The Systematic Position of the Cave Bear from

Mitt. Komm. Quartärforsch. Österr. Akad. Wiss., 13:197–200, Wien 2004
The Systematic Position of the Cave Bear from
Potočka zijalka (Slovenia)
Gernot RABEDER1), Michael HOFREITER2) & Gerhard WITHALM1)
RABEDER, G., HOFREITER, M. & WITHALM, G., 2004. The Systematic Position of the Cave Bear from Potočka zijalka
(Slovenia). — Mitt. Komm. Quartärforsch. Österr. Akad. Wiss., 13:197–200, Wien.
Summary
All metrical and morphological data derived from the
preserved bone and teeth material have been used for a
cluster-analysis. In comparison with other cave bear faunas
it is evident, that the bears from Potočka zijalka are closest
related to the bears from Gamssulzenhöhle (U. ingressus
RABEDER et al., 2004), which corresponds to the results of
the genetic analyses.
Keywords: Potočka zijalka, cave bear, Ursus ingressus,
Karawanken mountains, Upper Pleistocene
Zusammenfassung
Alle metrischen und morphologischen Daten, die aus
dem überlieferten Knochen- und Zahnmaterial gewonnen
wurden, sind für eine Cluster-Analyse verwendet worden.
Im Vergleich mit anderen Höhlenbärenfaunen stellt sich
eindeutig heraus, dass die Bären der Potočka-Höhle dem
Gamssulzenbären (U. ingressus RABEDER et al., 2004) am
nächsten stehen, was mit den genetischen Untersuchungen
übereinstimmt.
Schlüsselwörter: Potočka zijalka, Höhlenbär, Ursus ingressus, Karawanken, Jungpleistozän
Izvleček
Vse merske in morfološke podatke zob in kosti smo
uporabili za klastersko analizo. Po primerjavi enakih izsledkov raziskav jamskih medvedov iz drugih najdišč smo
1)
UNIV.-PROF. DR. GERNOT RABEDER & DR. GERHARD
WITHALM, both: Institute of Palaeontology, University of Vienna, Althanstr. 14, A-1090 Vienna, Austria.
e-mail: [email protected], gerhard.withalm
@univie.ac.at
2)
MICHAEL HOFREITER, Max-Planck-Institute for Evolutionary Anthropology, Deutscher Platz 6, D-04103 Leipzig,
Germany. e-mail: [email protected]
ugotovili, da so medvedje iz Potočke zijalke ﬁlogenetsko
zelo blizu “Gamssulzen medvedom” (Ursus ingressus
RABEDER et al., 2004). Te ugotovitve se ujemajo z genetskimi raziskavami.
Ključne besede: Potočka zijalka, jamski medved, Ursus
ingressus, Karavanke, mlajši pleistocen
1. Introduction
The afﬁliation of the bone and tooth material from Potočka
zijalka to the cave bear group is assured on the basis of the
morphology of the molars, premolars and incisors as well
as its big dimensions. Only 10 years ago this conclusion
would have been satisfying. Differences in metrical and
morphological data would have been regarded as the result
of a different level of evolution within a phylogenetic lineage, the “Ursus spelaeus-lineage”. At the most, there would
have been made an intraspeciﬁc differentiation, traced
back to ecological circumstances, like it is expressed, for
example, by the term “high-alpine small form”, which has
been brought in by K. EHRENBERG (1929) for small sized
cave bears of the high-alpine karst plateau.
Since the appearance of the monograph about the Gamssulzenhöhle (RABEDER, 1991) this is seen differently.
Comparative studies showed, that in two caves of the
Totes Gebirge, the Ramesch-Knochenhöhle and the Gamssulzenhöhle, which lie very close to each other, two cave
bear groups, different in metrics and morphology, had
been living at the same time (at least between 48.000 and
30.000 years BP) and had not intermingled (RABEDER,
1991:80-81…that leads to the hypothesis, that the bear
populations of both caves lived at the same time in the same
mountains, but without interbreeding, therefore they were
separated in breed (hypothesis of speciation): Following
this hypothesis the cave bear clade would have been split
up into two lineages, into the “high-alpine form” and into
the “normal form”).
Studies of other Alpine bears showed, that at least a third
cave bear group lived in the Alps at the same time in the
Conturineshöhle in the Dolomites.
A provisional family tree was drawn up in the year 2000
and presented at the cave bear symposium in A Coruña
(RABEDER & NAGEL, 2001). A possibility to test this hypo-
198
Mitt. Komm. Quartärforsch. Österr. Akad. Wiss., 13, Wien, 2004
Dendrogram 1: Cluster analysis of teeth only.
Dendrogram 2: Cluster analysis of teeth and metapodial bones.
U. deningeri
U. spelaeus
Dendrogram 3: Cluster analysis of teeth and metapodial bones
including Repolusthöhle.
Dendrogram 4: Cluster analysis of teeth and metapodial bones
with taxonomic names.
Figure 1: Cluster diagrams (Dendrogram 1-4) of tooth and metapodial bone data of ﬁve Alpine cave bear faunas completed by the
deningeri fauna from Hunas (Franconia).
thetic family tree arose 10 years after the termination of
excavations in the Gamssulzenhöhle as it turned out, that
bones of most cave bears contain “ancient DNA”. First
studies showed, that the cave bears from Ramesch-Knochenhöhle are actually genetically so far apart from the
type of the Gamssulzenhöhle, that they have to be assigned
to different species (HOFREITER et al., 2004; RABEDER et al.,
2004). A phylogenetic autonomy was not only afﬁrmed but
also reinforced through the DNA studies.
Taking the present standard of knowledge, it can be assumed, that the line of the Gamssulzen bear (Ursus ingressus RABEDER et al., 2004) already split from the cave bear
main line approximately 600.000 years ago, at a time when
all representatives of this group still were on the evolutionary level of Ursus deningeri. The new, now genetically conﬁrmed family tree, presented at the cave bear symposium
in Kirchheim / Teck 2002 (RABEDER et al., 2002), does not
correspond with the hypothesis of 2000 in one essential
point: the bear from Gamssulzenhöhle (Ursus ingressus
RABEDER et al., 2004) stands more far apart from the bear
from Ramesch-Knochenhöhle (Ursus eremus RABEDER et
al., 2004) than from the holotype Ursus spelaeus from the
Zoolithenhöhle near Gaillenreuth.
The holotype became also known from Alpine caves of the
Préalpes (Balme à Colomb). Therefore it can be assumed
at present, that at least four forms of cave bears inhabited
RABEDER, G. et al., The Systematic Position of the Cave Bear …
199
Figure 2: Genetic “family tree”: The cladogram shows the relationships of mtDNA sequences of cave bears from Potočka zijalka
and other Central European cave bears.
different, and partly also the same mountain ranges in
Europe at the same time, namely during the late Middle
Würmian (ca. 50.000 to 30.000 BP).
2. Methods
For systematic classiﬁcation of the Potočka bears all
available morphological data were compared to the
corresponding values from other cave bear faunas, in
particular to values from the faunas of the locus typicus
of the new species (Gamssulzenhöhle, Conturineshöhle,
Ramesch-Knochenhöhle, see RABEDER et al., 2004) on
one hand, and to values from two faunas of the Middle
Pleistocene (Hunas and Repolusthöhle) on the other hand.
Clusteranalysis allows determining the morphological
nearest cave bear form by using the principle of the
smallest distance. The result is compared to the results
of the DNA analysis.
3. Clusteranalysis
A clusteranalysis, based on a matrix of data with 21 variables - length and width of all molars, morphodynamic
indices of p4, P4, m1, m2, and M2, also an index of length
and plumpness of all metapodial bones (see RABEDER, this
volume and WITHALM, this volume) of ﬁve Alpine cave
bear assemblages and of the Deninger bear from Hunas,
was drawn up and resulted in the following dendrogram
(see ﬁg. 1): the bear population from Potočka zijalka is
closely related to the cave bears from Gamssulzenhöhle
and is consequently a dependant of the form with the high-
est evolutionary level, namely Ursus ingressus. The bear
from Potočka is therefore not closer related to the other
high-alpine cave bears from the Dolomites (U. spelaeus
ladinicus RABEDER et al., 2004) and from the Totes Gebirge
(U. spelaeus? eremus RABEDER et al., 2004).
4. DNA analysis
Clearly identiﬁable tooth and bone remains from the excavation campaigns 1997 - 2000 but also fragments from
artiﬁcially manufactured bone points (see HOFREITER &
PACHER, this volume) were used in order to determine
mtDNA-sequences. All cave bear samples from Potočka
zijalka produced the same, uniform results (see phylogram,
ﬁg. 2): the Potočka samples correspond in their mtDNA
sequences to the samples from Nixloch near LosensteinTernberg in Upper Austria and Lieglloch in Styria (see
DÖPPES & RABEDER, 1997), as well as to samples from
Gamssulzenhöhle (see RABEDER, 1995) and Vindija-cave
in northern Croatia. Thus it clearly showed itself that the
cave bears from Potočka zijalka are genetically closely
related to these faunas.
5. Conclusion
Due to corresponding evidences of the dendrograms and
the clusteranalysis obtained on morphological and metrical
data as well as the phylogram based on mtDNA-sequences
the cave bears from Potočka zijalka are assigned to the
“Gamssulzen bear”, for which recently the nova species
Ursus ingressus RABEDER et al., 2004 has been established.
Mitt. Komm. Quartärforsch. Österr. Akad. Wiss., 13, Wien, 2004
200
Figure 3: The new phylogenetic tree of
the cave bears and the position of the
Potočka bear (after RABEDER & HOFREITER, 2004; supplemented).
The new taxonomic scheme is given
in ﬁgure 3.
Partly considerable differences
(dimensions, evolutionary level,
sexual dimorphism) between the
populations from Potočka zijalka
and Gamssulzenhöhle are ascribed
to the more convenient living conditions on the southern exposure
of the Southern Alps compared to
the more rough conditions in the
Northern Alps (see RABEDER, this
volume).
Moreover, radiometric dates show
that cave bears of the U. ingressusline lived at the same time in Gamsulzenhöhle, as well as in Potočka
zijalka and Vindija (Croatia), which
is situated approximately 100 km
away from the latter.
Following the present stage of data
it is assumed, that U. ingressus
immigrated into the Alps around
50.000 BP where this species
replaced the other two forms of
cave bears (RABEDER & HOFREITER,
2004).
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