(bison priscus) in the iberian peninsula

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

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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


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


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


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


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


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).

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doi: 10.5710/AMGH.03.06.2016.2995 Submitted: March 4th, 2016

Accepted: June 3rd, 2016

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