultaneous crossing of the two subbands occurring at a slightly different field, namely 47 T, and its coincidence with the peak critical temperature of the fieldinduced phase.

Fig. 1 presents a sketch of the proposed scenario. As the magnetic field increases, the gap between the spin-up subband of electrons and the spin-down subband of holes changes sign. At a critical field of 47 T not only the gap vanishes but also the two subbands empty. For fields exceeding 47 T, the formation of electron-hole pairs costs a finite energy. When the field sufficiently larger than 47 T, electron-hole pairs break up. Tuning down the temperature, on the other hand, will increase the thermal de Broglie wavelength. The BEC condensation will occur when the exciton wavelength becomes comparable to the interexciton distance, set by carrier concentration (4). The critical temperature of 9.2 K would be the degeneracy temperature of bosonic excitons in our picture.

We carried out a study of the Nernst effect [START_REF] Behnia | Nernst effect in metals and superconductors: A review of concepts and experiments[END_REF] in pulsed fields on Kish graphite samples. The Nernst effect has proved to be an extremely sensitive probe of Landau spectrum in compensated semimetals like graphite. Quantum oscillations are most prominent in the Nernst response and dominate the nonoscillating background [START_REF] Zhu | Nernst quantum oscillations in bulk semi-metals[END_REF]. However, measuring a Nernst signal in pulsed fields is challenging and a study of URu2Si2 [START_REF] Levallois | On the destruction of the hidden order in URu 2 Si 2 by a strong magnetic field[END_REF] is the unique known case prior to the one pre- Fig. 2 shows our Nernst data near the critical point (see the supplement (35) for data extended up to 60 T). The signal smoothly evolves upon cooling. Below 4 K, the low-field anomaly becomes similar to what was reported in a previous study of the Nernst effect below 45 T [START_REF] Fauqué | Nernst response of the Landau tubes in graphite across the quantum limit[END_REF]. Our extended data reveal additional information.

First of all, the peak near 47 T is the only one detected up to 60 T. This indicates that the evacuation of the two Landau subbands occur simultaneously. In other words, the separation in magnetic field is too small to be detected by experiment. This interpretation is consistent with the vanishing Hall conductivity observed near 47 T [START_REF] Akiba | Possible excitonic phase of graphite in the quantum limit state[END_REF].

The second observation is that this peak suddenly splits to two distinct anomalies when T < 9.2 K (See Fig. 2B,D,E). Finally, it is remarkable that the Nernst peak disappears for T > 35 K (See fig. 2C ). This temperature dependence allows us to quantify the high-field effective mass.

Fig. 3 compares the temperature dependence of the Nernst peaks near 8 T and near 47 T. What sets the thermal evolution of the amplitude of a quantum oscillation is the effective mass and the B/T ratio. The heavier the electrons, the faster the decay of the oscillating signal with warming. The larger the B/T ratio, the slower the decay. In this context, it is sriking to see that the high-field peak vanishes faster with warming than D R A F T the low-field one. Quantitatively, using the Lifshitz-Kosevich formula for thermoelectric quantum oscillations, Ω(T)osc ∝ [αX coth(αX) -1]/ sinh(αX) [START_REF] Morales | Thermoelectric power quantum oscillations in the ferromagnet UGe 2[END_REF][START_REF] Xu | Quantum oscillations in the noncentrosymmetric superconductor and topological nodal-line semimetal PbTaSe 2[END_REF], where α = 2π 2 kB/e and X = m * T /B, we find that the effective mass m * 47T = 0.48 m0 and m * 8T = 0.05 m0 (Fig. 3B), where m0 is the bare electron mass. The low-field value is consistent with the mass extracted from Shubnikov-de Haas measurements [START_REF] Yaguchi | A high-magnetic-field-induced density-wave state in graphite[END_REF]. Thus, there is a 10-fold field-induced enhancement in cyclotron mass of carriers. Note that recent specific heat measurement (40) find a field-induced enhancement of the DOS.

BEC occurs when the interBoson distance falls below the thermal de Broglie wavelength. In liquid 4 He, for example, the interatomic distance is 0.358 nm and the thermal de Broglie wavelength can be estimated taking the mass of each He atom. The two length scales become equal at 5.9 K. The BEC condition which corresponds to dexciton = Λexciton/1.38 (4) occurs at 3.1 K (41) to be compared with the superfluid critical temperature (2.17 K). The difference can be quantitatively explained by taking to account interactions, which lead to an effective mass larger than the bare mass [START_REF] Vakarchuk | Effective mass of 4 He atom in superfluid and normal phases[END_REF].

In our case, the interexciton distance and the de Broglie length are both anisotropic. According to the most extensive set of de Haas-van Alphen data [START_REF] Schneider | Using the de Haas-van Alphen effect to map out the closed three-dimensional Fermi surface of natural graphite[END_REF], the radius of Fermi surface along the c-axis is seven to nine times longer than in the basal plane. The effective mass is also very anisotropic and the ratio of out-of-plane to in-plane masses is estimated to exceed 90 [START_REF] Mcclure | Band structure of Graphite and de Haas-van Alphen effect[END_REF]. In the basal plane, the two relevant length scales can be estimated unambiguously. The frequency of quantum oscillations yields the interparticle distance in the basal plane, which is 18 nm for both electrons and 21 nm for holes (Table 1). Thus, the interexciton distance in the basal plane is dexciton = 19.5 ± 2 nm. The exciton mass would be twice the cyclotron mass resolved at 47 T, therefore, mexciton=0.96 m0. Using these numbers, one finds that the BEC condition dexciton = Λexciton/1.38 ( 4) is satisfied when D R A F T T = 8 K. This is remarkably close to the critical temperature of 9.2 K detected by our experiment. As seen in Fig. 4C, in graphite, the two length scales are two orders of magnitude longer than in 4 He and in both the experimentally observed critical temperature is close to where the BEC expected.

E F E 3 E 1 H H ' H ' H H '
In a compensated semimetal, charge neutrality does not impede a concomitant evolution of the density of electrons and holes with increasing magnetic field across the quantum limit. This is indeed what happens in semimetallic bismuth at high magnetic fields: at 30 T, the carrier density increases to more than five times its zero field value [START_REF] Zhu | Emptying Dirac valleys in bismuth using high magnetic fields[END_REF]. However, this is unlikely to happen in graphite because of its band structure [START_REF] Slonczewski | Band Structure of Graphite[END_REF][START_REF] Mcclure | Band structure of Graphite and de Haas-van Alphen effect[END_REF] (see Fig. 4A,B). When carriers are confined to the lowest Landau subbands, the DOS steadily increases due to the degeneracy of Landau levels. In a compensated metal, this can occur either by an enhancement in the concentration of electrons and holes, by an enhancement in mass, or a combination of both. Now, in graphite (in contrast to bismuth), electron and hole ellipsoids are aligned parallel to each other and their dispersion is similar. Moreover, and crucially, both electron and hole bands are half-filled along the kz. As a result, the room for any significant modification of the Fermi wavevector along the orientation of magnetic field and a change in carrier density is small. Thus, our analysis safely assumed that carrier density does not change between 7 T and 47 T.

Carrier F ⊥ (T )

A ⊥ (10 The boundaries of the EI phase shown in Fig. 4D are strikingly similar to the theoretical expectations [START_REF] Jérome | Excitonic insulator[END_REF][START_REF] Kozlov | The metal-dielectric divalent crystal phase transition[END_REF]. The left (low-field) boundary evolves to a mean-field expression for critical temperature Tc(B) = T * exp(- [START_REF] Yaguchi | A high-magnetic-field-induced density-wave state in graphite[END_REF][START_REF] Iye | High-magnetic-field electronic phase transition in graphite observed by magnetoresistance anomaly[END_REF] and the evolution of the DOS with magnetic field governs the evolution of the phase transition. This expression fails as the critical point is approached, leading to the saturation of the critical temperature. In contrast, on the right (high-field) side, the destruction of the field-induced state is abrupt and the critical temperature is pinned to a magnetic field of 53 T. This field does not correspond to the evacuation of any Landau level, as shown by the absence of any anomaly in our data. In the BEC scenario, it corresponds to the unbinding of the electron-hole pair by magnetic field (see the sketches in the Fig. 4D).

B * B ), which is a BCS- type formula k B Tc(B) = 1.14E F exp(- 1 N (E F )V )
Note that only at B = 47 T the critical temperature corresponds to the degeneracy temperature of excitons and the transition is, strictly speaking, a BEC condensation . When the magnetic field exceeds 47 T, the exciton binding energy becomes lower than the band gap and the order is destroyed by unbinding. On the other hand, decreasing the magnetic field diminishes the DOS, the screened Coulomb attraction between electrons and holes and the transition occurs well below the degeneracy temperature.

A BEC picture of field-induced phase transition would explain its presence in graphite in contrast to its absence in other semimetals pushed beyond the quantum limit [START_REF] Zhu | Landau spectrum and twin boundaries of bismuth in the extreme quantum limit[END_REF]. While the one-dimensional spectrum is a generic feature of the three-dimensional electron gas confined to its lower Landau level, exciton formation is not. In most semimetals with heavy atoms, the electric permittivity is large. Therefore, Coulomb attraction between holes and electrons is attenuated, hindering the formation of excitons. The electric permittivity in bismuth, for example, is twenty times larger than in graphite (see the supplement for more discussions(35)). Another difference between graphite and bismuth is the evolution of mass and carrier density across the quantum limit. The unavoidable enhancement in DOS due to Landau level degeneracy leads to an increase in carrier density in bismuth and an increased mass in graphite. As a consequence, the latter becomes a strongly correlated electron system at high magnetic fields.

One open question is the origin of the larger Nernst signal in the EI state in the vicinity of the critical point. Any quantitative analysis, however, requires a more complete set of data in order to quantify the magnitude of the transverse thermoelectric conductivity αxy, whose amplitude reflects the ratio of entropy to magnetic flux [START_REF] Behnia | Nernst effect in metals and superconductors: A review of concepts and experiments[END_REF][START_REF] Bergman | Theory of dissipationless Nernst Effects[END_REF]. Availability of DC fields above the present ceiling of 45 T would lead to multiprobe studies of the critical point unveiled by the present study.

In summary, we carried out pulsed-field Nernst measurements in graphite up to 60 T. We found a 47 T anomaly in the Nernst response and identified it as the result of the simultaneous evacuation of two Landau subbands, an electron-like and a hole-like one. The Nernst anomaly suddenly bifurcates to two distinct anomalies marking the boundaries of the field-induced state below 9.2 K. We showed that the BEC condensation temperature of excitons is expected to occur close to this temperature.

Materials and Methods

Samples. The Kish graphite samples we used in the experiment were obtained commercially. The summary of sample information are listed in the following

D R A F T

Supplemental Material for "Critical point for Bose-Einstein condensation of excitons in graphite"

S1. The Nernst Measurement under Pulsed Field

The Nernst effect under pulsed field was performed in WHMFC in Wuhan. The signal was recorded by high-speed digitizer PXI-5922 made by National Instrument running at 2 MHz rate. . The Fig. S1 shows the temperature dependence of resistance of the two chip resistors used to measure temperatures and temperature differences. Before measuring the calibration curves, the chip resistors were worn as thin as possible and trained several times in liquid nitrogen to stabilize the chip resistors. The resistors then were cooled down and calibrated to a known thermometer.

To produce a large thermal gradient and reduce the eddy current effect, the samples were suspended on two types of insulator supports (See Fig. S2A). The heat sink was made of a good thermal conductor: sapphire (Al 2 O 3 ) and the heat source side was made of a bad thermal conductor: Nylon. Two RuO 2 -type chip resistors R 1 and R 2 were mounted on two sides of sample with silver paint to measure T 1 (the heat sink side) and T 2 (the heat source side), which are served as heater also thermometer. Before the Nernst effect measurement, the resistances of the resistors were calibrated up to 250 K (See Fig. S1). During the measurements, the whole system was regulated at desired temperatures. To prevail background signal, the temperature differences had to be large around 4 K in whole measurements. The estimated sample temperature was calculated by Tave = (T 1 + T 2 )/2 shown in the Fig. S2B. However, the actual temperature below 9.2 K was calibrated by the transition field of α(1) without heating during a pulsed field and it is proved to be quite accurate from the sharpness of the rising at α and falling at α instead of broad shapes. This temperature is close to the estimated temperature from thermometers on the setup. To reduce the loop signal caused by large dB/dt, 20 µm diameter twisted enameled copper wires were connected to transverse laterals of sample to measure Nernst signal. The photograph in Fig. S2C shows the actual setup. An in-plane thermal gradient was established before applying pulsed field along the c-axis of sample.

Fig. S2C shows a typical Nernst measurement result at 7.3 K. The authentic Nernst signal (green) is obtained by subtracting the signal with heating (black) from the signal without heating (red). The measured signal in pulsed field includes three parts: the authentic Nernst signal Vyx, the varying-field induced voltage (electromagnetic induction) and the Seebeck signal from a misalignment of potential contacts. We obtained the authentic Nernst signal by a two-round measurement: with and without heating sample during pulsed fields. With temperature gradient under pulsed field, the voltage signal can be expressed as V 1 = Vyx ±(AdB 1 /dt+I eddy R)±Vxx (the black curve in Fig. S2C ). A is the wire loop area perpendicular to the field, which can be reduced or cancelled by using twisted wires. I eddy is the eddy currents (Foucault's currents) and proportional to the field variation rate dB/dt, R is the resistance along the eddy current and Vxx is the Seebeck signal. Since the Seebeck effect is around one-order smaller comparing to Nernst effect in graphite(2), we can neglect this Seebeck part. Indeed, the Nernst signal is almost same by reversing the orientation of the pulsed magnetic field. Meanwhile, without temperature gradient under pulsed field, we can obtain only the second part: V 2 = ±(AdB 2 /dt + I eddy R) (the red curve in Fig. S2C ). dB 1 /dt and dB 2 /dt can be easily to make same if we pulse the magnet to a same field at a same condition for two pulses. Finally, the Nernst signal is deduced by Vyx V 3 = |V 1 -V 2 | which is the green curve in Fig. S2C . This well overlap of authentic Nernst part as the field is rising and falling indicates the validation of measurement and also the sample was not heated much during the pulses.

T ( K ) C V y x ( m V ) B ( T ) V 1 w i t h h e a t i n g V 2 w i t h o u t h e a t i n g V 3 = | V 1 -V 2 |

S2. Reproducibility of the measurements

A typical result up to 10 T of reversing the orientation of magnetic field is shown in Fig. S3. After symmetrization/asymmetrization, the V N ernst is well-overlapped and much larger than V Seebeck . So we can obtain the Nernst signal by two shoots( with and without heating) in the same direction .

The whole Nernst signal data of K3 are shown in Fig. S4C . We also measured Nernst effect in different Kish graphite samples named K1 and K2. We obtained similar results in the two samples shown in Fig. S4A and S4B. All data have been shifted for clarity. The 47 T peaks can be observed clearly in all three samples. Disappearance of the peak is also similar at 35 K and the critical temperature of

Fig. 2 .

 2 Fig. 2. Experimental signature of a critical point. (A) The sketch of the setup. (B) The Nernst signal presents a structure near 47 T. The peak above 9.2 K is substituted by two distinct anomalies below. There is a jump in the Nernst signal followed by a fall as a function of increasing magnetic field. (C) The broad peak centered at 47 T gradually fades away upon warming. Dashed lines are guides for the eyes. Curves are shifted for clarity. (D) The evolution of the Nernst signal with warming over a broader temperature range. (E) The Nernst anomalies in the (B,T ) plane bifurcate at B= 47 T and T = 9.2 K.

Fig. 3 .

 3 Fig. 3. Thermal evolution of the Nernst anomalies and the effective mass (A) The oscillatory component of the Nernst signal near quantum limit and 47 T. The low-field peak survives up to 35 K upon warming, in contrast to the high-field peak, which fades away quickly upon warming. (B) The temperature dependence of the magnitude of the two anomalies allows to extract the effective mass and reveals a 10-fold mass enhancement.

Fig. 4 .

 4 Fig. 4. Band dispersion, length scales and the boundaries of the EI phase. (A) Electron and hole Fermi pockets and the Brillouin zone in graphite. (B) Band dispersion in graphite along the kz axis. Note that electron and hole bands touch each other and extend over the thickness of the Brillouin zone along the kz axis. The system is half-filled along the kz axis and remains so in finite magnetic field. (C) The interboson distance (times a numerical factor) and the de Broglie wavelength in liquid helium and in graphite. The two length scales become equal near the critical temperature. (D) The phase diagram of the first field-induced phase in graphite. BEC triggers the bifurcation of the Nernst anomalies shown by green (black) symbols above (below) the critical point. At high temperature, it corresponds to the simultaneous evacuation of the (-1 h -and 0 e + ) Landau levels. The inset is the zoom near the critical point, where the critical temperature ceases to follow the BCS behavior and saturates to a value set by the Eq. 1. The EI phase is destroyed in two different ways at its lower and upper field boundaries. Along the higher boundary, the binding energy falls below the field-induced gap. Along the lower boundary, the condensate is weakened by decreasing DOS and evolves toward a BCS-type weak-coupling behavior.

  for holes and electrons in graphite with magnetic field along the c-axis (24). This allows the quantification of the areas of extremal orbit, A ⊥ , the electronic wavelengths, λ ⊥ and the inter-particle distances, d ⊥ by using d = λ F √ 2π , assuming that the hole and electron Fermi surfaces are cylinders. The deviation caused by their elongated ellipsoid geometry is small.

  Fig. S1. The R-T curves of the two chip resistors used as temperature calibration in the Nernst measurements for K3.

AFig. S2 .

 S2 Fig. S2. To obtain the authentic Nernst signal under pulsed field: (A) The sketch of the Nernst effect setup under pulsed field. The temperature gradient was established by applied current to the chip resistor on the Nylon side. (B) The estimated temperature difference and average temperature from the two calibrated resistors after applied heating current. (C) A typical result of Nernst data obtained at 7.3 K with a photo of the setup inserted. The authentic Nernst signal (green) is obtained by subtracting the signal with heating (black) from the signal without heating (red).

Table 1 . The de Haas-van Alphen Effect frequencies

 1

	Holes	4.7	4.49	52	21
	Electrons	6.45	6.15	45	18

12

cm -2 ) λ ⊥ (nm) d ⊥ (nm) , F ⊥

table :

 :

	Sample	type	dimension (mm 3 )
	K1	kish	1×0.95×0.04
	K2	kish	0.95×0.9×0.02
	K3	kish	1.1×1.2×0.06

Table 2 . A description of the samples used in this study.

 2 2016YFA0401704), the National Science Foundation of China (Grant nos.51861135104 and 11574097) and Fundamental Research Funds for the Central Universities (Grant no. 2019kfyXMBZ071). In France, it was supported by the Agence Nationale de la Recherche (ANR-18-CE92-0020-01; ANR-19-CE30-0014-04) and by Jeunes Equipes de l Institut de Physique du Collège de France.Z.Z. acknowledges useful discussions with Ryuichi Shindou and Yuanchang Li.

	The Nernst Measurement under Pulsed Field. The measurement of
	Nernst effect under pulsed field was performed in WHMFC in
	Wuhan. The signal was recorded by high-speed digitizer PXI-5922
	made by National Instrument running at 2 MHz rate. More detailed
	information to obtain the authentic Nernst signal is provided in the
	SI Appendix.
	Data Availability. All data are made available within the article or
	SI Appendix.

| www.pnas.org/cgi/doi/10.1073/pnas.XXXXXXXXXX Wang et al.

| www.pnas.org/cgi/doi/10.1073/pnas.XXXXXXXXXX Wang et al.

| www.pnas.org/cgi/doi/10.1073/pnas.XXXXXXXXXX Wang et al.

ACKNOWLEDGMENTS.

This work was supported by the National Key Research and Development Program of China (Grant no.

| B | ( T )

V 1 ( B u p , w i t h h e a t i n g ) V 2 ( B u p , w i t h o u t h e a t i n g ) V 3 ( B d o w n , w i t h h e a t i n g ) V 4 ( B d o w n , w i t h o u t h e a t i n g )

Fig. S3. A typical result up to 10 T. V N ernst (in cyan) is much larger than V Seebeck (in magenta).

. phase A is as well as consistent in the three samples.

S3. The EI scenario in other semi-metals

The concept of an excitonic insulator (EI) was proposed by Mott (3) in the context of semimetals. It was extended to semiconductors by Knox (4). This led to a search for the EI in semimetals. Semimetallic elements Bi, Sb, and As appeared promising at the first sight. However, the large electric permittivity in these semimetals hinders the formation of EI.

The binding energy of an exciton is E B = (µ/m 0 )(1/ 2 )Ry (5,6). Here µ, m 0 , and Ry are the reduced mass, the free electron mass, the dielectric constant and the Rydberg energy. With a typical mass of µ = 0.01m 0 and a dielectric constant of = 100 for these semimetals, the binding energy would be E B = 1.36 × 10 -5 eV ∼ 0.1 K


Hervé Monchot
email: herve.monchot@wanadoo.fr

Marjan Mashkour
email: mashkour@mnhn.fr

Fereidoun Biglari
email: fbiglari@gmail.com

Kamyar Abdi
email: kamyar.abdi@gmail.com

The Upper Pleistocene brown bear (Carnivora, Ursidae) in the Zagros: Evidence from Wezmeh

Keywords:

teaching and research institutions in France or abroad, or from public or private research centers.

Introduction

The brown bear ( Ursus arctos Linnaeus, 1758) is a terrestrial mammal species with a very broad geographic distribution across Europe, Asia and North America. The Syrian brown bear subspecies ( Ursus arctos syriacus Hemprich and Ehrenberg, 1928), which is

The majority of the Pleistocene sites in Southwest Asia that contain ursid bones are located along the Levantine coast from northern Palestine to the Lebanese mountains ( Table 1; Fig. 1). The earliest evidence of bear was found in the Lower Pleistocene site of Ubeidiya in the Jordan valley where 13 remains of Ursus etruscus (Cuvier, 1823) were recovered [START_REF] Martínez-Navarro | The large carnivores from Ubeidiya (early Pleistocene, Israel): biochronological and biogeographical implications[END_REF]. The only sites during the Middle Pleistocene with the presence of ursid remains are Cave Bear in the Upper Galilee [START_REF] Tchernov | Middle Pleistocene (early Toringian) carnivore remains from northern Israel[END_REF] and Darband Cave in the Alborz [START_REF] Biglari | The Lower Paleolithic occupation of Iran[END_REF][START_REF] Biglari | Darband cave: new evidence for Lower Paleolithic occupation at Western Alborz Range, Gilan[END_REF]. These comprise large assemblages of teeth and bones identified as Ursus deningeri (von Reichenau, 1904) (Argant unpublished report; interested readers may refer in particular to [START_REF] Argant | Les ours et leur évolution[END_REF] for an overview of the complex phylogeny of Plio-Pleistocene Eurasian Ursidae). Finally, during the Upper Pleistocene only U. arctos has been identified, from remains found in Mount Carmel, in northern Levant/Galilee, in the Lebanon Mount (e.g . [START_REF] Kurtén | The carnivora of the Palestine caves[END_REF][START_REF] Hooijer | The fossil vertebrates of Ksar'Akil, a palaeolithic rockshelter in the Lebanon[END_REF][START_REF] Garrard | Man-Animal-Plant relationships during the Upper Pleistocene and Early Holocene of the Levant[END_REF] and in Northwest Syria [START_REF] Griggo | Mousterian fauna from Dederiyeh cave and comparisons with fauna from Umm El Tlel and Douara cave[END_REF]. In the Zagros Mountains A. Vue générale de l'étroite vallée de la montagne Qazivand avec localisation de la grotte Wezmeh (indiquée par une flèche). B. Localisation de la grotte Wezmeh et carte du site avec l'emplacement de l'assemblage osseux (d'après [START_REF] Abdi | Islamabad project 2001: test excavations at Wezmeh cave[END_REF].

the brown bear has only been identified in Shanidar [START_REF] Evins | The fauna from Shanidar cave: Mousterian wild goat exploitation in Northeastern Iraq[END_REF], although a bear tooth was also collected on the surface of the Mar Koulian cave, an Upper Pleistocene/Holocene site near Rawansar [START_REF] Biglari | The discovery of Upper Paleolithic remains at Mar Kulian and Mar Dalan caves, Rawansar[END_REF]. To date, no remains of cave bear ( Ursus spelaeus Rosenmüller, 1794) have been described in Southwest Asia. Although common in the Caucasus (e.g. [START_REF] Baryshnikov | Cave bears from the Palaeolithic of the Greater Caucasus[END_REF][START_REF] Bar-Oz | Taphonomy and zooarchaeology of a high-altitude Upper Pleistocene faunal sequence from Hovk-1 Cave, Armenia[END_REF][START_REF] Van Der Made | The new material of large mammals from Azokh and comments on the older Collections[END_REF], it seems that this species never reached the Zagros-Taurus and Alborz mountain ranges.

Several recent studies have investigated the phylogeny of the Syrian brown bear ( Murtskhvaladze et al., 2010;[START_REF] Calvignac | Genetic diversity of endangered brown bear ( Ursus arctos ) populations at the crossroads of Europe, Asia and Africa[END_REF]. A genetic study on mitochondrial DNA shows that all brown bears occurring in the Caucasus appear to be monophyletic, and are a subspecies of the Eurasian brown bear ( U. arctos arctos ) ( Murtskhvaladze et al., 2010;[START_REF] Lortkipanidze | Brown bear distribution and status in the South Caucasus[END_REF]. However, the taxonomic status of the species in this region remains unclear and its distribution and our understanding of its evolution during the Pleistocene in this part of the world remains fragmentary. The discovery of 192 brown bear bones in Wezmeh cave in the Zagros Mountains is therefore an important contribution to the history of this taxon in Southwest Asia.

Wezmeh Cave

This site was discovered in 1999 during an archaeological survey under the direction of Kamyar Abdi in the area of Islamabad-e Gharb (Province of Kermanshah, Iran), a region with numerous Palaeolithic sites [START_REF] Smith | Paleolithic archaeology in Iran[END_REF]. The cave is located about 10 km southeast of the city of Islamabad-e Gharb and 3.5 km northeast of the village of Tajar-e Akbar. The cave is at an elevation of 1,430 m a.s.l., approximately 100 m lower than the summit of the Qazivand Mountains, and about 60 m above the valley floor on a steep slope with a 36 dip. The cavity is almost horizontal, formed between geological layers of karstic limestone. Its entrance faces north and is 2 m wide and 1.20 m high. The cave is about 27 m long and about 45 m 2 ( Fig. 2, [START_REF] Abdi | Islamabad project 2001: test excavations at Wezmeh cave[END_REF]. The cave was looted during the 1990s, resulting in piles of disturbed deposits in front of the cave entrance above the steep slope. After a careful inspection of the site, geoarchaeological studies indicate that the disturbed sediments came from the rear of the cave where sedimentation reaches a maximum of 3 m deep. These disturbed deposits on the exterior slope yielded an abundant faunal assemblage in excellent preservation, particularly rich in carnivoran and ungulate remains. An initial faunal list was established, and notably all of the carnivore families known to be present during the Upper Pleistocene in Southwest Asia are present in the Wezmeh assemblage: Hyaenidae (spotted hyena), Canidae (wolf and red fox), Felidae (lion, leopard, lynx/caracal and wild cat), Mustelidae (badger, stone marten and polecat), Herpestidae (mongoose) and of course Ursidae (brown bear). This last taxon represents 11.1% of the total NISP (carnivore and ungulate) and 13.9% of the carnivore remains found in Wezmeh site ( Table 2). The first series of palaeontological and taphonomical studies suggest an Upper Pleistocene origin for this assemblage, with a similar skeletal element profile to the Middle Palaeolithic assemblages from the southern Levantine [START_REF] Mashkour | Carnivores and their preys in the Wezmeh cave (Kermanshah Iran): a late Pleistocene refuge in the Zagros[END_REF].

Due to the disturbed context of this assemblage, and to the notable diversity of the mammalian species present, it was extremely important to run a direct dating analysis on the principal species present in the cave. Several bones were selected from the carnivore and herbivore remains, including bear, spotted hyena and wild boar. A series of U/Th dates (CNRS, Gif-sur-Yvette) were performed on tooth enamel, providing ages of 63.1 ± 12.6 and 67.3 ± 11.8 Kya for the brown bear, 14.4 ± 1.3, 23.4 ± 4.9 and 26.6 ± 9.4 Kya for the spotted hyena and finally 13.8 ± 3.4 Kya for the wild boar ( Mashkour et al., 2009 : tab. 2). The oldest occupation of the cave may therefore be dated to the Late Pleistocene. Three hyena coprolites were dated by the AMS radiocarbon method, producing calibrated 14 C ages of 19228 ± 352 cal BP and 12744 ± 370 cal BP, as well as a Holocene date of 405 ± 59 cal BP [START_REF] Djamali | Pollen analysis of coprolites from a late Pleistocene-Holocene cave deposit (Wezmeh Cave, west Iran): insights into the late Pleistocene and late Holocene vegetation and flora of the central Zagros Mountains[END_REF]. Several fragmented human bones and teeth were discovered in the cave among the faunal remains, including the Neanderthal premolar "Wezmeh 1" [START_REF] Trinkaus | Late Pleistocene human remains from Wezmeh cave, Western Iran[END_REF][START_REF] Zanolli | A Neanderthal from the Central Western Zagros, Iran. Structural reassessment of the Wezmeh 1 maxillary premolar[END_REF]. One of the human remains was dated to the early Neolithic, around 9300 cal BP [START_REF] Broushaki | Early Neolithic genomes from the eastern Fertile Crescent[END_REF]. No evidence of human activity (e.g. industry, burned bone, cutting traces) was observed.

The brown bear assemblage

The species identification of brown bear ( Ursus arctos ) versus cave bear ( Ursus spelaeus and Ursus deningeri ) was based on the shape (morphology and size) of certain skeletal elements (e.g. [START_REF] Kurtén | Sex dimorphism and size trends in the cave bear, Ursus Spelaeus Rosenmüller & Heinroth[END_REF][START_REF] Stiner | Mortality analysis of Pleistocene bears and its paleoanthropological relevance[END_REF][START_REF] Stiner | Reconstructing cave bear paleoecology from skeletons: a cross-disciplinary study of middle Pleistocene bears from Yarimburgaz cave, Turkey[END_REF][START_REF] Petronio | Morphological and biometrical differences in the limb bones of Ursus arctos and Ursus spelaeus and phylogenetical considerations on the two species[END_REF][START_REF] Schweizer | Etude d'une population d'ours des cavernes ( Ursus spelaeus Rosenmüller & Heinroth, 1794) provenant de la grotte de Vaucluse[END_REF][START_REF] Schweizer | Étude des caractères non-métriques dans une population d'ours des cavernes ( Ursus spelaeus Rosenmüller & Heinroth, 1794) provenant de la grotte de Vaucluse (Doubs, France)[END_REF]. All the bones (N = 192) present at Wezmeh cave undoubtedly belong to the brown bear ( Figs. 345).

The bear bone conservation

The surfaces of the bones are in an excellent state of preservation; in most cases they are smooth without cracks or fissures, characteristic of unweathered bones. Nevertheless, some pieces (n = 9) present very limited surface weathering with a cracking parallel to the fibre structure (longitudinal) ( Todisco and Monchot, 2008 : Tab. 2). While the small bones are complete or almost complete, the long bones exhibit recent fragmentation related to the looting activities or post-depositional processes. The fracture characteristics of the long bones clearly indicate a complete shaft circumference, a transverse fracture outline and a smooth fracture edge. All these criteria suggest that these long bones were broken when they were dry, i.e. an indication of post-depositional breakage [START_REF] Villa | Breakage patterns of human long bones[END_REF]. Finally, the bone surfaces show some traces of manganese (n = 3), concretions (n = 3) and carnivore tooth pits (n = 3).

The distribution of skeletal elements

Among skeletal elements there is a dominant presence of isolated teeth (20.8%) and extremity elements (45.8%) including carpal, tarsal, phalanxes and metapodial; trunk elements, i.e. vertebrae and ribs, contribute a smaller percentage (16.7%) ( Table 3, Fig. 6). The long bones are never complete and are very fragmented. Humerus elements are represented by two distal and one proximal fragment, tibia by one distal fragment, femur by four proximal and four distal fragments, the ulna by four proximal and one distal fragments, and the radius by three shafts, one distal and one proximal fragments. There is no marked predominance of a single anatomical region or module ( e.g. forelimb vs hindlimb) or in bone laterality ( 2 = 19 . 09; df = 22; p = 0.639), which excludes any selective transport (i.e. anthropogenic, carnivore, hydraulic) of brown bear bones within the cave (cf. [START_REF] Quiles | Les Ursidae du Pléistocène moyen et supérieur en Midi méditerranéen : apports paléontologiques, biochronologiques et archéozoologiques[END_REF]. This type of assemblage, which consists primarily of isolated teeth and extremities of the appendicular skeleton (i.e. carpal, tarsal, metapodial and phalanges), is found in many caves or dens of Ursus denigeri (e.g. Château, Scharzfeld, Westbury, Vertesszöllos II, Fosse et al., 2002 : Fig. 3) and Ursus spelaeus (e.g. Ekain, Azé 1-3, Divje Babe, Fosse et al., 2002 : Fig. 4) in Europe and in the Caucasus (Hovk-1 cave, [START_REF] Bar-Oz | Taphonomy and zooarchaeology of a high-altitude Upper Pleistocene faunal sequence from Hovk-1 Cave, Armenia[END_REF]. This type of skeletal representation is also found in the Mesolithic rock shelter of Kostias Klde in the Republic of Georgia where brown bears have been hunted by humans [START_REF] Bar-Oz | Bear in mind: bear hunting in the Mesolithic of the southern Caucasus[END_REF], highlighting the differential survivorship of different skeletal elements.

Thanatocoenosis structure and mortality profile

Two main methods were utilised in order to estimate the age at the time of death. The first method through the estimation of the stage of tooth eruption, and by analysing dental wear. Following sequential eruption, teeth become progressively more worn and distinctive wear patterns are formed on the different enamel facets. According to the work of [START_REF] Quiles | Les Ursidae du Pléistocène moyen et supérieur en Midi méditerranéen : apports paléontologiques, biochronologiques et archéozoologiques[END_REF][START_REF] Quiles | Tanières d'ours des cavernes (Carnivora Ursidae) du pourtour méditerranéen : étude taphonomique et paléobiologique de huit assemblages du Pléistocène supérieur[END_REF] , age estimation is correlated to nine age groups (see a description of each age group in Quiles, 2004 : tab. 5): infantile, juvenile, sub-adult 1, subadult 2, adult 1, adult 2, adult 3, old adult and very adult. These age-classes have unfortunately not yet been calibrated with real ages known for present day brown bear. The second method is based on the epiphyseal fusion [START_REF] Weinstock | Epiphyseal fusion in Brown Bears: a population study of Grizzlies (Ursus arctos horribilis ) from Montana and Wyoming[END_REF], and is less reliable as a result of the various taphonomical processes that affect skeletal remains, especially those of immature individuals and the types of skeletal elements with a high marrow and spongiosa content. Only four imprecise age-classes may be defined with the appendicular epiphyseal fusion method: foetus or newborn, juvenile, sub-adult and adult.

Thus, based on 40 identifiable teeth found at Wezmeh, 29 teeth (12 lower and 17 upper teeth) were assigned an age group, and when combined with tooth type a minimum of eight individuals were identified: one juvenile, three sub-adults and four adults ( Table 4). The only unfused bones present in the brown bear assemblage are one distal metapodial (an element which fuses around 4-5 years) and one proximal phalanx (which fuse around 2-3 years). This confirms the absence of young and old bears from the assemblage.

Osteometry: The Wezmeh brown bear

The Wezmeh brown bear metrical analysis is presented in Table 5 for the teeth and Tables 6-9 for the post-cranial material. The measurements were taken following the procedures outlined by [START_REF] Schweizer | Etude d'une population d'ours des cavernes ( Ursus spelaeus Rosenmüller & Heinroth, 1794) provenant de la grotte de Vaucluse[END_REF] . The tooth dimensions of U. arctos from the Wezmeh cave were compared with those from other Southwest Asian sites, and the post-cephalic dimensions were compared with material from various European assemblages. However, due to the small sample size and to the lack of published data on brown bears, the dimensions of the Wezmeh brown bear bones appear to fall within the size range of both present day and fossilised European brown bears [START_REF] Altuna | Hallazgos de oso pardo ( Ursus arctos , Mammalia) en cuevas del Pais Vasco[END_REF][START_REF] Ballesio | Le gisement pléistocène de la grotte de Jaurens à Nespouls, Corrèze, France : les carnivores (Mammalia Carnivora). III. Ursidae. Ursus arctos Linnaeus[END_REF]. For example, if we calculate the ratio of the length of the first metatarsal as a percentage of the length of the fifth, we obtain 72.3% for the Wezmeh material. This index ranges between 71.1% and 74.2% for Ursus arctos , and between 62.3% and 66.9% for Ursus spelaeus and Ursus deningeri ( Ballesio, 1983 : Tab. 19). The teeth ( Fig. 7) of the Wezmeh specimens are larger than those of the present day Syrian brown bear ( U. a. syriacus ) and are similar to those found at Middle Palaeolithic sites such as Ksar'Akil or Tabun C, which are prehistoric sites with later dates than the Acheuleo-Yabrudian site of Zuttiyeh [START_REF] Tchernov | Middle Pleistocene (early Toringian) carnivore remains from northern Israel[END_REF].

Like other ursids, the brown bear displays a strong sexual dimorphism, which is particularly notable in the transverse diameter of the canines [START_REF] Koby | Le dimorphisme sexuel des canines d'Ursus arctos et d'U. spelaeus[END_REF][START_REF] Kurtén | Sex dimorphism and size trends in the cave bear, Ursus Spelaeus Rosenmüller & Heinroth[END_REF]. However the overlap I: infantile; Juv: juvenile; SA1: sub-adult 1; SA2: sub-adult 2; A1: adult 1; A2: adult 2; A3: adult 3; OA: old adult; VOA: very adult/ I : infantile ; Juv : juvénile ; SA1 : subadulte 1 ; SA2 : sub-adulte 2 ; A1 : adulte 1 ; A2 : adulte 2 ; A3 : adulte 3 ; OA : adulte âgé ; VOA : adulte très âgé.

between the two sexes is such that it is impossible to separate them only from univariate and bivariate diagrams [START_REF] Kurtén | Sex dimorphism and size trends in the cave bear, Ursus Spelaeus Rosenmüller & Heinroth[END_REF][START_REF] Schweizer | Les ours des cavernes ( Ursus spelaeus Rosenmüller & Heinroth, 1794) de la grotte de Vaucluse (Doubs, France) : détermination du sexe et estimation du sex-ratio[END_REF]. It is therefore preferable to use more reliable statistical methods, such as mixture analysis [START_REF] Quiles | Sex ratio et analyse des mélanges d'Ursus spelaeus (Carnivora, Ursidae) du gisement pléistocène supérieur de Fate (Ligurie Italie). Implications paléobiologiques[END_REF][START_REF] Quiles | Mixture analysis: a new method for cave bear sex-ratio determination[END_REF]. However the small sample size and the general lack of data on sexual dimorphism in brown bears also prevents the types of analyses that are necessary in order to distinguish male and female remains.

Discussion

Structure of the Wezmeh bear population

The mortality profile indicates that sub-adult and adult males appear to have preferentially occupied the cave. The absence of very young juveniles may be explained by the fact that they do not hibernate and are exclusively nursed by their mother [START_REF] Nelson | Behaviour, biochemistry and hibernation in black, grizzly and polar bears[END_REF]. In addition, they are probably underrepresented because of the fragility of the tooth buds and bone remains, which are subject to extensive destruction, exacerbated by the absence of sieving during the excavation. The sub-adult period, which includes the first two hibernations, is critical to individual survival. Foraging by themselves, and still physically immature and inexperienced, young adults have the greatest probability of dying during hibernation [START_REF] Mcloughlin | Demography of barren-ground grizzly bears[END_REF]. An abundance of young adult remains is found for instance on the sites of Fate, Basura, Hortus and Arbreda [START_REF] Quiles | Les Ursidae du Pléistocène moyen et supérieur en Midi méditerranéen : apports paléontologiques, biochronologiques et archéozoologiques[END_REF]. The presence of adult 1 remains in the Wezmeh assemblage, mostly males (e.g. Badalucco), is probably due to these factors, especially as the dominant older Profil anatomique de l'ours brun de Wezmehétablie en % du nombre de spécimens identifiés (NISP). All measurements are in mm. L: Antero-posterior length near the base of the crown; B: transversal breadth near the base of the crown; HC: height of the crown; Bant: anterior transversal breadth near the base of the crown; Bpost: posterior transversal breadth near the base of the crown/ Toutes les mesures sont en mm. L : longueur antéropostérieure à la base de la couronne ; B : largeur transversale à la base de la couronne ; HC : hauteur de la couronne ; Bant : largeur transversale antérieure à la base de la couronne ; Bpost : largeur transversale postérieure à la base de la couronne .

males exert a social pressure that results in restricted access to food resources [START_REF] Stokes | Social behaviour of brown bear at Mac Neil River[END_REF][START_REF] Nelson | Behaviour, biochemistry and hibernation in black, grizzly and polar bears[END_REF]. Adult 1 males are also typically forced to live in less favourable environments (see the dispersion of sub-adult males, [START_REF] Mclellan | Natal dispersal of grizzly bears[END_REF], as dominant males already occupy the highest quality territories. Furthermore, it's interesting to note that bear MNI based on teeth is about twice the MNI based on post-cranial bone elements (3 to 8), emphasizing the dramatic loss of bone material in situ in Wezmeh Cave. Relatively speaking, the bones from Wezmeh Cave are in good condition, and the conditions for fossilisation appear to be favorable; the lack of certain skeletal parts therefore cannot be readily explained. The cavity has been used by many carnivores, including the spotted hyena ( Crocuta crocuta spelaea Goldfuss, 1823), which has the reputation of bone crusher [START_REF] Monchot | Des hyènes tachetées au Pléistocène supérieur dans le Zagros (grotte Wezmeh Iran). Archaeozoology of the Near East VIII, TMO 49[END_REF]. We may suppose that some of the bear bones were therefore consumed while the tissue was still fresh and edible. There is evidence of gnawing by carnivores on material from the cave, suggesting that some fragile bone elements (such as those that are thin, flat and trabecular) may have been partly or completely destroyed by biological agency soon after death ( Mashkour et al., 2009 : 689). All measurements are in mm. GL: greatest length; DT: transversal diameter; DAP: antero-posterior diameter; L art.: length of the articulation; DAP art.: articulation antero-posterior diameter; L manu: length of the manubrium ; DT facdist: transversal diameter of the distal articulation; DAP st: antero-posterior diameter of the sustentaculum tali; DDP st: dorso-plantar diameter of the sustentaculum tali ; DT troch: maximum transversal diameter of the trochlea; DT fossa: maximum transversal diameter of the fossa between the two condyles; DAP epimed: antero-posterior diameter of the medial condyle; DAP col: antero-posterior diameter of the diaphysis just under the proximal epiphysis; DT col: transversal diameter of the diaphysis just under the proximal epiphysis; Bcaput: Greatest breadth of the caput femoris ; Dcaput: greatest depth of the caput femoris; DT ol: olecranon transversal diameter/ Toutes les mesures sont en mm. GL : longueur maximale ; DT : diamètre transversal ; DAP : diamètre antéro-postérieur ; L art. : Longueur de l'articulation ; DAP art. : diamètre antéro-postérieur de l'articulation ; L manu : longueur du manubrium ; DT facdiste : diamètre transverse de l'articulation distale ; DAP st : diamètre antéro-postérieur du sustentaculum tali ; DDP st : diamètre dorso-plantaire du sustentaculum tali ; DT troch : diamètre transverse maximal de la trochlée ; DT fosse : diamètre transverse maximal de la fosse entre les deux condyles ; DAP epimed : diamètre antéro-postérieur du condyle interne ; DAP col : diamètre antéro-postérieur de la diaphyse juste en dessous de l'épiphyse proximale ; DT col : diamètre transverse de la diaphyse juste en dessous de l'épiphyse proximale ; Bcaput : la plus grande largeur du caput femoris ; Dcaput : plus grande profondeur du caput femoris ; DT ol : diamètre transverse de l'olécrane .

An update on the genetics of brown bears in the Southwest Asia

The genetics of Southwest Asian bears have not yet been studied in detail despite a large number of publications [START_REF] Taberlet | Génétique de l'ours brun des Pyrénées (Ursus arctos) : premiers résultats[END_REF][START_REF] Talbot | Phylogeography of brown bears (Ursus arctos) of Alaska and paraphyly within the Ursidae[END_REF]Shields, 1996 , Miller et al., 2006 ). Most recently, Calvignac and colleagues have analysed samples from Syria, Lebanon and Iran [START_REF] Calvignac | Genetic diversity of endangered brown bear ( Ursus arctos ) populations at the crossroads of Europe, Asia and Africa[END_REF]. The phylogenetic reconstruction based on mtDNA has revealed a high genetic diversity, with the presence of three distinct clades [START_REF] Miller | Phylogeography and mitochondrial diversity of extirpated brown bear (Ursus arctos ) populations in the contiguous United States and Mexico[END_REF]. The Syrian specimens display a genetic proximity to the east-European bears, whereas the Lebanese specimens are closer to the west-European bears [START_REF] Talbot | Phylogeography of brown bears (Ursus arctos) of Alaska and paraphyly within the Ursidae[END_REF]. Iranian bears form a third unique group that for the moment cannot be grouped with any of the others [START_REF] Miller | Phylogeography and mitochondrial diversity of extirpated brown bear (Ursus arctos ) populations in the contiguous United States and Mexico[END_REF][START_REF] Calvignac | Genetic diversity of endangered brown bear ( Ursus arctos ) populations at the crossroads of Europe, Asia and Africa[END_REF]. The history of the species in this area is therefore much more complex than previously thought, and it may have involved several Wezmeh may therefore have an important role in the future understanding of this phenomenon. A recent genetic study has indicated that there was also a coinciding emergence of several haplotypes [START_REF] Ho | The effect of inappropriate calibration: three case studies in molecular ecology[END_REF], corresponding with the oldest dates provided by the deposits in Wezmeh. However no viable DNA samples have yet been recovered from the Wezmeh bear remains in order to further investigate this relationship, despite several attempts [START_REF] Calvignac | Genetic diversity of endangered brown bear ( Ursus arctos ) populations at the crossroads of Europe, Asia and Africa[END_REF].

Conclusions

Ursid remains from Wezmeh Cave may be identified as brown bear, based on the metric and morphological characteristics of the skeletal elements present in the assemblage. The size of the Wezmeh specimens fall within the ranges of variation of both Upper Pleistocene and extant brown bears in Southwest Asia and can be attributed to a local species of brown bear (Ursus arctos ), but cannot be conclusively identified as the Syrian brown bear ( Ursus arctos syriacus ).

In respect to the palaeogenetic research, for the time being the absence of DNA prevents the identification of the brown bear bones of Wezmeh to a specific haplotype. The genetic analyses on modern bears show that Iranian and Syrian bears belong to different haplotypes. Any possible morphological markers of these genetic differences are as of yet unidentified. Unfortunately there are no large bone assemblages to our knowledge that are sufficiently representative of this small-sized subspecies of brown bear in order to provide the material to address this question. Only the re-analysis of the Ksar Akil site in Lebanon [START_REF] Hooijer | The fossil vertebrates of Ksar'Akil, a palaeolithic rockshelter in the Lebanon[END_REF], which contains ursid remains that are large in size, would allow for a direct morphometric or even geometric morphometric analysis. Unfortunately, to our knowledge this assemblage has been lost. It is unusual to find older bear remains that display such a distinctively small size, especially across the whole sample of elements. The small dimensions of these remains strongly exclude species identification as U. spelaeus . In addition, the dates obtained for the bear remains are notably older than those for Crocuta crocuta [START_REF] Mashkour | Carnivores and their preys in the Wezmeh cave (Kermanshah Iran): a late Pleistocene refuge in the Zagros[END_REF], despite spotted hyena becoming extinct in this region and brown bears persisting until the present day. However, there are only five dated samples (3 for the hyena and 2 for the bear), which may not be representative of the whole timerange of occupation of bears and hyenas in Wezmeh [START_REF] Monchot | Des hyènes tachetées au Pléistocène supérieur dans le Zagros (grotte Wezmeh Iran). Archaeozoology of the Near East VIII, TMO 49[END_REF][START_REF] Monchot | The spotted hyena (Crocuta spelaea) of the Wezmeh cave, Kermanshah, Central Western Zagros Mountains[END_REF]. Based on the present material it is not possible to fully understand the biogeographic chronology, especially when bearing in mind that very little is known of the evolution of Pleistocene carnivore fauna in the Zagros. Despite the lack of material, differential occupation of carnivores may have existed in the cave over about 65,000 years.

Fig. 1 .

 1 Fig. 1. Map of the main Pleistocene Ursid sites of Southwest Asia (not exhaustive). Carte des principaux sitesàursidés du Pléistocène en Asie du Sud-Ouest (liste non exhaustive).

Fig. 2 .

 2 Fig. 2. A. General view of the narrow valley at the Qazivand Mountain where the Wezmeh Cave is located. The site is indicated by an arrow. B. Location of Wezmeh cave and map of the site with bone assemblage location (after[START_REF] Abdi | Islamabad project 2001: test excavations at Wezmeh cave[END_REF]. A. Vue générale de l'étroite vallée de la montagne Qazivand avec localisation de la grotte Wezmeh (indiquée par une flèche). B. Localisation de la grotte Wezmeh et carte du site avec l'emplacement de l'assemblage osseux (d'après[START_REF] Abdi | Islamabad project 2001: test excavations at Wezmeh cave[END_REF].

Fig. 3 .

 3 Fig. 3. Teeth of Ursus arctos from Wemezh cave. Left upper P4 (WZ 109, 1-2 = lingual view; 3 = occlusal view); right upper M1 (WZ 104, 4 = lingual view; 5 = occlusal view); right upper M2 (WZ 101, 6 = lingual view; 7 = vestibular view; 8 = occlusal view); left upper canine (WZ 111, 9 = lingual view; 10-11 = buccal view); right lower M3 (WZ 1598, 12-14 = occlusal view; 13-15 = vestibular view); lower M2 right (WZ 1594, 16-18 = occlusal view; 17-19 = vestibular view). Dents d'Ursus arctosde la grotte de Wemezh. P4 supérieure gauche (WZ 109, 1-2 = vue linguale ; 3 = vue occlusale) ; M1 supérieure droite (WZ 104, 4 = vue linguale ; 5 = vue occlusale) ; M2 supérieure droite (WZ 101, 6 = vue linguale ; 7 = vue vestibulaire ; 8 = vue occlusale) ; canine supérieure gauche (WZ 111, 9 = vue linguale ; 10-11 = vue vestibulaire) ; M3 inférieure droite (WZ 1598, 12-14 = vue occlusale ; 13-15 = vue vestibulaire) ; M2 inférieure droite (WZ 1594, 16-18 = vue occlusale ; 17-19 = vue vestibulaire).

Fig. 4 .

 4 Fig. 4. Post-cranial elements of Ursus arctos from Wemezh cave. Right proximal radius (WZ 465, 1 = posterior view, 2 = proximal view); Right distal radius (WZ 466, 3 = posterior view, 4 = anterior view); right proximal ulna (WZ 471, 5 = lateral view); left proximal femur (WZ 1560, 6 = anterior view, 7 = posterior view); left distal femur (WZ 459, 8 = posterior view); left distal femur (WZ 460, 9 = posterior view); left calcaneus (WZ 456, 10-12 = anterior view, 11-13 = posterior view). Eléments post-crâniens d'Ursus arctosde la grotte de Wemezh. Radius proximal droit (WZ 465, 1 = vue postérieure ; 2 = vue proximale) ; radius distal droit (WZ 466, 3 = vue postérieure, 4 = vue antérieure) ; ulna proximale droite (WZ 471, 5 = vue latérale) ; fémur proximal gauche (WZ 1560, 6 = vue antérieure, 7 = vue postérieure) ; fémur distal gauche (WZ 459, 8 = vue postérieure) ; fémur distal gauche (WZ 460, 9 = vue postérieure) ; calcanéum gauche (WZ 456, 10-12 = vue antérieure, 11-13 = vue postérieure).

Fig. 5 .

 5 Fig. 5. Metacarpals (MC) and metatarsals (MT) of Ursus arctos from Wemezh cave: left MC II (WZ 325, 1 = lateral view; 2 = medial view), right MC III (WZ 324, 3 = medial view. 4 = lateral view), left MC IV (WZ 1930, 5 = lateral view; 6 = medial view), left MC V (WZ 323, 7 = lateral view, 8 = medial view); right MT I (WZ 327, 8 = medial view, 9 = lateral view); left MT II (WZ 326, 10 = lateral view, 11 = medial view); right MT III (WZ 1932, 13 = lateral view, 14 = medial view), right MT IV (WZ 328, 15 = medial view, 16 = lateral view). Métacarpiens (MC) et métatarsiens (MT) d'Ursus arctosde la grotte de Wemezh : MC II gauche (WZ 325, 1 = vue latérale, 2 = vue médiale) ; MC III droit (WZ 324, 3 = vue médiale, 4 = vue latérale), MC IV gauche (WZ 1930, 5 = vue latérale, 6 = vue médiale) ; MC V gauche (WZ 323, 7 = vue latérale, 8 = vue médiane) ; MT I droite (WZ 327, 8 = vue médiale, 9 = vue latérale) ; MT II gauche (WZ 326, 10 = vue latérale, 11 = vue médiale) ; MT III droite (WZ 1932, 13 = vue latérale, 14 = vue médiale) ; MT IV droit (WZ 328, 15 = vue médiale, 16 = vue latérale).

Fig. 6 .

 6 Fig. 6. Anatomical distribution of Wezmeh brown bear remains established as % of the number of identified specimens (NISP).

Fig. 7 .

 7 Fig. 7. Comparison of length and breadth of the upper M2 and lower M3 of brown bear (Ursus arctos) from Wezmeh Cave and specimens from southern Levantine sites (after Mashkour et al., 2009 ). Diagrammes bivariés longueur par largeur (en mm) des M2 supérieures et des M3 inférieures de l'ours brun(Ursus arctos)de la grotte de Wezmeh et de différents spécimens provenant de sites du Levant sud (d'après Mashkour et al., 2009 ) .

Table 1

 1 List of Pleistocene ursid remains from Southwestern Asia (not exhaustive). Liste des restes d'ours du Pléistocène du Proche et du Moyen-Orient (liste non exhaustive).

	Site (Level)	Country	Species	Age/Culture									NISP (MNI)	Reference
	Antelias	Lebanon	Ursus arctos	Upper Palaeolithic			Not counted	Garrard, 1980
	Amud (B2)	Israel	Ursus sp .	Upper Pleistocene/Middle Palaeolithic	1	Rabinovich and Hovers,
	Abri Bezez (B)	Lebanon	Ursus arctos	Upper Pleistocene/Middle Palaeolithic	3	Garrard, 1983
	Abri Zumoffen (9-3)	Lebanon	Ursus arctos	Middle Palaeolithic		3	Garrard, 1983
	Abri Zumoffen (21-11)	Lebanon	Ursus arctos	Middle	P	a	l	a	e	o	l	i	t	h	i	c	1	Garrard, 1983
	Azokh cave	Nagorno-Karabakh	Ursus spelaeus	Late Pleistocene						265 (8)	Van der Made et al., 2016
	Bear's cave	Israel	Ursus deningeri	Middle Pleistocene (Early Toringian)	63	Tchernov and Tsoukala,
	Darband cave	Iran	Ursus deningeri	Middle Pleistocene		62	Biglari et al., 2007
	Dederiyeh	Syria	Ursus arctos	Middle Palaeolithic		1	Griggo, 2004
	El-Wad (B-C)	Israel	Ursus arctos	Upper Palaeolithic-Epipalaeolithic	Not counted	Garrard, 1980
	Emirkaya (2)	Turkey	Ursus deningeri	Middle Pleistocene		Not counted	Sen et al., 1991
	Hayonim	Israel	Ursus arctos	Upper Paleolithic/Mousterian-Kebaran	2	Stiner, 2006
	Hovk-1	Armenia	Ursus spelaeus	Upper Pleistocene			186 (12)	Bar-Oz et al., 2012
	Karain (E)	Turkey	Ursus sp .	Upper Pleistocene/Middle Palaeolithic	73	Otte et al., 1998
	Kebara	Israel	Ursus arctos	Upper Pleistocene/Middle Palaeolithic	2	Dayan, 1994
	Kéoué	Lebanon	Ursus arctos	Upper Pleistocene/Middle Palaeolithic	Not counted	Garrard, 1980
	Ksar Akil	Lebanon	Ursus arctos	Upper Pleistocene/Middle Palaeolithic	Not counted	Hooijer, 1961
	Maslouk	Lebanon	Ursus arctos	Upper Pleistocene/Middle Palaeolithic	1	Monchot and Gauthier, unpublished
	Nahr Ibrahim	Lebanon	Ursus arctos	Upper Pleistocene/Middle Palaeolithic	31	Monchot and Gauthier, unpublished
	Nahr el Joz	Lebanon	Ursus arctos	Middle Palaeolithic		Not counted	Garrard, 1980
	Ras-El-Keb	Lebanon	Ursus arctos	Middle Palaeolithic		49 (11)	Garrard, 1998
	Shanidar	Iraq	Ursus arctos	Upper Pleistocene/Middle Palaeolithic	5 (3)	Evins, 1982
	Tabun	Israel	Ursus arctos	Midldle/Early Upper Palaeolithic	Not counted	Garrard, 1980
	Ubeidiya	Israel	Ursus etruscus	Early Pleistocene				13	Martínez-Navarro	et al.,
	Wezmeh	Iran	Ursus arctos	Upper Pleistocene			192 (8)	Mashkour et al., 2009 (this study)
	Zuttiyeh	Israel	Ursus arctos	Acheulo-Yabrudian complex	(3/4)	Bate, 1927
	Yarimburgaz	Turkey	U arctos/deningeri	Middle Pleistocene		3920	Stiner et al., 1998

NISP: number of identified specimens; MNI: minimum number of individuals/ NISP : nombre de restes déterminés ; MNI : nombre minimum d'individus .

Table 2 Species distributions of large mammals in NISP (Number of Identified Specimens) and MNI (Minimum Number of Individuals) for Wezmeh cave. Liste des grands mammifères en NISP (nombre de spécimens identifiés) et MNI (nombre minimal d'individus) pour la grotte de Wezmeh.

 2

			NISP	% NISP	MNI
	Carnivora
	Crocuta crocuta spelaea	Spotted Hyena	437	25.2	10
	Ursus arctos	Brown Bear	192	11.1	8
	Panthera leo	Lion	16	0.9	2
	Panthera pardus	Leopard	2	0.1	1
	Caracal/Lynx/Felis chaus	Caracal/Lynx/Jungle cat	4	0.2	2
	Felis silvestris	Wildcat	8	0.5	2
	Canis lupus	Wolf	176	10.1	6
	Vulpes vulpes	Red Fox	492	28.4	19
	Meles meles	Badger	42	2.4	7
	Mustela putorius	Polecat	7	0.4	2
	Martes martes (foina?)	Stone marten	2	0.1	1
	Herpestes sp.	Mongoose	5	0.3	3
	Perissodactyla
	Dicerorhinus	Rhinoceros	2	0.1	1
	Equus caballus	Horse	1	0.1	1
	Hemionus/E. asinus	Hemione/Donkey	4	0.2	1
	Artiodactyla
	Bos primigenius	Aurochs	13	0.8	2
	Sus scrofa	Wild boar	97	5.6	4
	Cervus elaphus	Red deer	59	3.4	3
	Gazella sp.	Gazelle	17	1.0	1
	Ovis orientalis	Mouflon	127	7.3	6
	Capra aegagrus	Wild goat	31	1.8	2

Table 3

 3 Census of brown bear (Ursus arctos) remains on Wezmeh cave in number of identified specimens (NISP) and in minimum number of individual of combination (MNIc)/ nventaires des restes d'ours brun(Ursus arctos)de la grotte de Wezmeh en nombre de spécimens identifiés (NISP) et en nombre minimal d'individu de combinaison (MNIc).

	Skeletal element	NISP				% NISP	MNIc
		Left	Right	Ind/Imp	Total
	Mandible	-	1	-	1	0.52	1
	Upper isolated teeth	8	9	1	18	9.38	5
	Lower isolated teeth	13	8	1	22	11.46	8
	Thoracic vertebra	-	-	13	13	6.77	-
	Lumbar vertebra	-	-	1	1	0.52	1
	Rib	-	-	17	17	8.85	-
	Sternebra	-	-	1	1	0.52	1
	Humerus	1	2	-	3	1.56	2 (D)
	Radius	1	2	2	5	2.60	3 (SH)
	Ulna	1	4	-	5	2.60	3 (P)
	Carpal	3	-	1	4	2.08	2
	Metacarpal II	2	1	-	3	1.56	2
	Metacarpal III	1	2	-	3	1.56	2
	Metacarpal IV	2	1	-	3	1.56	2
	Metacarpal V	2	1	-	3	1.56	2
	Pelvis	3	3	-	6	3.12	3
	Femur	4	4	-	8	4.17	3
	Patella	1	-	-	1	0.52	1
	Tibia	-	1	-	1	0.52	1 (SH)
	Fibula	2	-	-	2	1.04	2 (D)
	Tarsal	4	5	2	11	5.73	2
	Metatarsal I	1	1	-	2	1.04	1
	Metatarsal II	1	1	-	2	1.04	1
	Metatarsal III	-	2	-	2	1.04	2
	Metatarsal IV	1	3	-	4	2.08	3
	Metatarsal V	-	2	-	2	1.04	2
	Metapodial	-	1	6	7	3.65	-
	Phalange I	-	-	22	22	11.46	-
	Phalange II	-	-	9	9	4.69	-
	Phalange III	-	-	11	11	5.73	-
	TOTAL	51	54	87	192	100	8

SH: diaphysis; D: distal extremity; P: proximal extremity/ SH : diaphyse ; D : extrémité distale ; P : extrémité proximale .

Table 4

 4 Minimum number of individuals according to the 9 classes of age. Nombre minimum d'individus selon les 9 classes d'âge.

		I	Juv	SA1	SA2	A1	A2	A3	OA	VOA	Total
	Lower teeth	0	1	1	1	1	1	0	0	0	5
	Upper teeth	0	1	2	1	2	1	1	0	0	8
	Total (MNIc)	0	1	2	1	2	1	1	0	0	8

Table 5

 5 Tooth measurements of Ursus arctos from Wezmeh cave. Mesures des dents d'Ursus arctos de la grotte de Wezmeh.

			L	B	Bpost	HC	Wear stage
	WZ 101	Upper M2	35.7	19.4	15.8	8.1	4
	WZ 100	Upper M2	39.2	21.1	16.8	9.1	4
	WZ 102	Upper M2	38.2	20.7	-	8.5	5
					Bant
	WZ 104	Upper M1	24.6	17.1	16.7	9.0	6
	WZ 109	Upper P4	15.0	11.6		7.7	4
	WZ 1596	Upper P4	16.2	12.3		8.3	4
	WZ 1597	Upper P4	15.5	11.2		9.0	2
	WZ 116	Upper I3	9.3	10.2		-	4
	WZ 119	Upper I3	9.8	12.5		3.3	5
	WZ 118	Upper I3	10.4	13.1		7.9	4
	WZ 120	Upper I3	11.6	13.1		3.3	6
	WZ 1592	Upper I3	11.2	12.9		3.0	6
	WZ 125	Upper I1	7.0	8.1		8.1	3
			B	DT rac
	WZ 110	Upper C	17.1	21		-	2
	WZ 111	Upper C	16.3	19.1		-	3
			L	Bpost	Bant
	WZ 108	Lower M2	23.7	14.4	13.9	8.0	5
	WZ 105	Lower M2	26.6	17.0	15.7	7.5	4
	WZ 1594	Lower M2	24.7	15.1	14.4	9.3	4
			L	B	Bpost
	WZ 102	Lower M3	22.4	16.0	-	5.7	3
	WZ 106	Lower M3	20.7	15.7	-	8.7	4
	WZ 107	Lower M3	22.4	16.0	-	5.7	3
	WZ 1598	Lower M3	21.7	16.0	-	6.5	3
	WZ 113	Lower M1	-	-	13.9	-	5
	WZ 82	Lower P4	14.2	7.3	-	7.3	3
	WZ 1589	Lower P4	13.9	7.1	-	7.1	3
	WZ 126	Lower I2	6.6	9.3	-	4.7	3
	WZ 124	Lower I2	6.8	9.5	-	2.9	5
	WZ 1591	Lower I3	8.6	9.7	-	6.2	5

Table 6

 6 Measurements of the calcaneus, patella, femur and radius and ulna of Ursus arctos from Wemeh cave/ Mesures du calcanéum, de la patella, du fémur, du radius et du cubitus d'Ursus arctos de la grotte de Wemeh.

			GL	DT	DAP L manu DT facdist DAP st DDP st
	WZ 456 Calcaneus	76.5 50.9 38.5 42.9	26.5	19.4	15.5
				GL	DT		DAP	DAP art.	L art.
	WZ 458	Patella		49.7	35.8		22.6	20.5	38.9
				DAP	DT	DT troch	DT fossa	DAP epimed
	WZ 460	Femur distal	(66.2)	81	43.2		15.4	(50.5)
	WZ 459	Femur distal	55.1	80	42.0		15.9	50.3
					DAP col	DT col	Bcaput	Dcaput
	WZ 463	Femur proximal	21.6		33.9	-	-
	WZ 462	Femur proximal	-		-	49.0	48.1
	WZ 1560	Femur proximal	20.5		32.7	42.7	41.2
							DAP col	DT ol
	WZ 471		Ulna proximal		69.2	45.1
	WZ 1561		Ulna proximal		63.1	-
							DTd	DAPd
	WZ 468			Ulna distal			41.4	24.2
	WZ 472			Fibula distal		25.6	15.8
	WZ 1566			Fibula distal		25.9	16.0
	WZ 466		Radius distal		-	58.4

Table

  Measurements of metacarpal (MC) and metatarsal (MT) bones from Ursus arctos from Wezmeh cave. Mesures des métacarpiens (MC) et métatarsiens (MT) d'Ursus arctosde la grotte de Wezmeh.All measurements are in mm. GL: greatest Length; Bp: breadth of the proximal end; Dp: depth of the proximal end; Bdia: greatest breadth of the diaphysis; Dia: smallest breadth of the diaphysis; Bd: breadth of the distal end; Dp: depth of the distal end/ Toutes les mesures sont en mm. GL : longueur maximale ; Bp : largeur de l'extrémité proximale ; Dp : hauteur de l'extrémité proximale ; Bdia : plus grande largeur de la diaphyse ; Dia : plus petite largeur de la diaphyse ; Bd : largeur de l'extrémité distale ; Dp : hauteur de l'extrémité distale .

			GL	Bp	Dp	Bdia	Ddia	Bd	Dd
	WZ	MT I	58.1	20.0	20.1	8.7	9.1	15.9	13.7
	WZ	MT I	-	21.2	21.6	-	-	-	-
	WZ	MT II	68.0	13.8	23.2	12.5	8.8	18.6	15.1
	WZ	MT II	-	13.3	20.8	-	-	-	-
	WZ	MTT III	74.2	18.1	26.8	13.8	10.4	18.3	15.8
	WZ	MTT III	-	16.4	-	12.1	9.1	-	-
	WZ	MT IV	77.0	18.0	23.6	12.2	11.0	18.4	15.7
	WZ	MT IV	68.6	-	-	10.7	9.3	16.0	13.2
	WZ	MT IV	-	16.2	22.3	12.2	9.8	-	-
	WZ	MT IV	-	17.1	23.1	-	-	-	-
	WZ	MT V	80.4	22.6	20.6	11.4	11.5	20.2	13.5
	WZ	MC II	74.4	16.3	21.9	11.5	10.9	19.5	16.7
	WZ	MC II	-	15.7	22.1	10.8	11.0	-	-
	WZ	MC III	70.6	15.7	20.3	10.6	9.1	18.8	15.7
	WZ	MC III	-	-	-	11.4	10.5	20.4	16.8
	WZ	MC III	74.1	-	-	14.1	12.6	20.3	18.2
	WZ	MC IV	76.9	18.2	25.0	11.7	10.7	19.2	17.6
	WZ	MC IV	74.9	18.5	26.0	13.0	11.8	20.8	18.7
	WZ	MC V	79.8	23.1	25.6	14.1	12.0	22.4(16.5)
	WZ	MC V	76.3	24.1	25.8	13.9	11.8	22.3	17.0
	WZ1575	MC V	-	25.4	26.7	15.3	10.4	-	-
	Table
	Measurements of phalanx I and II of Ursus arctos from Wezmeh cave.
	Mesures des phalanges I et II d'Ursus arctosde la grotte de Wezmeh.
			GL	Bp	Dp	Bdia	Ddia	Bd	Dd
	WZ	Phalanx I	35.7	18.1	16.2	12.7	7.8	14.7	10.1
	WZ	Phalanx I	41.6	20.2	16.2	13.0 10.2	15.5	10.5
	WZ	Phalanx I	38.4	19.4	15.9	13.0	8.6	15.4	10.0
	WZ	Phalanx I	40.2	18.0	15.1	12.0	9.4	14.6	10.5
	WZ	Phalanx I	-	-	-	-	-	16.4	10.3
	WZ	Phalanx I	36.6	18.2	15.0	12.4	9.5	15.4	8.6
	WZ	Phalanx I	34.3	17.4	14.5	11.4 10	13.6	9.1
	WZ	Phalanx I	42.1	21.3	17.3	13.9	8.9	15.7	10.7
	WZ	Phalanx I	36.5	19.6	14.4	11.5	8.8	15.0	8.0
	WZ	Phalanx I	37.4	18.1	15.9	12.6	8.6	13.9	10.0
	WZ	Phalanx I	37.6	19.0	14.6	12.4	8.8	15.4	8.3
	WZ	Phalanx I	35.9	16.3	12.6	10.4	8.3	14.7	9.0
	WZ	Phalanx I	32.5	18.9	14.4	11.7	8.7	12.4	8.2
	WZ	Phalanx I	34.4	15.7	13.0	10.4	8.1	11.4	8.9
	WZ	Phalanx I	40.0	19.0	16.2	13.0	8.5	16.2	10.2
	WZ	Phalanx I	49.3	21.1	17.2	14.6 11.2	16.3	12.1
	WZ	Phalanx I	38.4	19.6	17.5	13.5	8.8	15.7	10.1
	WZ	Phalanx I	38.2	18.1	16.4	12.7	9.6	14.4	10.0
	WZ	Phalanx I	33.8	19.0	13.6	11.8	8.3	13.9	8.4
	WZ	Phalanx I	44.5	19.0	15.5	14.3 12.2	16.1	11.9
	WZ	Phalanx I	-	-	-	13.7	9.5	16.0	11.0
	WZ	Phalanx II	30.2	16.8	13.7	11.4	8.3	13.6	11.5
	WZ	Phalanx II	31.2	18.0	14.7	12.3	8.3	15.6	12.5
	WZ	Phalanx II	29.1	16.6	13.6	11.5	8.0	16.1	11.7
	WZ	Phalanx II	29.4	15.1	12.5	11.6	7.4	13.4	11.4
	WZ	Phalanx II	-	-	-	-	-	13.3	11.1
	WZ	Phalanx II	29.1	16.6	13.6	11.5	8.0	16.1	11.7
	WZ	Phalanx II	29.4	15.1	12.5	11.6	7.4	13.4	11.4
	WZ	Phalanx II	-	-	-	-	-	13.3	11.1
	WZ	Phalanx II	29.1	16.6	13.6	11.5	8.0	16.1	11.7

All measurements are in mm (Measurement references in Table

7

)/ Toutes les mesures sont en mm (Références de mesure dans le Tableau 7) .

Table 9

 9 Measurements of some carpal, tarsal and phalanx 3 of Ursus arctos from Wezmeh cave. Mesures de carpes, tarses et phalanges 3 d'Ursus arctosde la grotte de Wezmeh.

	DPD	DT	DDP

All measurements are in mm. DPD: proximo-distal diameter; DT: transversal diameter; DDP: Dorso-plantar (palmar) diameter; GL: greatest length; Bp: proximal breadth; Dp: proximal depth; Barti: articular breadth; Darti: articular depth/ Toutes les mesures sont en mm. DPD : diamètre proximo-distal ; DT : diamètre transverse ; DDP : diamètre dorso-plantaire (palmaire) ; GL : longueur maximale ; Bp : largeur proximale ; Dp : hauteur proximale ; Barti : largeur articulaire ; Darti : profondeur articulaire .

Acknowledgments

We are grateful to Sanaz Beizaee Doust (Bioarchaeology Laboratory, Central Laboratory University of Tehran), for her assistance in the preparation of the figures, and to Michèle Ballinger (CNRS-UMR 7041, Arscan) for the drawings. We also thank Elizabeth Kerr for the English copy editing of the paper.

Disclosure of interest

The authors declare that they have no competing interest.