En esta nota se realiza una revisión de los osos pleistocenos basada en el material español, que proviene, principalmente, de rellenos kársticos Se hace una revisión de la filogenia y distribución estratigráfica y geográfica de las diferentes especies

Torres Pérez-Hidalgo, Trinidad José

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Spanish karstic fillings: the key for pleistocene Ursids knowledge

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PALEONTOLOGIA
PALEONTOLOGIA
PALEONTOLOGY
13302
Spanish karstic fillings: The key for Pleistocene ursidae knowledge
T. de Torres (M Engineer Ph O.)
Empresa Nacional ADARa de InvestIgaciones Mmeras 5.A
RESUM
En aquesta nota es realitza una revisió deis óssos plistocenics. basada en malerial enpanyo/: material que pravé, lanamentalment,
de dipósits cárstlCS Es fa també una revisió de la filogenia i de la dlstnbucló estratigráfica i geográfica de les dtlerents especies.
RESUMEN
En esta nota se realiza una revisión de los osos pleistocenos basada en el material español, que proviene, principalmente, de
rellenos kársllCOS Se hace una revisión de la filogenia y distribución estratigráfica y geográfica de las diferenles especies.
SUMMARY
This paper deals of a review of Pleistocene bears, based upon 5panish material, which mainly anses lrom karslJC flllmgs We
make a review of phylogeny, and geographical and stratigraphical distribullons of the different specles.
Inlroduction
During Ihe lasl years. excavations made in Spaln have supphec
a very rlch material of fossil bears from dllferenl species and
ages, which have produced a great amounl 01 new knowledge
on phylogeny and strallgraphical and geographical dlstribulion.
Phylogeny
Ursus elruscus G CUVIER was the common ancestor of Ihe
mos! successlull evolutive trends (spetold and arctoid) 01
Pleislocene bears In EuraSIa
The spelold evolutlve trend is represented by two species
which show a more com pi icated denlal morphology, progressive
slmplilied denlal formula, and ske!eton also progressively more
heavily built The older is Ursus denin gen VON REICHENAU,
whlch is subs!lluted at the Riss by the cave bear -Ursus spelaeus
RQSENMÜLLER.HEINROTH- which dissapeared al the end 01
the Pleistocene times, probably because of paleolitic man and
brown bear demographic pressure,
The arctold evolutive trend is represented in Europa, until Ihe
Upper Riss time, by Ursus prearctos BOULE, which is not direetly
COnnecled with Ihe «true. brown bear - Ursus arctos lINNEO-
representing an specles closely relaled wilh ils dlfecl ancestor
Uelruscus
. Uprearctos appeared in Ihe Midd!e Pleistocene, in poputalions
WI!h locat dlfferences bul its skeleton was more heavlly built and
the dental morpflology was more complicated.
The polar bear -Ursus marilimus PHIPPS- which never was
found in Spaln, can be placed into the arclold group It probably
was derlvated from an ancestral populalion oi Uetruscus or U
Preare/os isolaled In an indelerminaled cold period,
The brown bear IS a lale emigrant whlch, al Ihe en 01 the~Ieistocene, moved southwrds Irom Asia, colonizing all Europa,
he Medllerranean border 01 Africa and N America.
Until Ihe Riss, Ihere IS an unfrequent bear in Ihe faunat
European panorama: Ursus medllerraneus F. MAYOR, closely
related wllh Ursus mlmmus DEVEZE&DEBOUtLLET.
Geographical and Stratigraphical distribution in Spain
(Fig.-2)
The oldest Spanish lossil bear lound In a karsllc syslem is
U minimus (sin Ursus arvemensis CR OIZ ET&JOBE RT) from Layna
(Guadalatara) 01 Lower Villafranchlan age No Spanlsh matellal 01
thls speCles, arises Irom no karslic deposlts Gavllle (Ilaly) and
Perpignan (France) The Spanish material could be nol 01 a true
cave has t, but alloctonous matenal lllllng karst ic lissures
Al the Middle-Upper Villalranchlan a true demographleal
explosion 01 Ursidae took place with the apparition 01 Uelruscus.
The oldest material has been lound at the lower Villafranchian
locality 01 Villarroya (Logroño). bul the boom 01 the species was
al Ihe Middle-Upper Villafranchian Val d'Arno (Italy), Tegelen
(Netherland), Saint Vallier (France) all of them non karsllCS
deposits In Spain, fossil remains 01 li etruscus have been found
at· Villar roya (Logroño), Puebla de Valverde (Ter uel), Ven ta Mlcena
(Granada) and Almenara (Castellón) The last is lhe oniy one with
a kars tic origln
The firsl true cave inhabilant has been found al Ihe Gran
Dolina (Burgos) 01 Günz age Relatively abundanl malerial,
comprising adults and cubs bones. have appeaderd. This bear,
a more evolulioned species bul closely related with ItS anceslor
Uelruscus appeared also in Mollet Racó, Barcelona (Mlndel) and
Pinilla del Valle, Madrid (Riss).
But In Europa Uelruscus evolved malnly in a spelold way
enormous increase 01 teelh size, skull volume (pneumatlzation)
and transversal dimension of postcraneal bones. This
phoenomena was probably related with Importanl changes in
leedlng habits and behaviour
The lirst representanl 01 Ihis evolutlve lrend is U denrngeri
215
---=:_.:=-o:::..=-:-e::._===:-_=--=--=_=:__:-:.==-=_.. ----1
o
z
'"ug
~
o
o
3
u
o
el
J, l) jl,MUIC ANl,J:5. i.,J iIIl.u:ITI""~ u AII(tOS ~ t(.IIl.AIt( .O~ ,t,L,vIOtOOA
lá '1 URSINAf . I 1
¡..;2=---++---I------r----t-------------T-- ~Ol>j~RINA~---+-----1--l
\ U.mn~us I 1-~l----\--l-~. '_ _1"'<;lO"'"
\ \ """,,"" ".."~.. I I /--~···~-\~.\-/t -j;!-- ---~
u.".., , "" u. ".u.an/;/ . II------+--..---~~~- - --t::-~~ --;;_1- --z¡A"""''''''''UMo HEMICYONINAf """ ........ u IfWSCUs. U' N,1l"tlllilLU:!I~ \ 'Z'OfClt. l~ jN~I'TO$ /
~ --~'7 "".., /-¿>o ---:::ii""'''''-------~----
~ S,to.VUS -~--== -----
u
o
i
Fig.- 1. Urs,dae PhY'ogeny Alrer Erdbrrn~ (1955) and Torres (1984), mod,/'ed
whlch mai nIy was fou nd In Europa assoclated wlth Ifuvio-Iacustrine
sediments But ,n Spaln there appeared 8xclusively in caves:
Lezetxiki (Guipuzcoa), Cueva Nueva 1·11 (Segovia), Santa Isabel
(Vizcaya) and Cueva Mayor (Burgos) -see Fig,·2-, Wilh the
exception 01 Lezetxlkl. of probabfy Rlss age, lhe other ones are
01 Mindef age
The more popular and besl known cave user is UspeJaeus
whose remains have been recovered In caves 01 2111 European
countnes, been unfrequents In open air paleonlological siles, wilh
the well konwn exceplions 01 the loess al Krasnodar and lhe
undersea sedimen!s 01 the Canal. In the au!hor's opinion the cave
bear appeared al lhe Upper Rlss Tres Simas (Burgos). Lezetxikl
(Guipuzcoa) and e) Troskaeta (Gulpuzcoa) -see Fig,·3-, where
there are small sized animafs This species. reached a true
ecofcgical boom al the Lower Wurm In Spain, despile of the lacl
thal traces of this speCles appeared In a great number o caves,
only live accummufallons 01 cave bear remains can be speclfied
Ekain (Deba, GUlpulcoa). El 1011 (Mora Barcelona) El Reguenflo
(Torrelaguna, Madrid), Raclau Viver (Bañolas Gcrona) and
Arrikrulz (Oñate, Guipllzcoa) With the excepllon 01 !he lasl one.
01 Wurm 111 age, the others are of Wurm I age
Spanish cave bear populatlons can be grouped In lhree
geographical areas Canlabflc. Medilerranean and Centraf
The first one is the best known and It IS eVldent that the cave
bear was a successfuf IfIhabltant of thls reglan ItS remalns
appeared in many caves, whlt the exceptlon 01 lhose sltualed in
lhe mounlalns.
From thls area, lhlS speele moved In \wo drfferenl dlrecllons
-Fig. -3- .. crossing the Cantabflan Cordillera reached the low lands
r-------- ---- - ..----...~- ..---.--.--------,
r------- ---- .-'. -----.
'--'--------------------------.,..-~
M,IIII_pio..[,g.,;:'t
~
§
~
Q
Fig.- 3. Cave be"" (fiSlllbur,,," ," rile Iberia" Peflmsu'a -1 Can'abrlc ",ea B
Med/lerranean Area C Central A/ea
'"
.,'
"'\...~ ;,
.,
Fig.- 2. Geographrcal d'slnb"l,on 01 less eomman bear s specleS Imds
e-lJ.el'useus I V'J'arroya'Logro~o (no I<arsl,e), 2.. Puebla de Valverde· Teruel (no
karslrcJ. 3· Venta Mrcena·Granada rno ka, si/eJ. 4 Almena/a Cas/ellón (karSI/C)
r·U 'usclnens,s layna·Guadaia¡" (~arsl/c) m U med"elfaneus l· Villav/ella·CaSlel!ón
(karsl/c;): 2.. Cau Bomjs·CaSlellÓn (I<ars/,c) d-U denll1gefl l· Cueva Nueva 1-11
Guadala¡ara (~ars¡,c). 2. Cueva Mayor-B,,,gos (karslie): 3 Cueva de Lezer.,k,
Gu'puzcoa (karsl/C). 4 . Parda~'·GUlp¡jzeoa (karSI/C) p-U p,eafClos· l.· P,mlla·Macrnd
r~MSrICJ, 2· MOlle! Racli·Gerofl~ r~~rSr¡cJ, 3 . P~I~clos de la Sterra-BurgOIi (karS!K:),
4 ' Gran Oc/ma 13'-"gos (~arsrICJ
of Nord Cas!lfha, cofonlzallng sorne caves of the North of lhe
province 01 Burgos Also, moved weslwards across non
karstifiabfe areas of GaliCia (metamorphlc formatlons) and in a
isolated slrata Intercafaled en a metamorphlC complex, where
sorne caves were devefopped,cave bear remalns have been lound
(G. del Purruñaf)
The Medlterranean populatlon was not connected wllh lhe
Canlabrian one, but probably was whll the meridional cave bear
populalions 01 France Thls group din nol move loo much
southwards. probably because 01 a kind 01 ecofogical barrier
216
¡lapllshed by the mouth and lower part 01 Ihe E:bro River. In the
pr:JVlnce cíí Castellon where malnly karstlllable rocks outcrop. tlle
calle bear has never lound And Ihls IS strange because lhls
speCles also hved in flver moulhs and marsh areas 01 Ihe
Netherlands
The Central population is Ihe mosl recenlly dlscovered, arld
1$ represented, almost excluslvely, by the Cueva de El Reguerillo
populatlon The cave bear probably moved soulhwards. Irom the
cantabric zone, gOlng around the eastern limils 01 lhe Nothern
Mesela, wh,ch probably acled as ecological barner. reachlng lIs
southern Iimit, which cOincides with lhe IImlt 01 karslifiable rocks,
al ¡he Cordillera Central And 11 did nol sUlpass the Soulhern
Meseta, bounded by no karsliliable rocks, towards lavourable
karslic areas ot Andalusla.
In conclusion: lhe Canlabric area was conlinually occupied
by cave bear populallon Irom de Upper Riss (Lezetxiki) unlil the
LJpper Wurm (arrikrul·Wurm 111), with a veslige populalion which
illssappeared al Ihe beginning 01 Holocene times Medilerranean
aod Cenlral populallon, probably were developped only durlng
Ihe Lower Wurm· Ihe lime 01 maximum successlul 01 Ihe specle
As It has been said belore, Ihere is also an arctoid evolulive
Irend Uprearclos, 01 large slratigraphical ad geographical
distribulion, probably because very low ecologlcal exigences (as
lis ancestor Uelruscus) This specie probably dissappeared under
the demographic pressure 01 the new appeared cave bear
populations. II was a Irue cave dweller and has been localed al
Gran Dolina and PalaCIOS de la Sierra (Burgos) 01 Günz age,
Mollel Racó (Gerona) 01 Mindel age and Pinilla del Valle Madrid
01 Riss age
Al the lower Wurm lhere IS a masslve emlgrallon Irom ASia
01 the «true" brown bear, Uaretos, under the pressure 01 a drasllc
climatic changemenl, colonlzallng N.Amerlca, aller crosslng Ihe
Behring Stralts, now Irozen, all Europa and the Mediterranean
borders 01 Asia and Afllca
Finally during Ihe Middle Pleistocene. a small sized species
appeared In lhe Medlteflar,ean border 01 Spaln U medilerrancus
Its remams have been lound ossocraded wlth karstlc sedlments
at Cau Borrás and Villavle¡a both 01 Rlss age
References
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ERD8RINK DP 1953 A review 01 fossil and recenl bears 01 Ihe
Old Worfd wllh r'Omarks on Iheir philogenese based upon thelr
dentltlon. J, de Lange Eds. 597 p
KURTEN B. 1975: The cave bear story Lile and death of a
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MIR A. SALAS R 1976 Tres nuevos carnivoros del yaCimiento
cuaternario de la Cueva d'en Mollel I Senñá Insl Inv Geol
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NOTARIO R. 1964 El oso pardo en España Serv Pesca Cont
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217


Spanish karstic fillings: the key for pleistocene Ursids knowledge

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