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Digitale Literatur/Digital Literature
Zeitschrift/Journal: Abhandlungen der Naturhistorischen Gesellschaft Nürnberg
Jahr/Year: 2005
Band/Volume: 45
Autor(en)/Author(s): Athen Kerstin, Pfretzschner Hans-Ulrich
Artikel/Article: Biometrical Analysis of Postcranial Elements of Fossil Ursids from Einhornhöhle, Germany 35-42 © Naturhistorische Gesellschaft Nürnberg e.V.download www.zobodat.at
Abhandlung Band 45/2005 Neue Forschungen Seite Naturhistorische Gesellschaft Nürnberg e. V. ISSN 0077-6149 zum Höhlenbären in Europa 35-42 Marientorgraben 8,90402 Nürnberg
Kerstin Athen & Hans-Ulrich Pfretzschner
Biometrical Analysis of Postcranial Elements of Fossil Ursids from Einhornhöhle, Germany
Zusammenfassung:
Ziel der in einem Poster präsentierten Untersuchung an Stylopodium, Zygopodium und Metapodium von Ursus deningeri und Ursus spelaeus ist die Benennung weiterer arttypischer Merkmale, mit denen die beiden Spezies gegeneinander abgegrenzt werden können. Ein weiteres Ziel besteht darin, das Fundmaterial aus der Einhomhöhle/BRD einer der beiden Arten zuzuordnen. Wie sich hier zeigt, liegt das Datenmaterial Einhomhöhle im Überschneidungsbereich zwischen Ursus deningeri und Ursus spelaeus.
Abstract:
A detailed study, which has been carried out on long bones and metapodia of Ursus deningeri and Ursus spelaeus, will be presented in this poster. One aim of this project is to make a distinction between these two species using further clear distinctive marks. A further aim is to classify the finds of fossil ursids from Einhomhöhle/FRG as belonging to one of the two species. It will be demonstrated that the material from Einhornhöhle shows characteristics of both species, which implies an intermediate state between Ursus deningeri and Ursus spelaeus.
Résumé:
Le but de cette étude des stylopodium, zygopodium et métapodium à’Ursus deningeri et à'Ursus spelaeus est l’identification de caractéristiques discriminatoires supplémentaires permettant de différentier clairement les deux espèces. Un autre but est d’attribuer le matériel en provenance de la Grotte de la Licorne (Einhomhöhle, Allemagne) à l’une des deux espèces. Il s’est avéré que les mensurations du matériel de la Grotte de la Licorne se situent dans la zone de recoupement entre Ursus deningeri et Ursus spelaeus.
Key words: long bones, metapodia, Ursus deningeri, Ursus spelaeus, Einhornhöhle/FRG
35 © Naturhistorische Gesellschaft Nürnberg e.V.download www.zobodat.at Introduction
At present it is assumed that the evolution of cave bears took the course from Ursus deningeri to Ursus spelaeus. One aim of this detailed study on long bones and metapodia is to make a distinction between these two species. Therefore, 249 measurements (D uerst 1926; von den D riesch 1976; G randal d ’A nglade 1993) on 16 different skeletal elements (humerus, ulna, radius, metacarpals I-V, femur, tibia, fibula, metatarsals I-V; Fig. 1, Fig. 2) of 2890 specimens Fig. 1: Metacarpalia I (Me I), 15 measurements, GL (greatest length), of exclusively adult individuals have SB (smallest breadth of diaphysis), SD (depth of diaphysis), GBp (greatest breadth proximal), SBp (smallest breadth o f proximal epiphysis), Dp been selected for this study. The (depth proximal i.e. thickness o f epiphysis), Bd (greatest breadth distal), samples originated from the sites Dd (greatest depth of distal epiphysis), SBdE (smallest breadth of (Fig. 3): Deutsch-Altenburg/Lower epicondyls), BdE (greatest breadth of distal epiphysis), Hd (distal height Austria (DA), Hundsheim/Lower from middle o f epiphysis to highest point ofjoint), BJp (greatest breadth o f proximal joint facet), DJp (greatest depth of proximal joint facet), PCm Austria (HH) and Repolusthohle/ (position of distal joint facet, distance between joint facet and medial Styria (RH) in Austria; Goyet/ margin), PCI (position of distal joint facet, distance between joint facet Condroz/Ardennes (Goy) in and lateral margin). All measurements were taken from: Duerst (1926), Belgium; Chateau/Saone-et-Loire von den Driesch (1976) and G randal d ’Anglade (1993). (CHA) in France; Einhornhohle/ Scharzfeld/Harz (EHH), Erpfingen/ Swabian Alb (Erpf), Mosbach-Sande close to Wiesbaden (MS) and Zoolithenhdhle/Franconian Alb (ZHSp) in Germany; Bacton/Norfolk (Bac), Banwell Bone Cave/Somerset (Ban) and Westbury-sub-Mendip/ Somerset (WSM) in the United Kingdom. The data sets have been statistically analysed using uni variate and multivariate tests like the i-test and Welch-i-test, and discrimi nant analysis in the programs SPSS and SAS in order to divide the two groups significantly. The second aim of this study is to classify the finds of fossil ursids from the Einhorn- Fig. 2 : Metatarsalia V (Mt V), 15 measurements, same as Me I. hohle (EHH) as belonging to one of the two species. The project was financially supported by the German Society of Research (DFG).
36 © Naturhistorische Gesellschaft Nürnberg e.V.download www.zobodat.at A Brief History of Einhornhohle The most EHH bear bone finds stemmed from old excavations of the years 1881 to 1907. This material was declared by R o d e (1935) to belong to the sub-species Ursus spelaeus var. hercynica, whereas S ch utt (1968) classified this material as belonging to the species Ursus deningeri. Further excavations followed between 1985 and 1988. N ielbo c k (1987) classified his finds as belonging to the species Ursus spelaeus. Datings of bone samples did not lead to any reliable results (R o se n d a h l et al. 2005). Alone a charcoal sample from the Jacob-Friesen-Gang in the Einhornhohle pointed to an age of the underlying bone layers as being older than 42,500 years.
Statistical Analysis: ¿-Test Results The aim of the i-test (also the Welch-t-test in some cases) on 2980 specimens was to track down differences between the fossil bear species Ursus deningeri and Ursus spelaeus. Requirements were met by forty-two variables, whose values followed an approximately Gaussian distribution in both groups. The EHH finds were left out of these tests for the time being. Significant differences were detected in the case of 36 of the 42 variables. (Table 1. Abbreviations cf. Fig. 1, Fig. 2). Fig. 3: Map with Statistical Analysis: Discriminant Analysis Results abbreviations for selected fossil sites; From altogether 602 EHH finds, 272 were classified as deningeri (Ud) and Ursus deningeri (black), Ursus 236 as spelaeus (Us), specimen by specimen, using discriminant analysis. spelaeus (red), Ninety-four specimens could not be classified. Out of 38 significant results classification case (blue).
37 © Naturhistorische Gesellschaft Nürnberg e.V.download www.zobodat.at
Fig. 4: Probability of in all, only 19 of high quality correct classification are cited in the Table 2. The to species Ursus deningeri. Skeletal EHH finds were in 11 cases element Me I, assigned to Ursus deningeri variables GL, SB, (Ud) and in 7 cases to Ursus SD, GBp, Bd, SBdE. spelaeus (Us) (italicized numbers). Only in one case could EHH material not be explicitly classified (ranged right). With all 38 results the ratio actually came to 25 Ud \\2Us\\ indeterminate case 78 59 (Fig. 4, Fig. 5 and Fig. 6 Ursus deningeri Ursus spelaeus show the probability of correct classification to one or other of the species on the
Fig. 5: Probability of 1,2 basis of an example). correct classification to species Ursus 1,0 Conclusions spelaeus. Skeletal element Me I, 0,8 O variables GL, SB, Fig. 6: Discriminant assumption that bear function values for evolution went from a tree individual groups, dwelling ancestor to Ursus Ursus deningeri, etruscus to Ursus deningeri, Ursus spelaeus and the EHH collection, and on to the purely ground depicted as box dwelling Ursus spelaeus, plots. Skeletal roaming over long distances, element Me I, which used its front paws to variables GL, SB, SD, GBp, Bd, SBdE. dig for food, e. g. roots (T h e n iu s 1945). Also in keeping with this adaptation to ‘economic locomotion’ are the dental findings, from N 57 80 40 Fig. 6 U.deningeri U.spelaeus EHH which it can be seen that 38 © Naturhistorische Gesellschaft Nürnberg e.V.download www.zobodat.at Ursus spelaeus was almost exclusively herbivorous, as opposed to the omnivorous species Ursus arctos and Ursus deningeri, and it attained bigger bodily proportions. EHH data in this investigation show the EHH finds to lie in an intermediate area between the two species, with predominant indications favouring membership of the species Ursus deningeri. There now ensue the following interpretation possibilities: Either this bear group was at a stage of development of high evolutionary level in the case of deningeri membership. This version speaks for a high geological age of the finds. The geological age though is not yet conclusively settled. Or the finds indicate an endemic living form with original ‘old’ characteristics, which coexisted con temporaneous with Ursus spelaeus. On the other hand the coherences expounded here still also allow of the interpretation, that it could be a question of a low evolutionary level in the case of spelaeus membership. References Duerst , J. U. (1926): Vergleichende Untersuchungsmethoden am Skelett bei Säugern.- Handbuch der biologischen Arbeitsmethoden, Abt 7, 125-530, Berlin, Wien. G randal d ’Anglade , A. (1993): El oso de las cavernas en Galicia: El yacimiento de Cova Eirós.- Nova Terra, 8, 1-285. N ielbock , R.-D. (1987): Holozäne und jungpleistozäne Wirbeltierfaunen der Einhornhöhle im Harz - Paläontologisch-biostratigraphische Untersuchungsergebnisse der Höhlengrabung 1985-87.- Dissertation, 1-194, Clausthal. Rode , K. (1935): Untersuchungen über das Gebiss der Bären.- Monographien zur Geologie und Paläontologie, Serie II, 1-163. Rosendahl , W., D öppes , D. D., Joger , U., Laskowski , R., L ópez C orrea , M., N ielbock , R.-D., W rede , V. (2005): New radiometric datings of different Cave Bear sites in Germany - results and interpretations.- Bull. Soc. Hist. Nat. de Toulouse et de Midi-Pyrénees, 104, 10, Toulouse (in press). Schütt , G. (1968): Die cromerzeitlichen Bären aus der Einhomhöhle bei Scharzfeld.- Mitteilungen aus dem Geologischen Institut der Technischen Hochschule Hannover, Diss., 7, 1-121. Thenius , E. (1945): Die Plantigradie der Bären. Zugleich ein Beitrag zur Klärung der primären und sekundären Plantigradie.- Dissertation, Universität Wien, 1-136, Wien. von den Driesch , A. (1976): A guide to the measurement of animal bones from archaeological sites.-Peabody Museum Bulletin, 1, 1-137. Addresses o f authors: Kerstin Athen, Marschnerstr. 45, D-30167 Hannover, Tel.: 0049/(0)511/1690-620; [email protected] Hans-Ulrich Pfretzschner, Universität Tübingen, Institut für Geowissenschaften, Sigwartstr. 10, D-72076 Tübingen 39 40 CO z *o *o £ to iff a o u « (/> mean value | bone / variable | teststatistics | p-value in SPSS | raw p-value in SAS | adjusted p-value, Bonferroni | adjusted p-value, Holm | d ti .c 5 CO CO COo ao CO st ■sto co ■st qj c O)a3 ID _C CD en o o o o o o Ö o o o o o o CNCN o st Ö o o o o p o o o o o Ö *- c ai O 3 ID -2 CDLO r-. CO ■sf CVjCO CD CD CD to Q.Q> 05 co b TO .c LOO If) r- CD00 p CNCO a> c CD <£ a p o o o d o o ■a CD h- CO d p d p o o o o o d o o o o o O o o b -2 CO CD f". in COCDto COst o CD & to 3 b TO .C CDCO CDLO r- CD p 05 CO CD CO CbC CD CD CO CÛ p in st CO o d p o o d p o o o o o p o o o o o d p o o o o o - d o Q_ b r-- h-. co 00 CDQ0LOr-- CNCO05 05 to a a3 CD0) 3 to b ■Q .C CO 05 COm COCN d p CD05 CO ■N" Cbc CD qj 2 a. O CD O CDCD CD d p o o o o o o o o d p o o o o O d p o o o o o d *- u E b ■2 st 05 d 03 COLOCO d CO o CNh- QJ a Cb3 b N- p LOLOo ~o .c LO CD CNCD CO qjc CD (b 2 > CLo d CO05 CN d p o o o o o o o o o d p o o o o o d p o o o o d u E d r- d CD r-. b 2 LO p •sf CN st CD to a cb (b to b TO .g CO d CO o sf p CDCN CO cb c CD CL CD LOst d o CD 05 CD d p o o o o o o o p o o o o o d p o o o o o - d © NaturhistorischeGesellschaftNürnberge.V.downloadwww.zobodat.at o E b LOCN h- LOD-h- 2 CN CNCDLO to a QJ CO LU cb CDCOCO CO■O 05 CD p d p o o o o o p o o o o o d o o d d p o o o o o b 2 r-- ■Sf h- CO5 LO CD05 LO GO to Q.Q) (b to 3 TO .c ■St CO•St COCN ■stin CN05 cb c CD 05 0 CO X _l ? A cb p 05 •St•st in r^ p ■St d p o o d o o o o o o d o o o o o d o p o o o o E D aj to a> o D b 2 COCO COCDst aq 00 CN05 CNst Do to a QJ E Q_ 0) o 3 CDo CO CDD-h- cb p o D»D- d p Q. to 3 3 *o CO .c CO d in to CO in D'o 05 CN 0) c 05 QJ I m co D^-o D- COo d p o o p o o o o o d p o o o o o d d p o o o o CN Û. 5 b - D 05 2 CN p o CNCO CDCNGOCN 8- q Cb3 3 b TO CO cb LOD-D- ,c COcb COCOO) 05 5 0 05 c X CO CD § cb D-D- A CD 00 CDTt CO d p o o d p o o o o o p d p o o o o o d o o o o CN E o3 3 3 if) CD a b LOD- CD LO •St CNCOao CD ■st CN 6 05 CD 0) to 3 b o * .c GO d st CO05 c CD 05 QJ 5 41 © Naturhistorische Gesellschaft Nürnberg e.V.download www.zobodat.at Tab.2: results of the probability classificati on of EHH to Species N bone / variables Table with list of of correct classification in % U d U s classification results U. deningeri 46 o f finds from Einhom- U. spelaeus 83 Ml V / SB, SD, SBdE 92,7 hohle, Harz/FRG EHH 43 16 2 3 (EHH). Out o f 38 U. deningeri 64 significant results in U. spelaeus 80 Ulna / BPcii, DPanc 91,5 all, only o f high EHH 33 7 3 quality are cited in U. deningeri 75 this table. U. spelaeus 90 Mcl/SD, BdE 85,6 EHH 42 20 8 U. deningeri 70 U. spelaeus 94 Ml IV / GBp, PCm 85,4 EHH 41 14 9 U. deningeri 80 Me v / (j L, bbt i>U, tjbp, bp, U. spelaeus 97 85,0 BJp, Dp, Bd, Hd, SBdE EHH 34 10 23 U. deningeri 70 U. spelaeus 97 Me II / GL. SB, SD, GBp, Bd, SBdE 84,7 EHH 43 25 15 U. deningeri 70 Me IV / GL, SB, SD, GBp, SBp, 82,9 U. spelaeus 94 Dp II, BJp, Bd, SBdE EHH 41 14 18 U. deningeri 49 U. spelaeus 83 Radius / D, DColl, Bd 82,9 EHH 26 4 1 U. deningeri 75 U. spelaeus 90 Me I / GL, SB, SD, GBp, Bd, SBdE 81,8 EHH 42 26 14 U. deningeri 80 U. spelaeus 97 McV/GL. SB, SD 80,7 EHH 34 9 2 4 U. deningeri 47 U. spelaeus 87 Tibia / SB, SD, BDd, Bd, Dd, BJd, DJd 79,8 EHH 39 10 2 0 U. deningeri 70 U. spelaeus 94 Mt IV/GL, SB, SD, Bd, SBdE 79,3 EHH 41 7 3 4 U. deningeri 69 U. spelaeus 97 Me III/SB, SD, Bd 79,1 EHH 42 28 7 U. deningeri 65 U. spelaeus 96 Mt II / Dd, PCm 78,6 EHH 36 15 4 U. deningeri 65 U. spelaeus 96 Mill/SBp, DJp 76,9 EHH 36 15 1 U. deningeri 38 U. spelaeus 63 Humerus / SB, BT, MT, LT, Bd. HO, BCse 75,0 EHH 25 14 3 U. deningeri 38 U. spelaeus 63 Humerus / SB, BT, MT, LT, Bd 75,0 EHH 25 15 2 U. deningeri 80 Mt III / GL, SB, SD. GBp, SBp, Dp, U. spelaeus 104 74,7 BJp. DJp, Bd, Dd, Hd, BdE, SBdE EHH 64 20 20 U. deningeri 64 Ulna / B, D, GDO, DPanc, U. spelaeus 80 BpPcor, OPcor, BPcor, BPcii, 74,3 Bd, Dd. Bpsty. Dpsly EHH 33 3 7 42