Departement du Rhone - Museum, Lyon
NEW TAXA OF ALPINE CAVE BEARS (URSlDAE, CARNIVORA)
Gemot RABEDERt, Michael HOFREITER2, Doris NAGEL1, and Gerhard WITHALM'
ABSTRACT Rt':sUMt Morphological and metrical differencesbetween seve Lesdifferences morphologiques et metriqucs entre plu ral alpine cave bear associations of the same age are sieurs populations d'ours des cavernes alpins de meme large enough to suspect more than one evolutionary age sont suffisamment importantes pour qu'on puisse line. The contemporaneous age of several morpholo supposer I'existence de plusieurs lignees evoiutives dis gically distinct forms is confinned by radiocarbon tinctes. Les datations par Ie mdiocarbone confinnent la dating. Moreover analyses of fossil DNA indicate at coexistence de fonnes morphologiquement differentes. least three different lineages within the cavebear De plus les analyses sur I'ADN fossile montrent group. For two of these groups, both morphological qu'existent au moins trois lignees d'ours des cavernes. and genetic data show evidence of reproductive isola Pour deux de ces groupes, les donnees morphologiques tion. Thus, we suggest that Ursus spelaeus comprised comme les donnees genetiques prouvent des isolats de at least two different species. Ursus illgressus n. sp. reproduction. Aussi nous suggerons qu' Ursus spelaeus occurred in the eastern parts of the Alpine region and correspond au moins a deux especes differentes. Ursus in the Dinarids of Slovenia and Croatia. For two other ingressus n. sp. est repandu dans les Alpes orientales et morphological forms, subspecific status with Ur.HI$ dans les Alpes Dinariques de Siovenie et Croatie . Pour spe/aeus fadillicus n. ssp. in the Dolomites and Ursu$ les deux autres fonnes morphologiques, nous propo $pelaeus eremlls n. ssp. in the Totes Gebirge is sug sons la distinction de deux sous-especes, Ursus spe iaeus ladinicus n. ssp. dans les Dolomites et Ursus spe gested, as they are genetically relatively close and no data about their reproductive relationship is yet laeus eremus n. ssp. dans Ie Massif des Totes Gebirge : available. elJes sont en effet relativement proches genetiquement mais il n'existe pour J'instant aucune donnee sur Icur croisement eventuel.
Mots-cl6i : Keywords phylogeny, eYolWion, uuonomy, cave bear. phylogenie. evolulion, /(uOIlOlllie, ours des DNA-analysis cal'emes, ADN.
INTRODUCTION on the excavation activity was intensified under the leadership of Gernot Rabeder; excavation campaigns
The study of the Alpine bear caves has a - in some cases lasting for many years - were carried out in the Ramesch Knochenhohle, the Brieglersberg long tradition aI lnstitute of Palaeontology at the cave, the Gamssulzen cave, the Brettstein cave and in University of Vienna. It began more than 80 years ago under the direction of Othenio Abel in the the Lieglloch (all of them in the Totes Gebirge). DrachenhOhle of Mixnitz, continued under Kurt Other excavations were undertaken in the Ehrenberg in the Schreiberwand cave on the Hartelsgraben cave (Gesause), the Herdengel cave Dachstein, the Merkenstein cave near Baden, the and (he Schwabenreith cave near Lunz am See bear cave of Wind en, and finally in the Salzofen cave (Lower Austria), in the Nixloch near Losenstein in the Totes Gebirge as well as in the Schlenken Ternberg in Upper Austria, the Sulztluh cave in the Durchgangshohle in !.heOstcrhom group. From 1980 R1itikon (Switzerland), the Conturines cave in the
'lnstitutfUr PaJaootologie dcr Universitill Wien, lMa;< Plaock-Gesellsc:haft, evo Inslilut rur lutionlire Anthropologie, leipzig
Cahiers scicntifiques - Departement du Rh6ne· Museum, Lyon, Hors sene n02 (2004) - p. 49-67. II fig., 5 tabl., 2 pI. Actes du 9< Symposium international sur I'ours des cavernes, Entremont-Ie-Vieux (Savoie, France), septcmbre 2003
,,°2 (2004) 49 Cahiers scienlifiques/ Hors serie Centre de Consermtiol! er d'£tude des Collectiolls
Dolomites (Italy) as well as in the Potacka zijalka in lution in this range of time (hypothesis of saltation) the Karawankes (Slovenia). Under O. Abel qu.estions or a bear population on a high level of evolution about the morphology of function and the life style of advanced from a so far unknown area to the eastem the cave bear became imponant as well as the reasons Alps (hypothesis of immigration):' for their extinction (ABEL & KYRLE 1931), while Another high alpine small fonn was discovered in the K. Ehrenberg was above all fascinated by the rela Dolomites. In the years 1998 to 200 I a large number tionship between bear and palaeolithic man (keyword: of fossils was excavated from the Conturines cave, cave bear cUlt). In the last two decades due to consi situated at an altitude of nearly 2800 m high that were derably advanced excavation methods we could go at first assigned to a smaller cave bear relative. deeper into questions of evolution and fine stratigra Because of the mixture of archaic attributes (small phy. because now statistically relevant numbers of fos dimensions. P3 in over 25% of the skulls still existing, sils are available and the possibility of radiometric il primitive, m2-trigonidtal primitive) and progressive dating gives totally different age classifications attributes (12, ml-and m2-enthypoconid on a high (DOPPF.s & RABEDER 1997). level of evolution. also m2 -mesolophid, M2-metha It was K. EHRENBERG (1929) who nOliced that the cave loph and above all P4) the Conturines cave bear must bear remains from different caves differ in size. based have an exceptional position. (RABEDER, 1999) : on the variability of the metric values. In the first des "'... this leads to the conclusion that this high alpine cription of the very small bear form from form was derived probably from a late Middle Schreiberwand cave he created the term "hochalpine Pleistocene U. dellillgeri-group... A genetic connec Kleinform" (high alpine pygmy form). The reduction tion with the high alpine cave bears of the nonh Alps of the dimensions was seen as an adaptation to life in ... can be excluded". the high Alpine region (shon summers, long winters). The work on the evolutionary statistics of metapodial Other authors explained the on average smaller bones done by G. WtTHALM (2001) and on tibiae by dimensions as sex-specific differences: the smaller M. FROEMEL (2001) clarified the exceptional position females would have preferred the higher situated of the high alpine cave bears also in respect to the caves for hibernation.This hypothesis,jokingly called postcranial skeleton. The more developed first meta "Pascha-theory" was taken to ad absurdum quite carpal- and metatarsal bones as well as the relatively recently (RABEOER, 2(01). longer and more slender tibiae are interpreted as an The excavation in the Ramesch bone cave from 1979 adaptation to tife in high alpine regions. to 1984 was the key to modern cave bear research in Now we can discuss the relationship between the the Alps. The until then unique site in an undisturbed, other cave bear faunas and the three aforementioned more than two meters thick profile of sediment which groups. Taking the paleontological data we can accept made it possible to ask questions about metric and with utmost probability that the bear from morphological details. The big difference to the Gamssulzen cave inhabited the following caves: remains from the Drachenhohle near Mixnitz leads at Nixloch near Losenstein-Ternberg, Lieglloch first to the conclusion that the "high 'alpine small (Austria), Schnurenloch (Switzerland), Potacka zijal foml" occurs also in the Ramesch bone cave (HtLLE & ka. Mokriska jama (Slovenia). Vindija (Croatia); the RABEDER, 1986). The taxonomic status of this fonn of bear from Ramesch cave is also documented in the the high alpine region could not be bener argued at Salzofen cave, whereas the bear from Conturines cave that time. But the conclusions drawn from the excava is so far only known from this cave in the Dolomites tion in the Gamssulzen cave brought a change. of South Tyrol. Finally, numerous cave bear faunas I.n the Totes Gebirge, two cave bear forms that were cannot be slotted into this scheme, for example the unequal in size and at extremely different levels of faunas of the Brieglersberg cave and the Brettstein evolution, lived contemporaneously for many thou cave (Totes Gebirge) or the caves of easl Switzerland, sands of years and apparently they did not interbreed. e.g. the Drachenloch near Vatlis and the Sulzfluh Already by then the possibility of a formal separation cave. (Ramesch bear and Gamssulzen bear as two different With the analysis of ancient DNA we have now a species) was taken into consideration (hypothesis of completely new possibility to investigate the phyloge speciation). netic relationship of glacial animals. It is well known Two hypotheses were presented (RABEDER, G., for a long time that in the sediments of caves organic
1995:81) : "There are abnormal differences of evolu molecules like collages are conserved much longer tion also to other absolute dated bear faunas that can than in fossils from open sites because of the constant only be explained that either there was an erratic evo- temperature and humidity in caves. This applies also
CaMers scie!JIijiqlles Nors sirie / 11°2 (2004) 50 Departement du RhOne - Museum, Lyon
to DNA. Although the preservation of PCR amplifia the University of Vienna and the Institute of ble genetic material differs from cave to cave, DNA Evolutionary Anthropology of the Max Planck analysis is possible on bones up to about 100.000 Gesellschaft in Leipzig - led to the investigation of years old. various aspects of the phylogeny and taxonomic posi The cooperation between palaeontology and genetics tion of cave bears. - in this case between the Institute of Palaeontology of
Fig 1 - Location map of the type localities: GS Gamssulten cave, RK - Ramesch KoochenhOh1e. CU - Conturines cave (Italy) and ZoolithenhOhle (Germany).
LOCATION OF THE TYPE LOCALITIES P4/4-lndex The so-called P4/4-index, the geometric mean from the index of the lower jaw-P4 and the upper jaw-P4, seems to be by far the best parameter to investigate the MATERIAL AND METHODS morphological level of evolution. The P4-indices are morphodynamic indices. Thus, the steps of evolution
are measured on the accessoric elements of the chew From all three alpine caves as well as from the additionally investigated sites significant numbers area like cusps and zygomatics compared to the ances of isolated teeth and postcranial elements exist. In the tral shape: every additional hunch (for example meta Conturines cave and the Gamssulzen cave this is the conulus, metaloph. protostyl. paraloph on the P4 sup., case for the material as a whole, whereas in the para-, meta- and hypoconide, meta- and hypolophid Ramesch Knochenhohle it was possible to collect ade on the P4 inf.) is adding to the index of a morphotype quate numbers of teeth from defined areas to discrete +1. Since it has been adopted, the P4/4 index has seve units. The metric and morphologic methods were des ral times proved to correlate with the age of the fossil (RABEDER, cribed quite recently in detail, for teeth see RABEDER assemblages 1983. 1999). (1999), for the metapodial bones see WITHALM Enthypoconid of infoand info (200I), and for the tibiae see FroEMEL (200 I). M I M2 From the numerous morphodynamic indices of the The one-piece or multi-part hunch lingual of the hypo teeth only those that proved to be significant were conid on the ml and m2 is called enthypoconid. It ari selected for taxonomic questions: ses step by step from the inner Ilank of the hypoconid
Cahiers scielllifiques Hor.Y $irie n02 (2004) 51 / Ce/l/re de Con.rervalion el d'E/we des Colleclions
with the help of length channels by cutting of an auto over time (RABEDER, 1999:28) but there is no linear nomous hunch that can be divided in two or three sub correlation with the P4/4-index. hunches. The reason for this complication of the che - - wing area is apparently an enlargement of the chewing , , - - - area to withstand the chewing pressure of the M 1- , "'. respective M2-exterior. The different morphotypes are ..< (II, . --!� ... �.,"" /�: shown in figure 2: The number of the developed ,. <. j{:, :\J, . enthypoconid-hunches affects the value of the factor , iPlO -\"- - • • assigned to a certain morphotype (0 = without enthy - , ,
poconid, I = one hunch, 2 = two hunches per enthy Fig 2 - TI>e morphotypes of M2 inr.-enthypoconid.shown on a M2 poconid and so on). The index of a population cor info dell\. in occlusal view. Morphotype A: without enthypoconid (Schwabenreith cave). morphotype B: one cusp (Gamssulzen cave). responds to the mean of the assessed factors multi morphotype C: lWO cusps (Gamssulzcn cave) and morphOlype . D: plied by 100. The enthypoconid values also change three cusps (Conturines cave).
DATA
MEASUREMENTS OF THE SKULLS FROM THE THREE TYPE LOCALITIES (TAB. 1-3)
amesch skulls holotype paratypes �ny. nr. RK332 RK.. RK 1001 RK 1004 i 1"<" sin. ,,�, sin. dex!. s n. ,,�, sin. dex!. otal lenglh 416,3 - · 426,0 odylobasal length 406,1 · 390,0 · po · · ngth of tooth row C-M2 165,0 167,0 170,0 . 163, �ngth of tooth row P4-M2 83,4 83,7 93,4 91,9 ,5 94,S 90, 94 �ngth of tooth row Ml-M2 66,. 67.9 74,2 75,0 74,2 73, 75,1 � • · · 3 alveole 041 041 . f l · · traOrbita width 76,2 91,7 rnterorbital width 84,1 106,0 · · FpraOrbital width 119,5 · · · storbitalwidth 61,9 · · · postral width on the P4 82,4 92,4 92,3 92,0 ro · stral width on the Csup. . 114,0 102,0 rougal width 249,5 · 291,0 · ion width 137,9 · 170,0 ·
p · poopital hight 126,4 · 129,0 · · reatest heigth 01 skull 152,4 166,0 stral heightbefore the P4 73,1 100,0 · ·
'" , RK 38 119 . 452 59. ��mesch mandibles " 44. kIo do, d" sin d" d" do, condylar length · · 295,3 · · 306,5 coronoid height · · 123,0 · · 136,5 condylo-coronoid length · · 104,3 · · · ight of mandible below m2 69,2 54,3 74,8 66,3 73,4 · �englh of tooth row C-m3 · · 160,6 190,6 (190 ·
· · ngth 01 tooth row p4-m3 89,9 100,0 102,5 (101,6 · len9th of tooth row ml-m3 85,2 n,O (85,2 84,1 (85,.
TABL. 1 - Skull measurements of Ursus sfNlaeus eremU$ n. ssp. from Ramesch KnochenhOhle. Upper Austria. Austria.
Cahiers sciellfijiques Hors serie I n02 (2004) 52 du RlI6nl! Museum, Dlpartemelll - Lyon
onturines skutls holotype paratypes 714 704 Cu703 nv. nr. Cu Cu Be. kIo sin. de�. sin. de�. sin. de�. ,,,. de�. otal length 340,4 347,0 398,3 340,2 fondylobasallength 303,8 309,1 363,8 340,2 �ngth of tooth row C-M2 140,5 141,0 · 140,0 158,9 157,3 138,0 134,2 P4-M2 95,' �ngth of tooth row 90,. 92,S 92,S 90,' 95,0 ",a ".1 Ml-M2 12,7 72,8 76,1 76,1 �ngth of tooth row 74,S 74,S ",3 ",a 3 alYeole • • • • · · · · wiclth 75,S 76,0 82,1 61,0 rfraOrbital 67,S n,s nterorb�al wiclth ",a 73,S �praOrbital Width ",' 92.2 101,3 105,3 po StOrbital width 77,S 76,2 77,2 76,6 rostral width on the P4 76,9 13•• 84,4 13,S rostral width on the Csup. 85,2 84.2 90,. 62,0 gal width " 195,0 212,5 202,0 �tion width 116,5 127,7 140,7 124,0 preatest helglh 01 slcull 130,7 136,0 142,2 143,5 rostral height before the P4 63,. 71,8 76,6 "
mandibles paratypes �onturines 715 72. 12 501-1 501-3 710 v.nr. Cu " H2 ...... de, d.. de, . de, "" �, ", 258.1 · 287,0 299 'condylar length 290,. 288.7 •• 25'� 250,' �ronoidheight 105,4 106,2 · · · · · · yIo-coronoid length 81,8 · 105,7 · 105.2 · · · fo� · · ight of mandible below m2 54.0 60,2 69,3 6O.S 54,S 54,7 r.ength of tooth row C-m3 161,3 161,2 174,2 180.0 178,1 175,9 164,4 160,9 � p4-m3 105,3 ",3 · ..,S · 96.4 lOS,S nglh of tooth row 93•• � , 83,1 82,4 84,3 82,4 IenQth of tooth row ml-m3 84.0 " . 61,9 ",a
• s�/(Jelll? /lJdinicus TABL.2 Skull measurements of Ur5us n. ssp. from Conturiroescave, Sooth Tyrol (IUlly).
Conturines skutls 00_""" paratypes Cu 714 Cu 704 Cu 703 v. nr . Be. klo sin . ..�. ., . ..� . sin. de�, sin. Oex\. otal length 340,4 347,0 398,3 340,2 :ondylobasallength 303,8 309,1 383,8 340,2 ngth of tooth row C-M2 140,5 141,0 · 140,0 158,9 157,3 138,0 134,2 P4-M2 ngth of tooth row 90.• 92,S 92,S 90,. 95.' 95,0 ",a ",1 Ml-M2 12,7 74,5 12,8 76,1 76,1 ngth of tooth row 74,5 ",3 ",a 3 alYeole • • • • · · · · 75,5 width 76,0 82,1 61,0 fraorbital n,s nterorbital width ",a 07,S 73,S supraorbital width 96,' 92,2 101,3 105,3 postorbital width n,s 78,2 n,2 76,6 ostral width on the P4 78,9 73,8 84,4 73,5 ostral wlcnhon the Csup. es,2 84,2 90,. 62,0 ugal width 195,0 212,5 202,0 ¥ " 6tion width 116,5 127,7 140,7 124,0 reatest heigth of skuW 130.7 136,0 142,2 143,5 slral heightbefore the P4 83,. 71,8 78,6 "
Conlurines mandibles paratypes v.nr. Cu 715 720 12 25 501-1 501-3 710 882 d" de, de, ., de, "" ... "" 258,1 · 288,7 287,0 259,2 ."",h 290,. 299,. 250,' ::oronoid�y,,, height 105,4 106,2 · · · · · · ::ondylo-coronoid length 61,6 · 105,7 · 105,2 · · · ighl of mandible below m2 54,0 · 60,2 ",3 · 60,S 54,S 54,7 �ngth of tooth row C-m3 161,3 161,2 174,2 180,0 176,1 175,9 164,4 160,9 �ngth ollooth row p4-m3 93,6 105,3 96,3 · 96,S · 96,4 105,5 � 84,0 , 61,9 ,1 82.4 82,4 �nQth 01 tooth row ml-m3 .. . 83 84,3 ",a TABL. 3 - Skull measurements of Unws ingl'r.UlI$ n. sp. from Garru;sulzen cave, Upper Ausuia. Alisuia.
53 Cohien scieflfijiqwuI Hon serie ,,"2 (2004) Celllre de COllservaliOlI el II'Elude des Col/ectiolls
STATISTIC PARAMETERS OF PREMOLARS AND MOLARS (TAB. 4)
m" • m' _ m1 '" RK4·5 '" '" " . � � RK.·5 '" '" " . � Imn' �n 28.83.3 211.� 2II . oeGoI 211.31155 �. � '3.i!11!5 .3.56.5 '4.'1oI ".i!II!51 .3._ . � . so .t. dev ." . - '.4' •.» '.� '.5 .. " ..- .� '.n 0.11' •." mu � .• n._ n.' n, n._ » .• '5.7 111.0 '0.0 '0.0 111.5 17 .2 min �.. �- �., �, �, �- .2.0 '" .2.0 .2.3 11.4 .2.3 , " ., � " ". . " " " � � ". . "
�. , �- m2 '" R"'''5 '" '" '" � '" R"'4' 5 '" '" " . " � � :N.17 '8.22 17)1' 17.0415 18.:l5 Im ean .»" 5 . 5� 211.'. � .� . .,... H.GO deY. . - '.� '.M '.M '.� '.� • • • e'37 ..� '.'11 1.17 •." , • • 9 a\., mu 3'.4 »0' m n._ n, ».• '11.11 �, �., '11.11 '11.11 2' . 5 . 15.4 '4.2 C' �, �- �., �.- �. 21.0 14 2 .., '5.5 .5.5 .. 5 n " � " � 5 » " � �
�. • �- m3 '" RK" S '" '" " . � '" R"'4·5 '" '" '" a, Im .," n.M 2• . 4' 2<1.51 � . � �. Q n.M 11.36 10 .38 ..,. '1.e8 >1 .e8 1 11 . " ,,- '.N '.� ..� ..� .M 1.14 us '.n '.M .n ,.'0 1 .14 m" n._ 27.1 �.' m n.o 3'.3 �.' �.' �.' '11.7 �.' 2'.4 ",in 111.4 10.0 2'.0 �.' 2'.' �.- 14.5 14.2 ." 10.3 IS.' 10.3 , N � ,. � � M � 5 n " " �
.. . • .. - M1 '" RK4.5 '" '" " . � � RK405 '" '" " . " 21.4«1 28.13 '8.2\1111 .11.22.0 111.m4J .11.1322 '8.1IeII' 11.1 I �- V. o4INIII V.SO' n._ ' 5 ndev.mn, '." '." '.n .." '.5 -.'$ '.� '.� '.� '.� '.� ' .� 5.' �.' 3'.0 3•• e 3 ••3 n.' �. 21.4 21., '" �.• n._ C" moo �., 24.7 �., �., 24.3 �.• \5.0 18.4 17.2 IT.O le.4 IT . l , " � � " . " .. " " � " ". �
W' • W_ M2 R"'''5 '" '" '" " '" R"'40S '" '" " . " Oo Im N • ., ., . .'.31122 4J.00I00! 42.2515:2 ., .I11III7 "" 21.21 21.� 2'.112 2 • •'10 . 2' . 5SO!\ n . � . " 1.31 1.111 '. " '.18 .. - " '.5 '.5 ,. '.N '.M '.5' '.5 'm .. 5.' "' 47.0 ".- M.' �.' 24 .• �., 24.8 ,., ,. . �. • moo 5.' �- �.- �, � .. 5.' ' 3.5 111.3 �.. III.' '11.3 �.- " M " " M M ., .. " " " " "
- "" ...mI>y�. m1, m2 � R "",·S ,� ,� '" RK40S ,� ,� '" " " ,as . 4 .n. '25.7 '20. 0 '22.5 • .s.5 1115.0 '$01.0 1311.7 .85.3 " n " " " N � » � " " .
,,_. P_. p4, P4 '" R """ 5 ,� ,� ". R"'.... 5 ,� ,� " . a, -" 1�.7 123.' 127.3 1:N.5 '27,8 ."' '32.� '47.3 "'.1 �, W " � n M � .. M � ."
PoII.-.
TAIIL.4 - Basic statistical parameters of premolars and r11Qlars from Ihe type localities.
MEASUREMENTS AND INDICES OF METAPODIAL BONES (TAB. 5)
�. , �. �OK mel ' " " "" ,.'" " '" ,."" ,.'" " ' " " "" ,.'" e, 6, . 22 �.� � . m �. � 31 .06 30 . 21 �.� 3CI .1� '.C 7.82 '. � 7.74 4.13 '.00 '.C '.� • . •3 '.M .� ..� .. � '.� _.m '.� M.' 73.6 73.7 7'.1 5.' �.' D.' D.' '" '.- '.' '.' �r �., �.- �.' � . ' V.' 21.0 �., 27.0 '.' '.' .' .. , ' � n � � � V " � , � � "
�, , �, �,. me2 '" " "" ,.'" e, � RKOIE "'" e, '" ,. "" ,.'" e, n . m � . " 16. 1 1 n ." " . M " .� " . M ... " ' . C 7 ••0 7.2' ,� ��' -.� _.M ••82 -. � ' " '.31 1.13 '." OJ' 0.71 0.9' 0.76 " :v. 87.2 ".' �. . � . - 37.0 .' ..., 31.0 .. , '.' '.' '.' , ... , " .• �.' .• ».• '.- .• -.' .' � �, M._ � � .• , ' �, " � " " " n n ., C " • ..
scielllifiq!les se i 11°2 (2004) C(Ilrier.\· / Hors r e 54 J(lrlemenl du Rhone Museum, Di - L 'Oil
"'" , "'" """ me3 c. "'" RK8JC G' c. RKDIE """ G' c. """ RK8JC G' 75.'12 81.19 79,15 N." "." 31,16 ". .. 33,11 8,62 '." '." '." " . dew. 4.12 ,. .. -. ,." 0,91 '." '.M '." • 00 '." '." , ... ":'�' ".' � 87,1 91,4 31.7 . 37.3 '.' ' ' .. 10,3 �. .' " - " , . , " " " " �, ".' 70.0 .' '" .' .' .. ". . ,., ,., ,. . '.' " " " '" " " .. .. " " " "
- , .... �" mc. Co R KDIE " "" G' " """ "'" G' c. """ "'" G' " �" n.M75 83.41 81.92 "." "." 31.85 30.81 D . " 7.67 •.ro '. B.91 " " ... 5.81 5,01 '.n 5.17 1.47 .. '." ' . '50 1.81 0,97 1.01 . ".' ..., 91.7 ro . , "., ". "-' 37,0 '.' . , '.' 11,4 , " ".' "" ro., 74,4 ".' ".- 27,S ., ,., '.- .. , '.- ::: ., � " " " " " " " " " "
"" , "" "'" , me' c. "'" RK8JC G' c. """ "'" G' c. "'" "'" G' 79.3152 83.97 82,47 ".00 " . " 12.20 13.14 12,55 13,07 r-" ".D D." .. ... " '.'" -." '.00 ,. . " ,... 2.18 I.Bl '." ,. .. '.n . " 1-'" .. ro.• . ., ., .. 18.5 17.9 15.0 18.3 " ro .• ro , " " " "J 71,9 m ". . 31,3 "., '" "., ., '.- 10.0 n' • •., � " " " " " " " " " " " " � . , �. "'' ' mt1 c. "'" """ G' c. RKDlE RK8JC G' c. " '" RKBIC G'
SO.16 "." 53.91 "." D.n 31,70 31.08 D . " •. ro 10.32 .. � 11,23 �" " " " " . dev. . . . 2,11 ' . '." ' . '.n ' ' , , "" >." '.00 " ." 81.1 ". 81.1 . 35,1064 36.7925 36.6279 11.6897 12'168; �3.0732 14.l805 , .,.. " .... " .... 47.4 47.1 30.6266 26.152 211.9151 ".- 7.1271 7,87879 7,4351 9.02409 ::: .. " .. '" .. " " " " 1 " " "
�, , �, """ ml2 Co '''''' """ G' c. "'" "'" G' c. """ " "" G' " ". .. 68.27 ... " 67.31 31.16 ". .. "." 31,82 5.10 5.41 '. ,. ro 3,81 ." 1.76 '." '. 1.51 ,.ro '.n 0.81 . .dev. "" "" - " , .. 74.1 74,8 ". 37.6 .' .. ,. ":'�' ro.• ' D.' D.- " , ••• , ••• 50.' 50' 57.3 50.' '" ".- -.' ' ' '.' -.' "J n., . ;:: � " � " " " " '" " .. .. "
ml3 . . �; . �:�; �� �,� . ::�. : i: ,:; �:d I :: :: j; � " � �::'" " �j" " .. " � ,. .:
mt.
. 1.41 1,� ��.. �r I �I � ' 31.11 31,8 ' ' �' 2 7 3 37 � �j " F ;; ;; ,; .:;' .; � rmr;, mI' ' -::�� " ; ,';' I!:.�I 0.71 .. '1 is.S I ;'Sil5.3; I ';96.2 'i;':XI.5 28.0 29.8 31.4 11,4 10,3 11,8 14,0�: �� �.� �:� 7 9 24.7 2 0 2 8 .' '!!:; r:jl �8 �o :; � �9 ' �: � �: �.: _ Measurements and indices of mctapo:xiial bones from the typelocalities. TABL. 5
Cahiers scielllijiques Hors serie nO] (2004) 55 / Celllre de COII.�er\!(/lioll el d'£wde ties CQlleCliolls
RADIOMETRJC DATA "" (FIG. 2) • :; 130 ComUriIles radiometric datatioo5 •
- r---'======-----, .,ro.£ " ,,,., ...... _...... 0 • 110 • -- � > F'
. Fig. 2 Radiometric dllta of cave bear bon�s from the three type localities and from Potocka'tijalka, "" COMPARISONS
Only elements that are available in an ade quate amount from all six localities were compared -- U,IP __ ' for teeth and metapodial bones. The conservation of "'_ U�I(J._o 6. the long bones and the skull is especially inadequate 40 in the Ramesch Knochenhohle. however, the number l of single teeth and small bones is so large that it was possible to compare the elements from different stra ••l:-c ·, .,.-,·"',.-, ·:-"' ·, :-"' ,·�-·, :-"' ·, :-·,, '"C , -',,, tigmphic levels. These four respective five units are, r.t1o: IIbl. IengWmf length ,.land., from the top to bottom: RK I (0-50 cm under surface).
2 (50-100 em) RK 3 (100-150 cm) RK 4 (150- Fig. - RK 5" TIbia and M I info length. 200 cm) RK 5 (200-250 cm); because the units RK 4 and RK 5 yielded only small numbers of bones for mOSI of the elements, they are summarised to the unit COMPARJSON OF THE MOLAR MEA RK 4+5 (150-250 cm). SUREMENTS (TAB. 4)
For the comparison with Ihe type fauna, '"' values of the cave bear fauna from the "Guloloch" from Zoolithenhohle were used. II has been known for I,., , a long lime that two time levels have 10 be distinguis g& �! J1rc, . =�.":""'c " c ",. c.- 'I, hed in the Zoolithenhohle (RABEDER 1989). The com plex "cleavage+Core" contained considerably more N" 10 � l:J. � '0 ·IZd��1 primitive cave bears than the units "Guloloch" and �!.• ,0 0, "Siirenkammer". We can assume that the type skull I�·�I o on which the species UrSI/S spe/aeus sensu I � • (1897) was established is from a strati �>---�--�----�------,, ., "' ROSENMOLLER � " " ' graphic younger area of the cave that is today known m11ength as "old part" and is therefore assigned to the Middle
n Wiirmian. The reason for this assumption lies in the Flg,3 - Propo ions of mondibular molar.!.
Cahiers scientifiqlle�' Hor.f serie I 11°2 (2004) 56 du RlI6ne Lyon Dtfpartemem . MWiium.
••
.. ' � '" • . .' I: & ... ' . . . . ' • I::.:--....·...... :.,··T J • I • • ...... / • • ...... M: "·Mac MM • �M • � � .. �:;: .. � . - - - - •• � � - �
. Fig Ii - Diagrom showing the differences betweco the standardized (GS::I()()%) on-al·erag.: si:tC-rel�lions (gL) within metacarpus (left) alld metlltarsus (rigllt) from the bear populations from Rarnescll-KnocllenMllle (RK DIE and SIC). Gamssulzenllt\lllc (GS) lind from Conturines cave (Cu). Please nOte tllm tile bears from tile deeper units (DIE) of tile Rmnescll-profile tend to be bigger than those from the upper units (B/C) anti the t�tremt shonness of tile bears from Conturines cave.
� •• •• i'- •• i'... • • � ..J , -' --- .. . .' ./I'\. • """,V • /' \'.... V . ....\ . • • ' . "·MM: ...... � '. • �M • �M � � � � .. .. �. - - - - • • � � - �
7 "'ig. - Diq:ram showing the differences between the standardized (GS::I()()%) on-'"'Crlseplumpness-relations (ip) within melaCarpus (kll) DIE and and mcl8lJU'SuS (right) from the bear populations from Ramesch-KnochenhOhle (RK and BIC). Ganw;ulunhOhle (GS) from Conturines cave (Cu). Please note that the frombean; the R:uncsch-profile are generally less plump !hanfrom those Garru;sulzenhOhlc. espe cially those from the upper unit!; (BIC). Please note also the: extreme plumpness of the: first metacarpal and melalanal bortes from the Conturines bears.
•• •• " " • • •• •• � � I • I • • • �� . �� . .. • ��-. ��CIIII • .. i .. .. • f � �
• • .. • • •• � - •• • • • • .M ....,l _ __ mil _ . of '*"'-' __ .of�_ -
and Fig. 8· Dia&ru" showing the: distribution of the avemged andstandanlized (GS",IOO%) inde� of plumpness of the: first metacarpal (left) bone �traord ary bear metatarsal (right) from fivedifferent sites.Please note the: e in plompness of theContorines . which even sopersedes those bl'ar see of the bigge5t and most developed members of the: ca\'c group. FQr abbreviations fig. 6.
•• '. � " • ' M ,. - - I • ! • • • . � � �K • • .�� MM' .. ..
• • I � I �
• • • • • • • •• •• •• " " • • •• •• ••
....,1 _ g.... I_length. _a,clized mil _ ___ .•1af>Cjardi_ 11eng!h
Fig. 9 - Diagram showing the distribution of the averaged and st.andardizcd (GS",I()()%)greatest length of thefirst metacarpal (left) and met.a· tarsal (right) bone from five different sites. Please note Ihe tlltraoroinary shortness of especially the first metatarsal bone of Ihe Conturines bear. which ;s even sooner than those of U. deningeroides. Forabbreviation.� sec fig. 6.
Cuhiers scit!nlijiqut!S Hors sirie n02 (2004) 57 I Centre dt! COl/sen'lItion et d'Etl/de des Col/ectiOiIS
topology of this cave. Comparison of the data from the clear, but it seems very likely that it is an adlptation different faunas leads to the following conclusions: for the high altitude and the continuous strolling the bears of the Conturines cave are the smallest in all around 00 more or less steep slopes. measurements whereas in the profile. of the Ramesch The aforementioned differences arc not the only ones cave the amplitude of the molars diminishes or increa but ofcourse the most Striking and therefore best usa ses at first but then diminishes. The diminishing ble indicators for diagnostic use. The fo llowing dia means of the length of the m2 and M2 are particular grams will give an overview on the differences within Iy obvious. metacarpus and metatarsus, see figs. 6 and 7. The values of the Gamssulzen cave exceed those of The trends within these members of the cave-bear the other alpine cavc.<> and also the measurements of group are quite different and the peculiarities of the the Zoolith cave with the exception of m2- and M2- bears from Conturines cave arc the most eye-catching length. as they show another pattern especially for the index of plumpness. The fo llowing scalter-plot diagrams, sec fig. 8 and 9, COMPARISON OF THE MORPHODY illustrate the differences between these quite different NAMIC AND METRIC INDICES OF bears with a spccial l'ocus on to greatest length and the METAPODIAL BONES (SEE FIGS. 6·9) index of plumpness of the first metacarp;11 and meta tarsal bones as these seem to be the most significant The evolution of the metapodial bones pro differences between the bear populations. voked some striking differences, which can be seen on the following distribution diagrams dealing with the index of plumpness and the greatest length. see I1gs. 8 It is obvious that there are many differences between and 9. On the y-axis the standardized P4/4-index is the proposed new taxa that are partly overlapping in plolted against the index of plumpness and the grea time so that there are sufficient arguments to plea for test length respectively. There is a typical succession this step of splitting a species that has remained beginning in the lower left comer where Ur�·lj.s deltin unchanged for more than 200 years. In combination gemides from the Repolust cave in SlY ria has its with the analysis of lOt-DNA, which supports up the place. II is an old, small and thus more primitive bear herein presented results from a totally different elo.�e to U. (/ellingeri. In the right upper quadrant, on methodological point of view, it seems to be reasona top. there are always the most developed bears, from ble to ereCI the fo llowing new taxa belonging to the the "standard" population of Gamssulzen cave and genus Urslls L. from POlocka zijalka in Slovenia (representing U. ingresstls n.sp. see below), which are even more developed and usually plumper and larger than those SYSTEMATIC CHARACTERISATION: in het midrange of the diagram. The range of dilTcren ce between the smallest and biggest bears lies within Genus Ursus L. 8-12%. In the midrange there are the bears from Ramcsch Knochcnhtihle and those from Conturines Ursus spelaem·? ludjll;ctls nov. ssp. cave in Italy. with their P4/4-indices around 50% but with big differences concerning the index of plump 1991 Urslt.\· spelaelu. RAHEDER. G., Die ness and the greatest length as well. H6hlenbiiren der Conturincs .. The most eye-catching fcalUre of the distributions of index of plumpness is t.he position of the "Conturioes Der-ivalio nominis: From the ladinic part of the bear". which has. despite its small dimensions. the by Dolomites (Italy) where the locus typicus is situ:Lted. far plumpest first metacarpal and metatarsaJ bones. In Holotype: Complete skuJl with nearly aU teeth, addition to this the distributions of the greatesl length inv.no.: Cu 714. show something unexpected: whereas the first meta Paratypcs: 2 skulls (Cu 703 and 704) and 3 lower tarsal bone proves the expectation to be the shortest in jaws (Cu 715. 720 and 12, see tab. 2). the fieldthe first metacarpal bone is nOi the shortest at Differentia) diagnosis: Considerably smaller than the all. It comes close to the length of the "Ramesch bear" geologically cQCval cave bears from the middle moun thus showing that its plumpness is not a result of a tains and the lowland, the P 3 sup. exists in atxJut shortening of the bone but of broadening. The reason 30 % of the skulls. the metapodials especially of the for this adaptation of the "Conturines bear" is not yet first fingers and toes (metacarpaJc I, melatarsale 1)
Cllhius serie scil'lItijiqlles I Hors ,,"2 (2004) 58 DiparWlllflll du Rhone - Mu.�eum. Lyon
[Ire considembly cruder than with the typical cave Differential diagnosis: considerably smaller than the bear. Phylogenetic position according to the analysis geologic coeval cave benrs of the G stUlds over the. noor. DNA, the Rameseh cave bear is closest \0 the Storage: Naturmuseum Bozen, Italy Conturines bear with the classical cave bear from Systematic position: The combination of primitive Zoolithcn cave falJingbasal to these two. (small dimensions, P3 sup.) and progressive attributes Locus typicus: Ramesch bone cave in the Totes (P4/4-indcx. cnthypoconid indices, plumpness of the Gcbirge, Warscheneck group. community Spital am metacarpalc I and metatarsale I) lead to an indepen Pyhrn, UpperAustria. dent development of the Conturines bearthat is confir Basis data: Entrance 1960 IT! above sea level. 14°17' med now by mcans of genetic analyses (HOFREtTER et E_ 47°39' N, aL 2002). Dating of lineage separation via genetic Stratum typicum: typical cave bear loam riddled data always results in large confidence intervals, with many small Stones. however, it is likely that this lineage separated earlier Storage; collection of Deparlment for palaeontology. than 300.000 yenrs from its closest relatives, see fig. University of Vienna 5. TIle morphological differences to the cave bear of Systematic position: The singularity of the Ramcsch the type loc:llity Zoolithen cave (UrSlis spe/atllls sensu bear is given wilh lhe decrease of the dimensions and StrictO) on the one hand and to the other alpine cave the morphologic level of evolution from top to bottom bears on the other leads to the erection of a new sub (from Ramesch 3 to Ramesch 2 respective I) in the species UrslIs spl'lllells /alJi"icus nov. ssp. middle Wlirmian profile of lhe cave. The morphologic Recognition lit the species level is possible, but mnnot and metric distances as well as the distances in the be finnly established based on the genetic distance mitochondrial DNA (see HOFREITER et al., 2002) com alone and in the nbsence of positive evidence for pllred to the Conturines and Zoolithen cave bears are reproductive isolation as �hown below for U. ifl8res consistent with subspecific re cognition. The fact that SIIS. The assignment to U,..ws .I-pe/aells results in the both morphological and mitochondrial DNA data composiLion of all morphological and metric dat:! strongly indicate long-ternl reproductive isolation bet from teeth compiled in the cluster analysis. By inclu ween the Ramesch and Garnssulzen cave bears shows sion of the metapodial data the assignment to thm a separation of these two cave bears as indepen U. dellillgeri would also be possible (see fig. 10), dent species dcst�ribes the situation best. Other sites: Brieglersberg cave in Totes Gebirge, Styria. Other sites: Salzofenhohle and Ochsenhalthohle (Totes Gebirge) UrSIiS speiaells erelllllS nov, ssp. Urs/ls ;lIgres.flIs nil\'. sp. 1986 Ursus spe/llells. - Hi lle, P. & Rabeder, G., Die Ramesch-Knothcnhohlc ... 1995 Ur.ws sp ell/ellS. - RABEDER, G.. Die Gamssulzenhohle... Oerivatio nominis: From the site "Ramesch bone cave". The name "Ramesch" is derived from the latin Derivatio nominjs: From the latin word ·'ingrcssus", word "eremus" , meaning detached. alone (compare to to wander into, with regard to the immigration of this hermjt). species in the Northern C:\lcareous Alps, probably at Holotype: Strong fragmented skull inv.no.: RK 84. about 60.000 years b.p. Paratypes: 3 fragmentary skulls (RK 1001, 1002. Hololype: We ll preserved skull with both mandibles, 10(4) and 6 lower jaws (RK 38, 119, 224, 440, 452 in.".no.: GS718. I). and 594, see tab. Paratypes: I skull (GS 524) and 4 lower jaws (GS Differential diagnosis: Preserved arc canine sin. and 150-2. 457, 512-2 and 513, see tab. 3). both tooth rows P4-M2. In the lcft maxilJare there is Din'crenlial diagnosis: Considerably bigger than the an alveolus of P3, missing on the right side. geologic coeval cave bears of the Rameseh bonc cave, 59 Celltrt! de COlIl'tn'a/ioll el d'£/Ude del' ColleCliulIJ" the P 3 sup., denture highly dcveloped. thc mctapodial lineages already about 60f).0CI0 years ago (HOFREITER bones especially of the first fingers and toes (metacar et al.. 2002). This shows that sufficient time for spc· palc I, metatarsale I) are less crude Ihan in the cies separation was available. Ramesch and the Conturines cave bear, The, level of Other sites: Potocka zijalka, Mokrinkajama and Divje evolution of the denture especially the premolars is babe (Slovenia), Vindija (Croatia). Kugelstcin· much higher than the levels of othcr cave bears, Tropfstcinhohle. DrachenhOhle of Mixnitz (paleoli Phylogenetic position according to thc analyses or thic hunting statioo). Nixloch near Losenstcin mitochondrial DNA see fig. 5 Ternbcrg and Licgloch (Austria), Schnurcnloch Locus typicus: Gamssulzen cave in the TOles (Switzerland). GciBcnklostcrle and Hohler Fels in the Gebirge. Warschenecli. Group. community: Spital am Achtal (Gennany). Pyhrn, Upper Austria. Basis data: Entrance 1300 m above sea level, 14"17' E. 47°41' N, Stratum typicum: loam of excavation area 3. Storage: Depanment of PaiaeunlOlogy. Univcrsity of DNA-ANALYSES Vi enna Systematic position: The bears of the Gamssulzen Cave bears arc probably the Pleistocene spe cave developed attributes (especially P4/4-index) in cies from which so far the most samples have been the dentition considerably more evolved than the cave analysed (HOFREITER et al. 2002, ORLANDO et al. 2002. bear of the type locality Zoolith cave (Ursus spelael/s HOFREJTER el al. in press). Unfortunately, the taxono senSu stricto}.These differences would only allow a mic sl - multiple evidence, such as genetic distance plus morphological data and I or proor of reproductive iso lation and it should be kept in mind that taxonomic Fig.10 · upper left diagram: cluster analysis of teeth only: upper right di agram: clu.ter anal�sis of teeth and rnctapodialbones; lower assignments are a working frame li able to later revi lefl diagl1lm: cluster analysis of teeth and Illctapodiul bones illc]u, ding Repolust cave (Slyria. Austria); lower right diagram: cluster sion. It should also be kept in mind that the topologics analysis of teeth and metapodial bones with laJlonomic names. oblaincd for phylogenetic trees of cave bear mtDNA Cahiers scielllijiques Hors serie ,,°2 (2004J / 60 Departemelll du Rhone - Museum. Lyon are currently based on short sequences and lhus. in plausible explanation is reproductive isolation bet most places only weakJy supported (HOFREmlR et al. ween the Ramesch and Gamssulzen cave bear popula 2002). Work is in progress to obwin considerably lon tions. This conolusion is funher supported by the fact ger mtDNA Sequences to establish a more re1illble that cave bears that come from a number of additional phylogeny, but a number of, at least preliminary caves from the Austrian Alps that lie above 1.600 III conclusions can nevertheless be drawn. First. the dif altitude lind thaI have been shown to be morphologi ferent mtDNA lineages separated comparatively long cally close to the Ramesch bear (RABEDER, 2001) also ago, probably about 600.000. but almost definitely yielded mtONA sequences close or identical to those more Ihan 300.000 years ago. Thus, even the more from Ramesch cave. Similarly, cave bears from lower closely related groups of cave bears would have had. altitude caves from the same geographical region, thai in the abscnce of gene flow between populations, suf were found to be morphologically close to the ficient time for population specific adllptations to Gamssulzen cave bears (RAlIEDf.R, 200 I) arc also develop. Second, independent of the analysis, the genetically close or identical to the Gamssulzcn cave. Gamssulzen cave bears are very distant from all bears. Such a result is highly unlikely to be.produced others, and often. as shown in figure 5, the deepest by chance and strongly indicates both ecological dif divergence falls between the Gamssulzen (Le. ferentiation and reproductive separatlon of the two U. ingl'esslIs) bears and all other cave bears, currently types of cave bears. and thus well justifies the reco treated as a single species, U. spelaells, wilh three sub gnition of these fonns as two independent species. species. Interestingly, the Zoolithen, the Ramesch and the Conturines cave bears each fall in a different one of three, albeit weakly separated mtDNA sequence groups (fig. II). Thus, the separation of morphologi cal groups is consistent, and not contradictory, with } .. -- Ursus spellJflUS the grouping of mtONA sequence groups. However, -- twO points need to be noted. First, the three mtDNA sequence groups within spelaells each contain a U. }.--� number of populations and extensive morphological ___ _'} L o-..... studies will be necessary to investiglllc whether all n.• I':: U'S US Ingfflssus p. populations within a group are for all comparisons o.or...... l closer to each Olher than to populations from a diffe l'ig. II - CllIdogrnm showing Ihc relatiollship of cn�e bear mito rent mtDNA sequence group. Second, mtDNA is inhe chondrial DNA sequences. rited strictly maternally. A tree based on mtDNA sequence data therefore reflects only the female histo ry and male mediated gene flow due to migmlion of PHYLOGENETIC CONCLUSIONS male cave bears could show a different picture. As male mediated gene. flow .� hould homogenize the The metric and morphologic differences bet morphology between populations, consistency bet ween thc three alpine cave bear faunas and the type ween phylogenies based on mlDNA and morphologi fauna from the Zoolith cave are consistent with the cal data. respectively. is a slrong argument that the results from DNA- analyses. Thus, in Ill!! Middle oblained topology is correct. The genetic evidence Wtinnian. at least in the time span between ca. 50.000 thai supports the separation of U. ingresslIs as a diffe and 30.000, at least three genetic lineages of cave bear inhabited the eastern Alps that scpar'dted in the early rent species from U. spelaell.f is. however, based nei ther on the topology of thc mlDNA tree. nor on the Middle Pleistocene. At least two of these represent age of the separation of different genetic lineages. autonomous species in the sense of the biological spe Rather, it is derived from the facl that the genetic data cies concept, leading to the separation of U. illgressus from Gamssulzen cave and Ramcsch Knochenhohle from all other cave bears which are maintained as sub fai l to show positive evidence for gene flow between species within U. spel(lell�'. the two caves. dl!Spite 15,000 years of contempora neous occupation (HOFREt'l'ER et aI., in press). The most likely explanation for this result is that barriers to gene flow existed between the two populations. As no geographical bamers exist and the two caves are less Iban 10 km distant fr om each other, the only 61 Callien sciemijiques / Hors sirie n� (2004) Cellfrt! de COIrsen'ul;mr el d·t:,ude des Col/eelimlS (1973) ACKNOWLEDGEMENTS KURTEN B. - Trnnsberingian relationships of Ur.nJS arctos Line (brown and grizzly bears). The excavations in many alpine caves were - Commell/atiull Biology, 65: I-I O. financially supported by lhe Austrian Academy of LOREILLE R ANDO PATOU-MATUIS M .. Science and the University of Vienna. 0., O L L., PHILIPPE M., & For the reviewing of the article we say many thanks to TAAERLET P. HANNI. C. (200 • Prof. Dr. David K. Ferguson (lnstilUle of I) Ancient DNA analysis reveals diver URSUS SPElAEUS, Palaeontology, University of Vienna) and to gence of the cave bear, and URSUS AII.CTOS. Jacqueline and Alain Argant for the correction of the brown bear, lineages. - Cllrr. Bioi. , 11. 200-3. French abstract. MCCUNE & LoVEJOY (1998) A.R. N.R. . I.n Endless Forms. Species and Specialion. (Eds. Howard, OJ. and Beriocher, S.H.) - OxfimJ ABEl. O. & KYRLE G. (1934) - Die Drachenhohle von University Press. Oxford, New York. Mixnilz. - Speliiolog. Monogr. 7-8, WIEN. ORLANOO L., D. BONJEAN, H. BOCHERENS, A. THENOT. R M. & NI (2002) DOPPES D. & RABEDER G. (1997) - Pliozane und A. A (;ANT, OrrE, c. HAN - Pieisioziine Faunen OSlerreichs. - Milt. Ancient DNA and the population genelics of Ko mm. QIIlIr1iirjorsch. asterr. Akad. IVi ss .. cave bears (Urslls spelaells) through space Mol. 10: 1-41 1. Verlag Osterr. Akad. Wien. and time. Bioi. £1'01., 19(11),'92G-1933. EURENBERG K. (1929) - Die Ergebnisse der RABEDER (1983) • Ausgrabungen in der Schreiberwandhohle G. Neues vom Hohlenbaren: zur 11.3:261-268. Morphogenelik der Bac-ken-zahne. am Dachstein. - Palaeon!. Z 3412: 67-85. Berlin. Die Hohle. Wien, RABEDER (1991) FROEMEL M. (2001) Die Evolution der G. - Die Hohlenbaren von Hohlenbiirenlibia aus sieben alpinen Hohlen. Conturines. Entdeckung und Er-forschung 2800 - Dipl. -Arbci/, pp. 1-80, Fak. Mmh. ciner Oolomitell-Hohle in m Hohe. - 125 Na/i/lwiss., Univ. Wien. Allresio-Ver/ag, Bozcn. p., RAaEDER (1995) • HIl.LE P. & RABEDER G. (eds.) (1986) - Die Ramesch G. (cd.) Die Gamssulzenhohle im Knochenhohle im Totcn Ge-birge. - Mil/. TO len Gcbirge. (gennan incl. English sum mary) Mill. KO lllm. QI/artiirjorsch. Osrer Ko",m. Qmll'tiirforsch. Osterr. Alwd. Wiss.• - r. Akad. 9:1·133,. 6:1-66. Wicn. Wiu .. Wien HOFREITER M .. KRINGS M., WAITS Rabeder G. (1999) Die Evolution des .. CAPELLI C L, NARD . MONZEL . RABEOER Hohlenbarengebisses. (gennan incl. English CO N. S . G .. NAGEl. A NOVIC M .. JAMBRESIC MEYER . summary) Mill. KO lllm. Qllonarforsch. D., P U G., S . - 02,. WEISS & . (2002) Os/err. Akad. Wil·S., Wien II:I-I G. . PAA8Q S . - Ancient DNA analyses reveal high mitochondrial RAAEDER. (2001) · DNA sequence diversity and paralic I G. Geschlechtsdimorpttismus und morphological evolution of laic Pleistocene Korpergro6e bci hochalpinen cave bears. Mol. BioI. £1101., Hohlenbarenfaunen. (Gennan, iocl. English 19(8): 1244-1250. summary)- Beitr. It/Iliom., Wien, 26: 1 17-132. RIDLEY M. (1996) • HOFHEITER M . JAENICKE . WITHALM Evolulion. 2o.l cd .. Blad.:weff . RAAEDER G .. V. 719 PAUNOVIC M .• B I Science, Oxford. p. Go, NAGEl. D .. JAM RES C G., and Pii.ii.ao S., (2003) - Evidence for repro SCHLIJJ'ER (2000) • ductive isolation belween cave bear popula D., The Ecology of Adaptive Cllrr. . R�djation, Oxfo rd Ulliversity Press, New tions. - Biology accepted. York. Cuhicrs licielllifiques I Hors s�rie ,,°2 (2004) 62 Deparlemelll du RhOne - Museum, Lyon SHIELDS G. E, ADAMS D., GARNER G., LABELLE M., TALBOT S. L. & SHIELDS G. E (1996) Phylogeography Pnorscu 1.. RaMSAY M., SCUWARTl c., TITUS of brown bears (Ursus arctos) of Alaska and K. & WILLIAMSON S., (2000) paraphyly within the Ursidae. - Mol. Phylogeography of mitochondrial DNA Phylogellet. £1'01. ,5:477-94. variation in brown bears and polar bears. Mol. Phylogellel. £1'01., 15:319-26. WtTHALM G. (200 t) - Die Evolution der Metapodien in der Hohlenbaren-Gruppe (Ursidae, SPAHN! J-c. (1954) - Les gisements 11 Ursus speJaeus Mammalia). - Be;lr. Pa laolll. , Wien, de I' Autriche et leurs problemes. - Bull. 26:169-249. Soc. Frallc. Le Mans, LJ,7;346-367. Ca hiers 63 scienlijiques I Hors serie n02 (2004) Celllre ae COIlServatiol1 et a'Etude des Collections Plates 1-2 The holotypes (skulls) of new alpine cave bear taxa Fig. I Ursus ingressus nov. sp. from Gamssulzen cave Fig. 2. Ursus spelaeus? eremus nov. subsp. from Ramesch cave Fig. 3. Ursus spelaeus? ladinicus nov. subsp. from Conturines cave a lateral, b dorsal, c ventral view, d tooth rows, occlusal view Scale: fig. la-c, 2a-c. 3a-c: 25% of natural size. fi g. Id, 2d, 3d: 55% of natural size C6hiers Hors serie sciemijiquesI ,,°2(2004) 64 Dip(Jf1t!�1l1du RMIlt! - Muslum, Lyoll PLATE 1 65 Cohius scit!lJlijiqut!s I HOTS slrit! n02 (2004) Cellfre de COI!.�en·atiOIl el d'Elllde des Col/celiO/IS Plates 1-2 The holotypes (skulls) of new alpine cave bear laxa Fig. I UrsIIs ingresstls nov. sp. from Gamssulzen cave Fig. 2. Urstls speiaells? eremus nov. subsp. from Ramesch cave Fig. 3. Ursus spelaells? ladiniClls nov. subsp. from Conlurines cave a lateral, b dorsal, c ventral view, d tooth rows, occlusal view Scale: fig. 1 a�c, 2a-c, 3a-c: 25% of nalural size, fig. I d, 2d, 3d: 55% of natural size Cahiers sdelllijiqlles I Horl' serie /1"2 (2004) 66 Dlpllrtt'lI1t'm du Rh6n� - Mlu'illln, Lyon PLATE 2 67 Cohi�rs Scit'llIijiqUt'S / Hon sirie 11°2 (2004)