FIRST COMPLETE SKULL OF A LATE PLEISTOCENE STEPPE BISON (BISON PRISCUS) IN THE IBERIAN PENINSULA
JONE CASTAÑOS1 PEDRO CASTAÑOS2 XABIER MURELAGA1
1 2
Universidad del País Vasco/EHU, Facultad de Ciencia y Tecnología, Departamento de Estratigrafía y Paleontología, Apartado 644, E-48080 Bilbao, España. Sociedad de Ciencias Aranzadi, Centro Geo-Q, Santimami Auzoa, E-48940 Leioa, España.
Submitted: March 4th, 2016 - Accepted: June 3rd, 2016
To cite this article: Jone Castaños, Pedro Castaños, and Xabier Murelaga (2016). First complete skull of a late Pleistocene steppe bison (Bison priscus) in the Iberian Peninsula. Ameghiniana 53: 543–551. To link to this article: http://dx.doi.org/10.5710/AMGH.03.06.2016.2995
PLEASE SCROLL DOWN FOR ARTICLE
Also appearing in this issue: Braincase and endocranial morphology of proterochampsids suggest high olfactory acuity, as in modern crocodiles.
New glossopterid fertile organs from the Permian of Australia and a revision of Ottokaria from across Gondwana.
Remains of red algae from the Danian and the post K/Pg coral reef communities in northern Patagonia.
AMEGHINIANA - 2016 - Volume 53 (5): 543 – 551
ARTICLES
ISSN 0002-7014
FIRST COMPLETE SKULL OF A LATE PLEISTOCENE STEPPE BISON (BISON PRISCUS) IN THE IBERIAN PENINSULA JONE CASTAÑOS1, PEDRO CASTAÑOS2 AND XABIER MURELAGA1 1
Universidad del País Vasco/EHU, Facultad de Ciencia y Tecnología, Departamento de Estratigrafía y Paleontología, Apartado 644, E-48080 Bilbao, España.
2
Sociedad de Ciencias Aranzadi, Centro Geo-Q, Santimami Auzoa, E-48940 Leioa, España.
[email protected]
[email protected];
[email protected]
Abstract. The Kiputz IX site has provided one of the best-preserved late Pleistocene bison populations in the southern Pyrenees and has yielded the first almost complete skull of a steppe bison (Bison priscus) in the Iberian Peninsula. This Bison priscus skull is compared on morphological and osteometric grounds with other specimens of steppe bison from Europe and North America. The skull from Kiputz IX falls within the range of the extinct subspecies Bison priscus mediator. Available data support the evidence of three chronological subspecies of Bison priscus (Bison priscus gigas, Bison priscus priscus, and Bison priscus mediator) during the Middle and late Pleistocene. Key words. Steppe Bison. Skull. Marine Isotope Stage 2. Northern Spain. Basque Country.
Resumen. PRIMER CRÁNEO COMPLETO DE BISONTE ESTEPARIO (BISON PRISCUS) EN EL PLEISTOCENO TARDÍO DE LA PENÍNSULA IBÉRICA.
El yacimiento de Kiputz IX ha proporcionado una de las colecciones de bisonte mejor preservadas al sur de los Pirineos durante el Pleistoceno tardío, además de aportar el primer cráneo completo de bisonte estepario (Bison priscus) encontrado en la Península Ibérica. Este trabajo presenta los resultados del estudio morfológico y osteométrico del cráneo de Bison priscus y compara las medidas con otros ejemplares de bisonte estepario de Europa y América del Norte. El cráneo de Kiputz IX ha dado valores cercanos a los de la subespecie extinguida Bison priscus mediator y proporciona datos que respaldan la evidencia de cronologías de tres subespecies sucesivas de Bison priscus (Bison priscus gigas, Bison priscus priscus y Bison priscus mediator) durante el Pleistoceno medio y tardío. Palabras clave. Bisonte estepario. Cráneo. Estadio Isotópico Marino 2. Norte de España. País Vasco.
THE steppe bison (Bison priscus) was a characteristic middle
European assemblages of steppe bisons include those from
end of the last Ice Age (McDonald, 1981; Pfeiffer, 1999;
La Vaissière (Brugal and Fosse, 2005), and Romain-la-Roche
and late Pleistocene species which became extinct at the
Jaurens (Guérin and Valli, 2000), Habarra (Prat et al., 2003),
Benecke, 2005). The geographical range of this eastern ex-
(Vercoutère and Guérin, 2010) in France and Kiputz IX in
Europe into Siberia and across Beringia into North America
Findings of complete steppe bison skulls in Europe are
tended from northern Spain through central and Eastern
Spain (Castaños et al., 2012).
(Kahlke, 1994). In this large area, the subgenus Bison evolved
rare and, at present, only three neurocrania with both
mental conditions (Von Koenigswald, 1999).
one from Movileni (Romania; Codrea and Ursachi, 2010) and
into different species that were adapted to various environThe literature on steppe bisons in Europe is abundant.
The assemblages from Ilford (Davies, 1874) in England,
horn-cores, one from Irtysh River (Russia; Flerow, 1977),
one from Romain-la-Roche (France; Vercoutère and Guérin, 2010) have been published, as well as three partial skulls
Châtillon-Saint-Jean (Mourer-Chauviré, 1972) in France,
from Habarra (France; Prat et al., 2003).
born and Taubach (Flerow, 1968, 1976) in Germany and
vided the richest collection of steppe bison remains in the
basic systematics of the species. There are also regional
in this sample is a complete steppe bison skull (Fig. 2). To
Mosbach and Mauer (Schertz, 1936), Ehringsdorf, SüssenIsernia-la-Pineta (Sala, 1987) in Italy, have established the
(Brugal, 1984–1985; Crégut-Bonnoure and Guérin, 1996) and
sexual dimorphism (Drees, 2005) studies. Recent records of AMGHB2-0002-7014/12$00.00+.50
Kiputz IX (Fig. 1) is a late Pleistocene site that has pro-
Iberian Peninsula (Castaños et al., 2010). The main specimen date, it is both the first finding with these features in the Iberian Peninsula and the southermost record of a complete
543
AMEGHINIANA - 2016 - Volume 53 (5): 543 – 551
skull in Europe. The scarcity of this anatomical element in
wide and 2.5 m long, to circa 6 m2 of floor area and a 4.2
the fossil record increases the interest of this discovery. The
m-thick sediment-filled bottom divided into eight strati-
teometrical study that could be used for more fragmented
bones (Cervus elaphus) establishes the time interval of the
aim of the present paper is to offer a morphological and os-
materials and which also allows these measurements to be
compared with other specimens of the steppe bison from Europe and North America.
GEOLOGICAL SETTING
graphic levels (Fig. 1.2). Radiocarbon dating of red deer
study site as 32,810 ± 390 BP to 11,750 ± 60 BP (Beta Analytic, Florida, USA; Castaños et al., 2012). Stratigraphic levels were classified and divided based on macroscopic sediment descriptions including colour, particle size and
sorting and morphology of the pebble and cobble fraction
Kiputz IX is a palaeontological site (Mutriku, Gipuzkoa)
(Castaños et al., 2006). Levels D and F show the highest
ern Iberian Peninsula) (Fig. 1.1). This site was discovered in
One of the most important features of the site is the
located in the Basque Country (southern Pyrenees, north2003 and excavated between 2004 and 2007. Kiputz IX is
an open karst shaft which functioned as a natural trap where the animals fell and could not escape. The hole is 2 m
density of bone remains.
excellent preservation of its fossil specimens. Therefore,
its characteristic as a trap, combined with the preservation of the site, has provided a representative sample of the
Figure 1. 1, Map showing the geographical location of the Kiputz IX site (Mutriku, Gipuzkoa, Southern Pyrenees); 2, Stratigraphic sequence of the upper Pleistocene Kiputz IX site (Mutriku, Gipuzkoa).
544
CASTAÑOS ET AL.: SKULL STEPPE BISON OF KIPUTZ IX
species that lived in that ecosystem during the late Pleistocene (Castaños et al., 2006).
The bison skull described in this paper was recovered in
July 2006 from Level F, at a stratigraphic depth of 37 dm. Radiocarbon dating of red deer bones (Cervus elaphus), from
the same stratigraphic depth, establishes the age of this
specimen as 18,850 ± 80 BP (BETA-355781, Beta Analytic, Florida, USA). The results were calibrated at 2 sigma based
on the Intcal09.14C calibration dataset, calculated with the
CALIB REV6.1.0 program (Reimer et al., 2009), to 22,188– 22,679 cal BP.
The faunal assemblage includes a variety of taxa among
which red deer (Cervus elaphus), reindeer (Rangifer tarandus)
and bison (Bison priscus) are the most common species.
Although red deer is clearly dominant, the bison remains
constitute the best-preserved populations in the Iberian Peninsula to date (Castaños et al., 2012). Additionally, the
bison skull preserved almost intact can be considered unique
Figure 2. The bison skull from Kiputz IX site (Mutriku, Gipuzkoa, Southern Pyrenees) in frontal view. Scale bar= 10 cm.
three indices refer only to the horn-cores. Index 2 indicates
the relative horn-core curvature through a ratio between
in the Iberian Peninsula for this species.
the lower curve length (Measurement 4) and the straight
MATERIALS AND METHODS
dorsoventral horn-core compression through a ratio be-
traction it needed to be protected with plaster reinforce-
6 and 7). Index 4 shows the relative lower horn-core length
until further restoration. The skull was held by a perspex
length of core on lower curve, tip to burr (Measurement 4)
facilitating measurements. It was thus handled and trans-
burr (Measurement 3) which is missing in two specimens.
The skull was found in a very delicate state, so for its ex-
ment in situ. The process was able to protect the remains
base and forks and later stored in a wooden box made for ported more safely. The restoration was performed in the Archaeological Restoration Service of the Provincial Council
line length (Measurement 5). Index 3 indicates the relative
tween the diameters of the horn-core base (Measurements in relation to the frontal width of the skull. In Index 4 the
is used to replace the horn-core length, upper curve, tip to Despite this change, the value of the index does not vary.
To compare the specimen from Kiputz IX, Simpson’s
of Gipuzkoa (Basque Country, Spain) by Giorgio Studer. The
Ratio Diagram is used (Simpson, 1941). This methodology
deposited in the Basque Government official repository
history between modern bison species and the extinct
All the skull measurements were taken in accordance
technique in which measurements can be compared to a
specimen was labelled as KI IX.2B.37.412 (Fig. 2) and was institution for the Province of Gipuzkoa in San Sebastián.
was applied in an attempt to establish an evolutionary
Bison antiquus occidentalis. The ratio diagram is a univariate
with those defined by Skinner and Kaisen (1947) and Mc-
standard. In this case we use Bison (Bison) bonasus measure-
measurements are shown in Figure 3, and also several in-
dard.
Donald (1981) (Tab. 1), and given in millimetres. These
dices following Skinner and Kaisen (1947: p. 142–143) were calculated, with certain modifications (Tab. 2). These provided
information on the proportional relationship of various
anatomical parts and were used to eliminate misleading impressions of size and shape. The first index referred to the
whole skull and related the postorbital width (Measurement
14) with the total skull length (Measurement 1). The other
ments given by Skinner and Kaisen (1947, p. 214) as a stan-
DESCRIPTION OF THE SKULL
The Kiputz IX skull is conserved in a very good condition.
It is only missing a portion of the frontal bone, some lateral
elements of the temporal and the vomer region. The skull
was found upside down (ventral side up) resting on a large
limestone clast. This limestone clast had broken into the
545
AMEGHINIANA - 2016 - Volume 53 (5): 543 – 551 TABLE 1 – Bison priscus skull measurements used in this work following to Skinner and Kaisen (1947) and McDonald (1981). Assignation in Figure 3
Skull measurements
2
Spread of horn-cores, tip to tip (SHTT)
1
Condylobasal length
3
Horn-core length, upper curve, tip to burr (CLUC)
5
Straight line distance, tip to burr, dorsal horn-core (TB)
7
Transverse diameter horn-core base (TD)
4 6
Length of core on lower curve, tip to burr Vertical diameter horn-core base (VD)
8
Minimum circumference, horn-core base (CHC)
10
Width of occipital condyles (CW)
9
Width of occipital at auditory openings (GWA)
11
Depth, nuchal line to dorsal margin of upper border of foramen magnum (DEP)
13
Width between bases of cores
12
Depth, nuchal line to dorsal margin of lower border of foramen magnum (DEP)
14
Least width of frontals, between horn cores and orbits (WHCO)
16
Antero-posterior internal diameter of orbit
18
Distance P2-M3
15 17 19 20
Greatest width of frontals at orbits (GPW) Dorso-ventral internal diameter of orbit Distance M1-M3
Width between P2
skull due to the pressure of the overlying sediments. As a
plane, the oval section in the bases (Sala, 1987) and the
but also the bones of the ventral part. For this reason, in
cided with a male individual. The horn-cores are long, ex-
result there was a gap affecting not only the dorsal region
dorsal view the entire nasal bone and the endonasal forma-
presence of large burrs, we deduce that the specimen coin-
panded laterally and backwards beyond the nuchal line, with
tions have disappeared. Neither the central portion or the
tips bending forward.
lacrimofrontal and lacrimomaxilary sutures are still in the
This character is presumably correlated with age as it in-
right side of the front were preserved. The supraoccipital, process of fusion. These sutures close late in life in extant
The orbits are less tubular and with normal protrusion.
creases with individual ontogeny. The supraorbital foramen
individuals of Bison (Skinner and Kaisen, 1947).
is roofed over. The two maxillaries are also preserved and
and very well conserved, especially the left one. The small
premaxillaries are complete. All molars, despite the three
The two horn-cores are almost complete, well ossified
losses of the right horn-core do not affect the evaluation of
its dimensions. At their base, they are bent downwards
conserve the infra-orbital holes and, unusually, the two missing premolars, were in use.
below the frontal surface. The surfaces show well marked
RESULTS AND DISCUSSION
Based on the angle between the horn-core and the sagittal
Pliocene of Asia under the names of Bison paleosinensis
furrows on their whole length, deeper on the ventral side.
546
The subgenus Bison appears for the first time in the late
CASTAÑOS ET AL.: SKULL STEPPE BISON OF KIPUTZ IX
and Bison sivalensis. The first dispersal to Europe is recorded with Bison tamanensis, with two subsequent evolutionary
lineages on one hand Bison schoetensacki with primitive
characters, small size and horn-cores small and thick, and
on the other hand the steppe bison (Bison priscus) with
long horn-cores and a larger size. The size and the great
development of the horn-cores of the skull of Kiputz IX justify their inclusion in the species Bison priscus discarding
its relation with other species of bison.
The steppe bison was a typical ungulate of middle and
late Pleistocene steppes of the Northern Hemisphere that extended from Spain through Europe to Siberia and across
Beringia into North America (Kahlke, 1994). For several millennia it evolved into different subspecies that were
adapted to various environmental and biogeographical
conditions. The patterns of its extinction at the end of the
Pleistocene and of its replacement by the current forms of American bison (Bison bison) and European bison (Bison
bonasus) are still unknown (Benecke, 2005).
The species Bison priscus is represented by three sub-
species in Eurasia (Flerow, 1976). Bison priscus gigas from
the first half of the middle Pleistocene is the largest
Eurasian bison. The tip-to-tip horn-core spread reaches 2
metres. Its geographical area extends from southern Siberia
and eastern Europe to the Volga (Flerow, 1977). The situation of its findings in the central area of the species distribution can justify the large size. Bison priscus priscus, from the second half of the middle Pleistocene, has a tip-to-tip
horn-core spread of between 0.9 and 1.36 metres. Its range extends from western Europe to the Ienisei River in the east
Figure 3. The bison skull showing the location of measurements. 1, dorsal view; 2, nuchal view; 3, ventral view.
and Kazakhstan in the south. Bison priscus mediator from the
late Pleistocene, is smaller than the nominal subspecies, with the horn-cores more curved and with a tip-to-tip
TABLE 2 – Indices used in this work following Skinner and Kaisen (1947). 1. Index skull: 100 x Least width of frontals, between horn cores and orbits (14) Over-all length of skull (1) 2. Index of curvature: 100 x Length of core on lower curve, tip to burr (4) Straight line distance, tip to burr, dorsal horn-core (5) 3. Index of compression: 100 x Vertical diameter horn-core base (6) Transverse diameter horn-core base (7)
4. Index of length: 100 x Horn-core length, lower curve, tip to burr (4) Least width of frontals, between horn cores and orbits (14)
547
AMEGHINIANA - 2016 - Volume 53 (5): 543 – 551
to in the following paragraph. In this species, there is a clear
spread of less than 0.9 metres, with the same geographical
distribution (Vercoutère and Guérin, 2010). Some authors
sexual dimorphism that manifests itself in the form and in
subspecies (Brugal, 1984–1985). The smallest size of Bison
et al., 2003). Comparing skulls without a prior separation of
the skull measurements, especially in the horn-cores (Prat
disagree on the chronological boundaries of the latter two
priscus mediator may be caused by the onset of evolutionary
genders, might lead to superimposing variation among sub-species and sexual dimorphism. To avoid any bias, only
changes resulting in the emergence of the European bison.
male skulls were included in this comparison.
Measurements and indices of this skull are compared
with those available on six Bison priscus specimens referred
The first specimen is a male skull recovered from the
TABLE 3 – Measurements of steppe bison skull from Kiputz IX (Mutriku, Gipuzkoa) and other European and American sites (see text). Measurement 1 2 3 4 5 6 7 8 9
10 11
Bison bonasus
Kiputz IX
608
970 425
526
605
226
375
268 187 71
106.5
238
331
79
243 95
12
142
14
254
13 15
16
316
17
1005
1118
487
590
600
430 390
107
107
280
297
136.4 115
139.5
158 295 334 81
440
136.5 140
410
150.5
139
128
150.5 320
353
Blue Babe
707
595
556
590*
565*
279
273
2000 540
940
922
120
430 326
Tsiigehtchic
335 314 363
130 305
197 347
85
74
103.5 74
Index 1
48.28
448.76
Index 3
89.87
99.53
Index 4
1050
Irtysh River
62.5
19
Index 2
Movileni
59
18 20
357
Romain- laRoche
Habarra
143.3 105.51
119.04 144.06
124.87
97.5
191.08
92.3
46.89
49.1
211.46*
203.66*
The first column indicates numerically the type of measurement described in Figure 3. All measurements are given in millimetres. Skull references in the text. * Measurements and indices from horn-cores with sheaths.
548
CASTAÑOS ET AL.: SKULL STEPPE BISON OF KIPUTZ IX
Figure 4. Simpson diagram applied to different bison skull samples from Kiputz IX, other European and North American sites.
French site of Habarra (Arudy, France) with an age bracketed
compared with the published measurements for various
BP (sector L). It has been attributed to Bison priscus media-
table provides several means from a sample of modern Eu-
la-Roche Cave (Doubs, France) dated from the latest middle
collected by Skinner and Kaisen (1947, p. 418) in their ex-
priscus (Vercoutère and Guérin, 2010). The next skull was
of including this information is to use them as a standard
between 14,500 ± 260 yr BP (sector E) and 28,200 ± 700 yr tor (Prat et al., 2003). There is another skull from Romain-
Pleistocene which has been attributed to Bison priscus
found in the late Pleistocene levels at the Movileni site,
Romania (Vaslui district; Codrea and Ursachi, 2010) and
assigned to Bison priscus. The fourth specimen is a skull re-
covered in the Irtysh River, Bashkirian Autonomous Republic of Russia, assigned to Bison priscus gigas, and
attributed to the Mindel/Riss stage (Flerow, 1977). Another
specimens (Tab. 3). In addition the second column of the
ropean bison (Bison bonasus). These measurements were
tensive work about the fossil bison of Alaska. The purpose sample in the Simpson diagrams (Fig. 4).
Figure 3 shows that the measurements of the Bison
priscus skull exceed those of the European bison. The
greatest differences occur in the length of the horns. These
data confirm the decrease in skull size between Pleistocene and extant bisons (Brugal, 1984–1985, p. 16; Be-
specimen was found in Tsiigehtchic, Northwest Territories,
necke, 2005, p. 422).
and dated at 11,830 ± 45 yr BP. The last skull, discovered
River skull are larger than in all other specimens. This con-
specimen is known as “Blue Babe” (Guthrie, 1990). The last
Pleistocene bison, and is consistent with its chronology. Its
Canada (Zazula et al., 2009). It is attributed to Bison priscus
north of Fairbanks (Alaska, USA), is 36,000 years-old. This two skulls are partially mummified.
Identification of Bison priscus skulls is usually based on
the horn-core character suite (McDonald, 1981). These
characters allow distinguishing the three described subspecies. Measurements and index values of this skull are
Table 3 and Figure 4 indicate that the values of the Irtysh
firms its attribution to Bison priscus gigas, the largest horn-cores are more compressed than in Kiputz IX and
Romain-la-Roche (Index 3).
The measurements of the Moliveni skull are similar to
those from Romain-la-Roche in the length and section of the
horn-cores and slightly lesser than the latter in the occipi-
549
AMEGHINIANA - 2016 - Volume 53 (5): 543 – 551
tal. However, the values of these two skulls exceed those
individual was an adult male. The good preservation of the
The Roman-la-Roche skull is attributed to the nominal sub-
indices. Metric data of the Kiputz IX skull were compared
of the Kiputz IX, Habarra, Tsiigehtchic and Blue Babe skulls.
species for its osteometrical characters (Vercoutière and
Guérin, 2010). The Movileni skull has no subspecific attri-
bution. According to the metric similarity with the Romain-
skull has allowed obtaining 20 measurements and four
using Simpson diagrams with those of other of Bison priscus fossil skulls recovered at sites in Europe and North America.
All Bison priscus skulls are larger than those of the Eu-
la-Roche skull, it might also be attributed to Bison priscus
ropean bison (Bison bonasus). The skull from Irtysh River
cores is larger than in the Irtysh River skull and slightly
measurements than the other five skulls, especially in the
priscus. The compression of the Romain-la-Roche horn-
(Russia) attributed to Bison priscus gigas presents higher
lower than the Kiputz IX skull (Index 3). In addition the
length of the horn-cores. The Romain-la-Roche and Movileni
to the frontal width than the Kiputz IX skull (Index 4).
skull and all the remaining skulls. Therefore the attribution
Movileni skull shows a greater horn-core length in relation The Habarra skull shows a trend very similar to that of
the Kiputz IX one with somewhat higher values (Fig. 4). In this sample it has only been possible to calculate the second
index which defines the curvature of the horn and it almost matches the Kiputz IX index (Tab. 4). These data and the attribution to Bison priscus mediator to the Habarra skull
can confirm the initial hypothesis (Castaños et al., 2012)
that the Kiputz IX sample corresponds to the same subspecies as Habarra. Also the curvature of the horn-cores of both skulls is very similar (Index 2).
The Tsiigehtchic skull measurements are slightly lower
than those of Kiputz IX, except the horn-core length (Measurement 4). However, it is important to note that this sample has been preserved mummified and consequently
retains the horn sheaths, so that the horn size is considerably overestimated. This event distorts both Measurement 4 and
Index 4. However, if we ignore this measurement and we
consider all of the above, this skull is similar to the Kiputz
IX specimen.
The Blue Babe skull is the smallest one within the com-
parison sample. The measurements are close to those from
Tsiigehtchic (Fig. 4). It also presents deviations in Measurement 4 and the last index (Tab. 4) because it is a mummified animal and therefore preserves the horn sheaths.
CONCLUSIONS
The Kiputz IX site has provided a nearly complete skull of
a steppe bison (Bison priscus) which is the first in the Iberian Peninsula and a unique finding to date. The skull is in a very
good condition, and is just missing a portion of the frontal
bone, some lateral elements of the temporal and the vomer
region. Dental and horn-core morphology indicate that this
550
skulls display intermediate values between the Irtysh River of the Romain-la-Roche skull to Bison priscus priscus can be
extended to the Movileni skull. The skulls from Kiputz IX and Habarra, because of their metric similarity, can be attributed
to Bison priscus mediator confirming the conclusion of the
metric study of the postcranial skeleton (Castaños et al., 2010).
ACKNOWLEDGEMENTS
We thank M. Sasieta and J. Mari Arruabarrena, members of Munibe Taldea (Azkoitia, Basque Country) for the discovery of the site of Kiputz IX and help in excavating it. We also wish to acknowledge the help of G. Studer, a member of the Archaeological Restoration Service of the Provincial Council of Gipuzkoa (Basque Country, Spain), who made the skull restoration. The authors would like to express gratitude to Benedetto, Sala, and Pierre-Olivier Antoine for critical remarks and recommendations that improved the manuscript. The study presented in this paper has been partially funded by the research project of the Spanish Science Ministry HAR201453536-P and by the Research team GIU15/34 of the University of the Basque Country UPV-EHU. Jone Castaños has a postdoctoral grant “Contratación para la especialización de personal investigador doctor” from the University of the Basque Country (UPV/EHU).
REFERENCES
Benecke, N. 2005. The Holocene Distribution of European bison: the archaeozoological record. Munibe (Antropologia-Arkeologia) 57: 421–428. Brugal, J.-P. 1984–1985. Le Bos primigenius Boj., 1827 du Pléistocène moyen des grottes de Lunel-Viel (Hérault). Bulletin du Musée d’Anthropologie préhistorique de Monaco 28: 7–62. Brugal, J.-P., and Fosse, P. 2005. Les grands bovidés (Bison cf. schoetensacki) du site Pléistocène moyen de la Vayssière (Aveyron, France). In: E. Crégut-Bonnoure (Ed.), Les ongulés holarctiques du Pliocène et du Pléistocène. Association française pour l’Étude du Quaternaire, Paris, p. 75–80. Castaños, J., Castaños, P., and Murelaga, X. 2006. Estudio osteométrico preliminar de los restos de ciervo (Cervus elaphus) del yacimiento paleontológico del Pleistoceno superior de Kiputz IX (Mutriku, Gipuzkoa). Geogaceta 40: 163–166. Castaños, J., Castaños, P., Murelaga, X., and Alonso-Olazabal, A. 2012. Kiputz IX: Un conjunto singular de bisonte estepario (Bison priscus) del Pleistoceno superior de la Península Ibérica.
CASTAÑOS ET AL.: SKULL STEPPE BISON OF KIPUTZ IX Ameghiniana 49: 247–261. Castaños, J., Murelaga, X., Castellanos, I., Alonso-Olazabal, A., Zuluaga, M.C., and Ortega, L.A. 2010. Evaluación del grado de diagénesis en huesos fósiles mediante espectroscopía de infrarrojos. Geogaceta 49: 11–14. Codrea, V., and Ursachi, L. 2010. The Pleistocene Steppe Bison (Bison priscus, Bovidae, Mammalia) from Movileni (Vaslui District). Oltenia 26: 281–286. Crégut-Bonnoure, E., and Guérin, C. 1996. Ordre des Artiodactyles; Famille des Bovidae. In: C. Guérin, and M. Patou-Mathis (Eds.), Les grands Mammifères Plio-Pléistocènes d’Europe. Masson, Paris, p. 62–106. Davies, W. 1874. Catalogue of the Pleistocene Vertebrata from the neighbourhood of Ilford, Essex, in the collection of Sir Antonio Brady. London (printed for private circulation). Drees, M. 2005. Sexual dimorphism in Pleistocene Bison priscus (Mammalia, Bovidae) with a discussion on the position of Bison schoetensacki. Senckenbergiana lethaea 85: 153–157. Flerow, K.K. 1968. Die Bison-Reste aus den Travertinen von Weimar-Ehringsdorf. Abhandlungen des Zentralen Geologischen Instituts 23: 171–199. Flerow, K.K. 1976. Die fossilen Bisonreste von Taubach und ihre Stellung in der Entwicklungsgesschichte der Gattung Bison in Europa. Quartärpaläontologie 2: 179–208. Flerow, K.K. 1977. Gigantic Bison of Asia. Journal of the Palaeontological society of India 20: 77–80. Guérin, C., and Valli, A.M.F. 2000. Le gisement pléistocène supérieur de la grotte de Jaurens à Nespouls, Corrèze: les Bovidae (Mammalia, Artiodactyla). Cahiers scientifiques du Muséum d’Histoire Naturelle de Lyon 1: 7–39. Guthrie, R.D. 1990. Frozen Fauna of the Mammoth Steppe: The Story of Blue Babe. University of Chicago Press, Chicago, 323 p. Kahlke, R.D. 1994. Die Entstehungs-, Entwicklungs- und Verbreitungsgeschichte des oberpleistozänen Mammuthus-Coelodonta-Faunenkomplexes in Eurasien (Großsäuger). Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft 546, Frankfurt-am-Main, 164 p. Koenigswald, W. von. 1999. Paläoökologie und Vorkommen des pleistozänen Auerochsen (Bos primigenius BOJANUS, 1827) im Vergleich zu den großen Rindern des Pleistozäns. In: G.-Chr. Weniger (Ed.), Archäologie und Biologie des Auerochsen. Wissenschaftliche Schriften des Neanderthal-Museums 1, Mettman, p. 23–33. McDonald, J.N. 1981. North American Bison: Their Classification and Evolution. University of California Press, Berkeley, 350 p. Mourer-Chauviré, C. 1972. Étude de nouveaux restes de vertébrés provenant de la carrière Fournier à Châtillon-Saint-Jean (Drôme) III – Artiodactyles, chevaux et oiseaux. Bulletin de l’Association Francaise pour l’Étude du Quaternaire 4: 271–305. Pfeiffer, TH. 1999. Systematic relationship within the Bovini with
special references to the fossil taxa Bos primigenius Bojanus and Bison priscus Bojanus. In: G.-Chr. Weniger (Ed.), Archäologie und Biologie des Auerochsen. Wissens chaftliche Schriften des Neanderthal-Museums 1, Mettmann, p. 59–70. Prat, F., Delpech, F., Cancel, N., Guadelli, J.L., and Slott-Moller, R. 2003. Le bison des steppes, Bison priscus Bojanus, 1827, de la Grotte d’Habarra à Arudy (Pyrénées-Atlantiques). Paleo 15: 1–102. Reimer, P.J., Baillie, M.G.L., Bard, E., Bayliss, A., Beck, J.W., Blackwell, P.G., Ramsey, C.B., Buck, C.E., Burr, G.S., Edwards, R.L., Friedrich, M., Grootes, P.M., Guilderson, T.P., Hajdas, I., Heaton, T.J., Hogg, A.G., Hughen, K.A., Kaiser, K.F., Kromer, B., McCormac, F.G., Manning, S.W., Reimer, R.W., Richards, D.A., Southon, J.R., Talamo, S., Turney, C.S.M., van der Plicht, J., and Weyhenmeyer, C.E. 2009. IntCal09 and Marine09 radiocarbon age calibration curves, 0-50,000 years CAL BP. Radiocarbon 51: 1111–1150. Sala, B. 1987. Bison schoetensacki Freud. From Isernia la Pineta (early Mid-Pleistocene, Italy) and revision of the European species of Bison. Paleontographia Italica 74: 113–170. Schertz, E. 1936. Der Geschlets-Untershied an Metapodien von Bison. Senckenbergiana 18: 357–381. Simpson, G.G. 1941. Large Pleistocene felines of North America. American Museum Novitates 1136: 1–27. Skinner, M.F., and Kaisen, C. 1947. The fossil Bison of Alaska and preliminary revision of the Genus. Bulletin of the American Museum of Natural History 89: 123–256. Vercoutère, C., and Guérin, C. 2010. Les Bovidae (Mammalia, Artiodactyla) du Pléistocène moyen final de l’aven de Romain-laRoche (Doubs, France). Revue de Paléobiologie 9: 655–696. Zazula, G.D., MacKay, G., Andrews, T.D., Shapiro, B., Letts, B., and Brock, F. 2009. A late Pleistocene steppe bison (Bison priscus) partial carcass from Tsiigehtchic Northwest Territories, Canada. Quaternary Science Reviews 28: 2734–2742.
doi: 10.5710/AMGH.03.06.2016.2995 Submitted: March 4th, 2016
Accepted: June 3rd, 2016
551