Ursus gr. spelaeus from Grotta del Bandito (Piedmont, Northern Italy). 1. Morphodynamics of teeth and indications of an evolutionary step

This study is focused on cave bears from the Grotta del Bandito (Cuneo Province, Piedmont Region, Northern Italy), especially their teeth. The morphodynamic analyses have been elaborated and compared with other cave bears from Northern Italy, and other cave bear taxa from particular European sites. The Grotta del Bandito bears are similar in size to other cave bears from Italy, and no differences were observed with the other cave bear species or subspecies (e.g. U. ingressus, U. s. ladinicus, U. s. eremus). Thus their inclusion in the U. gr. spelaeus is the most appropriate classification, and they show a simpler dental surface than the “modern” cave bears (i.e. ingressus). The question of whether a single species of spelaeus lived in North Italy, is also introduced. In fact, the differences in size are probably the result of population variability, and no clear evidence of other spelaeoid species and/or subspecies, i.e. ingressus, s. ladinicus and s. eremus, was discovered. delle Streghe) and in Mount Fenera, is an important site of cave bear remains in the Piedmont region near the boundary with France. In the past, ZUNINO & PAVIA (2005) conducted a su­ perficial survey of this cave which is geographically located on the right side of the Valley of Gesso (Fig. 1A). Its entrance is at 726 m a.s.l. and it is formed in the Jurassic limestone. The cave is composed of an E­W corridor (217 m) and three major cavities along with others of smaller dimensions. ZUNINO & PAVIA (2005) subdivided this cave into two parts with the same sedi­ mentary infills: the first, called the “eastern branch” is composed of a sand­clay with sedimentary pebbles, and the second one, very similar in the first part of the corridor to the “eastern branch”, occupies the western end (Fig. 1B). These authors also conducted three profiles (called L, M and N respectively) (Fig. 1C) in three different areas of the cave and discovered a vertebrate association listed in Table 1. 3. MATERIALS AND METHODS The studied teeth (Table 2) are stored in the Museo Civico di Sto­ ria Naturale of Milan (Lombardy). All dentition types are present in the sample: incisors, canines, premolars and molars. This study uses the morphodynamics developed by RABEDER (1999) and GRANDAL D’ANGLADE & LÓPEZ GONZÁLEZ (2004). In his paper RABEDER (1991) conceived and applied the morpho­ dynamic analysis on the P4/p4 only, and later elaborated it for the other teeth typologies, after having applied this method for the first time to the Arvicolidae of the Pliocene­Pleistocene (RABEDER, 1981). It was subsequently also proposed for the in­ cisors, premolars and molars of cave bears. The most important concept on which this morphodynamic method is based, is the evaluation over time of the development of the complication of the dental surface (from the simpler forms to the most compli­ cated and assigning each one a particular morphotype labelled Article history: Manuscript received Juny 24, 2019 Revised manuscript accepted January 23, 2020 Available online February 29, 2020


INTRODUCTION
Ursus gr. spelaeus is one of the most important members of the faunal assemblage during the continental Late Pleistocene in Eu rope and, consequently has been the subject of many publications. In order to establish the correct relationship among cave bears, ge netic analyses have been recently addressed by morphometric ana lyses with the result of creating some confusion in the elaboration of an accurate phyletic tree. In Italy the greatest number of contri butions to this debate have been based on morphometric and mor phodynamic analyses, although lately genetic and chronological studies have gained attention, albeit in a lesser number with respect to those from foreign populations of cave bears (PEREGO et al., 2001;ORLANDO et al., 2002;BONA, 2004;QUILES, 2004;ROSSI et al., 2006;, 2011, 2012, 2013ROSSI et al., 2018;, 2008ZUNINO & PAVIA, 2005;MARTINI et al., 2014;TER LATO et al., 2017. The most previously studied caves are those from Lombardy and Veneto in Italy, whereas those from Pied mont (Sambughetto Valstrona, Grotta del Bandito among the most important ones) have been only superficially analysed (e.g. ZUN INO & PAVIA, 2005;SANTI et al., 2005;. The aim of this study therefore is to analyse in detail the morphodynamics of cave bear teeth from the Grotta del Bandito (Cuneo Province) (bearing in mind that the morphodynamic indi ces are useful only to give an indication of the evolutionary level ROBU, 2016). This will also expand our knowledge on cave bears in an area of Piedmont near the cave of Basura (Liguria) where more definitely "modern" bears have been observed with respect to those of other cave bear populations in Northern Italy (QUILES, 2004).

GROTTA DEL BANDITO
The Grotta del Bandito (Cuneo Province) along with the other caves located in Valstrona (the most important being the Grotta Ursus gr. spelaeus from Grotta del Bandito (Piedmont, Northern Italy).

Morphodynamics of teeth and indications of an evolutionary step
delle Streghe) and in Mount Fenera, is an important site of cave bear remains in the Piedmont region near the boundary with France. In the past, ZUNINO & PAVIA (2005) conducted a su perficial survey of this cave which is geographically located on the right side of the Valley of Gesso (Fig. 1A). Its entrance is at 726 m a.s.l. and it is formed in the Jurassic limestone. The cave is composed of an EW corridor (217 m) and three major cavities along with others of smaller dimensions. ZUNINO & PAVIA (2005) subdivided this cave into two parts with the same sedi mentary infills: the first, called the "eastern branch" is composed of a sandclay with sedimentary pebbles, and the second one, very similar in the first part of the corridor to the "eastern branch", occupies the western end (Fig. 1B). These authors also conducted three profiles (called L, M and N respectively) (Fig. 1C) in three different areas of the cave and discovered a vertebrate association listed in Table 1.

MATERIALS AND METHODS
The studied teeth (Table 2) are stored in the Museo Civico di Sto ria Naturale of Milan (Lombardy). All dentition types are present in the sample: incisors, canines, premolars and molars. This study uses the morphodynamics developed by RABEDER (1999) and GRANDAL D'ANGLADE & LÓPEZ GONZÁLEZ (2004). In his paper RABEDER (1991) conceived and applied the morpho dynamic analysis on the P4/p4 only, and later elaborated it for the other teeth typologies, after having applied this method for the first time to the Arvicolidae of the PliocenePleistocene (RABEDER, 1981). It was subsequently also proposed for the in cisors, premolars and molars of cave bears. The most important concept on which this morphodynamic method is based, is the evaluation over time of the development of the complication of the dental surface (from the simpler forms to the most compli cated and assigning each one a particular morphotype labelled  with a letter from A to F which is associated with a morphody namic factor), and to correlate it with the cave bears evolutionary line (RABEDER, 1991;1999;RABEDER & TSOUKALA, 1990;ROBU, 2016 etc.). Figure 2 clearly explains the morphotype con cept for the P4/p4, the most important teeth in the evolutionary picture. Moreover, the methods do not give absolute results, but being a quantitative analysis advance an indication only of the evolutionary step reached by the studied specimens (RABEDER, 1991;ROBU, 2016). GRANDAL D'ANGLADE & LÓPEZ GONZÁLEZ (2004) proposed a similar morphodynamic elabo ration to that of Rabeder for m1. For the different components of this kind of tooth (paraconid, trigonid cusplets and talonid one) the so called "Convergence Index" is evaluated. As well as the morphodynamic index sensu Rabeder, the main target of the  "Convergence Index" is to indicatively evaluate the evolutionary step reached by the cave bears. For a more complete and detailed overview we included the data analysed by ZUNINO & PAVIA (2005) of part of the teeth discovered in this cave from and earlier preliminarily study. The following proposed considerations will be commented on and compared with the data of the cave bears from European caves in which recently different species and subspecies (Ursus ingressus RABEDER et al., 2004, U. spelaeus eremus RABEDER et al., 2004, U. spelaeus ladinicus RABEDER et al., 2004 have been identified. In these three caves, remains of U. ingressus and U. kudarensis BARYSHNIKOV, 1985 (the latter is described in LUBINE et al., 1985) were discovered and examined in greater detail. Referring to the teeth from the Bandito cave the morpho type "p" for the fossa lunaris-cingulum of I2 is the only one ob served; the standardized index (abbreviated "std") is low (58.91). (For meaning of the "morphotype" see above). This can be a con sequence of a low number of specimens (n=10), which makes it impossible to indicate the evolutionary step of these teeth. Also for I3 the low number of specimens (n=10) prevents any definite conclusions on their evolutionary level. However, some conside rations can be proposed: the kalyx distalis has "0" as the most frequent morphotype and only rarely (two teeth) the "1" one. The highest morphotypes are totally lacking. If this is confirmed, the evolutionary degree is rather low, but this deduction is thwarted by the scarcity of specimens. To conclude this short discussion on the morphodynamics of the incisors, all of the i2 have an "s" morphotype frequency typical of spelaeus (RABEDER, 1999). In i3 the lowest morphotype of the sulcus mesialis is B/C and in contrast to the other incisors, the main type most frequently ob served is the "D" morphotype. In this case also the scarce speci mens (n=8) prevents a definitive conclusion, but if this is con    Fourth upper/lower premolars (P4/p4). Figure 3. shows the distribution of the frequency of the morphotypes in the upper and lower fourth premolars. Considering also the analysis undertaken by ZUNINO & PAVIA (2005) for the p4, the frequencies of the morphotypes (sensu RABEDER, 1999) are: 1 A/B1, 1 B1, 1 B1/ C1, 1 C1/D1, 16 C1, 4 C2, 8 D1, 3 D2, 1 D3, 1 E1, while for P4 the morphotype frequencies are: 9 A (Fig. 4), 12 B, 1 C, 6 A/D, 5 D, 2 E. This distribution (the most frequent being that of the lowest morphotypes) is similar to that obtained for other cave bear popu lations in Northern Italy.

MORPHODYNAMICS OF TEETH
In summary, cave bear populations with variable sizes (from medium to large) lived in Northern Italy, but with very simple premolars; this is evidence that could indicate, a probable inter mediate step in the evolutionary tree. In Figure 5 we only present the results of our measurements for the upper and lower fourth premolars, as well as the relationship between the morphodyna mics and length of P4/p4 of bears from the Grotta del Bandito and other caves where different species and subspecies were found. Differences among the species are not noticed. A surprising view is shown in Fig. 6; in fact, the most important caves in Italy are placed in an interesting "cloud". A radiometric date of 29,130±90 yrs, was recently published for the Covoli di Velo by . In Figure 6, in which the morphodynamic index of P4 and p4 are in correlation, the Grotta del Bandito data are also in serted. We recall that the Fontana Marella and Mt. Generoso caves are located in western Lombardy near the boundary with Piedmont. In general, three aspects are noted: 1) both the Grotta del Bandito and Covoli di Velo (Veneto region) are placed along the "Austrian line", 2) the group composed by the levels FM an MG is substantially placed around the "French line", 3) the Grotta del Cerè (Veneto region) and the Grotta del Buco dell'Orso (La glio, Lombardy region) are placed in a position in which the Aus trian and French lines are not clearly separated. Data for Laglio (Buco dell'Orso Cave) premolars are too scarce to be able to ad vance any considerations. The exception is the S. Donà samples positioned along the "French line". Previously we indicated that the morphodynamic indices only give a relative indication of the evolutionary step reached by the bears (ROBU, 2016). The posi tion of the Bandito is between the MIS 5 (80130 Ky) and MIS 4 ( Fig. 6) and to our surprise we can see that these bears occupy an evolutionary step similar to that of the early "populations" of cave bears. The contradiction between the bears of the younger age and the relative simplicity of their dentition, is confirmed. Con sequently, we can also hypothesize that Italian territories were colonized by populations of bears coming from different areas of Europe, a mixed population that maybe separated rather early. Are the North Italian territories the zones of migration paths from west to east and vice versa? Is it possible that only a single phy letic line, the Austrian one, colonized Northern Italy and that the   influence of the "French line" (Basura (?) in the Liguria Region) may have arrived and tried to enlarge it in the last stages of evo lution? We need more data to confirm or reject this hypothesis. Finally, Figure 7 shows that the Bandito bears are placed within the "cloud" of the other Italian cave bears with the only exceptions of Cerè (most ancient deposit) and Basura (more ad vanced bears) which are separated from the main distribution. In Northern Italy we have a scarcity of radiometric dating, and only some populations of cave bears have been dated. In Table 3 we reported the radiometric data calculated up to the present.
First lower molars (m1). Considering the morphotypes pro posed by GRANDAL D' ANGLADE & LÓPEZGONZÁLEZ (2004) (Fig. 8), the m1s from the Grotta del Bandito correspond to those of advanced cave bears; the paraconids are equally sub divided in acute and straight and the protoconid from smooth posterior edge to rough posterior edge. Metaconids fall between the single with cusplet, to double, while the entoconids are fre quently double with cusplets and rarely, single with preceding cusplets (morphotype 13 sensu GRANDAL D' ANGLADE & LÓPEZGONZÁLEZ, 2004). Finally, the hypoconids exhibit an internal cusplet and hypoconulid. In the Bandito bears, teeth with only one metastylid (sensu RABEDER, 1999) are the most fre quent, although more derived morphotypes (with two or three metastylids) are also present. The entoconid area is composed of cusplets of about the same size, along with another of small size (most frequent) or two (one specimen). Regarding the enthypo conids only the simpler ones are found; comprising morphotype A (the simple one), A/B (with an enthypoconid but not yet divided from the hypoconid by a furrow) and B (enthypoconid clearly di vided from the hypoconid by a furrow). RABEDER (2014) studi ed the teeth from Križna jama (Slovenia) and classified them as U. ingressus; the simplicity of construction of the m1s from Bandito prevents their attribution to this species. The dis tribution of the main "Convergence Indices" (sensu GRANDAL  Fig. 9; these are rather homogenous, independent of the absolute length of the tooth. Also among these teeth a mix of "old" and advance features has also been noted; for example, considering the rela tionship between the "Total length -Talonid breadth" (Fig. 10) the point of the Bandito bears falls in the field of the "primitive" bears. In contrast, their relationship among the "Paraconid con vergence index -Total length", the "Trigonid convergence index ̶ Total length" and the "Talonid convergence index-Total length" suggests the Bandito bears be placed among the "advanced" cave bears. This is a confirmation that in Italy cave bears have a mix of features of tooth elements present in the finds of Middle and Late Pleistocene age.
Second lower molars (m2). Metrical data are reported in Fig. 11, while the calculated indices are shown in Fig. 12. A greater frequency of mediumlow indices for the development of the met alophids is observed (Fig. 12) (on 26 specimens, 11 have a "B" morphotype), whereas in the trigonid construction, the simpler form is lacking. The mesolophid area is characterized by mor photypes A (with a central crest and a first formation of a meso lophid) and B (with a continuous mesolophid developed as a crest or a line). Referring to the enthypoconids area, in the sequence of morphotypes, only the A (without enthypoconids) and the D (subdivision in three main parts) types are lacking. Intermediate indices are hence present in the Bandito "population", similar to that for others. Generally, a comparison with teeth from other caves shows, as in the Bandito specimens, that they have a low morphodynamic index; in the study on the cave bears from Križna jama (Slovenia), RABEDER (2014) calculated for the en thypoconid morphotype an index of 181.82 (std 98.12) from 11 specimens, and for the teeth from Risovača (Late Pleistocene, Serbia; CVETKOVIĆ & DIMITRIJEVIĆ, 2014)) the index is 181.3 (std 97.8). Also for the bears from Loutrá Aridéas (Greece) the enthypoconid index is 144.67 (std 78.08) (TSOUKALA et al., 2006). Furthermore, the "palaeopopulation" from Ajdovska jama (Slovenia) shows a very high morphodynamic index for the enthypoconid (std 166.03) that RABEDER (2011) considers at tributable to U. s. ladinicus, but in a followed control, the value of 166.03, (Tab. 7 in RABEDER 2011) is a mistake. In fact, RABEDER (2011: p. 36) quoted: "The enthypoconid-index is as tonishingly high and even supersedes the Gamssulzen population by more than 17%)". There is a mistakenly given standard value of 133 (instead of 185.3) so the correct standardized value is 117.4, which corresponds to the statement in the text (more than 17%, not more than 66%)". Here, considering the 28 elements in the specimens, it is difficult to distinguish these two species U. ingressus and/or U. s. ladinicus at the species or subspecies level for the Bandito bears.
Third lower molars (m3). A scarcity of specimens does not enable definite conclusions to be drawn, but observing the data the cave bears from the Bandito Cave have index values that are not completely low or high. The low number of specimens pre vent further discussion as the sample is too small. ̶ M2). Also for these teeth (M1s) the morphodynamic indices (14 specimens and re lated to the paracone) are low (index is 75, std is 39.89); this is important because it confirms that for the cave bears in Italy, M1s were simple and that in the evolutionary position they occupied an intermediate place. The other morphodynamic indices are de fined for the metacone (14 in specimens, index is 110.71, std is 62.90) and the proto?/metacone (14 specimens, index is 135.71, std is 43.43).

First and second upper molars (M1
Considering the main components of the tooth, the morpho dynamic indices most frequently have an intermediate range; in general the indices for the Bandito M2s are low when compared with those from the other caves, aligning them with the other an alysed teeth. In conclusion we propose in Fig. 13 a summary of the mor phodynamics of the teeth from the Bandito cave compared with other species of bears (RABEDER et al., 2005). In fact, the pre molars of the Bandito Cave fall inside the cloud formed by U. spelaeus, U. s. eremus and U. s. ladinicus; with only U. ingressus being well separated from the cloud (Fig. 13A). With these mor phodynamic indices, it is difficult to subdivide the different spe cies with the exclusion of ingressus, and consequently in our opinion it is better to indicate this distribution (with the Bandito bears) as gr. spelaeus. The same consideration can be advanced observing the Fig. 13B. The separation of the different species, however, is greater compared to that of the Fig. 13A, as well as the third one. Having utilised different indices for the different kind of teeth it remains uncertain whether the Bandito bears be long to one species or another.

MORPHODYNAMICS-ALTITUDE RELATIONSHIP
A relationship between the altitude of the cave and the morpho dynamics of the teeth has often been searched for in the past (RABEDER et al., 2005;LAUGHLAN, 2012). Here we inves tigate whether there is any possible relationship between the al titude and teeth (morphodynamic and morphometry) from the Bandito Cave. Fig. 14AE shows the relationship between the premolar and molar indices and the altitude of the cave; the data of the Bandito bears falls in between the spelaeus eremus and ingressus taxa. Moreover, a hypothetical relationship between the m2 size and the altitude of the cave is also shown (Fig. 14 D); there is not an equal position of the Bandito bears; in fact in the Fig. 14A the mark is placed next to some of those belonging to s. eremus and s. ladinicus, while in the second one the Bandito bears are approximated to ingressus. The picture is confused and a relationship between the size of the tooth and the altitude of the cave cannot be demonstrated. The same considerations could be proposed when considering the enthypoconid of m2s and the to tal length of these teeth. On the basis of these data, a relationship between the distribution of the different species of cave bears and altitude also does not exist. Concerning the Bandito bears another consideration can be postulated. It is very difficult to determine which species of cave bear inhabited this area of Piedmont dur ing the Late Pleistocene. Finally, a relationship between the P4/ p4 index and the altitude of the cave entrances referred to Ursus ingressus only, is also shown (Fig. 14E). Independently of the taxon considered (U. ingressus), the position of the Bandito point is interesting because it opens a discussion about the "migration path" and consequently about the colonization of the cave bears and also perhaps some phyletic considerations. In fact, the Ban dito is placed at the beginning of the distribution of the "Alpin gruppe" (sensu FASSL & RABEDER, 2015) nearly on the same altitude where the Arzberg cave is placed (the latter is included in their "colline gruppe" by FASSL & RABEDER, 2015). The distribution of the Italian localities is unexpected. From a com mon beginning, the lineage direction is very different to that of other bears. Is it true that the Italian cave bears are effectively a "separate" assemblage from others in Europe? Many signs would seem to confirm this (size, a low or medium morphodynamic in dex of the teeth, both the plumpness indices and the KI for the metapodia are similar to those of other European "populations" etc.). Further studies are necessary to solve this dilemma.

IS URSUS SPELAEUS THE SINGLE SPECIES THAT LIVED IN ITALY?
Often in the past we have supported the thesis of the presence of a single species Ursus spelaeus in Northern Italy (e.g. SANTI & ROSSI, 2014). Previous studies on fossils from different caves of Northern Italy have shown a strong homogeneity among the Ita lian "populations". The Grotta del Cerè (Verona, Veneto) is the exception, where in fact, among the fossiliferous material, re mains of U. deningeri VON REICHENAU, 1904 and U. arctos LINNAEUS, 1758 together with U. spelaeus have been found . In their research on the metapodia from selected caves of Italy, SANTI & ROSSI (2014) have shown that morphometrically Ursus spelaeus ROSENMÜLLER, 1794 is the single species that lived in the Italian Alps. The differences in size are probably a result of population variability and no clear evidence of other spelaeoid species and/or subspecies, i.e. ingressus, s. ladinicus and s. eremus, has been discovered. A relation ship between the size and the altitude of the cave is absent. In Italy therefore up to now, chronological data and mtDNA analy ses are lacking, a complete study, that includes the morphometry, morphodynamics, chronology and genetics, is missing. Until now, only the Basura assemblage (Liguria Region) seems to be very different from the other cave bears in Northern Italy; in fact, the morphodynamic indices of the fourth premolars are higher compared with those of other Italian cave bears (QUILES, 2004;SANTI & ROSSI, 2014). For this assemblage a possible higher step in the evolutionary scale could be hypothesized. Contrary to the Basura population, in the Pocala Cave (Friuli Venezia Giulia, Eastern Italy), the cave bears have a large size, but the fourth pre molars has a very simple structure (ROSSI & SANTI, 2015). Consequently, a confused picture on the morphodynamic indices is highlighted with the highest value (Basura) and the lowest ones for the Grotta delle Fate assemblage (Liguria). Two different ex planations could be proposed: either this represents the presence of more taxa of Ursus, or an intraspecific variability of a single species. The discussion is, at the present time still open.

CONCLUSIONS
The study of the teeth of the cave bear from the Grotta del Ban dito (Piedmont, Northern Italy) leads to the following conclu sions. The teeth are generally of medium size, although in some cases large specimens are also present; they are similar to other taxa of cave bears (ingressus, s. eremus, s. ladinicus). With these morphometric results the studied bears are determined as cave bears, but for the Bandito sample and the Italian cave bears in general it is not easy to distinguish which species they belonged to. The Bandito bears show the features of different "species"; in some cases these elements appear to be "primitive", in others they are more evolved. To indicate a dominant taxon is difficult, be cause the taxonomic determination sensu RABEDER et al. (2004) is mostly based on genetics, not on morphometrics. We prefer to indicate the Bandito cave bears as gr. spelaeus, not a single species, and no relationship with the altitude of the cave is noted.
From a general point of view, these bears occupied probably, an intermediate evolutionary step, and therefore more evolved individuals are also present. In fact, in the fourth premolars, ZU NINO & PAVIA (2005) found higher morphotypes than in the other "Italian" "populations" in which these are absent. The ex planation is not clear, but the closeness of the Basura Cave (Lig uria Region) in which more advanced bears are found (QUILES, 2004) can suggest postulation of for example, the hypothesis of the replacement of the "primitive" bears with "advanced" ones. If in the Basura Cave the disappearance of the primitive bears seems clearer, in the Bandito Cave it is less clear. In other words, during the Late Pleistocene, close to the cave bears extinction (in fact the assemblage of the Bandito Cave and that of Fontana Marella are very similar), two different "populations" probably cohabited for a short period in the Bandito Cave area, one of these composed of "primitive" bears and a second, formed of a few, but more evolved specimens. Therefore, the complete substitution did not occur at once because the cave bears vanished and became extinct. The problem is still open.