Geo.Aip - Science South Tyrol

Transcription

Geo Alp
A new yearly journal devoted to
· Alpine geology
Neue Jahreszeitschrift zur
AI pe·ngeolog ie
La nuova rivista per Ia Geologia
delle AI pi
Geo.Aip
Redaktionskomitee: Rainer Brandner, lnnsbruck, Diethard Sanders, lnnsbruck, Volkmar Mair, Bozen, Benno Baumgarten,
Naturmuseum Bozen
Technische Redaktion/Layout: Monika Tessadri-Wackerle
Herausgeber, EigentUmer und Verleger:
lnstitut fur Geologie und PaHiontologie, Universitat lnnsbruck, Naturmuseum Bozen
Chefredakteur Geo.Aip 2: Karl Krainer
Referentlnnen dieser Nummer:
M. Breda, Padova; H. Kerp, Munster; T. Kotsakis, Roma; S.G. Lucas, Albuquerque; D. Nagel, Vienna; Chr. Rupp, Vienna
B. Sala, Ferrara; R. Sardella, Roma; G. Tichy, Salzburg
Erscheinungsweise und Bezug:
Geo.Aip erscheint einmal jahrlich und kann bei beiden herausgebenden lnstitutionen im Abonnement oder einzeln bezogen werden :
lnstitut fUr Geologie und Palaontologie, lnnrain 52, A-6020 lnnsbruck, Austria
Naturmuseum SUdtiroi/Museo Scienze Naturali Alto Adige, Bindergasse/via Bottai 1, 1-39100 Bozen/Bolzano, Italy
©
lnstitut fUr Geologie and jlalaontologie, Universitat lnnsbruck; Naturmuseum SUdtiroi/Museo Scienze Naturali Alto Adige
Genehmigung des Landesgerichts Bozen Nr. 12/2004 vim 05/11/2004
Verantwortli~;:her
Direktor: Dr. Vito lingerie
ISSN 1824-7741
Umschlagbild: Monika Tessadri-Wackerle, verwendete Abbildung von Evely Kustatscher
Druck: Walser Druck KG
F
Geo.Aip
In halt
Herbert Scholz, Karl-Heinz Bestle & Sebastian Willerich: Ouartargeologische Untersuchungen im Oberetsch
Beitrage zu ,Giornate della Paleontologia der Societa Paleontologica ltaliana 2004", 20-23. Mai 2004:
Raffaele Sardella, Claudia Bedetti, Luca Bellucci, Nicoletta Conti, Danilo Coppola,
Emmanuele Di Canzio, Marco Pavia, Carmela Petronio, Mauro Petrucci & Leonardo Salari:
The Late Pleistocene vertebrate fauna from Avetrana (Taranto, Apulia, Southern Italy) : preliminary report.............
25
Evelyn Kustatscher & Johanna H.A. van Konijnenburg-van Cittert: The Ladinian Flora (Middle Triassic)
of the Dolomites: palaeoenvironmental reconstructions and palaeoclimatic considerations ........................................
31
Cristiana lata & lassos Kotsakis: Italian fossil chiropteran assemblages: a preliminary report ......................................
53
Gabriella Mangano: Cervus elaphus siciliae from Pleistocene lacustrine deposits
of Acquedolci (North-Eastern Sicily, Italy) and its taphonomic significance.........................................................................
61
Gabriella Mangano, Laura Bonfiglio & Daria Petruso: Excavations of 2003 at the
S. Teodoro Cave (north-eastern Sicily, Italy): preliminary faunistic and stratigraphic data
71
Giuseppe Santi: Lower Permian paleoichnology from the Oroboc basin (northern Italy) ................................................
77
Maria Teresa Curcio, Longino Contoli, Emanuele Di Canzio & lassos Kotsakis: Preliminary analysis of the first
lower molar variability in Late Pleistocene and living populations of Terri cola savii (Arvicolidae, Rodentia) ...........
91
Davide Mana: A test application of the SHE method as a biostratigraphical parameter .......................... .......................
99
Cinzia Galli, Mario Rossi & Giuseppe Santi: Ursus spe/aeus Rosen muller, 1794 from the Venetian region
of Northern Italy: Preliminary notes on its evolutionary path .................................................................................................. 107
Alessandro de Carlis, Enrico Alluvione, Alessandro Fonte, Mario Rossi & Giuseppe Santi: Morphometry
of the Ursus spelaeus remains from Valstrona (Northern Italy) ................................................................................................ 115
Abstracts zu ,Giornate della Paleontologia der Societa Paleontologica ltaliana 2004", 20-23. Mai 2004:
Francesco Garofalo, Fabrizio Bizzarini & Federica Ferrieri: The activities of the Ligabue Study Research Centre
on the thirtieth anniversary of its foundation ................................................... ........................................... ................................. 127
Nicola Daii'Oiio : The origin of the palaeontological fossil concept ......................................................................................... 131
INSTRUCTIONS TO AUTHORS
Articles may be submitted in English, German or Italian. In case of a German or Italian text, the captions to all figures, plates and tables must be also in English, and an English abridged version (1000-1500 words) and abstract are
to be delivered.
Articles shall be submitted in
th~e
copies to:
Karl Krainer, Diethard Sanders, Institute of Geology and Palaeontology, University of lnnsbruck, lnnrain 52, A-6020
lnnsbruck, Austria. E-mail: [email protected]; [email protected]
or to:
Benno Baumgarten, Naturmuseum Si.ldtiroi/Museo Scienze Naturali Alto Adige, Bindergasse 1Nia Bottai 1, 1-39100
Bozen/Bolzano, Italy: E-mail: [email protected]
Articles must be typed double-space. The quality of line-drawings must be ready for print. In line-drawings and figures of any sort, all labellings, numbers and letters should be readable upon 50% reduction in size.
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Referencing:
Articles:
Author 1, X. Y., Author 2, Z. A. (2002): Title of article. - International journal abbreviation (e.g. Sediment. Geol.), vol. :
pp-pp.
.
Articles in books:
Author 1, X. Y., Author 2, Z. A. (2002): Title of article.- In: Person A, Person B. (eds.): Title of book, pp-pp, publisher,
place of publication.
Books:
Author 1, X. Y., Author 2, Z. A. (2002): Title of book. - no. of pages, publisher, place of publication.
Reprints: 50 reprints are free of charge
Geo.Alp, Vol. 2, S. 1–23, 2005
QUARTARGEOLOGISCHE UNTERSUCHUNGEN IM ÜBERETSCH
Herbert Scholz, Karl-Heinz Bestle & Sebastian Willerich
Mit 8 Abbildungen und 1 Fototafel
With 8 figures and 1 plate
Herbert Scholz, Karl-Heinz Bestle und Sebastian Willerich,Lehrstuhl für Ingenieurgeologie der Technischen Universität München,
Arcisstr. 21, D-80290 München; e-mail: [email protected]
Zusammenfassung
In der weiten Talung von Eppan-Kaltern im Südtiroler Überetsch bei Bozen ist ein ganzes System von kiesigen Lateralmoränen, breiten Kamesterrassen sowie in die mächtige, komplex aufgebaute pleistozäne Talfüllung erosiv eingeschnittenen, kastenförmigen Schmelzwasserrinnen erkennbar, mit deren Hilfe sich unterschiedliche Eisstände einer „Kalterer Zunge“ aus dem ausgehenden Hochglazial rekonstruieren lassen, einer
Teilzunge des Etschgletschers. An den Osthängen des Mendel-Roèn-Kammes sind zudem zertalte Erosionsreste von Murfächern nachweisbar, hier als „Murkames“ bezeichnet, die direkt gegen den absinkenden
Westrand dieses Gletschers geschüttet wurden. Sehr kalkund dolomitreiche Mursedimente, durch Eisauflast
überkonsolidiert und damit vermutlich älter als der letzte Eishöchststand, aber auch Ablagerungen von
deutlich jüngeren Muren, die aus dem Spät- und Postglazial stammen dürften, bedecken große Flächen an
der Ostflanke des Mendel-Roèn-Kammes bis ins Tal hinunter, besonders im Gebiet zwischen Margreid, Penon
und Kurtatsch. Gerade auf diesen von den Einheimischen „Kampferde“ genannten diamiktischen Ablagerungen liegt ein Großteil der Weinberge rund um Tramin und Kurtatsch.
Summary
Within the wide vale of Eppan-Kaltern (Appiano-Caldaro) at Überetsch (Oltradige, Sella di Appiano-Caldaro) close to Bozen (Bolzano) in South Tyrol (Alto Adige) a complicated system of gravelly lateral moraines,
large kame terraces as well as erosive fossil meltwater valleys can be identified, which are deeply incised into
a thick and complex sequence of Pleistocene sediments. Most of these structures are remnants of the
„Kaltern lobe“, a late Pleniglacial tongue of the Etsch (Adige) valley glacier. Moreover erosional remnants of
debris fans can be identified at the eastern slopes of Mendola-Roèn-Ridge, obviously deposited in the gap
between the mountain slope and the western rim of this recessing glacier (“fankame“). The eastern slopes of
the Mendola-Roèn-Ridge, especially the region between Margreid (Magrè all’Adige), Penon (Penone) and
Kurtatsch (Cortaccia), are widely covered with debris flow deposits which are extremely rich in limestone and
dolomite fragments. They are partly older and partly younger than the “fankame“. Some are obviously overconsolidated by the extra load of glacier ice and therefore presumably older than the Last Glacial Maximum,
some are obviously younger and may have a Late to Postglacial age. Many vineyards around Kurtatsch (Cortaccia) and Tramin (Termeno) are situated right on the surface of these diamictic deposits, called
“Kampferde” by the local people.
1
Einleitung
Die hier vorgestellten Ergebnisse wurden im Rahmen dreier Kartierungsübungen mit fortgeschrittenen Geologiestudenten der TU München
sowie bei eigenen Geländebegehungen im Gebiet
zwischen Oberplanitzing und Margreid in Südtirol
erarbeitet. Diese von Prof. Dr. Herbert Scholz betreuten Kartierungsübungen waren vom 31. März
bis zum 11. April 2002, vom 24. März bis zum 4.
April 2003 und vom 23. März bis zum 3. April 2004
durchgeführt worden und hatten vor allem das Ziel,
die quartären Sedimente im Überetsch zu erkunden,
genetisch zu klassifizieren sowie zeitlich zu ordnen.
Alle Geländeübungen wurden seitens des Amtes für
Materialprüfung und Geologie in Bozen (Dr. Volkmar Mair), des Südtiroler Amtes für Gewässerschutz
und der Gemeinde sowie der freiwilligen Feuerwehr
von Kurtatsch (Bürgermeister Oswald Schiefer,
Kommandant Albert Terzer) unterstützt.
An den drei Kartierungsübungen haben folgende
Studenten bzw. Studentinnen teilgenommen: W.
Bäumel, I. Baumann, K.-H. Bestle, A. Dargel, M. Döhner, M. Elsner, Chr. Gampe, G. Ghon, R. Hohlfeld, J.
Kadlcakova, F. Köppl, M. Lammel, F. Meyer, Chr.
Minet, Chr. Mögele, I. Monsorno, S. Suckfüll, I. Thiele, B. Weiher, Chr. Weber, K. Wendl und S. Willerich.
Eingehendere Untersuchungen zur Quartärgeologie des Gemeidegebietes von Kurtatsch wurden
im Rahmen zweier Diplomarbeiten von Karl-Heinz
Bestle und Sebastian Willerich am Lehrstuhl für Ingenieurgeologie der TU München angestellt (Bestle
2005 und Willerich 2005). Diese Diplomarbeiten
wurden von Prof. Dr. Herbert Scholz im Gebiet zwischen Tramin, Graun, Fennberg und Margreid vergeben und betreut. Sie wurden vom Amt für Geologie
und Baustoffkunde in Kardaun sowie von der Gemeinde Kurtatsch unterstützt. Allen, die wissenschaftlich, logistisch oder finanziell zum Gelingen
dieser Untersuchungen beigetragen haben, sei an
dieser Stelle herzlich gedankt.
1. Geologischer Aufbau des Überetsch
1.1 Die Gesteine des Untergrundes im Überblick
Die westliche Talflanke der Etschtalfurche südwestlich von Bozen, das Gebiet von Überetsch und
dem Mendelkamm, wird vor allem von permotriassischen Gesteinen aufgebaut. Die Schichtfolge, die
2
mit dem unterpermischen Bozener Quarzporphyrkomplex beginnt, fällt generell leicht nach SW ein.
Die Mächtigkeit des Bozener Quarzporphyrs dürfte
weit über 1000 m betragen, denn das Gebiet befindet sich noch innerhalb der permischen „Caldera von
Bozen“ (Bosselini 1998: 82), in der besonders mächtige Ignimbritfolgen akkumuliert wurden. Über den
permischen Vulkaniten folgt der terrestrische, mit 40
bis 70 m vergleichsweise geringmächtige Grödner
Sandstein (Perm), eine teilweise kohleführende, bunt
gefärbte Folge von Sandsteinen und Tonschluffsteinen (Brandner & Mostler 1982, Gwinner 1971).
Die Gröden Formation geht zum Hangenden in
die flach-marine Werfen Formation (Skyth) über. Die
oberpermische Bellerophon Formation, die schon
wenige Kilometer östlich der Etsch weit verbreitet ist
(Heissel 1982: 22, 28), fehlt hier hingegen völlig. Die
hier aufgeschlossene, regelmäßig gebankte tonigschluffige Schichtfolge der Werfen Formation enthält zahlreiche feste Bänke aus Schluffsteinen,
Sandsteinen, Dolomiten, Mergelkalken, Kalken und
Oolithen. Sie lässt sich nicht ohne weiteres mit dem
Werfener Standard-Profil im Schlerngebiet oder in
der nur wenige Kilometer entfernten Bletterbachschlucht parallelisieren (vergl. Brandner & Mostler
1982, Moser 1996). Das Unteranis wird durch den
Unteren Sarldolomit und stellenweise durch brennend rote Sandsteine (evtl. Äquivalente des VoltagoKonglomerates) repräsentiert, die sich ohne scharfe
Grenze aus der unterlagernden Werfen Formation
entwickeln. Diese bunten Sandsteine werden von
einer grau gefärbten, kalkig-mergeligen Schichtfolge überlagert, bei denen es sich um Äquivalente der
oberanisischen Morbiac-Kalke handeln dürfte. Diese
gehen zum Hangenden hin in den Contrindolomit
über, dolomitischen und gebankten gelblich anwitternden Flachwasserkarbonaten des Oberanis.
An den steilen Schluchthängen, die vom Mendelkamm zum Etschtal hinunterziehen, sind immer wieder grobblockige Konglomerate mit sandig-tonigem,
rotem Bindemittel, rötliche Sandsteine und Tone
aufgeschlossen. Diese konglomeratischen, teilweise
riesige Blöcke enthaltenden Sedimente stellen offensichtlich Füllungen klammartiger fossiler Erosionsrinnen dar, die mehrere Dekameter tief in die
Schichtfolge der Werfen Formation eingeschnitten
sind. Sie werden sicher vom Contrindolomit, teilweise wohl auch von den Morbiac-Kalken überlagert
und sind z.B. an der Forststraße im Höllental aufgeschlossen, die von Graun nach Söll führt. Diese
Konglomerate enthalten vor allem gelblich gefärbte
Geo.Alp, Vol. 2, 2005
Komponenten der Werfen Formation, daneben aber
auch kleine Geröllchen aus hellgrauem Dolomit. Ob
es sich um Äquivalente des Richthofen- oder des
Voltago-Konglomerates handelt, ist unbekannt. Die
oben genannten Konglomeratvorkommen füllen im
Höllental fossile Erosionsrinnen auf, die klammartig
bis in den Unteren Sarldolomit eingeschnitten sind.
An der Anis-Ladin-Grenze entwickeln sich im
Hangenden des Contrindolomits zwischen Margreid
und Tramin die kalkig-mergeligen, teilweise auch
sandigen „Zwischenschichten“, eine bituminöse
Beckenfazies mit Tuffiteinschaltungen. In diese
Schichtfolge sind Dolomite, gebankte Kalksteine
und chaotisch gelagerte Brekzien aus Flachwasserkalken (Olistostrome) eingeschaltet, denn die „Zwischenschichten“ verzahnen sich nach S hin offen sichtlich mit den Flachwasserablagerungen des basalen Schlerndolomits. Sie haben sich in einem kleinen, aber vermutlich recht tiefen, im Norden durch
Störungen begrenzten Becken gebildet, im sogen.
„Kurtatscher Loch“. Der Mendelkamm selbst wird
von mächtigem Schlerndolomit (Ladin) und Hauptdolomit (Karn/Nor) aufgebaut, der im Norden direkt
dem Contrindolomit, im S auch den „Zwischen schichten“ auflagert. Südlich von Margreid, wo die
gesamte Mittel- und Obertrias in ähnlicher dolomitischer Fazies entwickelt ist, lässt sich die Dolomitfolge nur schwer untergliedern und wird als „Mendeldolomit“ bzw. „Etschtaldolomit“ zusammengefasst (Geyer 1993).
An der Straße von Penon nach Fennberg und in
Fennberg selbst werden die Dolomite von geringmächtigen, teilweise bunt gefärbten pelagischen Kalken überlagert, die schon dem Jura und
der Oberkreide angehören.
1.2 Tektonik im Überblick
Zwischen Bozen und Neumarkt folgt das Etschtal
vermutlich einer N-S-orientierten Störungszone, an
der die östliche Talflanke gegenüber der westlichen
deutlich herausgehoben zu sein scheint. Die Auflagerungsfläche des Grödner Sandsteins auf dem
Quarzporphyr liegt bei Kaltern mindestens 500 m
tiefer als auf der gegenüberliegenden Seite des
Etschtales bei Branzoll. Parallel dazu dürfte wohl
auch – unter mächtigem Quartär verborgen – eine
Störungszone durch die weite Talung von EppanKaltern im Überetsch verlaufen, denn der Quarzporphyr zwischen Gaid und Tramin passt nicht zu dem
auf der anderen Seite dieser Talung. Die Oberfläche
des Quarzporphyrs am Montiggler Wald scheint
mehr als 230 m tiefer zu liegen als am gegenüberliegenden Gandberg bei Oberplanitzing, der am
Mitterberg knapp 100 m tiefer als am gegenüberliegenden Seeberg bei Altenburg. Insgesamt scheint
die Etschtalstörung hier also die Struktur eines Grabens mit etwas ungleich hoch liegenden Grabenschultern zu besitzen.
Außerdem ist die Schichtfolge durch mehrere
quer dazu verlaufende, vor allem E-W- bis SE-NWorientierte Störungen in einzelne Schollen zerlegt.
An solchen Abschiebungen verspringt die Oberkante des Quarzporphyres erkennbar, etwa nördlich von
Söll oder unmittelbar nördlich des Bergsturzes von
Eppan, wo sich zwischen Matschasch und dem
Schloss von Englar eine Sprunghöhe von rund 480
m ergibt! Viele dieser Störungen werden durch
große Täler und Schluchten nachgezeichnet, die
vom Überetsch zum Mendelkamm hinaufziehen,
z.B. das Höllental.
Oberhalb von Penon und Graun ist eine größere,
über weite Abschnitte subhorizontal verlaufende
Überschiebungsbahn kartierbar (Vigo-di-Ton-Termeno-Linie), entlang der die Dolomite des Mendelkammes auf unterschiedliche Trias- und Jura-Gesteine in südöstlicher Richtung überschoben sind.
An dieser Überschiebungsbahn sind die Dolomite
der Deckenbasis extrem stark beansprucht und in
einer teilweise dekametermächtigen Zone kleinstückig zerbrochen worden. Diese jedenfalls postjurassisch entstandene Überschiebungsbahn ist sicher
alpidisch. Sie wird von einigen der oben erwähnten
Querstörungen versetzt, die gleichfalls alpidisch
oder jünger sind. Andere Störungen, etwa die E-Wverlaufende große Abschiebung mit einer Sprunghöhe von mindestens 430 m, die zwischen Graun
und Kurtatsch den Nordrand des „Kurtatscher Loches“ markiert, müssen dagegen schon in der Trias
aktiv gewesen sein, da sich die Mächtigkeit der
„Zwischenschichten“ an dieser Störung sprunghaft
ändert. Diese Störung scheint allerdings abschnittsweise den Charakter einer Aufschiebung zu besitzen, also wohl durch die alpidische Einengungstektonik überprägt zu sein.
1.3 Das Quartär im Überblick
Die permotriassischen Festgesteine des Überetsch
werden großflächig von lockeren Ablagerungen des
3
Abb. 1: Stark vereinfachtes Übersichtskärtchen des Etschtales zwischen Bozen und Auer an der Wende vom Hoch- zum Spätglazial (Eisstand von Auer). Eingezeichnet sind eine Reihe von
Phänomenen, auf die in den folgenden Kapiteln näher eingegangen wird: Schwemmfächer, Felssturzmassen, Blockgletscher, Verbreitung von Mursedimenten, „Murkames“ und der Eisstand von Auer (Fuschgalai-Stadium). Kamesterrassen und pleistozäne Erosionsrinnen, die für die Rekonstruktion des Rückmelzens der Kalterer Zunge herangezogen wurden, sind hier der
Übersichtlichkeit halber weggelassen und auf einer eigenen geomorphologischen Karte dargestellt (Abb. 6).
Fig. 1: Simplified map showing the Etsch Valley between Bozen (Bolzano) und Auer (Ora) at the beginning of late Würmglacial times (stage of Auer). The phenomena shown on this map are
described more thoroughfully in the following chapters: alluvial fans, masses of rock fall debris, rock glaciers, distribution of debris flow sediments, „fankame“ and the stage of Auer
(Fuschgalai-substage). Due to clearness Pleistocene erosional valleys and kame terraces are ignored here, although they are important for the reconstruction of the deglaciation. These phenomena are shown on a separate geomorphological map (fig. 6).
4
Quartärs verdeckt, die in der weiten Talung von
Eppan-Kaltern erhebliche Mächtigkeiten erreichen
können. Es handelt sich vor allem um Geschiebelehme, Schmelzwasserkiese, Seesedimente und Murablagerungen, deren Mächtigkeiten mit zunehmender
Höhenlage generell abnehmen. Der größte Teil dieser Sedimente ist während des Pleistozäns, vor
allem während der Würmeiszeit entstanden und
steht in direktem oder mittelbarem Zusammenhang
mit dem Etschgletscher, der in den kältesten Abschnitten des Eiszeitalters zeitweise das ganze
Etschtal ausfüllte.
Während des Vereisungsmaximums der letzten
Eiszeit, vor ca. 20.000 Jahren, dürfte das Etschtal
südlich von Bozen bis in Höhen von über 2000 m
mit Gletschereis erfüllt gewesen sein (Hantke 1983:
197). Nur noch die höchsten Teile des Mendelkammes, am Roèn (2116 m), überragten noch die Eisoberfläche (Klebelsberg 1949, Husen 1982). Südtirol
dürfte damals ähnlich im Eis ertrunken gewesen
sein wie die Gebirge Ostgrönlands (Scholz 1984,
1986). Über dem Mendelpass stand das Eis des Etschgletschers mit dem im Nonstal liegenden NoceGletscher in Verbindung (Hantke 1983: 197). Der
Etschgletscher stirnte in dieser Zeit noch südlich des
Gardasees südlich Solferino (Habbe 1969). Die Geschiebe, die der Etschgletscher transportierte, stammen größtenteils aus den zentralalpinen Nährgebieten dieses Gletschers, vor allem aus den ÖtztalStubaier Alpen, der Silvretta, dem Ortler-Gebiet,
den Sarntaler Alpen, westlichen Zillertaler Alpen
und westlichen Dolomiten.
Beim Rückschmelzen des Eises im ausgehenden
Hochglazial sank die Eisoberfläche der großen
Talgletscher – natürlich auch die des Etschgletschers – langsam ab. Dadurch wurden die übersteilten Talhänge freigegeben und waren zunehmend der Erosion ausgesetzt. Auf der Höhe von
Auer muss sich der Etschgletscher beim Dünnerwerden des Eises in zwei Eisloben aufgespalten haben
(Abb. 7): eine Eiszunge floss über Bozen und folgte
dem Etschtal abwärts (Etschtalzunge), eine zweite
Eiszunge drang bei Missian ins Überetsch ein und
folgte der weiten Talung von Kaltern (Kalterer
Zunge). Große Felskuppen, die vom Wilden-MannBühel über den Großen Priol, Jagenberg, Mitterberg,
Unterberg und über die Leuchtenburg zum Piglon
ziehen (insgesamt teilweise als „Mittelberg“ bezeichnet), wirkten dabei als Eisteiler (Abb. 1, 7). Das
Eis der Etschtalzunge muss um ein Vielfaches mächtiger gewesen sein als das der Kalterer Zunge. Be-
sonders die Kalterer Zunge und ihr langsames Rückschmelzen lässt sich anhand entsprechender
Ablagerungen gut rekonstruieren.
Der Etschgletscher hat das Etschtal zum weiten
Trogtal umgeformt, dessen trogähnlicher Talquerschnitt aber nicht sichtbar ist. Der heutige Talboden, die landwirtschaftlich intensiv genutzte Etschtalebene, ist eine Akkumulationsfläche, die erst
während und nach dem Rückschmelzen der Gletscher entstanden ist und bei Andrian rund 240 m,
bei Tramin 215 m über dem Meer liegt. Das Etschtal
ist, wie alle großen Alpentäler, mit mächtigen quartären Ablagerungen aufgefüllt, vor allem mit fluviatilen Kiesen und Seesedimenten. Am Aufbau der
quartären Talfüllung sind zwischen Bozen und Salurn entsprechend einer mündl. Mitt. von Herrn Dr.
W. Sadgorski (vormals LfW, München) auch mächtige Torfe mit geringmächtigen Auelehm-Zwischenlagen beteiligt (insgesamt 30 und 60 m).
Randlich dürften auch Rutschmassen und Mursedimente am Aufbau der Talfüllung beteiligt sein. Der
Felsuntergrund ist bei einer Bohrung südlich von
Andrian erst in einer Teufe von über 670 m unter
der Oberfläche erreicht worden (Werth 2003). Bei
Auer hat eine Bohrung den Felsuntergrund in einer
Tiefe von 200 m dagegen noch nicht erreicht
(mündl. Mitt. Dr. Volkmar Mair).
2. Landschaftselemente im Überetsch
2.1 Rundhöcker und Gletscherschliffe
Weit verbreitet sind im Überetsch eisüberschliffene Rundhöckerlandschaften. Große Felder mit
Rundhöckern sind fast ausschließlich auf Quarzporphyr-Oberflächen ausgebildet, z.B. in der Umgebung der Montiggler Seen, am Seeberg bei Altenburg oder am Kalvarienberg in St. Michael (Eppan).
Allerdings scheint die Ausbildung ideal geformter,
walrückenartiger Rundhöcker, mit flachen, geschrammten Luv- und steilen, gebrochenen Leeseiten durch die engständige Klüftung vielfach verhindert worden zu sein. Einige ideal geformte Rundhöcker sind am Trimm-dich-Pfad östlich des Sportplatzes von Kaltern zu finden. Schöne Rundhöckerfelder sind auch auf dem Plateau von Unterfennberg südlich von Margreid auf Contrin-Dolomit
entwickelt. Die anderen Gesteine des untersuchten
Gebietes sind offenbar nicht hinreichend isotrop
5
und fest, um die Entwicklung von Rundhöckern zuzulassen. Geschrammte Gletscherschliffe sind gewöhnlich nur dort erhalten geblieben, wo die
Gesteinsoberflächen durch eine hinreichend mächtige Auflage von Geschiebelehmen vor der
Verwitterung geschützt waren. Trotz einer anzunehmenden Eisüberlagerung von 1500 bis 1800 m
im Überetsch, die an sich zur Ausbildung von Sichelmarken und Parabelrissen ausreicht, wurden auf
den eisüberschliffenen Gesteinsoberflächen keine
entsprechenden Strukturen beobachtet.
Abb. 2: Schema der Genese von Kamesterrassen am Westrand
des Etschgletschers. Die Kamesterrassen wurden durch
Schmelzwässer zwischen Berghang und Eisrand aufgeschüttet,
teilweise auch unter Beteiligung von Murmaterial, das den
Schmelzwassersedimenten vom Berghang her seitlich zugeführt wurde (unten). Nach dem Abschmelzen des Gletscher eises wurden die Kamesterrassen zertalt (oben).
Fig. 2: Simplified sketch showing how kame terraces at the
western rim of the retreating Etsch valley glacier may have
formed. They have been generated by accumulation of meltwater sediments within the gap between the mountain slope
and the glacier. Gravel derived from the slope above has been
added by debris flows (below). After the glacier ice has vanished these kame terraces have been cut by erosional valleys
(above).
6
2.2 Tille (Geschiebelehme, Geschiebesande)
Stellenweise treten im Überetsch schluffig-sandige und stark verdichtete Geschiebelehme auf
(lodgement till, „Grundmoräne“), die teilweise so
wenig Schluff enthalten, dass sie besser als Geschiebesande bezeichnet werden sollten. Diese Tille enthalten vor allem Kristallingeschiebe, auch viel
Quarzporphyr, aber vergleichsweise wenige und
kleine Karbonatkomponenten (Abb. 3). Die westliche Hälfte des riesigen, fast 10 km breiten Talgletschers, die den Überetsch erreichte, dürfte vor allem
aus Eis bestanden haben, das dem Etschgletsacher
aus dem W des Einzugsgebietes zugeführt worden
ist, vor allem aus dem Val Müstair, Martelltal und
Ultental. Ein Großteil der Geschiebe im Überetsch
dürfte demnach vor allem aus der relativ nahe gelegenen Ortlergruppe stammen. Gelegentlich sind
auch Serpentinit-Komponenten zu finden, die aus
dem Oberengadin stammen und über eine Tansfluenz am Reschenpass ins Etschtal gelangt sein dürften (Ebers 1972: 114).
Die in den Tillen enthaltenen Geschiebe sind gewöhnlich recht gut gerundet, aber nur die Karbonate sind deutlich gekritzt. Lokal dünnen diese Ablagerungen stark aus und bilden einen geringmächtigen Geschiebeschleier, doch sind Aufschlüsse selten,
in denen sich die Mächtigkeit dieser Geschiebelehme ermitteln lässt. Der teilweise ausgezeichnete
Rundungsgrad der Kristallinkomponenten ließe sich
durch die Annahme erklären, dass das Eis ältere fluviatile Kiese im Etschtal und im Überetsch aufgearbeitet haben könnte.
Oft liegen Geschiebelehme dem eisüberschliffenen Felsuntergrund in wechselnder Mächtigkeit direkt auf. Insgesamt sind richtige lodgement tills, die
wohl aus Zeiten mit hoher Eisbedeckung stammen,
weit verbreitet. Geschiebelehme mit einem eindeutig lokalen Geschiebespektrum, also Ablagerungen
von Lokalgletschern des Mendelkammes, waren
nicht zu finden.
2.3 Eisrandablagerungen (Moränenwälle und
Kames)
Schon Penck (in Penck & Brückner 1909: 924)
war am Westhang des Mitter- und Unterberges gegenüber von Kaltern ein großer Moränenwall aufgefallen, der südlich von Girlan beginnt, die Montiggler Seen abdämmt und bis gegen den Kalterer
See hinziehen soll. Nach Penck (in Penck & Brückner
1909: 924) markiert er einen längeren Gletscherhalt. Weniger zusammenhängend sieht er die Moränenwälle an der Westseite von Eppan. Er gibt an,
dass sie sich oberhalb St. Pauls an den Fuß des Buchberges lehnen, bei Planitzing durch das Trümmerwerk
eines Bergsturzes und bei Kaltern durch einen großen
Schuttkegel unterbrochen sind (Penck in Penck &
Brückner 1909: 924 f.). Die Existenz dieser Eisrandablagerungen, Moränenwälle und Kamesterrassen,
konnte durch die Kartierungen tatsächlich bestätigt
werden.
Im E der Talung gibt es am Westhang des Mitterund Unterberges gegenüber von Kaltern nicht nur
einen einzigen großen Moränenwall, sondern ein
ganzes System von kiesigen Lateralmoränen und
Kamesterrassen (Abb. 2, 6), mit deren Hilfe sich
mindestens zwei unterschiedliche Eisstände einer
„Kalterer Zunge“ rekonstruieren lassen, die in der
Talung von Eppan-Kaltern gelegen haben und
knapp südlich des heutigen Kalterer Sees gestirnt
haben muss (Abb. 1, 7). Die am höchsten gelegene
und deutlichste dieser Strukturen ist ein Wall, den
man auf über 1,5 km Länge verfolgen kann. Er hat
ein deutliches Gefälle in südlicher Richtung und
liegt an seinem N-Ende um ca. 60 m höher als an
seinem S-Ende (Taf. 1). Ursprünglich scheint es sich
wohl eher um eine Kamesterrasse gehandelt zu
haben als um einen Wall. Bei sinkendem Eisstand
wurde durch ein sich bergseitig eintiefendes
Schmelzwassertal (Fuschgalai) ein wallartiger
Rücken abgetrennt (Abb. 6). Weiter im S lässt sich
der Eisstand von Fuschgalai mit Kamesterrassen am
Falzig weiterverfolgen, die am Kreithof wieder in
einem deutlichen Wall auslaufen (Abb. 6). Dieses
Wallstück ist inzwischen größtenteils einem Kiesabbau zum Opfer gefallen. Obwohl die in den 60er
Jahren ausgebeutete Grube inzwischen völlig verwachsen ist, lässt sich immer noch erkennen, dass
das Material, aus dem der Wall besteht, stark kiesig
und sehr kristallinreich ist und zahlreiche metergroße Kristallinblöcke enthält. Castiglioni & Trevisan (1973: 6 ff.) rechnen diese groben, auf einer
ihrer Abbildungen erkennbar geschichteten Kiese
freilich zu den glazifluvialen Schottern des „Conglomerato di Caldaro“. Diese Kiese sind aber in unmittelbare Nähe des Eisrandes entstanden, da sie
große Mengen gekritzter Geschiebe enthalten.
Anders als Penck (in Penck & Brückner 1909:
924) glaubt, sind die Wallsysteme in Richtung Montiggler Seen und Girlan nicht weiter zu verfolgen.
Das auf dem Moränenwall abgreifbare Gefälle
spricht eher dafür, dass sich der Eisrand der Kalterer
Zunge in der Zeit des Fuschgalai-Stadiums an den
NE-Hang des Jagenberges und sich südlich des
Großen Priol mit der Etschtalzunge vereinigt hat.
Zwischen dem Wilden-Mann-Bühel und dem
Großen Priol müssen damals mehrere Quarzporphyr-Kuppen das Eis als Nunatakker knapp überragt
haben (Abb. 1, 7). Dieses Stadium könnte zum Eisstand von Auer gehören, der nach Hantke (1983:
234) demjenigen von Kufstein auf der Alpennordseite entsprechen soll. Nach Jerz (1993: 95) entspricht das einem Alter von etwa 15.000 bis 16.000
Jahren vor heute.
Im W der Talung Eppan-Kaltern gibt es, anders
als Penck (in Penck & Brückner 1909: 924) vermutet, kaum Moränenwälle, wohl aber ein System von
breiten Kamesterrassen zwischen Kaltern und St.
Josef am Kalterer See (Taf. 1, 2), die einen Eisstand
nachzeichnen, den wir hier als Stadium von Kaltern
bezeichnen wollen (Abb. 6, 7). Die ursprünglich
wohl zusammenhängenden, bis zu 500 m breiten
Terrassen mit ebenen oder leicht welligen Oberflächen sind durch jüngere, W-E-orientierte Erosionstäler, die dem generellen Gefälle des Hanges folgen, in mehrere Teilstücke zerlegt worden (Abb. 6).
Am Barleitherhof ist ein N-S-orentiertes, wallartiges Teilstück der Kamesterrasse durch ein Erosionstälchen vom bergwärtigen Rest der Terrasse
abgetrennt worden (Abb. 6). Die Zertalung muss
schon unmittelbar nach der Entstehung dieser Terrassen begonnen haben, denn viele der Erosionsrinnen sind Trockentäler. Ein besonders großes Teilstück der Kamesterrassen, auf dem der Ortskern von
Kaltern steht, ist von der Bergseite her durch den
komplexen Schwemmfächer des Pfusser Baches
überschüttet worden (Abb. 6). Penck (in Penck &
Brückner 1909: 924) glaubt die Kamesterrassen in
Richtung Oberplanitzing und Eppan weiterverfolgen zu können, was sich jedoch als unmöglich
herausstellte.
Das Gefälle dieser Eisrandterrassen ist etwas geringer als das des Walles auf der Gegenseite. Sie liegen auch deutlich tiefer und entsprechen von ihrer
Höhenlage her wohl eher den Kamesterrassen an
den Bergflanken unterhalb von Fuschgalai (Abb. 7).
Mit dem Stadium von Fuschgalai der Kalterer Zunge
dürften wohl eher drei kleine Terrassenreste oberhalb des Barleither Weges korrespondieren (Abb. 6).
In den Kamesterrassen gibt es zahlreiche Aufschlüsse, die Einblicke in ihren inneren Aufbau erlauben.
7
Abb. 3: Gegenüberstellung der Texturen genetisch unterschiedlicher Sedimente mit diamiktischer Kornverteilung im Etschtal. Auf
den Bildern sind die wichtigsten im Aufschluss sichtbaren Eigenschaften dieser Sedimente sowie deren genetische Deutung schematisch dargestellt.
Fig. 3: Comparison of genetically different diamictic sediments in the Etsch valley. Textures and some other important macroscopic
visible features of these sediments are shown here, together with their genetic interpretation.
Zum größten Teil bestehen sie aus gut ausgewaschenen, geschichteten Kiesen, die teilweise sehr
grob sind und große Mengen gekritzter Geschiebe
enthalten, also sehr eisrandnah abgelagert worden
sind. Daneben spielen geschichtete Sande und
Schluffe eine wichtige Rolle. Die Kiesgrube vom Voglmeierhof westlich des Kalterer Sees, die bei Castiglioni & Trevisan (1973: Abb. 7) abgebildet ist, zeigt
keine Schotter, die zum glazifluvialen „Conglomerato di Caldaro“ gehören, sondern eisrandnah entstandene Kameskiese, wie sie in allen Kamesterrassen auf der Westseite der Kalterer Zunge akkumuliert worden sind.
Verglichen mit den Kiesen innerhalb des Walles
auf der Ostseite der Talung ist das Material hier
deutlich reicher an Karbonatkomponenten. Stellenweise konnten glazialtektonisch bedingte Schichtstörungen beobachtet werden. Obwohl zahlreiche
gekritzte Geschiebe zu finden sind, treten tillartige
8
Sedimente stark in den Hintergrund. Dafür sind in
die Kamesterrassen stellenweise schluffreiche Sedimente mit lokalem Schutt integriert. In diesen Sedimenten, die als Bestandteile der Kamesterrassen z.B.
am Barleither Weg 500 m NNW‘ des Barleitherhofes
oder im Tal oberhalb von Schloss Kaltenburg aufgeschlossen sind, dominieren eckige Komponenten aus
Schlerndolomit sowie aus Karbonaten, Schluff- und
Sandsteinen der Werfen Formation. Nur ganz
untergeordnet finden sich auch Kristallingerölle. Bei
diesen Sedimenten handelt es sich definitiv nicht
um Lokalmoränen (siehe unten).
2.4 Mursedimente
Weit verbreitet sind im Untersuchungsgebiet Sedimente, deren Habitus auf den ersten Blick an Tille
(„Moränen“) erinnert, die aber von den Komponen-
tenspektren, den Kornformen und den Kornoberflächen her keine glazigenen Sedimente sein können. Diese Sedimente haben eine diamiktische
Korngrößenverteilung (Taf. 4) und sind von daher
Tillen ähnlich (Abb. 3). Es handelt sich um matrixgestützte Sedimente mit einer sandig-schluffigen
Grundmasse, in der zahlreiche grobe Komponenten
schwimmen. Die Korngrößen des Grobmaterials liegen im Bereich von Kies bis Blockwerk; gelegentlich
kommen auch metergroße Blöcke vor. Die groben
Komponenten sind eckig, weisen vielfach scharfe
Bruchkanten auf, doch sind auch kantengerundete
Bruchstücke zu finden. Gut gerundete und/oder gekritzte Komponenten, kristallines Material und andere Fremdgesteine fehlen oder sind zumindest selten. Die Hauptmasse der Komponenten besteht aus
Schlern-, Haupt- bzw. Contrindolomit sowie Bruchstücken der Hartbänke aus der Werfen Formation.
Doch die Zusammensetzung schwankt in weiten
Grenzen. Es gibt Bereiche, in denen diese Gesteine
fast nur aus Schlern- und Contrindolomit-Bruchstücken bestehen, an anderen Stellen nur aus Fragmenten der Werfen Formation, manchmal auch aus
einer Mischung aus beidem. Die Farbe der feinerkörnigen Matrix ist grau, häufig auch rötlich oder
gelblich, letzteres vor allem dort, wo viele Werfener
Komponenten in der Grobfraktion zu finden sind.
Deutliche Schichtungsgefüge sind meist nicht zu
erkennen, selbst dann nicht, wenn man meterhohe
Aufschlüsse begutachten kann. Selten kommen
aber doch Lagen mit deutlich weniger Grobmaterial
oder schluffige, sandige oder kiesige Einschaltungen vor.
Im Aufschluss sind diese Gesteine überraschend
standfest; fast vertikale Straßen- und Wegan schnitte erweisen sich seit Jahrzehnten ohne Sicherungsmaßnamen als standfest (Taf. 4). Diese Gesteine finden sich im Untergrund vieler Weinberge zwischen Kaltern und Margreid. Die steinigen Sedimente sind auf den Feldern nur schwer zu bearbeiten, so dass sie die Weinbauern als „Kampferde“
oder „Kampf“ bezeichnen, ein Ausdruck, der anderenorts in Südtirol auch für lodgement-till („Grundmoräne“) verwendet wird (mündl. Mitt. Dr. Volkmar
Mair, Bozen). Die Sedimente bilden oft mächtige
Decken über dem Felsuntergrund, deren basale
Auflagerungsflächen oft geneigt sind und parallel
zum Hang einfallen. Mitunter kommen sogar fast
vertikale Kontaktflächen an Stellen vor, wo die Sedimente offensichtlich alten, verschütteten Felsstufen angelagert sind. Die Mächtigkeiten sind meist
nur schwer abschätzbar. Oft lassen sich aufgrund
der Tiefe von Erosionstälern Mächtigkeiten von
mehreren Dekametern schätzen; in Einzelfällen
kommt man auf 60 bis 80 m.
Im Überetsch sind Sedimente dieses Typs weit
verbreitet (Abb. 1). Als fast geschlossene Decken
von erheblicher Mächtigkeit treten diese Ablagerungen an den Hängen oberhalb von Kurtatsch,
Entiklar und Margreid auf, wo sie bis über Penon
hinauf die tonig-kalkigen „Zwischenschichten“ des
Unterladin zusammen mit ihren mächtigen Kalkund Dolomiteinschaltungen überlagern. Nur in besonders tief eingeschnittenen Erosionstälern wird
hier das Quartär durchschnitten. Hier bilden diese
Ablagerungen eine fast geschlossene Decke mit
einer Gesamtfläche von fast 5 km2. Weiter im N sind
diese Sedimente weniger geschlossen verbreitet,
nehmen jeweils kleinere Flächen von immerhin
noch vielen Hektar Größe ein. Auch hier können die
Vorkommen mehrere Dekameter mächtig werden.
Auffällig ist, dass die Verteilung der Vorkommen
eine klare Beziehung zu den bedeutenderen, tief
eingeschnittenen Rinnen zeigen, die zum Mendelkamm hinaufziehen. Ein besonders mächtiges Vorkommen dieser Sedimente bildet z.B. die markante
Kuppe am Ausgang des Höllentales in Tramin, auf
der St. Jakob in Kastellaz liegt (Abb. 1). Ein anderes
Vorkommen ist beispielsweise an der Straße von
Kaltern nach Altenburg aufgeschlossen, genau unterhalb des tief eingeschnittenen Val della Lavine.
Manche dieser merkwürdigen Sedimente zeigen
eindeutige Beziehungen zu jungen Oberflächenformen. „Kampferde“-Sedimente, die z.B. NW’
Penon, zwischen Altenburg und Kaltern oder oberhalb von Pfuss bei St. Nikolaus in Kaltern vorkommen (Taf. 3), bauen jeweils mehrere parallel orientierte, schmale Rücken auf, die von tief eingeschnittenen Erosionstälern voneinander getrennt
werden. Die Oberflächen benachbarter Rücken weisen ein identisches Gefälle von 15 bis 30° auf (Taf.
3). Talwärts sind diese Rücken durch einen Gefälleknick begrenzt; unterhalb davon hören die Rücken
mit einer kräftigen Versteilung des Hanges auf (Taf.
3). Dieser Gefälleknick liegt bei benachbarten
Rücken ungefähr auf der gleichen Höhe; die Strukturen erscheinen dadurch wie abgehackt. Bei diesen
Rücken könnte es sich um Erosionsreste von fächerartigen Gebilden zu handeln, wohl um die Reste
alter Murfächer, die von parallel orientierten Tälern
zerschnitten worden sind (Abb. 6). Aufgrund günstiger Aufschlussverhältnisse am anerodierten Mur-
9
Abb. 4: Schema der Genese von „Murkames“ am Westrand des
Etschgletschers. Die Murkames entstanden als Murfächer und
enthalten ausschließlich Material, das aus Erosionsrinnen im
Hang gegen den Rand des Etschgletschers vorgeschüttet
wurde (unten). Nach dem Abschmelzen des Eises wurden die
Murkames, die talwärts primär durch eine steile Sackungskante begrenzt sind, erosiv zerschnitten (oben).
Fig. 4: Simplified sketch showing how a "fankame“ at the western rim of the Etsch valley glacier may have been formed.
Originally they have been generated as alluvial fans by accumulation of debris flows at the glacier rim, the debris deriving
entirely from the hillslope above (below). These "fankame“ expose a typical steep edge at their lower part and have been cut
by erosional valleys since the glacier ice has vanished (above).
fächer von Pfuss ist zu erkennen, dass die Hauptmasse der Höhenrücken tatsächlich aus Ablagerungen dieses Typs aufgebaut wird. Schon Penck (in
Penck & Brückner 1909: 924) hat diese Vorkommen
bei St. Nikolaus in Kaltern gekannt, in ähnlicher
Weise als „Schuttkegelrudimente“ gedeutet und sie
ins „Spätglazial“ gestellt. Am Fuß der Versteilungen
unterhalb des Gefälleknicks scheinen die Mursedimente durch eine Zunahme des Kristallinmaterials,
des Rundungsgrades der Komponenten und dem
vermehrten Auftreten gekritzter Geschiebe in kar-
10
bonatreiche Geschiebelehme überzugehen, was die
unmittelbare Nähe des Eises am talwärtigen Ende
der Strukturen anzeigt.
Hier besteht also der begründete Verdacht, dass
es sich um Murfächer handelt, die gegen den Eisrand des zurückschmelzenden Etschgletschers geschüttet worden sind; wir wollen sie hier „Murkames“ nennen (Abb. 1, 6). Neben diesen „Murkames“ gibt es auch, wie oben schon dargelegt, gewöhnliche Kamesterrassen mit ebenen Oberflächen,
die außer kiesigen oder schluffig-sandigen, gut geschichteten Schmelzwassersedimenten auch abschittsweise „Kampferde“-Sedimente enthalten.
Solche Kamesterrassen sind z.B. NE‘ von Penon oder
südlich von Kaltern am Barleiter Weg zu finden.
Die meisten Vorkommen von Sedimenten dieses
Typs lassen indes keinerlei Beziehungen zu irgendwelchen charakteristischen Oberflächenformen erkennen. An einigen Stellen ist zu beobachten, dass
derartige Ablagerungen eindeutig von kristallinreichen Geschiebelehmen überlagert werden. Das ist
z.B. an Ablagerungen im Hügel von St. Jakob in Kastellaz in Tramin ganz in der Nähe des Bungalows
der Wildbachverbauung zu sehen. Dieses und einige
andere Vorkommen scheinen zudem rundliche,
drumlinähnliche Geländeformen zu bilden und sollten folglich vom Gletschereis überfahren worden
sein. Deshalb muss zumindest ein Teil dieser Sedimente vor dem Höchststand des Eises der letzten
Eiszeit entstanden sein. Ähnlich sieht das auch
Penck (in Penck & Brückner 1909: 921). Er argumentiert, dass sie zeitlich zwischen zwei
aufeinanderfolgende Vergletscherungen zu stellen
wären, da sie gelegentlich auch (umgelagerte)
Fremdgeschiebe enthalten. Auch bei Meran hat
Penck (in Penck & Brückner 1909: 921) solche
Schuttablagerungen gefunden, zwischen Gardasee
und Meran will er gar Reste von vier verschieden
alten Schuttkegeln nachgewiesen haben.
Dafür, dass es sich bei den „Kampferde“-Sedimenten um Ablagerungen von debris flows handelt,
spricht vor allem die praktisch fehlende Rundung
der Komponenten und die äußerst schlechte Sortierung des Materials (Johnson & Rodine 1984: 315).
Warum sind die „Kampferde“-Ablagerungen, wenn
man sie als Mursedimente deutet, kaum oder gar
nicht geschichtet, obwohl postglaziale mudflowSedimente, genauso wie rezente Murkegel, immer
eine wenn auch undeutliche Schichtung aufweisen
(Costa 1984, 1988, Davies 1988)? Der typische Aufbau junger Mursedimente kann beispielsweise im
Nussental am Hang oberhalb Kuenburg am Kalterer
See studiert werden, wo ein steiler Murkegel durch
eine kleine Grube angeschnitten ist. Das hier
aufgeschlossene diamiktische Material, sehr reich
an eckigen Quarzporphyr-Komponenten, ist undeutlich geschichtet. Der geschichtete Eindruck
wird durch einen Wechsel in der Korngröße und in
der Zusammensetzung der Mursedimente erzeugt,
wie sie für Ablagerungen typisch sind, die von debris flows aufgebaut werden (Coussot & Meunier
1996).
Vielleicht hängen die Unterschiede zu den fossilen Mursedimenten damit zusammen, dass die
heute noch aktiven, mehrere Dekameter mächtigen
Murkegel im Laufe von vielen einzelnen Murereignissen akkumuliert worden sind. Bei jedem Murgang werden hier jeweils nur wenige Meter Sediment auf einmal abgelagert, da sich die Mure über
einen Teil des Fächers flächenhaft ausbreiten kann.
Gleiches gilt auch für die rezenten Beispiele, die bei
Johnson & Rodine (1984: 266 ff.) angeführt werden. Die viele Dekameter mächtigen „Kampferde“Sedimente sind im Gegensatz dazu wohl alle kaltzeitlich und bei sinkenden Eisständen abgelagert
worden. In den Kaltzeiten gab es auf den frisch vom
Eis freigegebenen Steilhängen, wo das Lockermaterial für die Muren mobilisiert werden konnte, keine
Vegetation, die den hier liegenden Hangschutt und
Geschiebelehme hätte stabilisieren können, und
auch der sich nach dem Eisrückzug aufbauende Permafrost dürfte bald in der ausgehenden Eiszeit zusammengebrochen sein (Haeberli 1996). Dadurch ist
bei einem einzelnen Ereignis offenbar ungleich
mehr Material umgelagert worden als heute. Noch
dazu konnten sich die Muren auf den Fächern nicht
ausbreiten sondern stauten sich am Eisrand (Abb. 4),
was schon bei einem einzigen Ereignis zur Akkumulation von dekametermächtigen, intern weitgehend
ungeschichteten Mursedimenten führte (Abb. 5).
2.5 Blockgletscher, Lokalgletscher und
Gehängebrekzien
Seit dem Abschmelzen der Gletscher haben sich
vor allem unter den Dolomit-Steilwänden bedeutende Hangschuttmassen akkumuliert. Große
Schuttmassen haben sich vor allem im oberen Teil
einer mehr als 1 km breiten Hangverflachung gebildet, die oberhalb von Kurtatsch zum Tal hin durch
eine markante Geländestufe aus Contrindolomit be-
Abb. 5: Schematische Schnitte durch moderne Murfächer und
„Murkames“, die während des Rückschmelzens des Etschgletschers entstanden sind. Durch den Rückstau am Rande des
Talgletschers waren die Sedimente, die ein einziger Murgang
bzw. ein einzelnes Murereignis hinterließ, bedeutend mächtiger (unten) als in heutigen Murfächern (oben). Dadurch erscheinen die Schichtfolgen in „Murkames“ weitgehend ungeschichtet.
Fig. 5: Schematic cuts through a modern fan in comparison to
a late Pleistocene "fankame“, which was generated when the
Etsch valley glacier retreated. Due to the damming effect of
the glacier rim, the sediment succession from a single debris
flow is much thicker within a "fankame“ (below) than in a recent alluvial fan (above). Therefore the successions within
"fankame“ are poorly stratified.
grenzt wird. Diese Hangverflachung, auf der auch
der Ort Graun liegt, ist letztlich durch die hier vorkommenden kalkig-mergelig „Zwischenschichten“
bedingt, die besonders leicht erodiert werden konnten. Etwa 1 km nördlich von Graun, im Oberen Gemeindewald westlich des Hofes Locherer, liegt eine
nach drei Seiten steil abfallende, einige hundert
Meter breite Hangnase, deren Oberfläche ein auffällig unruhiges Relief trägt. Das dicht bewaldete
Gelände, dessen höchster Punkt 1018 m hoch liegt,
zeigt ein kompliziertes System von Wällen mit tiefen, abflusslosen Depressionen dazwischen, die an
Toteislöcher erinnern. Ein Teil der wallartigen
Rücken scheint sich zu zungenartigen Loben zusammenzuschließen. Das Gebiet, das hangaufwärts
in die Schutthalden unter den SchlerndolomitWänden übergeht, besteht selbst ausschließlich aus
hoch porösem Dolomitschutt. Fremdmaterial und
gerundete Komponenten fehlen praktisch völlig. Ein
etwas kleineres und ca. 50 Höhenmeter tiefer liegendes Areal mit morphologisch vergleichbaren
Strukturen wird vom Traminer Höhenweg etwa 1
km weiter im N gequert.
Bei beiden Strukturen dürfte es sich um Blockgletscher handeln, also ehemals gefrorene Schutt-
11
Abb. 6: Geomorphologisches Übersichtskärtchen des Gebietes zwischen Kalterer See und Oberplanitzing im Überetsch. Die Karte
wurde auf der Grundlage von geologisch-geomorphologischen Detailkartierungen im Maßstab 1:10 000 im Gebiet zwischen Eppan
und Margreid erstellt.
Fig. 6: Simplified geomorphological map showing the region between Kalterer See (Lago Caldaro) and Oberplanitzing (Pianizza di
sopra) at Überetsch (Oltradige, Sella di Appiano-Caldaro). The map was created on base of detailed geological and geomorphological
mapping in the region between Eppan (Appiano) and Margreid (Magrè) at a scale of 1:10 000.
12
massen, die sich kriechend wie ein Gletscher bewegen (Abb. 1). Diese Blockgletscher sind fossil und
bewegen sich heute mit Sicherheit nicht mehr aktiv,
denn in Höhen um 1000 m ist in den Südalpen
unter den heutigen Klimabedingungen (Weinbau
bis in über 800 m Höhe!) mit Sicherheit kein Permafrost mehr zu erwarten. Sie dürften sich nach
dem Rückschmelzen des Etschgletschers an der
Wende vom Hoch- zum Spätglazial gebildet haben,
vor allem während der spätglazialen Klimadepressionen. Blockgletscher ,aber auch richtige kleine Lokalgletscher, die sich gleichzeitig in Karen unterhalb
des Mendelkammes gebildet haben könnten, sind
denkbare Auslöser für große Murgänge, die für die
Genese der oben beschriebenen pleistozänen Mursedimente verantwortlich waren.
Am Nordhang des Höllentales oberhalb von Tramin liegt ein auffälliger Hangvorsprung, der durch
das Vorkommen einer calcitisch zementierten, hoch
porösen quartären Brekzie bedingt ist. Diese weitgehend ungeschichtete Gehängebrekzie, die fast
ausschließlich aus eckigem Dolomitschutt besteht,
lagert der Werfen Formation in einer Mächtigkeit
von mindestens 10 m auf, in die die Höllentalschlucht eingeschnitten ist. Über das genaue Alter
der Brekzie lässt sich nichts aussagen, doch weisen
Erosion sowie starke Zementierung des Vorkommens darauf hin, dass es sich möglicherweise um
präwürmglaziale Bildungen handelt. Weitere Vorkommen von ähnlichen Gehängebrekzien sind auch
nahe dem Hof Steiner am Hang oberhalb des Höllentales gegenüber von Tramin oder westlich von St.
Nikolaus bei Kaltern zu finden. Stacul (1980) stellt
die Bildung des Karbonatschuttes, aus dem die
Gehängebrekzie von St. Nikolaus besteht, in eine
Kaltzeit, unmittelbar nach dem Rückschmelzen des
Etschgletschers. Ihre Verkittung durch „Kalksinter“
soll hingegen in einem Interglazial oder einem Interstadial erfolgt sein.
2.6 Kalterer Schotter
Nach Hantke (1983: 233) ist die weite Talung von
Eppan-Kaltern mit mächtigen quartären Kiesen erfüllt, die ihrerseits von würmeiszeitlichen Geschiebelehmen bedeckt sein sollen. Die Gesamtmächtigkeit der Schotter von Eppan beträgt nach Blaas
(1892) bis zu 200 m. Die Schotter werden dem
„Konglomerat von Kaltern“ gleichgesetzt, obwohl
sie größtenteils nicht verfestigt sind. Nach Ebers
(1972) sind die „Überetscher Schotter“ nicht älter
als Eem. Nach Castiglioni & Trevisan (1973) ist das
„Conglomerato di Caldaro“ von Schmelzwässern des
vorstoßenden Etschgletschers aufgeschüttet worden. Seine Aufschüttung soll im Val-Caldaro-Interstadial erfolgt sein, das mit einem radiometrisch ermittelten Alter von rund 30.000 Jahren (Fuchs
1969) dem Interstadial von Baumkirchen in den
Nordalpen entsprechen könnte. Auch Klebelsberg
(1926, 1935) und Ebers (1972) gehen davon aus,
dass alle größeren Kiesvorkommen im Überetsch
genetisch identisch sind, eine einheitliche Bedeckung von Geschiebelehmen aufweisen und deshalb vor dem Gletscherhöchststand der Würmeiszeit entstanden sind.
So einfach ist die Sache allerdings nicht. Ebers
(1972) und Castiglioni & Trevisan (1973) subsummieren unter den Begriffen „Überetscher Schotter“
und „Conglomerato di Caldaro“ viele Kiese, die hier
zu unterschiedlichen Zeiten und unter ganz unterschiedlichen Bedingungen entstanden sind. Castiglioni & Trevisan (1973) stellen beispielsweise die
groben Kiese zum „Conglomerato di Caldaro“, die
früher am Kreithof („Maso Kreit“) westlich des Kalterer Sees in einer Kiesgrube abgebaut worden sind
(Castiglioni & Trevisan (1973: 6 ff.). Diese Kiese sind
aber Teil eines komplexen Systems von Kamesterrassen und Wällen auf der Ostseite der Kalterer
Zunge (siehe oben). Auch die westlich des Kalterer
Sees gelegenen Kiese vom Vogelmeierhof (Castiglioni & Trevisan (1973: 6 ff.) gehören zu einem System
von komplexen Kamesterrassen, die auf der Westseite der Kalterer Zunge im ausgehenden Hochglazial der Würmeiszeit akkumuliert worden sind. Daneben sind aber auch tatsächlich eindeutig prähochglaziale Bildungen zu finden.
Tatsächlich ist die weite Talung von Eppan-Kaltern von kristallinreichen, teilweise sehr grobkörnigen, abschnittsweise kaum geschichteten und oft
schluffreichen Kiesen erfüllt, die größtenteils sehr
schlecht aufgeschlossen sind. In den hangenden
Abschnitten der Kiese sind gekritzte Geschiebe häufig; fleckenweise tragen sie sogar eine Decke von
Geschiebelehmen; östlich von Kaltern sind im Hangenden dieser Kiese sogar wallähnliche Strukturen
entwickelt. Da die Karbonat- und Kristallinkomponenten dieser Kiese kaum Verwitterungserscheinungen zeigen, dürften sie vergleichsweise jung sein.
Womöglich handelt es sich wenigstens teilweise um
Vorstoßschotter, vor allem in der Umgebung der
Montiggler Seen, wo die Oberfläche kiesiger Abla-
13
gerungen drumlinisiert ist. Vielfach dürfte es sich
aber wohl auch um Schmelzwasserschotter aus der
ausgehenden Eiszeit handeln, die vor der zurückschmelzenden Kalterer Zunge akkumulierten und
bei einer Eisoszillation nochmals überfahren wurden. Sie könnten in einem Totraum abgelagert worden sein, der sich zwischen der nach Norden
zurückschmelzenden Kalterer Zunge und dem
Becken des Kalterer Sees befand (Abb. 6).
Die Kiese sind gewöhnlich locker und nicht oder
kaum verfestigt und enthalten immer wieder Einschaltungen von sandig-schluffigen Laminiten, bei
denen es sich um Stillwasserablagerungen handelt.
Nur in der kleinen Schlucht zwischen Festplatz und
Kalvarienberg in Kaltern, über die der Bach aus dem
Tröpfeltal das Lavasontal erreicht, kommen auf der
orographisch linken Talseite durch calcitische Zemente fest verbackene, kristallinreiche Konglomerate heraus. Diese mit Höhlen und Kavernen durchsetzten Ablagerungen sind wohl das „Konglomerat
von Kaltern“ im ursprünglichen Sinne. Es handelt
sich um gut sortierte, ausgewaschene Schmelzwassersedimente, die zahlreiche Rollkieslagen enthalten. Die Imbrication der Gerölle weist auf einen
generellen Sedimenttransport von N hin. Deutliche
Verwitterungserscheinungen an den Dolomitkomponenten des Konglomerates lassen Zweifel aufkommen, ob es mit den weit verbreiteten Kiesen der
Umgebung etwas zu tun hat oder ob es nicht doch
älter ist.
Die fraglichen Vorstoßschotter und die Konglomerate sind jedenfalls in der Talung Eppan-Kaltern
nur bis zu einer Linie flächenhaft verbreitet, die von
der Kirche von Kaltern nach Montiggl zieht. Weiter
im S sind diese und vielleicht auch jüngere Ablagerungen teilweise ausgeräumt und durch ein System
von Kiesterrassen ersetzt, die keine Bedeckung von
Geschiebelehmen tragen und während des Rückschmelzens der Kalterer Zunge entstanden sein
müssen. Es lassen sich hier zumindest drei unterschiedliche Terrassenniveaus auskartieren und einerseits miteinander, andererseits aber auch mit
einem System von Trockentälern in Beziehung bringen, aus denen diese Kiese offenbar zu unterschiedlichen Zeiten herausgeschüttet worden sind
(Abb. 6). Es gibt auch eine deutliche Beziehung dieser Terrassen mit dem Kalterer See: Je höher diese
Terrassen liegen, desto weiter liegen sie vom nördlichen Seeufer entfernt. Die niedrigsten (und vermutlich jüngsten) Terrassen liegen dem See am
nächsten (Abb. 6).
14
2.7 Trockentäler
Die gesamte Talung von Eppan-Kaltern wird von
einem ganzen System von tief eingeschnittenen,
breiten, kastenförmigen Trockentälern durchzogen
(Abb. 6). Abschnittsweise werden die Trockentäler
auch von heutigen Gewässern benutzt, die die alten
Talböden teilweise durch Schwemmfächer verschüttet, in einigen Fällen auch ältere Talgenerationen
anerodiert und zerstört haben.
Die Trockentäler bilden ein mehrfach verzweigtes
Talsystem, dessen Talachsen größtenteils N-S oder
NE-SW-orientiert sind. Das größte und am wenigsten von jüngeren Schwemmfächern aufgefüllte
Trockental, das Lavasontal, lässt sich von den Reitwiesen am Kalterer See über 6 km nach N verfolgen
(Abb. 1, 6). Mehrfach zweigen seitlich einmündende
Trockentäler in nordöstlicher Richtung davon ab
(Abb. 6), deren Talböden teilweise vom Haupttal unterschnitten sind. Nördlich des Feldhofes zweigt ein
breites Tal in NNW’ Richtung vom Lavasontal ab, das
durch junge Schwemmfächer teilweise stark aufgefüllt und dadurch undeutlich geworden ist. Dieses
Tal lässt sich über den alten Bahnhof von Kaltern
hinaus nach N verfolgen, wo es sich in mehrere Rinnen aufspaltet. Diese Verzweigung des Trockentales
ist teilweise durch die dichte Bebauung, teilweise
aber auch wegen der Erosion durch den Bach aus
dem Tröpfeltal undeutlich geworden. Die am weitesten nach N verfolgbare Rinne dieses Systems ist
diejenige, die von Kaltern nach Oberplanitzing
zieht, das Oberplanitzinger Trockental (Abb. 6).
Folgt man den Tälern aufwärts, steigen sie mit
meist gleich bleibendem Gefälle an, werden undeutlich und streichen schließlich in die Luft aus,
was für Schmelzwassertäler typisch ist. Wenn diese
Rinnen abschnittsweise von modernen Gewässern
verwendet werden, fließen diese von der Seite zu;
die Quellen liegen niemals am Beginn der Rinnen.
Besonders schön ist das am schluchtartig eingeschnittenen Oberplanitzinger Trockental zu sehen,
das im Dorfzentrum von Oberplanitzing plötzlich
undeutlich wird und verschwindet. Auch im N des
Lavasontales ist das undeutlich Werden und Verschwinden der Rinne sehr gut zu beobachten.
Die jüngste Terrasse läuft nach S hin, an den
Reitwiesen, auf Seeniveau aus, setzt sich aber nach
N hin ins weithin trockene Lavasontal fort, das sich
erst 6 km weiter im N bei St. Michael verliert. Die
Trockentäler, die auf die älteste der drei Terrassen
auslaufen, Frühlingstalele und Val Fusca, lassen sich
kaum mehr als 1 km nach N verfolgen. Die dazwischen liegende Terrasse korrespondiert mit dem
Fondatal und anderen Trockentälern, die weiter im
N enden aber nicht so weit zu verfolgen sind, wie
das Lavasontal (Abb. 6). Um die Gesetzmäßigkeit
noch mal auf den Punkt zu bringen: je älter die
Täler sind, desto weniger weit reichen sie nach N,
desto höher lag offensichtlich auch der Vorfluter im
Bereich des Kalterer Sees. Das zuletzt aktive Tal, das
Lavasontal, erhielt sein Wasser auch so weit von N
wie kein anderes, der Vorfluter, der das Wasser aufnahm, war damals schon fast so tief wie der Kalterer See.
Penck (in Penck & Brückner 1909: 924) nimmt
an, dass der Überlauf eines Stausees bei St. Pauls
über ein „heute trocken daliegendes Tal, das sich
östlich von Kaltern zum Kalterer See zieht“ erfolgt
sein soll, also wohl über das Lavasontal. Bei Kaltern
soll dieser Ausfluss nach Castiglioni & Trevisan
(1973: Abb. 26) in einen weiteren, etwas niedriger
liegenden Stausee gemündet haben, der südlich des
Kalterer Sees vom Etschgletscher abgedämmt worden sein soll, also immer noch deutlich höher gelegen haben muss, als der heutige Seespiegel. Das
kann aber nicht sein, wie oben ausführlich dargelegt wurde. Zudem kann diese Annahme nur die
Entstehung eines der Trockentäler erklären, für alle
anderen bleibt sie eine Deutung schuldig.
Viel plausibler ließen sich sämtliche Beobachtungen interpretieren, wenn man annimmt, dass die
Bildung aller Trockentäler und die Entstehung des
Terrassensystems am Kalterer See im Zuge des Rückschmelzens der Kalterer Zunge entstanden sind. Bei
den Trockentälern würde es sich demnach um ein
System peripherer und terminaler Rinnen handeln,
über die die Schmelzwässer der zurückschmelzenden Kalterer Zunge abgeflossen sind (Abb. 8). Mit
dem Rückschmelzen waren immer neue Täler in
Funktion, während andere trocken fielen. Mit dem
weiteren Rückzug der Zunge nach N, in Richtung St.
Michael, war zuletzt nur noch das tiefst gelegene
und die Achse der Talung nachzeichnende Lavasontal in Funktion. Als die Gletscherzunge schließlich
über den Sattel bei St. Michael zurückgeschmolzen
war, suchten sich die Schmelzwässer neue Wege
und erreichten den Kalterer See nicht mehr (Abb. 8).
Wie groß war die Menge des hier erodierten Materials? Das hängt unmittelbar mit der Frage nach
der Dimension dieser Erosionstäler zusammen. Das
Lavasontal ist über 6 km lang, auf 5 km Länge ist es
um 50 bis 75 m tief in die Umgebung eingeschnit-
ten, mit einer Breite des ebenen Talbodens zwischen
50 und 110 m. Man kann abschätzen, dass alleine in
dieser Rinne mindestens 50 Mill. m3 erodiert und
nach S verfrachtet worden sind. Angesichts der
Größe der Erosionstäler und der Menge des in den
tief eingeschnittenen Tälern erodierten Materials ist
es eigentlich unverständlich, dass der kleine Kalterer
See nicht schon während des Eisrückzuges zugefüllt
worden ist. Das Material, das in allen Rinnen zusammen erodiert worden ist, dürfte ausreichen, um
einen See, der um ein Vielfaches größer ist als der
Kalterer See, restlos aufzufüllen. Dabei ist noch
nicht einmal berücksichtigt, dass die erodierenden
Schmelzwässer sicher nicht nur das in den Tälern
erodierte, „alte“ Material transportiert haben, sondern sicher auch vom Eisrand her mit „frischem“
Kies, Sand und Schluff überfrachtet waren.
Um erklären zu können, warum das Becken des
Kalterer Sees trotzdem nicht aufgefüllt worden ist,
benötigt man eine weitere plausible Annahme: Das
Seebecken könnte durch eine im Seebecken liegende große Toteismasse, einem abgetrennten Teil
der zurückschmelzenden Kalterer Zunge, solange
vor dem Sedimenteintrag geschützt worden sein,
bis es nicht mehr durch Schmelzwasser erreicht
werden konnte (Abb. 8). Ursprünglich könnte diese
Toteismasse auch die weite Senke nördlich des heutigen Sees ausgefüllt haben. Die Annahme einer
solchen langsam abschmelzenden und immer kleiner werdenden Toteismasse würde auch zwanglos
erklären, warum der Vorfluter sich ständig abgesenkt hat (Abb. 8/ 3-5). Bei dieser Annahme hätten
die Schmelzwässer einen Teil der mittransportierten
Grobstoffe seitlich um die Toteismasse herum
führen und im Etschtal selbst ablagern müssen. Das
aber sollte sich durch entsprechende Bohrungen
nachweisen lassen.
2.8 Seesedimente
An einigen Stellen zwischen Eppan und Kaltern
treten geschichtete, sandig-schluffige Ablagerungen auf, die von Penck (in Penck & Brückner 1909:
924) als „glaziale Mehlsande“ von St. Pauls bezeichnet wurden. Sie bedecken vor allem den Nordteil
des Überetsch, zwischen Unterrain, Frangart und St.
Pauls und überlagern hier ältere quartäre Ablagerungen bzw. Gesteine der Permotrias. Nach Penck
(in Penck & Brückner 1909: 924) wurden diese stellenweise viele Dekameter mächtigen Sedimente in
15
einem vom Eis aufgestauten See abgelagert. Gleiches gilt auch für ähnliche Bildungen, die sich östlich des Kreither Sattels beiderseits der Laimburg
oberhalb des Etschtales (am Stadlhof) erhalten geblieben sind, ein Vorkommen, das von Castiglioni &
Trevisan (1973: 19 f.) als das von „Novale al Varco“
oder „Maso Stadio“ bezeichnet wird. Ausführlich
werden diese und die glazilakustrinen Sedimente
von St. Pauls durch Castiglioni & Trevisan (1973: 18
ff.) beschrieben. Obwohl die Ablagerungen stellenweise durch Eisauflast etwas verdichtet und durch
das Eis glazialtektonisch teilweise gestört erscheinen, müssen sie nach Castiglioni & Trevisan (1973:
19) ins Spätglazial, also genauer ins Bühl-Stadium
gestellt werden (Hantke 1983: 234).
Der Überlauf des Stausees bei St. Pauls soll nach
Penck (in Penck & Brückner 1909: 924) über ein
„heute trocken daliegendes Tal, das sich östlich von
Kaltern zum Kalterer See zieht“ erfolgt sein, also
wohl über das Lavasontal. Bei Kaltern soll dieser
Ausfluss nach Castiglioni & Trevisan (1973, Abb. 26)
in einen weiteren, etwas niedriger liegenden Stausee gemündet haben, der südlich des Kalterer Sees
vom Etschgletscher abgedämmt worden sein soll.
Wie oben schon dargelegt wurde, ist das Lavasontal
eher als normales Schmelzwassertal angelegt worden und hat, selbst wenn es später als Überlauf für
einen solchen Schmelzwassersee gedient haben
sollte, jedenfalls nicht in einen größeren Schmelzwasserstausee im S des Überetsch gemündet. In der
Umgebung des Kalterer Sees gibt es, abgesehen von
den Stauseesedimenten östlich des Kreither Sattels,
keine See- oder Deltaablagerungen, die die Annahme eines solchen Sees rechtfertigen würden.
Tatsächlich gibt es Hinweise auf einen Stausee im
Becken des Kalterer Sees, der aber deutlich älter
sein muss und eher mit dem frühwürmeiszeitlichen
Eisaufbau des Etschgletschers als mit dessen Rückschmelzen im Spätglazial etwas zu tun hat. Beim
Hotel Leuchtenburg in Kreit am Kalterer See sind
oberhalb der Straße Aufschlüsse in schluffig-feinsandigen, feinschichtigen, etwas eisenschüssigen
Stillwassersedimenten zu finden, die von kaltzeitlichen, sehr eisrandnah entstandenen, groben Schottern überlagert werden. Die feinkörnigen Sedimente
sind überkonsolidiert und deshalb mit Sicherheit
eisüberfahren. Bei den überlagernden Schottern
könnte es sich um Vorstoßschotter handeln, vielleicht sind es aber auch Kiese, die zu den Eisrandablagerungen von Fuschgalai gehören und somit als
spätglazial einzustufen sind. Stellenweise sind in
16
diesen Seeablagerungen schlecht erhaltene Pflanzenreste zu finden, offenbar Abdrücke von Stengeln, Zweigen und Blättern. Das Einschwemmen
von Pflanzenresten in glaziale Stauseen erscheint
im Zuge des Eisaufbaues eher vorstellbar als
während des Rückschmelzens der Gletscher. Vergleichbare Seeablagerungen wurden übrigens auch
in einem künstlichen Aufschluss oberhalb eines Erosionstales am Westhang des Lavasontales bei Kaltern beobachtet.
3. Rückschmelzen der Kalterer Zunge –
ein Rekonstruktionsversuch
Der hier vorgestellte Rekonstruktionsversuch des
„Eisrückzuges“ in der Umgebung von Kaltern
(Abb. 8) wurde auf der Grundlage von geologischen
Detailkarten erarbeitet, die bei den drei vom Erstautor betreuten Kartierungsübungen mit Geologiestudenten der TU München in Südtirol entstanden
waren (siehe oben). Die hier dargestellten Rückzugsstände (Abb. 8/ 1-5) sind wohl mit dem Eisstand von Auer parallelisierbar, der nach Hantke
(1983: 234) demjenigen von Kufstein auf der Alpennordseite gleichzusetzen sein soll. Nach Jerz
(1993: 95) entspricht dies einem Alter von etwa
15.000 bis 16.000 Jahren vor heute. Das Rückschmelzen der Zunge von Kaltern muss also insgesamt im ausgehenden Hochglazial bzw. an der
Wende zum Spätglazial der Würmeiszeit erfolgt
sein. Was man zur Bestätigung der Annahmen und
zur Abrundung des Bildes allerdings noch bräuchte,
ist die Auswertung von hinreichend tiefen Bohrungen in der Talebene südlich des Kalterer Sees.
1. Die Stirn des Etschgletschers ist im Haupttal bis
etwa nach Auer zurückgeschmolzen. Ein Seitenast,
die Kalterer Zunge, bedeckt große Teile des Überetsch, die weite Talung von Eppan-Kaltern und
stirnt etwas südlich des Kalterer Sees. Das Etschtal
ist teilweise von Schmelzwasserseen erfüllt. Die
Kalterer Zunge wird von Eis genährt, das über
Transfluenzen von N her bei Eppan und von NE
her über die Montiggler Seen vom Hauptgletscher
her überquillt (Abb. 7). In dieser Zeit entstehen die
höchsten Kamesterrassen an der Barleit südlich
von Kaltern (Abb. 6) und die Lateralmoräne von
Fuschgalai am Westhang des Unterberges gegenüber von Kaltern (Fuschgalai-Stadium, Abb. 6).
Abb. 7: Rekonstruktionsversuch des Etschtales zwischen Bozen
und Neumarkt im ausgehenden Hochglazial der letzten Eiszeit.
Deutlich ist zu erkennen, dass sich das Eis des Etschgletschers
in zwei Eisloben aufgespaltet. Die Etschtalzunge (ETZ) im E
folgt dem eigentlichen Etschtal abwärts, die Kalterer Zunge
(KLZ) im W dringt bei Missian ins Überetsch ein, folgt der weiten Talung von Eppan-Kaltern und stirnt südlich des Kalterer
Sees (punktierte Linie). Für die Kalterer Zunge lassen sich zwei
Eisstände besonders gut dokumentieren: ein älteres Fuschgalai-Stadium (dick) und ein jüngeres Stadium von Kaltern
(dünn). Unterhalb des Überetsch war das Etschtal in dieser Zeit
vermutlich von rasch verlandenden Schmelzwasserseen erfüllt
(schwarz).
Fig. 7: Attempt to reconstruct the situation within the Etsch
(Adige) Valley between Bozen (Bolzano) and Neumarkt (Egna)
at the transition from the Pleniglacial to Late Glacial Period.
Two separate glacierlobes at the front of the Etsch valley glacier are clearly visible. The Etsch valley lobe (ETZ) to the east
flows down the Etsch Valley, the ice front of the Kaltern lobe
(KLZ) in the west invading the vale of Eppan-Kaltern (AppianoCaldaro) at Missian (Missiano) is situated directly south of
Kalterer See (Lago di Caldaro, dotted line). Two different ice
margins of the Etsch Valley lobe are clearly traceable: an older
Fuschgalai-substage (thick line) and a younger Kaltern substage (thin line). The Etsch Valley south of these retreating
glacier tongues has presumably been filled with rapidly vanishing meltwater lakes (black).
2. Der Etschgletscher schmilzt weiter zurück, der Eisspiegel der Kalterer Zunge sinkt etwas ab. Der
größte Teil der Kamesterrassen zwischen Kaltern
und dem Kalterer See entsteht, außerdem Kamesterrassen unterhalb der Lateralmoräne von
Fuschgalai und im Leuchtenburger Wald (Stadium
von Kaltern, Abb. 6, 7). Beim Absinken des Eisspiegels werden durch Schmelzwässer parallel zur Lateralmoräne bzw. parallel zur Kamesterrasse südlich von Kaltern die Erosionstäler des Fuschgalai
bzw. am Barleiter Weg eingetieft.
3. Die Kalterer Zunge schmilzt zurück. Durch das Absinken des Eisspiegels dünnt das Eis bei Kaltern so
weit aus, dass sich von der Kalterer Zunge eine
große Toteismasse im Kalterer See abtrennt. Zwischen der Toteismasse und dem aktiven Eisrand
bei Unterplanitzing akkumulieren flächenhaft
Kiese, die bei einem kurzen Vorstoß dieser Zunge
nochmals überfahren werden. Die Schmelzwässer
fließen um die Toteismasse herum und münden
südlich des Kalterer Sees ins Etschtal. Hier entstehen im Niveau des Etschtales vermutlich Deltakiese.
4. Mit dem Rückschmelzen der Kalterer Zunge, dem
allmählichen Kleinerwerden der Toteismasse und
dem dadurch bedingten Tieferlegen des Vorfluters
schneiden sich die Schmelzwässer in die zuerst gebildeten Kiesflächen ein. In den Rinnen des Lavasontales, Val Eusca, Frühlingstalele etc. werden
Schmelzwassersedimente erodiert und nördlich
der Toteismasse auf tieferen Niveaus erneut abgelagert. Zunehmend sind weniger Schmelzwasserrinnen aktiv, am längsten die des Lavasontales und
die tief eingeschnittene Rinne von Oberplanitzing.
Die Schmelzwässer fließen immer noch um die
Toteismasse herum und münden südlich des Kalterer Sees ins Etschtal.
5. Während die Kalterer Zunge langsam nach Eppan
zurückschmilzt, ist zuletzt nur noch die Schmelz-
17
wasserrinne des Lavasontales aktiv. In dem Maße
wie sich die Toteismasse im Becken des Kalterer
Sees verkleinert, vergrößern sich die Kiesflächen
nördlich und südlich davon. Das Eis im Becken des
Kalterer Sees verschwindet erst, als kein Schmelzwasser mehr von N her zufließt. Dadurch bleibt
ein Teil der Hohlform bis heute als See erhalten.
Die eiszeitlichen Ablagerungen werden stellenweise erodiert, teilweise auch durch junge Schwemmund Murfächer überdeckt.
4. Schlussfolgerungen
In der weiten Talung von Eppan-Kaltern ist ein
ganzes System von kiesigen Lateralmoränen, breiten
Kamesterrassen (Taf. 1, 2) und peripheren Rinnen
erkennbar (Abb. 1, 6), mit dessen Hilfe sich unterschiedliche Rückschmelzstadien einer „Kalterer
Zunge“ rekonstruieren lassen. Sie muss während des
Eisstandes von Auer im ausgehenden Hochglazial in
der Talung von Eppan-Kaltern gelegen und knapp
südlich des heutigen Kalterer Sees gestirnt haben
(Abb. 7). Die Kamesterrassen bestehen vor allem aus
sehr kristallinreichen Schmelzwasserkiesen und sanden, Stillwassersedimenten und zu einem kleinen Teil auch aus einer Vielzahl von diamiktischen
Sedimenten, darunter Geschiebelehme (Tille) und
Mursedimente (Abb. 3).
Bergwärts gehen die den Eisrand begleitenden,
leicht nach Süden hin einfallenden Terrassen stellenweise tatsächlich in stärker geneigte alluviale
Fächer aus karbonatreichem Murschutt über, der
von den Hängen unterhalb des Mendelzuges
stammt. Neben Murfächern, die mit diesen Eisrandterrassen direkt verbunden sind (Abb. 2), treten
auch Strukturen auf, die hier „Murkames“ genannt
werden. Es handelt sich um Erosionsreste von stark
geneigten Murfächern, die offensichtlich direkt
gegen den absinkenden Eisrand des Etschgletschers
geschüttet wurden. Diese „Murkames“ besitzen auf
ihrer talwärtigen Seite einen deutlichen Gefälleknick (Taf. 2), eine Sackungskante, die ihre Entstehung dem Eisrand verdankt, gegen den die Sedimente ursprünglich geschüttet worden waren
(Abb. 4). Daneben gibt es auch jüngere, aktive und
inaktive Murfächer, aber ebenso Erosionsreste von
deutlich älteren, die offensichtlich vom Eis überfahren und dadurch überkonsolidiert sind (Abb. 6).
Diese müssen aus der Zeit vor dem Eishöchststand
der Würmeiszeit stammen. Stellenweise tritt extrem
18
matrixarmer Karbonatschutt auf, der bei Graun
Oberflächenstrukturen zeigt, wie sie für einen (sicher nicht mehr aktiven) Blockgletscher typisch
sind (Abb. 1). Ähnliche Ablagerungen sind im Höllental und oberhalb Kaltern bei St. Anton durch
karbonatische Zemente zu festen Brekzien verfestigt worden. Mursedimente unterschiedlichen Alters bedecken in überraschend großer Mächtigkeit
weite Flächen an der Ostflanke des Mendelzuges bis
hinunter ins Tal, besonders in der Umgebung von
Kurtatsch. Tille, diamiktische Sedimente (Taf. 4) und
Brekzien unterschiedlicher Zusammensetzung und
Genese werden im Rahmen dieser Arbeit ausführlich beschrieben (Abb. 3).
Die gesamte Talung von Eppan-Kaltern wird von
tief eingeschnittenen, breiten, kastenförmigen
Trockentälern durchzogen (Abb. 1, 6). Abschnittsweise werden diese Trockentäler auch von heutigen
Gewässern benutzt, die die alten Täler teilweise
anerodiert und zerstört, in einigen Fällen auch mit
ihren Ablagerungen aufgefüllt haben. Die
Trockentäler bilden ein verzweigtes Talsystem, das
in südlicher Richtung zum Kalterer See hin entwässert. Das größte und am wenigsten von jüngeren
Schwemmfächern zugeschüttete Trockental, das Lavasontal, lässt sich von den Reitwiesen am Kalterer
See über 6 km Richtung N bis nach Eppan (St.
Michael) verfolgen. Die Talböden der hiervon abzweigenden Trockentäler werden teilweise vom
Haupttal deutlich unterschnitten. Bei allen diesen
Trockentälern handelt es sich um Schmelzwasserrinnen, die zu einem Zeitpunkt entstanden, als die
„Kalterer Zunge“ nach Norden in Richtung Eppan
zurückschmolz (Abb. 8).
Das komplexe System aus mächtigen Schmelzwassersedimenten, erosiven Schmelzwasserrinnen,
Kamesterrassen und Lateralmoränen in der Talung
von Eppan-Kaltern lässt sich nur dann zwanglos
deuten, wenn man eine große, langsam abschmelzende Toteismasse im Gebiet des Kalterer Seebeckens annimmt (Abb. 8/ 3-5). Diese Toteismasse
muss während des Rückschmelzens des Etschgletschers dafür gesorgt haben, dass sich im Norden
davon zunächst mächtige Schmelzwassersedimente
akkumulieren konnten (Abb. 8/ 3), die mit dem
langsamen Zurückschmelzen des Toteises und dem
dadurch bedingten Absinken des Vorfluters allmählich wieder ausgeräumt wurden (Abb. 8/ 4). Zudem
sorgte sie offensichtlich dafür, dass das Becken des
Kalterer Sees, trotz erheblichen Sedimenteintrages,
nicht restlos aufgefüllt werden konnte.
Abb. 8: Das Rückschmelzen der Kalterer Zunge des Etschgletschers an der Wende vom Hoch- zum Spätglazial der Würmeiszeit, dargestellt in 5 Etappen. Rekonstruktionsversuch auf der Grundlage von geologischen und geomorphologischen Detailkartierungen im
Maßstab 1:10.000. Stand 1 entspricht dem Fuschgalai-Stadium, Stand 2 dem Stadium von Kaltern. Nähere Erläuterungen zu den
Rückzugsetappen im Abschnitt 3.
Fig. 8: An attempt to reconstruct 5 substages of the ice recession at the transition from the Pleniglacial to Late Glacial Period: the
Kaltern lobe (ETZ), part of the Etsch (Adige) svalley glacier. Based on detailed geological and geomorphological mapping in the region between Eppan (Appiano) and Margreid (Magrè) at a scale of 1:10.000. The sketch on the left (1) corresponds to the Fuschgalai
substage, the next one (2) to the Kaltern substage. For more information concerning the different substages of ice recession see
chapter 2.
5. Conclusions
Within the wide vale of Eppan-Kaltern (AppianoCaldaro) at Überetsch (Oltradige, Sella di AppianoCaldaro) close to Bozen (Bolzano) in South Tyrol
(Alto Adige) a complex system of gravelly lateral
moraines, large kame terraces (plate 1, 2) as well as
erosive peripheral meltwater valleys can be identified (fig. 1, 6). With the help of these structures it is
possible to reconstruct different substages of the
„Kaltern lobe“, a late Pleniglacial tongue of the
Etsch (Adige) valley glacier. Originally the vale of
Eppan-Kaltern was filled with the glacier ice of this
lobe. At the transition from the Pleniglacial to the
Late Glacial Period, in a time roughly corresponding
to the stage of Auer (Ora), the front of this glacier
was situated directly south of present Kalterer See
(Lago di Caldaro, fig. 7). The kame terraces are built
up of meltwater sands and gravel extremely rich in
crystalline material, lake sediments and a variety of
different diamictons, for example tills and debris
flow deposits (fig. 3).
The kame terraces which dip gently downvalley,
gradually change into steeper inclined fossil alluvial
fans to the west (fig. 2), built up entirely by angular
fragments of limestone and dolomite, deriving from
the steep slopes below the Mendola-Roèn-Ridge
above the terraces. Apart from these structures
connected with kame terraces isolated erosional
remnants of steeply inklined debris fans can be
identified, obviously deposited in the gap between
the mountain slope and the western rim of the
shrinking glacier. These „fankame“ expose a typical
steep edge at their lower parts (plate 2), generated
by the glacier which formerly served as an abutment for these sediments (fig. 4). They have been
deeply cut by erosional valleys since the glacier ice
has disappeared.
Apart from these fossil alluvial fans younger active and inactive fans can be identified. Beyond
that there are erosional remnants of debris fans,
which are overconsolidated (fig. 6), because they
have been overridden by the glacier ice and therefore are clearly older. Structures of this type as for
example the hill of St. Jakob in Kastelaz at Tramin
(Termeno) should have formed in the time before
the LGM. Thick diamictons of this type which are
obviously no tills at all cover great areas around
19
Kurtatsch (Cortaccia) and Tramin (Termeno). Many
vineyards are situated on these rigid stony deposits,
called „Kampferde“ (which means „soil to fight
with“) by locals. Most of these sediments are presumably debris flow deposits of different ages.
In places coarse grained sediments with an extremely low content of silt and sand occur, consisting mostly of angular fragments of carbonate rocks.
Close to Graun (Corona) sediments of this type
show morphological surface structures characteristic for rock glaciers (fig. 1). Due to their comparably
low altitude of only 1000 m above sea level this
rock glacier is probably fossil and not active at present. Similar sediments within the Höllental (Valle
del Inferno) and close to St. Anton near Kaltern
(Caldaro) were transformed to breccias by carbonate cementation. Tills, diamictons and breccias of
different composition, origin and age are mentioned above in detail (fig. 3).
Within the large vale of Eppan-Kaltern a variety
of erosive meltwater valleys can be identified,
deeply incised into a thick and complex sequence of
Pleistocene sediments, forming a branched fossil
drainage system (fig. 1, 6). In places modern creeks
use parts of these valleys and have destroyed them
both, by erosion and infill of sediments. The Pleistocene dry valleys mostly show flat bottoms and
drain roughly to the south into the basin of Lake
Kalterer See. The Lavason Valley is the largest and
the best preserved of these meltwater valleys,
tracable from Reitwiesen just north of Lake Kalterer
See to St. Michael at Eppan (Appiano) over a distance of 6 km. The bottoms of its tributary valleys
are clearly cut by the main valley, and therefore
seem not to have been active for such a long time
as the Lavason Valley itself. All these valleys were
formed by meltwater streams of the „Kaltern lobe“
in a later substage (fig. 8), when this glacier tongue
melted slowly back to Planitzing (Pianizza) and
Eppan (Appiano).
The formation of the whole complicated system
of lateral moraines, kame terraces as well as erosive
fossil meltwater valleys within the vale of EppanKaltern, can only be interpretated in a simple and
satisfying way, if a large and slowly vanishing mass
of stagnant ice is postulated to have existed within
the basin of Kalterer See (fig. 8/ 3-5). This stagnant
glacier ice may have been an obstacle for the meltwaters, streaming from the retreating glacier
tongue in the north towards the lake basin in the
south. North of this hypothetical abutment of stag-
20
nant ice a thick sequence of glaciofluviatile and
glaciolacustrine sediments was accumulated for a
while (fig. 8/ 3). With the slow downmelting of the
stagnant ice the meltwater rivers rather began to
erode and several generations of erosional drainage
systems were formed here (fig. 8/ 4). The mass of
stagnant ice may also have prevented the lake basin
from infill of meltwater sediments. Otherwise
Kalterer See would not have survived.
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21
Tafelerläuterungen / Explanation of plates
1: Blick über die Talung von Eppan-Kaltern nach SE, von der Barleite zum Unterberg. Die begrünte Verebnungsfläche ist
die Kamesterrasse von Kaltern, am Hang des bewaldeten Berges ist die Lateralmoräne von Fuschgalai als leicht nach
rechts geneigte gerade Linie zu erkennen. Das Tal dahinter ist das Etschtal.
1: View to Unterberg from Barleite in the northwest, across the vale of Eppan-Kaltern (Appiano-Caldaro). The green plain
is the kame terrace of Kaltern, the slightly inclined line at the slope of the wooded mountain is the lateral moraine of
Fuschgalai. The valley behind that is the Etsch (Adige) Valley.
2: Blick auf Kaltern von S her. Bei der breiten Verebnungsfäche handelt es sich um die Kamesterrasse von Kaltern (vgl. Abb.
2).
2: View to Kaltern from the south. The large green plain is the kame terrace of Kaltern (see fig. 2).
3: Blick von der Mendelpassstraße nach S in Richtung St. Nikolaus. Der bewaldete, nach links (E) gleichmäßig geneigte
Rücken ist der Erosionsrest des „Murkames“ oberhalb von Pfuss, das von rechts (W) vom Hang her gegen den Rand des
Gletschers geschüttet wurde. Deutlich ist der Gefälleknick an seinem unteren Ende zu erkennen, die Kante, an der das
Murkames ursprünglich ans Eis grenzte (vgl. Abb. 4).
3: View to St. Nikolaus (S. Nicolo) to the south from the road from Eppan (Appiano) to Mendelpass (Passo della Mendola).
The wooded hill gently dipping from rigt (W) to left (E) belongs to the „fankame“ above Pfuss, originally supplied with
debris from the slope on the right hand side (W). There is a typical steep edge at its lower end generated by the glacier
which formerly served as an abutment for these sediments (see fig. 4).
4: Straßenaufschlüsse in karbonatreichen, diamiktischen und nahezu ungeschichteten Mursedimenten an der Straße von
Kurtatsch nach Penon. Es sind zahlreiche, größere, eckige Dolomitblöcke zu erkennen.
4: Roadside exposures of diamictic and nearly not stratified debris flow sediments ritch in carbonate fragments, at the
road from Kurtatsch (Cortaccia) to Penon (Penone). Some of the angular dolomite bolders can be identified.
22
1
2
3
4
23
Geo.Alp, Vol. 2, S. 25–29, 2005
THE LATE PLEISTOCENE VERTEBRATE FAUNA
FROM AVETRANA (TARANTO, APULIA, SOUTHERN ITALY): PRELIMINARY REPORT
Raffaele Sardella1,2, Claudia Bedetti1, Luca Bellucci1, Nicoletta Conti1, Danilo Coppola3,
Emmanuele Di Canzio2, Marco Pavia4, Carmelo Petronio1,2, Mauro Petrucci1 & Leonardo Salari1
With 3 figures
1 Dipartimento di Scienze della Terra, Università di Roma “La Sapienza”; e-mail: [email protected]
2 CNR-IGAG Istituto di Geologia Ambientale e Geoingegneria
3 Dipartimento di Storia, Università “Tor Vergata” di Roma
4 Dipartimento di Scienze della Terra, Università di Torino
Riassunto
In una cava inattiva, nei pressi della cittadina di Avetrana, a Est di Taranto (Puglia, Italia meridionale),
è stata rinvenuta un ricca associazione faunistica a vertebrati contenuta nel riempimento di una cavità
di origine carsica. Nella cava affiora la formazione delle “Calcareniti di Gravina”, compatta di colore giallastro, con ricca malacofauna ed echinidi, ascrivibile al Pleistocene inferiore. A seguito di un saggio di
scavo condotto nell’ottobre 2003 è stato possibile condurre una prima analisi stratigrafica del riempimento carsico, con campionamento dei diversi livelli riconosciuti e recupero di alcuni degli abbondanti
reperti fossili affioranti. Attualmente i resti di vertebrati recuperati sono conservati presso la
Soprintendenza di Taranto. Nel riempimento della fessura carsica indagata sono stati riconosciuti nove
livelli fossiliferi e due tasche. Le specie presenti nel deposito sono riferibili al Pleistocene superiore.
Abstract
In an abandoned quarry near Avetrana (Taranto, Apulia, Southern Italy) a fossiliferous karst filling deposit
rich in vertebrate remains has been discovered. This deposit fills a wide karst fracture crossing a massive, yellow marine bio-calcarenite termed “Calcareniti di Gravina” Formation. This formation contains a rich macrofauna and echinids, referable to the Early Pleistocene. In October 2003, field activities including sampling of
the sediments and a preliminary excavation of the fossiliferous levels started. Within the karst filling deposit
nine levels and two pockets could be determined. The collected fossils are stored at the “Soprintendenza per
i Beni Archeologici per la Puglia” (Taranto, Apulia).
This fossiliferous karst deposit can be referred to the Late Pleistocene.
Introduction
The Salento Peninsula is well known in the
palaeontological literature for its Late Pleistocene
vertebrate faunas, mainly in its southern part
(Blanc 1920, De Giuli 1983, Corridi 1987, Di Stefano
et alii 1992, Bologna et alii 1994, Rustioni et alii
1994 among others). Data available for the Ionian
northern part of Salento are quite rare. Here we
report the discovery of a new fossiliferous locality
in the area of Taranto. It is a karst filling deposit
rich in fossil bones located in an abandoned calcarenite quarry, in the area of Avetrana, not far
from Manduria (Fig. 1).
After a preliminary survey during May 2003, carried on by some of the authors (D. Coppola and C.
Petronio, in particular), a team of palaeontologists
of the “La Sapienza” University, leaded by Prof.
25
C. Petronio, started the field campaign, in accordance with “Soprintendenza per i Beni Archeologici
per la Puglia”. Field work was continued in October 2003, with
activities of sampling sediments
and a partial excavation of the
fossiliferous levels. The collected
fossils are stored at “Soprintendenza per i Beni Archeologici per
la Puglia” (Taranto, Apulia).
In the present paper we present
a preliminary analysis of the collected material and some general
outlines of the fossiliferous karst
deposit.
Stratigraphy
Fig. 1: Location of Avetrana
The “Calcareniti di Gravina” Formation is a massive bio-calcarenite, rich in molluscs and echinids,
that widely outcrops in the central-southern
Apulian Peninsula; its age spans the Late Pliocene
(Adriatic side) and Early Pleistocene (Ionian side)
(Ciaranfi et alii 1988). In the considered quarry near
Avetrana this formation is exposed in a section
which is approximately 10 m thick. The vertebrate
fossil bones occur in a karst fissure filling (Fig. 3).
The sediments containing fossil vertebrates are
divided into two parts: the upper part fills the main
cavity (layers 1 to 9), the lower part fills a network
of small fissures which opened under the main one.
The small fissures (layer 0 in Fig. 3) are filled with
orange-yellow nonlaminated sandy clays rich in
small and medium-sized vertebrate remains. The
main cavity is filled with laminated sediments 4,5 to
5,5 m thick. From the bottom to the top the following levels have been determined (Fig 3):
5) 140 cm of clayey sand with very abundant bones
and rare calcareous pebbles. This layers is characterised by a level of bones and pebbles at its
base and also by a sandy lens with rare bones 20
cm above the base of this layer.
6) 20 cm of sand with abundant bones and calcareous pebbles.
7) 40 cm of clay with abundant bones and large
calcareous boulders, especially at the base of the
layer (Fig. 2).
8) 75 cm of clay and bones, the bones are also concentrated at the base of the layer, separating it
from the underlying layer.
9) 70 cm of clay with sparse bones, most of them
decalcified.
Palaeontology
1) 30 cm of clayey sand with rare altered calcareous pebbles and bones. A continuous level of
calcareous pebbles constitutes the basal part of
the layer.
2) 20 cm of sandy clay very rich in fossil remains.
3) 20 cm of clayey sand with some bones and rare
calcareous pebbles.
4) 20 cm of sandy clay rich in fossil bones and calcareous pebbles. This layer is separated from
layer 3 by an erosional surface which is marked
by a level of calcareous pebbles and bones.
26
Most of the fossil remains found at different levels of the main cavity are of medium to large size,
mainly referable to Bos primigenius (Fig. 3). The preliminary analysis of the fossil material enables us to
present the following faunal list:
Layer 0: AVES: Perdix perdix, Columba livia, Athene
noctua, Pyrrhocorax graculus; MAMMALIA:
Erinaceus europaeus, Lepus europaeus,
Oryctolagus cuniculus, Hystrix cf. H. cristata,
Terricola savi, Felis silvestris.
Layer 1: Bos primigenius.
Layer 2: Vulpes vulpes, Canis lupus, Crocuta crocuta, Lynx lynx, Stephanorhinus sp., Bos primigenius, Bovidae indet., Dama dama, Cervus elaphus.
Layer 3: Lepus europaeus, Vulpes vulpes, Canis
lupus, Bos primigenius, Dama dama, Cervus elaphus.
Layer 4: Lepus europaeus, Vulpes vulpes, Canis
lupus, Bos primigenius, Dama dama, Cervus elaphus.
Layer 5: Vulpes vulpes, Canis lupus, Bos primigenius,
Cervus elaphus, Dama dama.
Layer 6: Vulpes vulpes, Canis lupus, Bos primigenius,
Cervus elaphus, Dama dama.
Layer 7: Vulpes vulpes, Canis lupus, Crocuta crocuta, Stephanorhinus sp., Bos primigenius, Cervus
elaphus, Dama dama, ?Megaloceros giganteus.
Layer 8: Vulpes vulpes, Canis lupus, Lynx lynx,
Panthera leo, Bos primigenius, Cervus elaphus,
Dama dama.
Layer 9: fossil bones are absent.
Layer 0 is characterised by the occurrence of
small vertebrate remains. Bird remains are represented by some limb bones, while mammals are
mainly represented by teeth. In particular, two
lower molar teeth (M2 and M3) of Erinaceus
europaeus, 7 well preserved mandibles and some M1
of Terricola savi, M1 and two M2 of a porcupine,
slightly smaller than the living Hystrix cristata, have
been recorded. Among lagomorphs the hare and the
rabbit occur with some fragments of skull and
mandible and some limb bones. Such taxa are also
recorded from the main cavity deposit (from layers
2 to 8) but are poorly represented. The wild cat is
represented by fragmentary limb bones of peculiar
size.
In the main cavity filling, Bos primigenius is the
best-represented taxon in each fossiliferous level
(1–8); layers 5 and 6 are very rich in limb bones, in
some cases in anatomical connection. Layer 8 is also
characterised by the occurrence of skull fragments
and mandibles with jugal teeth.
Cervids are represented by some isolated teeth
and limb bones (layers 2–8); two large-sized first
phalanxes recorded in level 7 can probably be
ascribed to Megaloceros giganteus. Among carnivores, the occurrence of the wolf and the red fox is
testified by some isolated teeth, occurring from layers 2 to 8, the lynx and the cave lion are recorded
from layer 8 (some teeth and a talus respectively).
Fig. 2: Fossil bones of layer 7 (scale bar: 20 cm).
The occurrence of a rhino is testified in layers 2 and
7. This taxon is represented by one fragmentary
molar tooth and one pisiform in every layer.
The faunal assemblage on the whole can be
referred to the Late Pleistocene. The occurrence of
the fallow deer (in particular of the modern subspecies Dama dama dama) and of a rhino, generally
referable to Stephanorhinus sp., recorded from layers 2 and 7, gives important biochronological constraints. In fact, the modern fallow deer was widespread in Italy at the beginning of the Late
Pleistocene, while rhinos referable to the genus
Stephanorhinus survived until the beginning of the
Pleniglacial (MIS 3) (Gliozzi et alii 1997).
At the moment, only general considerations on the
palaeoenvironmental conditions can be pointed out.
In layer 0, the occurrence of Terricola savii and
Hystrix cf. H. cristata suggests the presence of temperate climatic conditions with dry and open
palaeoenvironments. Moreover, such a general
framework is supported also by the occurrence of the
avifauna including Perdix perdix and Athene noctua,
while Columba livia and Pyrrhocorax graculus suggest the presence of rocky cliffs. In the sequence filling the main cavity (layer 1 to 8), large mammal
species of a wider ecological significance occur.
Preliminary taphonomical observations indicate
that the fossil bones seem to be not oriented. This
27
Fig. 3: Stratigraphy of the fossiliferous deposit
fact suggests quick deposition of the fossil-bearing
sediments, which is also supported by sedimentological observations. In fact, the different layers are
characterised by a normally graded distribution of
the sediment, with the heaviest material like big
bones and calcareous pebbles concentrated in the
lowest part of the layer. Frequently such bones and
pebbles constitute a well defined level at the base
of the layer to separate one layer from the underlying one.
A detailed analysis of the fossil remains has just
begun, in accordance with the “Soprintendenza ai
Beni Archeologici per la Puglia”, with the aim of
providing further palaeontological information and
a framework of the palaeoenvironmental evolution
of the area during the Late Pleistocene.
Acknowledgments
All phases of fieldwork were financially supported by CNR – IGAG and by the Municipality of
Avetrana. A special mention is due to Francesco
28
Nigro, responsible for the Cultural Heritage. The
support of Maria Antonietta Gorgoglione, responsible for “Soprintendenza per i Beni Archeologici per
la Puglia”, is warmly acknowledged. Moreover, the
Earth Science Department of the University of Turin
and the “Museo delle Civiltà Preclassiche delle
Murge Meridionali” provided facilities for research.
We wish to thank Giuseppe “Pippo” Arcidiacono,
Francesco Ciminelli, Vincenza Montenegro and
Michael Giagnoni for participating the field work
and, finally, Karl Krainer and Marzia Breda for their
comments and suggestions on the manuscript.
References
Blanc, G. A.(1920): Grotta Romanelli. – Arch. Antrop. Etn.,
50(1-4): 1-39.
Bologna, P., Di Stefano, G., Manzi, G., Petronio, C.,
Sardella, R., Squazzini, E. (1994): Late Pleistocene
mammals from the Melpignano (Le) „Ventarole“: preliminary analysis and correlations. - Boll. Soc. Paleont.
It., 33 (2): 265-274.
Ciaranfi, N., Pieri, P., Ricchetti, G. (1988): Note alla Carta
Geologica delle Murge e del Salento (Puglia centromeridionale). - Mem. Soc. Geol. It., 41(1): 449-460.
Corridi, C. (1987): Faune pleistoceniche del Salento: 2. La
fauna di fondo Cattìe, Maglie, Lecce. - Quaderni del
Museo Comunale di Paleontologia, 3: 5-74.
De Giuli, C. (1983): Le faune pleistoceniche del Salento: 1.
La fauna di S. Sidero 3. - Quaderni del Museo
Comunale di Paleontologia, 1: 45-84.
Di Stefano, G., Petronio, C., Sardella, R., Savelloni, V.,
Squazzini, E. (1992): Nuove segnalazioni di brecce
ossifere nella costa fra Castro Marina e Otranto
(Lecce). - Il Quaternario, 5 (1): 3-10.
Gliozzi, E., Abbazzi, L., Argenti, P., Azzaroli, A., Caloi, L.,
Capasso Barbato, L., Di Stefano, G., Esu, D., Ficcarelli,
G., Girotti, O., Kotsakis, T., Masini, F., Mazza, P.,
Mezzabotta C., Palombo, M. R., Petronio, C., Rook, L.,
Sala, B., Sardella, R., Zanalda, E., Torre, D. (1997):
Biochronology of selected Mammals, Molluscs and
Ostracods from the Middle Pliocene to the Late
Pleistocene in Italy. The state of the art. – Riv. Ital.
Paleont. Strat., 103(3): 369-388.
Rustioni, M., Mazza, P., Abbazzi, L., Delfino, M., Rook, L.,
Petrucci, S., Vianello, F. (1994): The Würmian fauna
from Sternatia (Lecce, Apulia, Italy). - Boll. Soc.
Paleont. It., 33 (2): 279-288.
Manuscript submitted: December 17, 2004
Revised manuscript accepted: February 22, 2005
29
Geo.Alp, Vol. 2, S. 31–51, 2005
THE LADINIAN FLORA (MIDDLE TRIASSIC) OF THE DOLOMITES:
PALAEOENVIRONMENTAL RECONSTRUCTIONS AND PALAEOCLIMATIC CONSIDERATIONS
Evelyn Kustatscher1 & Johanna H.A. van Konijnenburg-van Cittert2
With 7 figures and 5 tables
1 Dipartimento di Scienze della Terra, Università degli Studi di Ferrara, C.so Ercole I d’Este 32, 44100 Ferrara, Italy,
e-mail [email protected]
2 Laboratory of Palaeobotany and Palynology, Budapestlaan 4, 3584CD Utrecht, Netherlands,
e-mail [email protected]
Abstract
The study of several plant fossils from the Ladinian of the Dolomites, that either had been described a long
time ago or had never been described at all, has led to a revision of this flora. The Ladinian flora now consists of the following taxa: Annalepis zeilleri (Lycophyta), Equisetites arenaceus (Sphenophyta), Cladophlebis
leuthardtii, C. ruetimeyeri, Neuropteridium elegans, Scolopendrites sp., Gordonopteris lorigae (Pteridophyta),
Ptilozamites heeri (Pteridospermae), Bjuvia dolomitica, Dioonitocarpidium moroderi, Pterophyllum jaegeri,
?Pterophyllum sp., Sphenozamites wengensis, Sphenozamites cf. bronnii, Taeniopteris sp. (Cycadophyta),
Voltzia dolomitica, V. ladinica, V. pragsensis, V. zoldana, Voltzia sp., Pelourdea vogesiaca und Elatocladus sp.
(Coniferophyta).
The flora, and especially the large number of specimens housed in the Natural History Museum at Bolzano
(I), indicates a dominance of conifers over (in this sequence) seedferns, cycads, ferns and horsetails. Several
factors may have caused this: climatic (an arid climate on the mainland), edaphic (immature soil) or taphonomic (caused by selection during transport). Quantitative palynological analyses of three localities (Ritberg
near Wengen, and Seewald and Innerkohlbach near Prags, indicate a generally warm and humid climate. The
dominance of the conifers and seedferns may, therefore, have been caused by their larger resistance during
transport rather than by climatic factors.
Ladinian palaeoclimatic reconstructions and the plant fossils studies indicate that during the late Ladinian
the Dolomites consisted of carbonate or volcanic islands of various sizes, which were covered with several
biotopes: coastal and ‚hinterland’; the latter divided into a more humid and a more arid zone.
Zusammenfassung
Das Studium verschiedener historischer, ebenso wie bisher unbeschriebener Pflanzenfossilien aus dem
Ladin der Dolomiten, die in italienischen und ausländischen Museen aufbewahrt werden, führte zu einigen
Erstbeschreibungen und systematischen Revisionen. Die Ladinflora setzt sich nunmehr aus folgenden Arten
zusammen: Annalepis zeilleri (Lycophyta), Equisetites arenaceus (Sphenophyta), Cladophlebis leuthardtii, C.
ruetimeyeri, Neuropteridium elegans, Scolopendrites sp., Gordonopteris lorigae (Pteridophyta), Ptilozamites
heeri (Pteridospermae), Bjuvia dolomitica, Dioonitocarpidium moroderi, Pterophyllum jaegeri, ?Pterophyllum
sp., Sphenozamites wengensis, Sphenozamites cf. bronnii, Taeniopteris sp. (Cycadophyta), Voltzia dolomitica,
V. ladinica, V. pragsensis, V. zoldana, Voltzia sp., Pelourdea vogesiaca und Elatocladus sp. (Coniferophyta).
Die Flora, insbesondere die, die sich im Naturmuseum Südtirol (BZ) befindet, weist eine Dominanz der
Koniferen über Samenfarne, Cycadeen, Farne und Schachtelhalmen auf. Eine derartige Zusammensetzung
kann auf verschiedenen Faktoren beruhen: (i) Klimatische (arides Klima auf dem Festland), (ii) edaphische
31
(unreife Böden) oder auch (iii) taphonomische (Selektion während des Transportes). Die quantitativen
Analysen der Palynofloren der drei Fundorte Ritberg (Wengen), Seewald und Innerkohlbach (Prags) weisen
auf ein generell warmes und feuchtes Klima hin. Aus diesem Grund scheint die Dominanz der Koniferen und
Samenfarne eher auf die größere Resistenz dieser Pflanzen gegen Zerstörung während des Transports, als auf
klimatische Auslese zurückzuführen zu sein.
Paläoklimatische Rekonstruktionen aus dem Ladin sowie die Analyse der Pflanzenfossilien weisen darauf
hin, dass die Dolomiten im oberen Ladin von karbonatischen oder vulkanischen Inseln verschiedener Größe
bedeckt waren, wo sich verschiedene Biotope gebildet hatten: das Küstengebiet und das Hinterland, das sich
wiederum in feuchtere und trockener Zonen unterteilen lässt.
Riassunto
Recenti studi sistematici della flora ladinica delle Dolomiti, condotti su collezioni sia storiche che inedite
di musei italiani e stranieri hanno portato a nuove segnalazioni e ad alcune revisioni sistematiche. La flora
ladinica risulta essere composta dai seguenti taxa: Annalepis zeilleri (Lycophyta), Equisetites arenaceus
(Sphenophyta), Cladophlebis leuthardtii, C. ruetimeyeri, Neuropteridium elegans, Scolopendrites sp.,
Gordonopteris lorigae (Pteridophyta), Ptilozamites heeri (Pteridospermae), Bjuvia dolomitica, Dioonitocarpidium moroderi, Pterophyllum jaegeri, ?Pterophyllum sp., Sphenozamites wengensis, Sphenozamites cf.
bronnii, Taeniopteris sp. (Cycadophyta), Voltzia dolomitica, V. ladinica, V. pragsensis, V. zoldana, Voltzia sp.,
Pelourdea vogesiaca ed Elatocladus sp. (Coniferophyta).
La flora, in particolare quella depositata nel Museo di Scienze Naturali dell’Alto Adige (BZ), presenta una
generale dominanza delle conifere e pteridosperme, sulle cicadee, felci e sfenofite. Una simile composizione
può essere imputabile a vari fattori: climatici (aridità delle terre emerse), edafici (suoli immaturi) e tafonomici (selezione tassonomica causata da un trasporto prolungato). Le analisi quantitative della palinoflora,
effettuate nei tre affioramenti di Ritberg (La Valle), Seewald ed Innerkohlbach (Braies), indicano un clima
complessivamente caldo umido. Pertanto, la dominanza delle conifere e pteridosperme sembra essere dovuta ad una maggiore resistenza di queste piante ai processi putrefattivi, che avvengono durante il trasporto,
piuttosto che a cause climatiche.
Sulla base delle ricostruzioni paleogeografiche del Ladinico superiore e sui resti macrofloristici determinati, le Dolomiti presentavano una serie di piccole piattaforme carbonatiche emerse e isole vulcaniche, sulle
quali si dovevano esistere biotopi differenti: aree costiere, e l’ambiente di entroterra, a sua volta suddivisa
in zone più umide e più aride.
1 Introduction
The first Ladinian plant remains from the
Dolomites have been figured by Wissmann and
Münster (1841). Afterwards several authors mentioned and figured plant fossils from the
“Buchensteiner Schichten” and “Wengener Schich ten” of various areas in the Dolomites (Mojsisovics,
1879; Arthaber,1903; Ogilvie Gordon, 1927, 1934;
Mutschlechner, 1932; P. Leonardi, 1953, 1968;
Calligaris, 1983, 1986; Jung et al., 1992) and from
Sappada (G. Leonardi, 1964) (for more detailed
information see also, Wachtler & van Konijnenburg
– van Cittert, 2000a, b; Kustatscher, 1999, 2001,
2004). On this account, a high number of different
plant remains have been described from the
Dolomites at the end of the last century (Table 1).
32
However, an extended search and study of local
and international plant collections with both
already described and unpublished material from
the Dolomites provided material for a first report
and some taxonomic revisions of the material
(Kustatscher, 2004; Kustatscher et al., 2004).
Also several palynological studies have been
applied during the last 25 years regarding successions of Ladinian age from the Dolomites (Cros &
Doubinger, 1982; van der Eem, 1982; Blendinger,
1988; Roghi, 1995a, 1995b; Broglio Loriga et al.,
1999). However, most of the articles use palynological data only on a biostratigraphic point of view
(Blendinger, 1988, Roghi, 1995a, 1995b; Broglio
Loriga et al., 1999). Only in one of them (van der
Eem, 1982) the palynomorphs are considered also as
a source for paleoclimatic data.
2 Material and methods
The historical and often unpublished plant fossil
collections are stored in several local and international museum and universities. In detail, the plant
remains figured by Ogilvie Gordon (1927) are kept
in the “Paläontologisches Museum” (Munich, D),
Mutschlechner’s (1932) material in the “GeologischPaläontologisches Institut” of the University of
Innsbruck. The plant fossils discussed in Leonardi
(1953) are treasured at the “Museum de Gherdëina”
(Ortisei, I) and at the “Museo di Geologia e
Paleontologia” of the University of Padova (I). Some
specimens are kept at the “Museo di Paleontologia e
Preistoria P. Leonardi” of the University of Ferrara (I)
as also the fossil plants from Sappada figured by G.
Leonardi (1964) and the plants figured in Leonardi
(1968) and Bosellini (1989, 1996). The neuropteridian leaf fragment, figured by Zardini (1980) is
exposed in the “Museo Paleontologico Rinaldo
Zardini” (Cortina, I). The material discussed and figured by Calligari (1986) is stored in the Museo di
Scienze Naturali (Trieste, I). Finally, the material discussed by Wachtler & van Konijnenburg – van
Cittert (2000a, 2000b) and Kustatscher (1999, 2001,
2004 p.p.) is stored in the “Museo di Scienze
Naturali Alto Adige / Naturmuseum Südtirol”
(Bolzano / Bozen, I), in the Museum de Gherdëina
(Ortisei) and in the “Museo Paleontologico Rinaldo
Zardini” (Cortina). Unfortunately the material mentioned by Mojsisovics (1879) seems to have been
lost. Additionally unpublished material is stored in
the Museums discussed above and also at the
Museo Ladino Fodom (Livinallongo del Col di Lana,
I), the Naturhistorisches Museum, the Geologische
Bundesanstalt (Vienna, I) and at the Geologisches
Landesamt (Munich, D).
For paleoclimatic considerations palynomorph
analyses have been carried out for 6 samples collected at two plant localities near Braies / Prags
(Seewald and Innerkohlbach) and one near La Valle
(Ritberg), belonging respectively to the upper part
of the Fernazza Formation (Ritberg and Seewald)
and to the base of the Wengen / La Valle Formation
(Innerkohlbach) (see Fig. 1).
The samples have been crushed into small fragments and treated with the standard palynological
techniques, including HCl (37%), HF (40%) and saturated ZnCl2 solution (D ≈ 2,3 g/ml). Afterwards,
the slides have been mounted in Canadian balsam.
Fig. 1: Geographic distribution of the studied sections and fossil plant localities cited in the article. 1. Prags / Braies, Seewald,
Innerkohlbach; 2. Gadertal / Val Badia, Wengen / La Valle, Ritberg; 3 St. Leonhard in Abtei / S. Leonardo in Badia, St. Kassian/
San Cassiano, 4. Grödental / Val Gardena; 5. Pufels / Bulla, Puflatsch / Bullaccia, Schgaguler Alm / Malga Scagul, Seiser Alm /
Alpe di Siusi; 6. Grödner Joch / Passo Gardena, Corvara; 7.
Monte Sief, Arabba; 8. Forcella Giau, Corvo Alto, Mondeval; 8.
Laste (Livinallongo); 9. Cercenà, Spiz Agnelessa; 10. Sappada.
For the quantitative analyses at least 300 palynomorphs have been counted for each sample; the
material has been divided into the main groups as
pollen, spores, fungal remains, algal cysts, acritarchs
and foraminiferous lignins. For the quantitative
data the palaeoclimate methods proposed by
Visscher & van der Zwan (1981) and Abbink (1998)
have been applied. The frequencies of each group
has been plotted with the aid of a specialised program, named Graph4win.
All the material (macrofossil and palynological)
from the plant localities of Ritberg, Seewald and
Innerkohlbach is stored at the Museo di Scienze
Naturali dell’Alto Adige / Naturmuseum Südtirol
(Bolzano / Bozen).
3 Macrofloral composition
The Ladinian flora from the Dolomites is composed
of the following taxa. The synonymy includes only all
references from the Ladinian of the Dolomites, not
from other areas. The localities from which material
has been recovered, are indicated as well.
33
DIVISION LYCOPHYTA
Order Isoetales
Annalepis zeilleri Fliche, 1910
2004 Annalepis zeilleri Fliche – Kustatscher, p. 157,
pl. 10, fig. 1.
2004 Annalepis zeilleri Fliche – Kustatscher et al.,
p. 58, pl. 1, fig. 1.
Localities: Wengen / La Valle.
DIVISION SPHENOPHYTA
Order Equisetales
Family Equisetaceae
Equisetites arenaceus (Jaeger, 1827) Schenk, 1864
1999 Equisetites arenaceus - Avanzini & Wachtler,
p. 118.
2000a Equisetites arenaceus (Jaeger) Schenk Wachtler & van Konijnenburg - van Cittert, p.
107, pl. 1, fig. 1, 2.
2000b Equisetites arenaceus (Jaeger) Schenk Wachtler & van Konijnenburg - van Cittert, p.
116, pl. 1, fig. 1, 2.
2004 Equisetites arenaceus (Jaeger) Schenk –
Kustatscher, p. 158, pl. 10, fig. 2.
Localities: Wengen / La Valle, Sappada.
cf. Equisetites
1953 Equisetites vel Calamites? – Leonardi, pl. 4,
figs. 4–5.
1964 impronta riferibile probabilmente ad
Equisetale - Leonardi, pl. 5, fig. 10.
1964 frammento di fusto di Equisetale, forse
Neocalamites sp. - Leonardi, pl. 5, fig. 11.
2004 cf. Equisetites – Kustatscher, p. 159, pl. 10,
fig. 3.
Localities: Pufels / Bulla, Wengen / La Valle,
Seiser Alm / Alpe di Siusi, Arabba, Cercenà,
Sappada.
DIVISION PTERIDOPHYTA
Order Filicales
Family Osmundaceae or indet.
Cladophlebis leuthardtii Leonardi, 1953
1841 Fahrenwedel – Wissmann & Münster, p. 22,
pl. 16, fig. 10.
34
1953 Cladophlebis leuthardti Leonardi, p. 11, pl. 2,
figs. 1-5.
1953 Cladophlebis rütimeyeri Heer n.var. heeri –
Leonardi, p. 11, pl. 1, fig. 1.
1964 Cladophlebis sp. - Leonardi, p. 201 pl. 5, fig. 7.
1968 Cladophlebis cfr. denticulata Brongniart –
Leonardi p. 179, pl. 28, fig. 7.
1986 Cladophlebis leuthardti – Calligaris, p. 9, fig.
B29.
1993 Cladophlebis leuthardti – Pozzi, p. 82, fig. 103.
1998 cf. Pecopteris reticulata (Leuthardt) - Stingl &
Wachtler, p. 82.
1999 ?Anomopteris mougeotii Brongniart, 1828 Kustatscher, p. 43, pl. 1, fig. B; pl. 2, fig. A.
2000a Cladophlebis leuthardtii Leonardi - Wachtler
& van Konijnenburg – van Cittert, p. 109, pl.
1, fig. 3.
2000b Cladophlebis leuthardtii Leonardi - Wachtler
& van Konijnenburg - van Cittert, p. 117-8, pl.
1, fig. 3.
2004 Cladophlebis leuthardtii Leonardi –
Kustatscher, p. 160, pl. 10, fig. 5; pl. 11, fig. 1.
Localities: Prags / Braies, Wengen / La Valle, Seiser
Alm / Alpe di Siusi, Pufels / Bulla, Grödner
Joch / Passo Gardena, Corvo Alto, Corvara,
Monte Sief, Laste (Livinallongo), Cercenà,
Sappada.
Cladophlebis ruetimeyeri (Heer, 1877)
Leonardi, 1953
1953 Cladophlebis rütimeyeri Heer - Leonardi, p.
10, pl. 1, fig. 15, pl. 3 figs. 6.
1953 Cladophlebis sp. - Leonardi, pl. 1 figs. 3-4.
1994 Pecopteris – Costamoling & Costamoling, p.
47, fig. 19.
2004 Cladophlebis ruetimeyeri (Heer) Leonardi –
Localities: Seiser Alm / Alpe di Siusi, Col Alto,
Cercenà.
Neuropteridium elegans (Brongniart, 1828)
Schimper, 1869
1993 Cladophlebis sp. – Pozzi, p. 85, fig. 107.
1998 Neuropteridium sp. - Stingl & Wachtler, p. 82.
1999 Neuropteridium grandifolium (Schimper et
Mougeot) Schimper - Kustatscher, p. 44, pl. 2,
fig. B.
Fig. 2: Relative abundance of the main plant groups present in the three main macrofloral localities (Seewald, Innerkohlbach,
Ritberg).
2000a Neuropteridium grandifolium (Schimper et
Mougeot) Schimper - Wachtler & van
Konijnenburg - van Cittert, p. 108, pl. 2, fig. 1.
2000b Neuropteridium grandifolium (Schimper et
Mougeot) Schimper - Wachtler & van
elegans
(Brongniart)
2004 Neuropteridium
Schimper – Kustatscher, p. 161, pl. 11, fig. 3.
elegans
(Brongniart)
2004 Neuropteridium
Schimper – Kustatscher et al., p. 59, pl. 1,
fig. 2.
Localities: Forcella Giau.
Scolopendrites sp.
2004 Scolopendrites sp. – Kustatscher, p. 162, pl.
11, fig. 4.
2004 Scolopendrites sp. – Kustatscher et al., p. 60,
pl. 1, fig. 3.
Localities: St. Kassian / San Cassiano.
1953 cf. Pecopteris sulzensis Schimper - Leonardi,
p. 10, pl. 1, fig. 14.
?1986 Pecopteris sp. - Calligaris, p. 9, fig. A48.
1998 Anomopteris mougeotii - Stingl & Wachtler, p. 81.
1999 Anomopteris mougeotii – Avanzini &
Wachtler, p. 117.
2000a Anomopteris mougeotii Brongniart Wachtler & van Konijnenburg - van Cittert,
p. 108, pl. 1, figs. 4-5.
2000b Anomopteris mougeotii Brongniart Wachtler & van Konijnenburg - van Cittert,
p. 116, pl. 1, figs. 4-5.
2001 Anomopteris mougeotii - Kustatscher,
p. 3.
2004 ?Filicales indet. – Kustatscher, p. 162-3, pl. 10,
fig. 4.
2004 Fern incertae sedis – Kustatscher et al.,
p. 60-1, pl. 1, fig. 4.
Localities: Wengen / La Valle, Mondeval, Corvo Alto,
Cercenà, Sappada.
Gordonopteris lorigae van Konijnenburg –
van Cittert et al. (name in submitted manuscript)
DIVISION PTERIDOSPERMATOPHYTA
Order indet.
Ptilozamites heeri Nathorst, 1878
1953 felce indeterminata - Leonardi, p.13, pl. 1,
figs. 9.
1953 Pecopteris cf. (Lonchopteris) reticulata
Leuthardt - Leonardi, p. 10, pl. 1, fig. 10.
1927 Pterophyllum brevipenne Kurr - OgilvieGordon, pl. 8, fig. 1.
1980 cfr. Pterophyllum venetum - Zardini, pl.1,
fig. 8.
35
1985 Cladophlebis cf. denticulata Brongniart Moroder, p. 27, fig. 21.
1993 Cladophlebis cfr. denticulata – Pozzi, p. 83,
fig. 105.
1999 Ptilozamites heeri - Avanzini & Wachtler, p.
118.
2000a Ptilozamites heeri Nathorst - Wachtler & van
Konijnenburg - van Cittert, p. 108, pl. 2, figs.
2-9.
2000b Ptilozamites heeri Nathorst - Wachtler & van
Konijnenburg - van Cittert, p. 118, pl. 2, figs.
2-9.
2004 Ptilozamites heeri Nathorst – Kustatscher, p.
163, pl. 11, fig. 5; pl. 12, fig. 1.
Localities: Prags / Braies, Wengen / La Valle, Gadertal / Val Badia, Grödental / Val Gardena, Corvo
Alto.
DIVISION CYCADOPHYTA
Order Cycadales
Bjuvia Florin, 1933
Bjuvia dolomitica Wachtler et van Konijnenburg van Cittert, 2000
1927 Zamites sp. - Ogilvie-Gordon, p. 68, pl. 8,
fig. 4.
1953 Pterophyllum sp. - Leonardi, p. 13, pl. 3, fig. 2.
1999 Bjuvia dolomitica Wachtler et van
Konijnenburg - van Cittert (in stampa) Kustatscher, p. 45, pl. 1, fig. C; p. 49, pl. 4, fig. A.
1999 Bjuvia dolomitica - Avanzini & Wachtler, p.
113.
2000a Bjuvia dolomitica Wachtler et van
Konijnenburg - van Cittert, p. 110-111, pl. 4,
fig. 1-3; pl. 5, fig. 1-5.
2000b Bjuvia dolomitica Wachtler et van
Konijnenburg - van Cittert, p. 120-1, pl. 4, fig.
1-3; pl. 5, fig. 1-5.
2004 Bjuvia dolomitica Wachtler et van
Konijnenburg - van Cittert – Kustatscher, p.
165, pl. 12, fig. 3.
Localities: Wengen / La Valle, Grödental / Val
Gardena, Schgaguler Alm / Malga Scagul, Mondeval.
cf. Bjuvia
1927 “Zamites sp.“ - Ogilvie-Gordon, p. 68, pl. 8,
fig. 4.
36
1927 Nilssonia sp. - Ogilvie-Gordon, p. 68, pl. 8,
fig. 6.
2004 cf. Bjuvia – Kustatscher, p. 165.
Localities: Schgaguler Alm / Malga Scagul, Grödner
Joch / Passo Gardena, Corvara, Sappada.
Sphenozamites wengensis Wachtler et
van Konijnenburg - van Cittert, 2000
1999 Sphenozamites - Avanzini & Wachtler, p. 118.
2000a Sphenozamites wengensis Wachtler et van
Konijnenburg - van Cittert, p. 109, pl. 3,
figs. 1-2.
2000b Sphenozamites wengensis Wachtler et van
Konijnenburg - van Cittert - Wachtler & van
figs. 1-2.
2004 Sphenozamites wengensis Wachtler et van
Konijnenburg - van Cittert – Kustatscher,
p. 166, pl. 12, fig. 4.
Localities: Prags / Braies, Wengen / La Valle.
Sphenozamites sp. cf. S. bronnii (Schenk)
Passoni & van Konijnenburg - van Cittert, 2003
2004 Sphenozamites cf. bronnii (Schenk) Passoni &
van Konijnenburg - van Cittert – Kustatscher,
p. 166, pl. 13, fig. 2.
2004 Sphenozamites sp. cf. S. bronnii (Schenk)
Passoni & van Konijnenburg - van Cittert –
Kustatscher et al., p. 62, pl. 2, fig. 2-6.
Localities: St. Leonhard in Abtei / S. Leonardo in
Badia, Laste (Livinallongo).
Dioonitocarpidium moroderi (Leonardi)
Kustatscher et al., 2004
1953 Cycadeoidea (?) moroderi Leonardi Leonardi, p. 14, pl. 2, figs. 6-8.
1968 Cycadeoidea (?) moroderi Leonardi Leonardi, p. 176, pl. 28, fig. 5.
1999 Dioonitocarpidium sp. - Kustatscher, p. 49,
58, pl. 3, fig. A-B.
2000a Dioonitocarpidium sp. - Wachtler & van
2000b Dioonitocarpidium sp. - Wachtler & van
fig. 2.
2004 Dioonitocarpidium moroderi (Leonardi) nov
comb. – Kustatscher, p. 168, pl. 13, fig. 5.
2004 Dioonitocarpidium moroderi (Leonardi) nov
comb. – Kustatscher et al., p. 61-2, pl. 2, fig. 1.
Localities: Schgaguler Alm / Malga Scagul.
Order Bennettitales
Pterophyllum jaegeri Brongniart, 1828
1953 Pterophyllum jaegeri Brongniart - Leonardi,
p. 13, pl. 2, fig. 12.
1968 Pterophyllum jaegeri Brongniart - Leonardi,
p. 176, pl. 28, fig. 4.
1989 Pterophyllum – Bosellini, p. 19, fig. 2.1.
1999 Pterophyllum jaegeri - Kustatscher, p. 57, pl.
4, fig. B.
1999 Pterophylliium jaegeri - Avanzini & Wachtler,
p. 118.
2000a Pterophyllum jaegeri Brongniart - Wachtler
& van Konijnenburg - van Cittert, p. 112, pl.
3, figs. 3-4.
2000b Pterophyllum jaegeri Brongniart - Wachtler
& van Konijnenburg - van Cittert, p. 122-3,
pl. 3, figs. 3-4.
2001 Pterophyllum jaegeri - Kustatscher, p. 6.
jaegeri
Brongniart
–
2004 Pterophyllum
2004 Pterophyllum sp. – Kustatscher, p. 169.
Localities: Prags / Braies, Wengen / La Valle, St. Kassian / San Cassiano, Corvara, Cercenà.
?Pterophyllum sp.
2004 ?Pterophyllum sp. – Kustatscher, p. 170,
pl. 13, fig. 3.
Localities: Laste (Livinallongo).
Order indet.
Taeniopteris sp.
1927 Taeniopteris angustifolia Schenk - OgilvieGordon, p.67, pl. 8, fig. 2.
1953 cfr. Taeniopteris sp. - Leonardi, p. 12, pl. I,
fig. 18.
1964 Taeniopteris (Nilssonia ?) - Leonardi, pl. 4,
fig. 3.
1999 Taeniopteris sp. - Kustatscher, p. 57, pl. 2,
fig. C; pl. 3, fig. C.
2000a Taeniopteris sp. - Wachtler & van
2000b Taeniopteris sp. - Wachtler & van
Konijnenburg - van Cittert, p. 122, pl. 6, fig.
1.
2004 Taeniopteris sp. – Kustatscher, p. 171, pl. 13,
fig. 1.
Localities: Prags / Braies, Grödental / Val Gardena,
Gadertal / Val Badia, Corvara, Cercená, Sappada.
DIVISION CONIFEROPHYTA
Order Coniferales
Elatocladus sp.
1968 Pterophyllum sp. - Leonardi, p. 176, pl. 28, fig.
2.
1985 Pterophyllum - Moroder, p. 31, fig. 26.
1989 Pterophyllum sp. - Bosellini, p. 89, fig. 12.9.
1993 Pterophyllum sp. – Pozzi, p. 85, fig. 108.
1996 Pterophyllum - Bosellini, p. 121, fig. 13.8.
1999 Elatocladus sp. - Avanzini & Wachtler, p. 119.
1999 Elatocladus sp. - Kustatscher, p. 51, pl. 5,
fig. A.
2000a Elatocladus sp. - Wachtler & van
2000b Elatocladus sp. - Wachtler & van
2004 Elatocladus sp. – Kustatscher, p. 172, pl. 14,
fig. 2.
Localities: Puflatsch / Bullaccia.
Pelourdea vogesiaca (Schimper et Mougeot, 1844)
Seward 1917
1953 Yuccites vogesiacus Schimper et Mougeot Leonardi, p.15, pl. 2, fig. 9, 11; pl. 3, figs. 3-4.
1986 Yuccites sp. - Calligaris, p. 15, figs. B21, 42.
1999 Yuccites vogesiacus - Avanzini & Wachtler,
p. 119.
2000a Yuccites vogesiacus Schimper et Mougeot Wachtler & van Konijnenburg – van Cittert,
p. 113, pl. 6, figs. 4, 5.
2000b Yuccites vogesiacus Schimper et Mougeot Wachtler & van Konijnenburg – van Cittert,
p. 121-2, pl. 6, figs. 4, 5.
2004 Pelourdea vogesiaca (Schimper et Mougeot)
Seward – Kustatscher, p. 172-4, pl. 13, fig. 4.
2004 Pelourdea vogesiaca (Schimper et Mougeot)
Seward – Kustatscher et al., p. 63, pl. 1, fig. 5.
37
Localities: Prags / Braies, Wengen / La Valle, Schgaguler Alm / Malga Scagul.
?Pelourdea sp.
1953 Yuccites sp. – Leonardi, pl. 3, fig. 5.
2004 ?Pelourdea sp. – Kustatscher, p. 174.
Localities: Seiser Alm / Alpe di Siusi, Cercenà.
Order Voltziales
Family Voltziaceae
Voltzia dolomitica Wachtler et van Konijnenburg van Cittert, 2000
1927 Voltzia recubariensis Schenk - OgilvieGordon, p. 67, pl. 8, fig. 7.
1932 Voltzia sp. - Mutschlechner, p. 31.
1953 Pagiophyllum (?) massalongi Zigno Leonardi, p. 18, pl. 3, figs. 8, 10; pl. 4, fig. 2.
1968 Brachyphyllum sp. - Leonardi, p. 176, pl. 28, fig. 1.
1986 Pagiophyllum cf. massalongi Zigno Calligaris, p. 16, figs. A64, B6, B7, B11, B19,
B27, B31.
1995 Voltzia recubariensis Schenk - Jung et al., p.
171, fig. 8.3.
1999 Voltzia dolomitica - Avanzini & Wachtler,
p. 117, 119.
2000a Voltzia dolomitica Wachtler et van
Konijnenburg - van Cittert 2000, p. 113-14,
pl. 7, fig. 1-4; pl. 5, fig.1-6.
2000b Voltzia dolomitica Wachtler et van
fig. 1-4; pl. 5, fig.1-6.
2001 Voltzia dolomitica - Kustatscher, p. 4.
2004 Voltzia dolomitica Wachtler et van
p. 175, pl. 14, fig. 1.
Localities: Prags / Braies, Wengen / La Valle, Schgaguler Alm / Malga Scagul, Puflatsch / Bullaccia,
Sappada.
2000a Voltzia ladinica Wachtler et van
figs. 1-5; pl. 11, figs. 1-4
2000b Voltzia ladinica Wachtler et van
figs. 1-5; pl. 11, figs. 1-4
2004 Voltzia ladinica Wachtler et van
p. 176-7, pl. 14, fig. 3.
Localities: Prags / Braies, Wengen / La Valle, Grödental / Val Gardena.
Voltzia pragsensis Wachtler et van Konijnenburg van Cittert, 2000
1953 Pagiophyllum cfr. foetterlei Stur - Leonardi,
p.19, pl. 4, fig. 6, 7, 9.
1986 Pagiophyllum cf. foetterlei Stur - Calligaris, p.
17, figs. A58.
1998 Voltzia sp. - Stingl & Wachtler, p. 79.
1999 Voltzia - Avanzini & Wachtler, p. 119.
2000a Voltzia pragsensis Wachtler et van
1-2.
2000b Voltzia pragsensis Wachtler et van
1-2.
2004 Voltzia pragsensis Wachtler et van
Konijnenburg - van Cittert – Kustatscher, p.
177-8, pl. 14, fig. 4.
Localities: Prags / Braies, Wengen / La Valle, Schgaguler Alm / Malga Scagul.
Voltzia zoldana Leonardi 1953
1953 Voltzia zoldana - Leonardi, p. 19, pl. 4, fig. 1
1968 Voltzia zoldana Leonardi - Leonardi, p. 176,
pl. 28, fig. 3.
2004 Voltzia zoldana Leonardi – Kustatscher, p.
178, pl. 14, fig. 5.
Localities: Spiz Agnelessa.
Voltzia ladinica Wachtler et van Konijnenburg van Cittert, 2000
Voltzia sp.
1999 Voltzia ladinica Wachtler et van
Konijnenburg - van Cittert (in stampa)Kustatscher, p. 52, pl. 4, fig. C.
38
1927 Voltzia sp. - Ogilvie Gordon, p. 69, pl. 8, fig. 8.
1953 Voltzia sp. - Leonardi, pl. 4, figs. 3, 8.
1953 Pagiophyllum (?) massalongi Zigno Leonardi, p. 18, pl. 4, fig. 2.
1964 Ramoscello di Brachyphyllum o Pagiophyllum
sp. - Leonardi, pl. 4, fig. 4.
1994 Ullmannia Broni – Costamoling &
Costamoling, p. 47, fig. 20.
2004 Voltzia sp. – Kustatscher, p. 178.
Localities: Prags / Braies, Wengen / La Valle, Seiser
Alm / Alpe di Siusi, Pufels / Bulla, Cercenà, Sappada.
4 Palaeoclimatic considerations
Macroflora
Most of the studied plant fossil collections are
composed of a few specimens only, collected in various and often not well-defined localities. However,
the main composition shows a dominance of
conifers, whereas cycads, pteridosperms, ferns and
horsetails occur only occasionally. Only one collection (in Bolzano) is composed of a higher number of
specimens (more than 150 specimens). Those plant
remains have been collected at two plant localities
near Braies / Prags (Seewald and Innerkohlbach) and
one near La Valle (Ritberg), belonging respectively
to the upper part of the Fernazza Formation
(Ritberg and Seewald) and to the base of the
Wengen / La Valle Formation (Innerkohlbach).
The pollen samples collected at those fossil-bearing horizons, attribute them to the secatus – vigens
phase sensu Van der Eem (1982), or to the
pseudoalatus-baculatus phase sensu Roghi (1995a,
b). Moreover, the plant deposits of Ritberg and
Innerkohlbach (Fig. 1) belong to the
Conbaculatisporites mesozoicus zone sensu Roghi
(1995), referred to the upper part of Neumayri
Subzone and to the base of Regoledanus Subzone
(Protrachyceras Zone, uppermost Longobardian).
The ammonoids (Lecanites glaucus, Protrachyceras
cf. ladinum, cf. Protrachyceras, “Eoprotrachyceras”
neumayri, cf. Joannites, cf. Mepinoceras and
Megaphyllites sp., det. P. Mietto) collocate the
localities to the Neumayri Subzone of the
Protrachyceras Zone (sensu Mietto & Manfrin,
1995). On the other hand, at Seewald no palynomorph zonal marker of Roghi’s scale has been
found.
Also
the
collected
ammonoid
(Macleanoceras sp., det. P. Mietto) permits to refer
the locality only to the Protrachyceras Zone
(Longobardian). However, the lithostratigraphic
attribution of the deposit to the Fernazza
Formation, narrows its age down to the upper
Longobardian (De Zanche et al., 1993) (for more
information see also Kustatscher, 2004).
The macrofossil collections, discussed already
partly in Kustatscher (1999, 2001, 2004), Wachtler &
van Konijnenburg – van Cittert (2002a, 2002b) and
Kustatscher et al. (2004), permit us to take a closer
look at the quantitative composition of the Upper
Ladinian macroflora (Fig. 2). All three plant localities
show a distinct dominance of the conifers (Voltzia,
Pelourdea). Also the pteridosperms (Ptilozamites) are
well represented in all three floras, whereas horseferns
(Cladophlebis,
tails
(Equisetites),
Gordonopteris) and cycadophytes (Pterophyllum,
Sphenozamites and Taeniopteris) are rare and occur
often only in one or two of the plant deposits.
This composition may be due to various factors
such as climate (aridity), edaphic (immature soils)
and taphonomy (i.e. selection due to transport).
Conifers are generally referred to arid environments due to their reduced leaf-surface, the thickness of their cuticles and the protection of their
stomata by papillae. On the base of these considerations, the composition of the Ladinian Flora from
the Dolomites might be referred to an arid climate
which the slightly imbricate pinnules of
Cladophlebis might indicate as well.
On the other hand, the fossil material is preserved within basinal sediments, and therefore, has
been subject to selection due to transport previously to its deposition. The high abundance of conifers
compared with the other groups (Innerkohlbach
and Seewald above 80%, Ritberg ca. 50%) could be
referred to selection caused by transport, as only
the more woody and resistant plants preserved after
the biostratinomic processes. However, the floral
composition cannot be explained exclusively by
means of taphonomy. The thickness of the cuticles
suggests also a certain degree of environmental
stress, related to adverse palaeoenvironment. This
could correspond to climatic or edaphic conditions.
The latter would suggest immature soils and shallow
water level. In this case the papillae on the stomata
might protect the stomata from salted sprays. On
the other hand, the presence of rare specimens of
ferns (Cladophlebis, Gordonopteris) and horsetails
(Equisetites), suggests the presence of restricted
humid microenvironments in the terrestrial habitats
as understorey and small ponds.
39
Microflora
The hypothesis of an arid climate during the
upper Ladinian is also in conflict with palynological
data available from literature. Van der Eem (1982)
suggests a progressive increase in humidity during
the Ladinian, opposed to the arid environmental
conditions at the end of the Anisian. These environments are however considered to be local, due to
the considerable amount of elements derived from
xerophytic plant-communities often present as well
(van der Eem, 1982, p. 72).
Additionally palynological data are known also
from the plant deposits (Kustatscher, 2004); in the
small outcrops of Seewald (SW) and Innerkohlbach
(IK) one pollen sample each has been studied, while
from the more extensive outcrop of Ritberg (RI) four
samples have been analysed.
Observing the main groups (spores, pollen grains,
algal cysts, acritarchs), Seewald is clearly dominated
by pollen grains, Innerkohlbach by spores whereas
in the Ritberg section an upwards increase of the
pollen fraction is observed (Table 2). These quantitative palynomorph fluctuations could be interpreted
both as climatic oscillations, and as variations in the
distance between the coast and the marine sedimentary environment, caused by sea level changes.
Applying the proposal of Visscher & Van der
Zwan (1981) for palaeoclimatic analysis, the palynomorphs have been divided into 15 groups (Table
3). Some of the groups such as A - monolete acavate spores, F – Porcellispora complex and J –
Samaropollenites complex are absent. Taxa, such as
Vallasporites ignacii and Enzonalasporites vigens,
referred by Visscher & Van der Zwan (1981) and van
der Eem (1982) to the vesicate pollen grains (M) are
now attributed to the (proto)monosaccate pollen
grains (N).
The pollen sample from Seewald (SW) is dominated by the Triadispora complex (L), trilete acavate
laevigate or apiculate spores (B) and alete
(proto)bisaccate pollen grains (I). Trilete laevigate or
apiculate spores (B), on the other hand, dominate
the Innerkohlbach (IK) sample. This would suggest a
more arid climate during deposition of the sediments corresponding with the Seewald plant
deposit, and a more humid climate when the
Innerkohlbach flora has been deposited.
Trilete laevigate or apiculate spores (B) dominate
also in the Ritberg outcrop. Furthermore, from
40
the bottom to the top of this section, the B
group, while still dominating, decreases in abundance. A concomitant increase of the Ovalipollis
complex (H, especially in RI 3), the Triadispora
complex (L) and alete (proto)bisaccate pollen
grains (I) can be observed. This would suggest an
increase of the aridity from the bottom to the top
of the section.
Also Abbink’ s palynomorph quantitative analysis
(1998) has been applied to the plant fossil deposits
(Table 5). Seewald shows a dominance of the
Coastal SEG, whereas Upland, Lowland, River and
Tidal SEGs are less abundant. At Innerkohlbach, on
the other hand, the more hygrophytic SEGs, such as
River and Lowland, dominate. However, as there is
only one sample per outcrop, no extended considerations can be deduced.
More information can be obtained from the
Ritberg section. This outcrop shows an upwards
increase of the Coastal and Tidal SEGs, while the
Lowland and Upland SEGs decrease. This trend can
be interpreted as an increase of the distance
between the coastal line and the area of plant
deposition (a transgression event) and thus it seems
to support that the palynomorph fluctuations may
be mostly due to sea level changes.
Observing in detail the Lowland SEG, the most
sensible one to climatic changes (Abbink 1998),
almost only taxa considered to be “more humid” can
be distinguished (Table 5). This suggests a prevailing
humid climate during the late Ladinian.
The hypothesis of sea level changes seems to be
confirmed also by the marine palynomorphs.
Although acritarchs and algal cysts are only additional elements (less than 20%), the acritarchs, considered as elements of open marine environments,
increase from the bottom to the top of the Ritberg
section, while algal cysts decrease (Table 2).
The hypothesis of taphonomic selection interacting with the Ladinian macrofloral deposition is supported also by the comparison between the abundance of the main groups (divisions) on macrofloristic and microfloristic levels (Table 4). The conifers,
represented by 50 to more than 80% in the
macroflora, never exceed 45% in the microflora
(max. 42,3 % at Seewald). Also pollen attributed to
the pteridosperms (2,6-17,9%) and cycads
(microflora 0-1,3%) are less abundant than the
macrofloral remains of these groups (respectively
7,8-28,3% and 0-10,9%).
On the other hand, ferns are much more important in the microflora (20,9 – 50,8%) than in the
macroflora (0-8,7%), becoming the most important
sporomorph group. This may be due to the high
fragility of the pinnate fern leaves, which are easily
destroyed during transport. Considering on the
other hand, that spores are generally underestimated in basinal sediments (Neves effect, Chaloner &
Muir, 1968) this dominance is even more important.
Additionally, the lycophytes are quite abundant
in the microflora with 3,3 to 17,2 %, while only one
macrofloral species attributed to the lycophytes
(Annalepis zeilleri) is known from the Ladinian of
the Dolomites. Spores (especially Uvaesporites),
however, are often preserved in tetrads probably
due to environmental stress of the mother-plants
(Looy et al., 2001). In any case, this abundance suggests that the lycophytes were better represented in
the Ladinian of the Dolomites than suggested by
the macrofloral remains alone.
Very abundant is also the genus Ovalipollis,
which botanical attribution is still unknown, as it
has been never found in situ.
Observing the separate plant localities in detail
(Table 4), Seewald is dominated by conifers (42,3%),
followed closely by ferns (20,9%) and pteridosperms
(17,9%). At Innerkohlbach, on the other hand, ferns
(50,8%) dominate among the lycophytes (17,1%)
and conifers (13,8%). At Ritberg, from bottom to
top lycophytes and ferns decrease in number
(respectively12,2 - 7,1% and 35,5 – 21,7%), whereas pteridosperms (5 – 13,2%) and conifers increase
(19,7 – 31,%).
Concluding, it can be suggested that the plants
grew in a general warm and humid local climate.
The high abundance of conifers and pteridosperms
and respectively low abundance of horsetails, ferns
and lycophytes in the macroflora seem to be more
due to local edaphic conditions and taphonomic
selection than to climate.
5 Palaeoenvironmental reconstructions
During the late Ladinian, the Southern Alps were
characterized by wide carbonate platforms bounded by more or less extended basins and were located north of an emerged land now buried under the
Po Plain („Southern Mobile Belt“ of Brusca et al.
1981).
Following the palaeogeographic reconstructions
of the uppermost Ladinian known from the literature (Assereto et al., 1977; Brusca et al., 1981;
Gianolla, 1993; Bosellini, 1996), Ritberg is situated
in a basin surrounded to the west by the carbonate
platforms of Putia / Peitler and Odle / Geißler and to
the northeast by the carbonate platform which
forms today the Piz da Peres. Southwards this basin
was bounded by the carbonate platforms of
Sassolungo / Langkofel, Sella, Tofane and
Marmolada. Additionally the volcanic complex of
Monzoni and Predazzo were exposed southwards as
well (Fig. 3). Some of these carbonate platforms and
the volcanic complex were subaerically exposed
during the time of deposition of the Fernazza
Formation and, therefore, subject to erosion (i.e.
Gianolla, 1993). The plant remains could have been
transported from the carbonate islands in the
northeast or west, or together with the volcanoclastic turbidites from the south.
Seewald and Innerkohlbach, on the other hand,
are positioned in a basinal environment west of the
Tre Cime di Lavaredo / Drei Zinnen and east of the
Piz da Peres platform. These platforms produced
carbonate sediments, whereas the terrigenous
material came from the south, from the volcanic
complex of Predazzo/Monzoni and perhaps also
from source areas more southwards than the
Valsugana line.
Considering the palaeogeographic reconstructions known from the literature and the paleoclimate discussed also in this article, the Ladinian
plants grew probably on more or less expanded carbonate or volcanic islands. On these islands various
environments developed: the coastal belt and the
so-called ‘hinterland’. The latter can be distinguished in more humid and more arid areas (Fig. 4).
The coastal environment (Fig. 5) was occupied
mainly by lycophytes (Annalepis) and pteridosperms
with thick cuticles (Ptilozamites). The Annalepis
scales were probably inserted on the top of some
centimetres high and thick stems with robust roots
(Grauvogel-Stamm & Lugardon, 2001), whereas
Ptilozamites was likely a shrubby plant, although no
reconstruction is so far known for this genus.
The hinterland, on the other hand, might have
been composed of ferns (Neuropteridium, Gordonopteris, Cladophlebis), cycads (Bjuvia, Spheno-
41
Fig. 3: Palaeogeographic reconstruction of the Dolomites during the late Ladinian (after Gianolla, 1993; Bosellini, 1996, mod.). RIposition of Ritberg, BR- position of the outcrops of Seewald and Innerkohlbach near Braies/Prags.
zamites), Bennettitales (Pterophyllum) and conifers
(Voltzia, Pelourdea).
Bjuvia is probably an arborescent form as discussed in the literature (Florin, 1933; Taylor &
Taylor, 1993), just as Pterophyllum (Mägdefrau,
1948; Kräusel & Schaarschmidt, 1966). Therefore,
these two taxa might have formed the canopy (Fig. 6)
of the more arid hinterland flora together with the
arborescent Voltzia, which, following Gall &
Grauvogel-Stamm (2000) could reach a height of
several meters. The shaded and more humid microenvironment of the understorey might have been
occupied by ferns of small to medium dimensions
such as Neuropteridium, but also some herbaceous
42
cycads such as Sphenozamites (Mägdefrau, 1948).
Additionally, also some shrubby conifers such as
Pelourdea might have grown in the understorey
(Mägdefrau, 1948; Seward, 1917, 1959).
In the more humid local environments (Fig. 7),
surrounding temporary ponds and swamps or along
a small river, larger ferns (Gordonopteris) with up to
50 cm long leaves could have grown together with
the above mentioned ferns of small to medium size
(Neuropteridium, Cladophlebis). Shrubby cycads
(Sphenozamites) and Bennettitales with higher
stems might also have inhabited the more humid
areas. Exclusively in this environments horsetails
(Equisetites), with heights of up to 6-8 m, might
Fig. 4: Reconstruction of a hypothetical environment of the Ladinian plants from the Dolomites. 1 – coastal belt, 2 – ‘hinterland’, 3
– more humid environments.
Fig. 5: Reconstruction of the coastal belt vegetation
with halophytic lycophytes such as Annalepis (1) and
shrubby pteridosperms such as Ptilozamites (2).
43
However, the outcrops of Seewald
and Innerkohlbach, since they consist of one horizon only, do not permit to extrapolate any climatic considerations. It is possible that the
increase of the spores and algal cysts
and decrease of pollen and
acritarchs at Innerkohlbach compared to Seewald is due to an
increase of humidity, or an approach
of the coastal line to the depositional area. The reduction of the
acritarchs in favour of the algal
cysts, however favours more the second hypothesis, variations of the sea
level, as would a comparison with
the sequence stratigraphy. The
Seewald outcrop is positioned at the
top of the Fernazza Formation, corresponding to the HST (Highstand
Systems Tract) of the depositional
sequence La3, composed of the basinal Zoppè Sandstone, the Acquatona
and the Fernazza Formation and the
Sciliar 3 platform (De Zanche et al.,
1993; Gianolla, 1993). Innerkohl bach, on the other hand, belongs to
Fig. 6: Reconstruction of the more arid ‘hinterland’ vegetation with herbaceous the base of the La Valle / Wengen
(Sphenozamites, 1) and arboreous cycads (Bjuvia, 4), high stemmed Bennettitales Formation, and is, therefore, corre(Pterophyllum, 3), shrubby (Pelourdea, 5) and arborescent conifers (Voltzia, 2).
sponding to the LST (Lowstand
Systems Tract) and TST (Transgressive
Systems Tract) of the following depositional
have grown as well (Frentzen, 1933; Mägdefrau,
sequence (Car1, sensu De Zanche et al., 1993;
1948, 1953; Kelber & Hansch, 1995; Kelber, 1999;
Gianolla, 1993), to which also the base of the S.
Gall & Grauvogel-Stamm, 2000).
Cassian Formation and the Cassian Dolomite 1 platform belong. The lowering of the sea level between
these two depositional events could be, therefore,
6. Discussion
the principal factor of the observed quantitative
variation between these two outcrops.
Quantitative variations of organic material (both
At the outcrop of Ritberg, on the other hand,
plant fossils and palynomorphs) within an outcrop
the four samples indicate an increase of pollen
depend on various factors. For those observed
grains throughout the section (Table 2), and also
between the three studied plant deposits two difan increase of the Coastal SEG, corresponding to a
ferent hypotheses have been proposed; climatic
decrease of the Lowland and River SEGs (Table 5).
oscillations of reduced time extension, or oscillaAlso in this case the most accredited hypothesis is
tions of the sea level and, therefore, of the relative
a transgression. This hypothesis is confirmed by
distance between the coast line and the point of
the increase of the acritarchs, especially at Ritdeposition.
berg 4 (Table 2). These (para)autochtonous marine
Throughout the Ritberg section and between the
palynomorphs seem quite sensible to bathymetric
Seewald and the Innerkohlbach sections, composiand salinity variations, but not to climatic variational variations have been observed (Tables 2-5).
tions.
44
7. Conclusions
The study of historical and inedited
material stored in various collections
of Italian and international Museums
and Institutions gives new insights
into the composition of the Ladinian
macroflora of the Dolomites.
The palaeoenvironmental reconstruction based on both macro- and
microfloral data shows more or less
expanded carbonate or volcanic
islands divided into various environments: the coastal belt and the socalled ‘hinterland’; the latter subdivided into more humid and more arid
areas.
Additionally, the integrated quantitative analyses (macro- and microfloral) suggest that the dominance of the
conifers results mostly from taphonomic selection. The flora probably
grew under environmental stress due
to salted spray, immature soils and
shallow water level, but in a locally
humid climate.
Quantitative palynological analysis
suggests also that the variations in
frequency between spores and pollen
or algal cysts and acritarchs are probably closer related to sea level changes
than to climatic changes. At present
the limited extensions of the fossil
horizons do not permit to exclude the
possibility of climate changes.
Fig. 7: Reconstruction of the more humid flora of the ‘hinterland’ environment
with high stemmed Bennettitales (Pterophyllum, 3), arboreous horsetails (Equisetites, 4) and herbaceous ferns (Neuropteridium, 1; Gordonopteris, 2) and cycads
(Sphenozamites, 5).
Acknowledgments
The systematic revision would not have been possible withouth the assistence of the various museums and institutions visited by one of the authors,
particularily by B. Baumgarten from the
Naturmuseum Bozen / Museo di Scienze Naturali
Alto Adige (Bolzano), the family Moroder from the
Museum de Gherdëina (Ortisei), P. Fedele and A.
Menardi from the Museo Paleontologico “R. Zardini”
(Cortina), F. Deltedesco from the Museo Ladino
Fodom (Livinallongo del Col di Lana), R. Pancaldi
from the Museo di Paleontologia e Preistoria P.
Leonardi (University of Ferrara), M. Fornasiero from
the Museo di Geologia e Paleontologia (University
of Padova), W. Resch and R. Brandner from the
Geologisch-Paläontologisches Institut (University of
Innsbruck), H.A. Kollmann from the Naturhistorisches Museum, F. Stojaspal from the
Geologische Bundesanstalt (both Vienna), H. Mayr
from the Paläontologisches Museum and T. Sperling
from the Geologisches Landesamt (both Munich).
Alberto Riva and Stefano Furin assisted during the
field work, Paolo Mietto determinated the
ammonoids found in the plant localities. We are particulary thankful to Renato Posenato and Guido
Roghi for ample discussions which permitted to
improve noticable the PhD-thesis on which this work
is based. The paleoenvironmental reconstructions
have been drawn by Mattia Guberti.
45
This work was supported by the “Progetto Giovani
Ricercatori 2001” with the titel “The terrestrial flora
from the Middle Triassic of the Dolomites: systematic, biostratigraphy and palaeoclimate”.
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Manuscript submitted: August 25, 2004
Manuscript accepted: March 11, 2005
47
48
Kustatscher, 2004
Kustatscher et al.,
2004
Wachtler & Van
Konijnenburg Van Cittert 2000a, b
Calligaris, 1986
Calligaris, 1983
G. Leonardi, 1964
Ogilvie Gordon,
1927
Mutschlechner, 1932
P. Leonardi,
1953, 1968
Mojsisovics, 1879
Annalepis zeilleri Fliche
Anomopteris mougeotii Brongniart
Asplenites roesserti Münster
Bjuvia dolomitica Wachtler et Van Konijnenburg-Van Cittert
cf. Bjuvia
Brachyphyllum sp.
Braiescycas leonardii Calligaris
Calamites meriani Brongniart
Chiropteris lipoldi Stur
Chiropteris pinnata Stur
Cladophlebis gaillardoti Brongniart
Cladophlebis leuthardti Leonardi,
Cladophlebis ruetimeyeri Heer
Cladophlebis ruetimeyeri Heer var. heeri Leonardi
Cladophlebis sp.
Dioonitocarpidium moroderi (Leonardi) Kustatscher et al.
cf. Cycadeoidea
Cordaicarpus sp.
Cycadeospermum sp.
Cycadites rectangularis Brauns
Danaeopsis marantacea (Presl) Schenk
Dioonitocarpidium sp.
Elatocladus sp.
Equisetites arenaceus (Jaeger) Schenk
cf. Equisetites
Equisetites sp.
?Equisetostachys
Fern incertis sedis
?Filicales indet.
Frenelopsis hoheneggeri Schenk
Ginkgo sp.
Lomatopteris sp.
Lycopodites sp.
?Neocalamites
Neuropteris elegans Brongniart
Neuropteris gaillardoti Brongniart.
Neuropteridium grandifolium (Schimper et Mougeot)
Schimper
Neuropteris ruetimeyeri Heer
Neuropteridium sp.
cf. Neuropteridium
Nilsonia sp.
Odontopteris sp.
Pagiophyllum foetterlei Stur
Pagiophyllum massalongi De Zigno
Pagiophyllum peregrinum (Lindley et Hutton) Seward
Pagiophyllum sp.
Pecopteris (Lonchopteris) reticulata Leuthardt
Pecopteris gracilis Heer
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
cf.
x
x
x
x
x
x
cf.
x
x
x
x
x
cf.
cf.
x
x
x
cf.
x
x
x
x
x
?
x
x
x
x
x
x
x
x
x
cf.
cf.
x
x
x
cf.
x
x
x
x
x
x
cf.
x
cf.
x
x cf.
x cf.
x
x
cf.
x
Pecopteris sulzensis Schimper
Pecopteris triascia Heer
Pecopteris sp.
Pelourdea vogesiaca (Schimper et Mougeot) Seward
Pelourdea sp.
Pterophyllum brevipenne Kurr
Pterophyllum giganteum Schenk
Pterophyllum jaegeri Brongniart
Pterophyllum sp.
?Pterophyllum sp.
Ptilozamites heeri Nathorst
Sagenopteris lipoldi Stur
Scolopendrites sp.
Sphenozamites wengensis Wachtler et Van Konijnenburg-Van
Cittert
Sphenozamites cf. bronnii Passoni et Van Konijnenburg-Van
Cittert
Taeniopteris angustifolia Schenk
Taeniopteris sp.
?Taeniopteris sp.
Thinnfeldia richthofeni Stur
?Thyrsopteris
Tingia sp.
Voltzia dolomitica Wachtler et Van Konijnenburg-Van Cittert
Voltzia cf. dolomitica Wachtler et Van Konijnenburg-Van
Cittert
Voltzia ladinica Wachtler et Van Konijnenburg-Van Cittert
Voltzia cf. ladinica Wachtler et Van Konijnenburg-Van Cittert
Voltzia pragsensis Wachtler et Van Konijnenburg-Van Cittert
Voltzia cf. pragsensis Wachtler et Van Konijnenburg-Van
Cittert
Voltzia recubariensis Schenk
Voltzia zoldana Leonardi
Voltzia sp.
?Voltzia
Zamites sp.
Sporofillo di cicadea o bennettitale
cf.
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
cf.
x
x
x
x
x
x
x
cf.
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
?
x
x
x
x
x
x
x
x
x
x
x
Tab. 1. Plant fossils of Ladinian age described and figured in the literature (Mojsisovics, 1879; Ogilvie Gordon, 1927; Mutschlechner, 1932; P. Leonardi, 1953, 1968; G. Leonardi, 1964; Calligaris, 1983, 1986; Wachtler & van Konijnenburg - van Cittert
2000a, b; Kustatscher 2004; Kustatscher et al., 2004).
SW %
IK %
RI 1 % RI 2 % RI 3 % RI 4 %
spores
40.42
68.86
44.52
46.13
33.28
35.95
pollen
59.58
31.14
55.48
53.87
66.72
64.05
Algal cysts
1.48
3.99
4.75
7.83
3.35
2.85
acritarchs
7.94
0.74
5.73
1.60
5.71
15.25
Tab. 2: Relative abundance of the main palynomorph groups (SW = Seewald, IK = Innerkohlbach, RI 1-4 = Ritberg).
49
A - monolete acavate spores
SW %
IK %
0.00
0.00
0.00
0.00
0.00
0.00
66.53
47.20
47.09
30.74
29.49
1.77
1.44
2.69
0.67
1.15
0.82
2.24
0.00
0.40
1.98
B - trilete acavate laevigate
24.84
or apiculate spores
C - trilete acavate murornate
1.27
spores
D - trilete cingulate and
0.85
zonotrilete spores
RI 1 % RI 2 % RI 3 % RI 4 %
E - Aratrisporites group
0.21
0.14
0.64
0.45
0.13
0.33
F - Porcellispora complex
G - monosulcate pollen
grains
0.00
0.00
0.00
0.00
0.00
0.00
0.21
0.14
0.00
0.15
0.00
1.48
H - Ovalipollis complex
13.59
13.47
22.72
20.48
28.05
18.95
I - alete (proto)bisaccate
pollen grains
21.02
5.58
7.52
7.92
12.21
12.36
J - Samaropollenites
0.00
0.00
0.00
0.00
0.00
0.00
K - taeniate (proto)bisaccate
pollen grains
3.61
0.14
1.28
0.60
1.48
2.47
L - Triadispora complex
31.00
6.53
9.12
13.90
20.40
26.36
M - vesicante pollen grains
N - (proto)monosaccate
pollen grains
0.00
0.00
0.00
0.00
0.00
0.00
2.97
2.18
4.64
5.08
3.49
3.13
O – Circumpolles group
0.42
2.72
3.20
1.64
2.42
2.31
Tab. 3. Palynological composition of the palynomorph groups proposed by Visscher & van der Zwan (1981); SW = Seewald, IK =
Innerkohlbach, RI 1-4: Ritberg.
50
SW 1%
Upland
8.4
Lowland
11.0
Coastal
34.0
River
13.4
Tidal
12.6
Ovalipollis
13.9
Not attributed
6.7
Lowland
“more humid”
“more arid”
SW
10.82
0.22
IK 1%
5.5
29.1
9.0
37.9
1.9
13.5
3.0
RI 1%
7.4
23.7
14.6
23.9
2.9
22.8
4.8
RI 2%
7.3
23.8
16.3
24.9
5.1
20.5
1.9
RI 3%
6.6
14.5
25.7
16.3
6.2
28.1
2.6
RI 4%
4.5
16.4
29.4
17.9
7.9
19.0
5.0
IK
27.22
1.92
RI 1
19.23
4.49
RI 2
20.39
3.45
RI 3
13.32
1.21
RI 4
13.22
3.14
Tab. 4. Abundance of the main floral groups within the microflora; SW = Seewald, IK = Innerkohlbach, RI 1-4: Ritberg.
Lycophyta
Sphenophyta
Pteridophyta
Pteridospermae
Cycadophyta
Ginkgophyta
Ovalipollis
Coniferophyta
altro
SW %
3.3
0.0
20.9
17.9
0.0
0.0
13.9
42.3
1.7
IK %
17.1
0.1
50.8
2.6
0.1
0.1
13.5
13.8
1.8
RI 1 % RI 2 % RI 3 % RI 4 %
12.2
16.9
7.4
7.1
0.5
0.1
0.1
0.2
35.5
32.5
23.1
21.7
5.0
5.7
9.4
13.2
0.8
0.0
0.1
1.3
0.3
0.1
0.1
1.2
22.8
20.5
28.1
19.0
19.7
22.9
30.4
31.8
3.2
1.0
1.1
4.5
Tab. 5. Relative abundance of the different SEGs within the plant deposits; SW = Seewald, IK = Innerkohlbach, RI 1-4: Ritberg.
51
Geo.Alp, Vol. 2, S. 53–60, 2005
ITALIAN FOSSIL CHIROPTERAN ASSEMBLAGES: A PRELIMINARY REPORT
Cristiana Tata & Tassos Kotsakis
With 2 figures and 1 table
Dipartimento di Scienze Geologiche, Università Roma Tre, L.go S. L. Murialdo 1, 00146 Roma Italy;
e-mail: [email protected], [email protected]
Abstract
This work is a preliminary report on Italian fossil chiropteran faunas. During the Paleogene just one sample of Early Oligocene age, pertaining to an extinct species, has been reported. A few findings have been
reported from the Neogene. Just one complete assemblage from the Late Miocene site of Brisighella has been
examined and has allowed palaeoecological inferences, whilst specimens from Late Miocene localities of
Baccinello V0 (Tuscany) and Gargano peninsula (Apulia) need a revision. A Late Pliocene assemblage has been
collected in Montagnola Senese (Tuscany) but it still needs a systematic revision.
During the Quaternary and most of all since the Middle Pleistocene the fossil record becomes richer. Some
assemblages testify a Mediterranean climate analogous to the present one. The most significant are: the Early
Pleistocene ones from Pirro Nord (Apulia) and Ghar Dalam Cave (Malta), the early Middle Pleistocene ones
from Slivia (Venezia Giulia) and Spinagallo Cave (Sicily) and the Late Pleistocene ones from Punta Padre Bellu
(Sardinia) and Breuil Cave (Latium). In other cases the species represented in the assemblages are typical of
colder climate and then they make it possible to infer cooler conditions in Italy during some periods. Good
examples in this sense are the Middle Pleistocene assemblage from Vento Cave (Marche) and the Late
Pleistocene one from Cittareale Cave (Latium). A distributuion chart of all fossil bats from Italy and Malta is
also presented.
Introduction
Nowadays it is quite common to support
palaeoenvironmental reconstructions using samples
from fossil mammal (especially micromammal)
assemblages as palaeoecological and/or palaeoclimatic markers.
Among micromammals bats are really meaningful in this respect but, especially in Italy, although
when they are found they are really abundant
(especially from Pleistocene sites), they are often
lacking. The lack of interest in this group is caused
by the bradytelic evolution of these animals that
makes them useless for biochronological studies
that, in the past decades, have been attracting
palaeontologists attention.
Anyway it has to be underlined that bats,
because of their peculiar ecological habits are
strongly influenced in their distribution by climatic and ecological parameters and this is why they
can be considered as good environmental markers.
In addition, just because of the low rates of evolution, living species are mostly analogous to fossil
ones. Since the present distribution and the climatic context of their life are known it is reasonable to make palaeoclimatic and palaeoecological
inferences from studying species pertaining to fossil assemblages. However first of all it is necessary
to review chiropteran assemblages and this work
represents a preliminary approach to this research
project.
Tertiary chiropteran assemblages
and their palaeoecological meaning
Just one species of bat is known from Paleogene
sediments in Italy: Archaeopteropus transiens
Meschinelli, 1903. It has been collected in the early
Oligocene (MP 21) lignites of Monteviale (Veneto)
53
Fig.1: Map of main fossiliferous localities of Italy and Malta.
1- Monteviale (Veneto); 2- Baccinello V0 (Tuscany); 3- Brisighella (Romagna); 4- Gargano (Apulia); 5- Montagnola Senese
(Tuscany); 6- Pirro Nord (Apulia); 7- Ghar Dalam Cave (Malta);
8- Spinagallo Cave (Sicily); 9- Slivia (Venezia Giulia); 10- Vento
Cave (Marche); 11- Punta Padre Bellu (Sardinia); 12- Breuil
Cave (Latium); 13- Monte Cucco Cave (Marche); 14- Cittareale
Cave (Latium).
(Meschinelli, 1903; Kotsakis et al., 1997) (Figs. 1, 2).
It is a large chiropteran classified in its own subfamily, Archaeopteropodinae, and considered by
some authors (Russel & Sigé, 1970) to belong to the
suborder Microchiroptera and by others (Smith &
Storch, 1981) to the suborder Megachiroptera.
Unfortunately the original sample has been lost
during the Second World War and only some rather
good casts are available. Archaeopteropus was part
of an assemblage that, if considered as a whole,
shows a tropical character.
In the Italian Miocene the presence of Chiroptera
indet. has been signalled from clays of Baccinello V0
(Tuscany - MN11) (Kosakis et al., 1997). Just one
Miocene assemblage is known, coming from
Monticino Quarry (Brisighella, Romagna) karst fissures. The assemblage is of Late Turolian age (MN13)
and it is composed of six species: Megaderma cf. M.
mediterraneum Sigé, 1974, Rhinolophus cf. R.
kowalskii Topál, 1979, Rhinolophus sp., Hipposideros
(Syndesmotis) cf. H. (S.) vetus (Lavocat, 1961),
Asellia cf. A. mariaetheresae Mein, 1958 and Myotis
cf. M. boyeri Mein, 1964 (Kotsakis & Masini, 1989).
Three species, the two rhinolophids and the vespertilionid, are similar to living forms now inhabiting
54
this same site. On the other hand the remaining
three genera, Megaderma, Hipposideros and Asellia
now live in tropical and subtropical areas. In particular the presence of Megaderma is indicative of
minimum temperatures higher than 14-15°C all
around the year, while the presence of Asellia is
indicative of subdesertic conditions (Sigé, 1974).
From the species represented here a littoral sandy
habitat has been inferred (Kotsakis & Masini, 1989).
Another finding from the Italian Miocene is from
Gargano Peninsula (Apulia) from karst fissure fillings characterized by the Hoplitomeryx and
Microtia assemblage and ascribed to the Late
Miocene – ?Early Pliocene. Here a single species, not
definitely studied yet, has been collected and previously ascribed to the genus Megaderma but an attribution to other megadermatid genera is possible
(Kotsakis et al., 1997). As in the preceding case the
presence of this genus, now inhabiting hot regions,
has suggested tropical temperatures in this area.
Another Neogene assemblage is from the Late
Pliocene (MN17, Middle Villafranchian Mammal Age
or Late Villanyian Micromammal Age, Costa San
Giacomo Faunal Unit/Olivola Faunal Unit; Kotsakis
et al., 2003) fissure fillings of Montagnola Senese
(Tuscany) (Fondi, 1972). It includes four species:
Myotis blythii (Tomes, 1857), M. gr. schaubi Kormos,
1934 – rapax Heller, 1936, Myotis sp. and ? Tadarida
sp. The fauna needs a systematic revision and it is
impossible to infer palaeoecological informations.
Quaternary chiropteran assemblages
and their palaeoecological meaning
During the Pleistocene, particularly in the Late
Pleistocene, an increase in the Italian fossil record is
observed. Among various sites under study at present the most meaningful are Pirro Nord (Gargano,
Apulia), Spinagallo Cave (Sicily), Punta Padre Bellu
(Alghero, Sardinia), Breuil Cave (Monte Circeo,
Latium), Monte Cucco Cave (Perugia, Umbria) and
Cittareale Cave (Rieti, Latium). One assemblage from
the late Early Pleistocene/earliest Middle Pleistocene from the Ghar Dalam Cave (Malta) is strongly
related to Italian faunas.
The oldest chiropteran assemblage is Pirro Nord,
ascribed to the early Pleistocene (Late Villafranchian
M.A. or Early Biharian Micromammal Age, Pirro F.U.)
(Gliozzi et al., 1997). A rich assemblage has been collected from one of the karst fissures in the area. It is
Fig. 2: Biochronological scheme of localities bearing fossil bats of Italy and Malta.
55
composed of six species: Rhinolophus ferrumequinum (Schreber, 1774), R. birzebbugensis Storch,
1974, Myotis blythii, M. capaccinii (Bonaparte,
1837), Miniopterus schreibersi (Kuhl, 1819) and
Miniopterus n. sp. (Masini et al., 1996; Tata, 2003).
Among them the living species R. ferrumequinum,
M. blythii, M. capaccinii and M. schreibersi are present in Europe in the central and southern part of
the continent (except for R. ferrumequinum that
extends to more northern latitudes). The remaining
two species R. birzebbugensis and Miniopterus n. sp.
cannot be considered as strong palaeoclimatic markers since the rhinolophid is known as a fossil only
from a few localities (Malta, Bulgaria and probably
Spain) (Storch, 1974; Popov, 2004; Tata & Kotsakis,
in prep.) while the miniopterid has been collected
here for the first time. Considered as a whole the
assemblage has a strong Mediterranean character. A
different assemblage including three species has
been collected from another fissure filling in the
same area (De Giuli & Torre, 1984): Rhinolophus gr. R.
euryale Blasius, 1853, Myotis cf. M. blythii and Myotis
sp. (small size). From a climatic point of view the
assemblage does not differ from the previous one.
Close in age to the previous assemblage is that
collected in the Ghar Dalam Cave (strata with Leithia
cartei; Storch, 1974) including ten species:
Rhinolophus hipposideros (Bechstein, 1800), R.
birzebbugensis, R. blasii Peters, 1866, Myotis exilis
Heller, 1936, M. bechsteini robustus Topál, 1963, M.
ghardalamensis Storch, 1974, M. capaccinii,
Eptesicus praeglacialis Kormos, 1930, Pipistrellus
pipistrellus (Schreber, 1774) and Miniopterus
schreibersi. The assemblage shows a Mediterranean
character with forested and open habitats and the
presence of fresh water.
Also the assemblage from the Spinagallo Cave,
ascribed to the early Middle Pleistocene (Elephas falconeri Faunal Complex) (Bonfiglio et al., 2003) is
quite rich including ten species: R. ferrumequinum,
R. hipposideros, R. mehelyi Matschie, 1901, R. cf. R.
blasii, Myotis mystacinus (Leisler in Kuhl, 1819), M.
bechsteini (Leisler in Kuhl, 1819), M. capaccinii,
Eptesicus serotinus (Schreber, 1774), Barbastella barbastellus (Schreber, 1774) and M. schreibersi (cfr.
Kotsakis & Petronio, 1980). Very probably the new
species of Miniopterus collected in Pirro Nord is also
present in the Spinagallo Cave assemblage.
The species are derived from two different strata;
in the lower one only three species are represented:
M. schreibersi, R. ferrumequinum and M. capaccinii,
56
the first being decisely dominant. In the higher stratum all the species, except M. capaccinii, are represented. M. schreibersi is always dominant although
less numerous than in the lower stratum. On the
whole the assemblage has a Mediterranean character although forms such as M. mystacinus and B.
barbastellus usually have a more northern distribution. All but one species are still living in Sicily, M.
bechsteini and B. barbastellus are less widespread
then the others (Kotsakis & Petronio, 1980). R. blasii
has a recent eastern Mediterranean distribution and
is present in the easternmost province of Italy near
the Italian-Slovenian boundary (Lanza & Agnelli,
1999). Palaeoecological conditions similar to those
of Ghar Dalam can be inferred.
At several fossil localities a small number of bat
species has been collected; usually the specimens
represented there belong to recent species now living in the same areas. A good example is the early
Middle Pleistocene (Early Galerian M.A., Slivia F.U.)
assemblage collected in Slivia karst fissure (Trieste,
Venezia Giulia) where two species have been recognized: Rhinolophus ferrumequinum and Miniopterus schreibersi (cfr. Ambrosetti et al., 1979). In
other cases some elements indicating colder conditions have been recognized in mammal assemblages
as for example in the late Middle Pleistocene (Early
Aurelian M.A.) deposit from Vento Cave (Ancona,
Marche), where two species are found: Rhinolophus
ferrumequinum and Myotis dasycneme (Boie, 1825)
(Esu et al., 1990). Among them particularly significant in this respect is the presence of M. dasycneme,
typical of cold conditions, known at present by just
one erratic individual in the north-eastern part of
the Italian peninsula .
In the Late Pleistocene the findings are abundant, isolated remains are often reported
(Lombardy, Santi, 2000; Sardinia, Abbazzi et al.,
2004), but systematic analysis has been conducted
only on a few cave deposits.
The assemblage from Punta Padre Bellu, collected in a destroyed cave near Alghero, has been
ascribed to the Late Pleistocene and is composed of
six species: R. ferrumequinum, R. hipposideros,
Myotis myotis (Borkhausen, 1797), M. capaccinii,
Nyctalus cf. N. lasiopterus (Schreber, 1780) and M.
schreibersi (cfr. Kotsakis, 1987). All the species with
the exeption of N. lasiopterus (that however is quite
rare in the peninsula today) are still living in
Sardinia suggesting a climatic context similar to the
present one.
X
Recent
X
X
X
Late Pleistocene
Middle Pleistocene
Late Pliocene
Late Miocene
Early Pleistocene
Archaeopteropus transiens+
Megaderma mediterraneum+
Megaderma (s.l.) sp.+
Rhinolophus kowalskii+
Rhinolophus ferrumequinum
Rhinolophus euryale
Rhinolophus birzebbugensis+
Rhinolophus mehelyi
Rhinolophus blasii
Rhinolophus hipposideros
Rhinolophus sp.+
Hipposideros vetus+
Asellia mariaetheresae+
Myotis boyeri+
Myotis bechsteini
Myotis bechsteini robustus+
Myotis myotis
Myotis ghardalamensis+
Myotis blythii
M. nattereri
Myotis gr. M. schaubi-M. rapax+
Myotis emarginatus
Myotis exilis+
Myotis mystacinus
Myotis brandti
Myotis daubentoni
Myotis capaccinii
Myotis dasycneme
Myotis sp.
Barbastella barbastellus
Plecotus auritus
Plecotus austriacus
Plecotus sp.
Pipistrellus pipistrellus
Pipistrellus nathusii
Pipistrellus kuhlii
Pipistrellus sp.
Hypsugo savii
Nyctalus leisleri
Nyctalus noctula
Nyctalus lasiopterus
Amblyotus nilssonii
Eptesicus praeglacialis+
Eptesicus serotinus
Vespertilio murinus
Miniopterus n. sp. +
Miniopterus schreibersi
Tadarida teniotis
Tadarida sp.
Early Oligocene
Bat species
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
?
X
X
X
X
X
X
X! *
X
cf.
X
cf.
X
X
X
X
X
X
X
cf.
cf.
cf.
* It has been reported just one specimen captured in Northern Italy in 1881.
X
X
?
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
?
X
cf.
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
?
X
X
14
Tab. 1: Distribution chart of fossil bats from Italy and Malta. + = extinct species or subspecies. Pipistrellus pygmaeus, Myotis punicus,
Plecotus alpinus and Plecotus n. sp. are not included among living species.
57
The rich assemblage from Breuil Cave (Monte
Circeo, Latium) collected in two strata (stratum “e”
and stratum “d”) must be referred to the Late
Pleistocene (OIS 3). Among micromammals bats are
well represented with five species: R. ferrumequinum, M. myotis, Nyctalus noctula (Schreber,
1774), M. schreibersi and Tadarida teniotis
(Rafinesque, 1814). All these species are present in
the lower part of the stratum “e”, while only R. ferrumequinum is represented in the upper part of the
stratum “d” (Kotsakis, 1989). The assemblage
derived from stratum “e” is constituted partly by
typical Mediterranean species such as M. schreibersi and T. teniotis (that are more aboundantly represented) and partly by species having a more northern distribution such as N. noctula. If this assemblage is considered in the general faunal context it
becomes quite clear that its interstadial character
denotes a woodland environment with moist areas
in the neighbourhood of the cave.
Another Late Pleistocene (OIS 2) chiropteran
assemblage is that from the Monte Cucco Cave
(Perugia, Umbria) (Capasso Barbato & Kotsakis,
1986), including five species: R. ferrumequinum, M.
myotis, M. blythii, M. bechsteini and M. emarginatus (E. Geoffroy, 1806). The absence of Miniopterus
schreibersi is interesting because it is a usual component of Italian cave- dwelling faunas. The assemblage does not show any peculiar characteristics
that allow palaeoclimatic inferences.
The chiropteran assemblage from Cittareale Cave
(Rieti, Latium) is clearly colder and ascribed to the
Late Pleistocene (OIS 2, Younger Dryas?); five
species are present: R. ferrumequinum, R. hipposideros, M. myotis, M. bechsteini and M. dasycneme (cfr. Argenti et al., in press). Particularly
meaningful in a climatic sense is the presence of M.
dasycneme that suggests the attribution of the
assemblage to a cold interval, presumably to the
Younger Dryas period. In addition all the species,
with the exeption of M. myotis that usually prefers
open and slightly wooded terrain, are common in
wooded areas suggesting then, for the assemblage,
a forested environment with open space and ponds.
been increased in the last years by new researches:
Pipistrellus pygmaeus (Leach, 1825) (Russo & Jones,
2000); Myotis punicus Felten, 1977 (Castella et al.,
2000; Beuneux, 2004); Plecotus alpinus Kiefer &
Veith, 2001 (Trizio et al., 2003); Plecotus n. sp. from
Sardinia (Mucedda et al., 2002) have been added in
the list of bats of Italy. However for an attribution
of Italian fossil material to these species a complete
systematic revision is necessary.
The number of Italian Tertiary fossil species is
much less; it has been calculated to include 12
species, among them 11 are surely extinct, but also
the twelwth, which has been attributed to a living
species, needs a systematic revision.
During the Quaternary an increase in the number
of species is observed with at least 31 represented;
5 of this number are extinct (R. birzebbugensis, M.
ghardalamensis, M. exilis, E. praeglacialis and
Miniopterus n. sp.). A fossil subspecies has also been
reported M. bechsteini robustus. Another species
has to be mentioned pertaining to the genus
Rhinolophus, R. botegoi Regàlia, 1893 described by
Regàlia (1893), from fossil remains collected in
Colombi Cave (Palmaria Island, Liguria). Its validity
seems to be improbable, but in any case the material needs to be revised. In the fossil record of Italian
bats the presence of troglophilous species is dominant, whilst non-cave dwelling species are not well
represented (see Table 1).
The analysis of the chiropteran assemblages confirms that during the time span between the
Miocene and Pleistocene the Italian peninsula has
been subjected to a general decrease of temperature. This inference comes from the observation
that species typical of tropical and subtropical environments present in the Neogene assemblages are
completely lacking from more recent assemblages.
It has to be emphasised that in some cases the presence of a single species with peculiar ecological
requirements gives clear palaeoecological information whilst in other cases it is the assemblage as a
whole (considering the percentage composition of
each single species) that allows palaeoecolgical
inferences.
Conclusions
Acknowledgments
Among the recent mammalian faunas of Italy
and Malta, Mitchell-Jones et al. (1999) indicate the
presence of 28 species of bats. This number has
We wish to thank Prof. G. Tichy of Salzburg
University for reviewing the manuscript and Dr.
D. Harrison of Harrison Institute for the helpful dis-
58
cussions about bat systematics with the first author
and the correction of the English.
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Revised manuscript accepted: March 30, 2005
Geo.Alp, Vol. 2, S. 61–70, 2005
CERVUS ELAPHUS SICILIAE FROM PLEISTOCENE LACUSTRINE DEPOSITS
OF ACQUEDOLCI (NORTH-EASTERN SICILY, ITALY) AND ITS TAPHONOMIC SIGNIFICANCE
Gabriella Mangano
Dipartimento di Scienze della Terra, Università degli Studi di Messina; e-mail: [email protected]
Abstract
Systematic excavations carried out on the Pleistocene lacustrine deposits of Acquedolci (North-Eastern
Sicily, Italy) yelded a very rich fossil vertebrate assemblage, containing thousands of remains of
Hippopotamus pentlandi, the endemic hippo of the Siculo-Maltese archipelago, associated with remains of
Cervus elaphus siciliae and scarce remains of Ursus cf. arctos, Canis lupus, Testudo cf. hermanni, Elephas
mnaidriensis and Aves. This paper presents a morphological, biometrical and taphonomical analysis of Cervus
elaphus siciliae remains. Morphological and biometrical features are in the range of the variability of Cervus
elaphus siciliae Pohlig, the endemic deer of Sicily which is characterized by a slightly smaller size compared
to the populations of the Italian peninsula. Taphonomical features, such as spatial distribution and orientation of the remains, composition of the skeletal part, age distribution, degree of skeletal articulation, fragmentation and bone modification, indicate that Cervus elaphus siciliae remains did not accumulate “in situ”,
unlike the autochthonous remains of Hippopotamus pentlandi, but probably they were occasionally deposited in the lacustrine basin as fragments of carcasses belonging to the animals inhabiting the surrounding
area.
Introduction
The lacustrine sediments of Acquedolci are
located on the northern flank of the Nebrodi range
(North-Eastern Sicily), at the base of the high vertical cliff of the Pizzo Castellaro carbonatic massif,
on which the well-known S. Teodoro Cave opens.
The deposit is composed of silt, gravel and pebbles
of variable size, probably fallen from the adjacent
cliff. It is superimposed on a Late Pleistocene
marine terrace located 131 m a.s.l. and represents
the remains of a late Middle Pleistocene lacustrine
basin (Bonfiglio, 1985; 1987; 1989; 1992). During
the years 1982-1987 systematic excavations were
carried out and seven trenches of different width
and depth have been excavated over an area of
104 m2 (Fig. 1). About 130 m3 of sediments were
removed and the entire succession of the deposit,
which was originally about 14 m thick, was investigated. In trench G the lacustrine sediments containing fossil remains are absent.
Most of the collected remains come from the
trench F, which was deepened for 6 m. In the other
trenches, which have a maximum depth of about 2
m, the fossil bones have been partially preserved in
situ because of their spectacular abundance, in
order to establish a field Museum.
A total number of 3.016 remains of Hippopotamus pentlandi, the endemic hippo of the
Siculo-Maltese archipelago, together with 104
remains of the endemic deer of Sicily, Cervus elaphus siciliae, and very scarce remains of Ursus cf.
arctos (15), Canis lupus (7), Testudo cf. hermanni
(6), Elephas sp. (1) and Aves (2) were collected
(Bonfiglio, 1995). One of the two remains of Aves
belongs to Gyps melitensis Lydekker, an extinct vulture (griffon) species (Pavia, 2001).
This faunal assemblage belongs to the “Elephas
mnaidriensis Faunal Complex”, one of the five
Pleistocene faunal complexes recognized in Sicily,
referred to the late Middle Pleistocene-early Late
Pleistocene (Bonfiglio et al., 2001; 2002). Amino-
61
Skeletal element
skull
antler
vertebrae
ribs
scapula
humerus
radius
femur
tibia
podials
metapodials
phalanges
Total
N.R.
3
23
7
15
1
4
11
2
4
5
22
7
104
Tab. 1: Composition of the skeletal part of Cervus elaphus
siciliae remains from Acquedolci.
Fig. 1: Topography of the Acquedolci area and location of the
excavation trenches (A-G) (modified from Bonfiglio, 1987).
acid racemization dating yielded an age of 200 + 40
Ky for the Hippopotamus pentlandi remains of
Acquedolci (Bada et al., 1991).
Morphological and biometrical descriptions
Fig. 2: Right shed antler of Cervus elaphus siciliae, internal
view.
A total number of 104 strongly fragmented
remains of Cervus elaphus siciliae were collected.
The only complete and well preserved bones are
represented by two metacarpals. Antlers and
metapodials are the most frequent skeletal elements
(Tab. 1). A morphological and biometrical comparison with the remains of Cervus elaphus siciliae
Pohlig from different Pleistocene deposits of Sicily,
described by Gliozzi et al. (1993), is presented. At
present, the data published by Gliozzi et al. (1993)
about the remains of Cervus elaphus siciliae from
Sicily are the only available ones. The remains from
Acquedolci do not have a catalogue number.
ANTLER
transverse diameter of the burr
antero-posterior diameter of the burr
transverse diameter of the beam above the bez-tine
antero-posterior diameter of the beam above the bez-tine
right
47
63
37
42
(Gliozzi et al., 1993)
–
min 51 – max 67
–
min 34 – max 44
Tab. 2: Measurements (mm) of the antler of Cervus elaphus siciliae from Acquedolci compared with the dimensions of antlers
described by Gliozzi et al., 1993.
62
SCAPULA
transverse diameter of the glenoid cavity
antero-posterior diameter of the glenoid cavity
antero-posterior diameter of the neck
antero-posterior diameter of the articulation surface
right
33
37
31
49
Tab. 3: Measurements (mm) of the scapula of Cervus elaphus siciliae from Acquedolci.
Fig. 3: Right distal humerus of Cervus elaphus siciliae; a) anterior view, b) posterior view.
Skull. The skull remains are represented by 3
pedicle fragments only. The most complete of these
bones is a left pedicle, which is rather short and
strong. The antero-posterior diameter is 44 mm,
while the transverse diameter is 41 mm. The skulls
of Cervus elaphus siciliae collected in the Puntali
Cave (Palermo) have antero-posterior diameters of
the pedicles varying between 34.8 and 40.6 mm,
and the transverse diameter of the pedicles ranging
HUMERUS
transverse diameter of the distal end
antero-posterior diameter of the distal end
transverse diameter of the trochlea
between 33.4 and 41 mm (Gliozzi et al., 1993).
Another fragment of skull from the Villafranca
Tirrena deposit (Messina) has an antero-posterior
diameter of the pedicle measuring 43 mm
(Mangano, 2000).
Antlers. A total number of 23 antler fragments
were recovered: 7 fragments of tines, 9 fragments
of beams and 7 shed antler fragments with burr. The
only measurable remain is a right shed antler fragment, which was strongly fractured and reconstructed by restoration (Fig. 2). The burr and the
first portion of the beam are preserved, the browtine and bez-tine are broken. The burr is moderately developed and formed by little pearls. The
approximate measurements of this specimen are
listed in Tab. 2. The dimensions of the antero-posterior diameter of the burr and of the beam above the
bez-tine are in the range of the variability of Cervus
elaphus siciliae (Gliozzi et al., 1993).
Vertebrae. 6 vertebrae are present. They are fractured and incomplete. Two fragments belong to
young individuals.
Ribs. 15 fragments of ribs lacking the articulation
surface were recovered.
Scapula. The scapula is represented only by one
proximal right fragment (Tab. 3). The glenoid cavity
is slightly ovoidal in shape with a well developed
concave surface. The glenoid tubercle is very strong.
The neck is rather short and slender. The remains of
scapula of Cervus elaphus siciliae recovered in the
Fata Donnavilla Cave (Messina) display the same
morphological features (Gliozzi et al., 1993).
Humerus. The humerus is poorly represented by 4
fragmentary specimens: 2 distal fragments preservright
48
49
45
left
42
43
40
min 40 – max 49.2
min 37.5 – max 46
–
Tab. 4: Measurements (mm) of the humerus of Cervus elaphus siciliae from Acquedolci compared with the dimensions of the remains
63
Fig. 5: Tibia of Cervus elaphus siciliae; a) left proximal fragment, posterior view; b) left distal fragment, posterior view.
Fig. 4: Semicomplete right radius of Cervus elaphus siciliae;
a) anterior view, b) posterior view, c) proximal articulation.
ing the articulation surface, 1 small fragment of the
distal articulation and 1 fragment of the shaft. The
diaphysis seems to have a great torsion. The olecranon fossa is deep and triangular in shape; the
trochlea is developed and medially inclined (Fig. 3,
a-b). The transverse diameter of the distal end is
within the range of the values of Cervus elaphus
siciliae (Gliozzi et al., 1993) while the antero-poste-
64
rior diameter of the distal end is slightly larger
(Tab. 4).
Radius. 10 remains of radius were recovered,
including 1 semicomplete right radius with a broken
distal end (Fig. 4, a-b-c), 5 proximal fragments and
4 distal fragments including one juvenile remain.
The posterior face of the diaphysis has a deep radioulnar groove. The proximal articulation surface is
sub-rectangular with a wide sigmoid notch which
separates it into two very unequal articulation
facets, whose medial one is very large. Most of the
remains, particularly the semicomplete right radius,
have a larger size than those described by Gliozzi et
al. (1993) (Tab. 5). Since other biometric data on
Cervus elaphus siciliae are lacking in the literature,
these differences in dimensions at present cannot
be correctly evaluated.
Femur. Only 2 femur fragments are present: 1
fragment of the proximal articulation (head) and 1
fragment of the distal articulation. The head is not
fused. The condyles of the distal articulation are less
RADIUS
greatest length
transverse diameter
of the proximal end
antero-posterior diameter
of the proximal end
transverse diameter
at half length of the shaft
transverse diameter
of the distal end
of the distal end
right
270
54
right
44
right
42
left
50
right
-
right
-
left
-
min 206 – max 237
min 39 – max 44.7
29
23
25
27
-
-
-
min 21 – max 24.5
31
20
-
-
-
-
-
min 22 – max 26.5
19
10
-
-
-
-
min 12.5 – max 16
-
-
-
-
43
46
45
min 27 – max 39
-
-
-
-
29
33
30
min 25.5 – max 29.1
Tab. 4: Measurements (mm) of the humerus of Cervus elaphus siciliae from Acquedolci compared with the dimensions of the remains
RADIUS
greatest length
transverse diameter
of the proximal end
of the proximal end
transverse diameter
transverse diameter
of the distal end
of the distal end
right
270
54
right
44
right
42
left
50
right
-
right
-
left
-
min 206 – max 237
min 39 – max 44.7
29
23
25
27
-
-
-
min 21 – max 24.5
31
20
-
-
-
-
-
min 22 – max 26.5
19
10
-
-
-
-
min 12.5 – max 16
-
-
-
-
43
46
45
min 27 – max 39
-
-
-
-
29
33
30
min 25.5 – max 29.1
Tab. 5: Measurements (mm) of the radius of Cervus elaphus siciliae from Acquedolci compared with the dimensions of the remains
developed and separated by a wide intercondylar
fossa. The medial condyle is strongly laterally
inclined. The remains are not measurable.
Tibia. Tibia remains are represented by 1 proximal, 1 medio-proximal and 2 distal fragments (Fig.
5, a-b). The proximal articulation surface is wide
and two very concave condylar facets are present.
The edges of the condylar facets bordering the
intercondylar area, which is narrow, are raised into
two prominent crests. The tuberosity of the diaphysis is well developed and shows a great torsion. The
distal articulation surface is irregularly trapezoidal
in shape. The edge of the lateral cochlea ends with
a prominent hook. The morphological and biometrical features of the remains are in the range of the
variability of Cervus elaphus siciliae (Gliozzi et al.,
1993) (Tab. 6).
Podials. Only 5 podial bones are present: 2 carpal
bones (scaphoid, lunar) and 3 tarsal bones (1
cuneiform, 2 astragali). The two astragali are broken.
The lateral lenght and the lateral antero-posterior
diameter of the two astragali are within the range of
the values reported by Gliozzi et al. (1993) (Tab. 7).
Metapodials. 22 metapodial fragments were collected: 9 metacarpal remains, 6 metatarsal remains
and 6 undeterminable metapodial remains.
Metacarpal remains include 2 complete and well
preserved bones (Fig. 6, a-b-c-d), 1 proximal fragment, 1 distal fragment and 5 shaft fragments.
Metatarsal remains are represented by 2 distal frag-
65
left
63
26
24
-
TIBIA
transverse diameter of the proximal end
antero-posterior diameter of the proximal end
transverse diameter at half length of the shaft
antero-posterior diameter at half length of the shaft
antero-posterior diameter of the distal end
left
59
57
-
left
23
20
37
30
right
46
32
min 48.5 – max 63.5
min 50 – max 64
min 21 – max 28.6
min 19 – max 27.5
min 33 – max 49
min 24.5 – max 35
Tab. 6: Measurements (mm) of the tibia of Cervus elaphus siciliae from Acquedolci compared with the dimensions of the remains described by Gliozzi et al., 1993.
PODIAL BONES
transverse diameter
lateral lenght
medial lenght
lateral antero-posterior diameter
medial antero-posterior diameter
SCAPH.
LUNAR
CUNEIF.
ASTR.
ASTR.
right
27
-
right
23
-
left
30
-
right
47
44
28
25
25
left
42
27
21
–
min 41.3 – max 47.8
–
–
min 24 – max 32
–
Tab. 7: Measurements (mm) of the podial bones of Cervus elaphus siciliae from Acquedolci compared with the dimensions of the remains described by Gliozzi et al., 1993.
METACARPAL
greatest lenght
transverse diameter
of the proximal end
of the proximal end
transverse diameter
transverse diameter
of the distal end
of the distal end
right
221
33
left
222
34
left
33
left
-
min 195 – max 226
min 29 – max 35.6
23
24
24
-
min 19 – max 26
18
20
-
-
min 16 – max 23.7
21
22
-
-
min 18 – max 22.4
34
35
-
34
min 27 – max 38.8
22
22
-
22
min 19 – max 24.4
Tab. 8: Measurements (mm) of the metacarpal of Cervus elaphus siciliae from Acquedolci compared with the dimensions of the remains described by Gliozzi et al., 1993.
METATARSAL
transverse diameter
of the distal end
of the distal end
right
35
left
34
min 29 – max 35
24
22
min 18 – max 23.4
Tab. 9: Measurements (mm) of the metatarsal of Cervus elaphus siciliae from Acquedolci compared with the dimensions of the remains described by Gliozzi et al., 1993.
66
ments and 4 shaft fragments, including one juvenile
specimen. The metacarpals are very slender. The palmar surface of the diaphysis is poorly channelled;
the ventral surface has a wide furrow along the
entire length of the diaphysis. At the proximal end,
the articulation facet for the magnum is wide, while
the articulation facet for the unciform is very small;
at the distal end, the two lateral condyles are separated by a narrow intercondylar notch. On the contrary, the palmar surface of the metatarsals diaphysis has a well developed central channel, and the
lateral condyles of the distal end are separated by a
wide intercondylar notch. The morphological features and the dimensions of the metapodials (Tabs.
8-9) are comparable with those detected by Gliozzi
et al. (1993) on other Sicilian specimens of Cervus
elaphus siciliae.
Phalanges. 7 remains were recovered: 4 fragmentary first phalanges and 3 complete second phalanges,
including one juvenile specimen. The dimensions of
the remains (Tabs. 10-11) are in the range of the variability of Cervus elaphus siciliae (Gliozzi et al., 1993).
teeth are absent; skulls, short bones and phalanges
are rare. The minimum number of individuals, based
on the most abundant long bone (the radius), is 5.
Taphonomical observations
Some taphonomical features, such as spatial distribution and orientation of the fossil remains, composition of the skeletal part, age distribution,
degree of skeletal articulation, fragmentation and
bone modification have been considered in order to
determine the biological processes that influenced
the accumulation of Cervus elaphus siciliae bones
(Badgley & Behrensmeyer, 1980; Behrensmeyer,
1975; Behrensmeyer Dechant Boaz, 1980).
In the lacustrine deposits of Acquedolci the number of Cervus elaphus siciliae fossil remains is very
low, with respect to the number of the remains of
Hippopotamus pentlandi.
The remains of deer were collected in all the
excavated trenches, with the exception of trench G
which is sterile, and about half of them come from
trench F. In each trench the remains are distributed
over the entire thickness of the sediments. The
bones are not concentrated and their spatial distribution is absolutely random, without preferential
orientation. Almost all the skeletal remains are very
fragmentary and fractured; complete specimens are
very rare. Articulated skeletal elements are absent.
Adult specimens are absolutely prevailing over juvenile remains, which are very scarce. Mandibles and
Fig. 6: Left metacarpal of Cervus elaphus siciliae; a) anterior
view; b) posterior view; c) proximal articulation; d) distal articulation.
67
FIRST PHALANX
transverse diameter
of the proximal end
of the proximal end
transverse diameter
transverse diameter
of the distal end
of the distal end
18
-
-
17
–
22
-
-
22
–
13
15
12
-
min 11 – max 15.7
17
18
-
-
–
-
17
15
-
–
-
10
13
-
–
Tab. 10: Measurements (mm) of the first phalanx of Cervus elaphus siciliae from Acquedolci compared with the dimensions of the
remains described by Gliozzi et al., 1993.
SECOND PHALANX
greatest lenght
transverse diameter
of the proximal end
of the proximal end
transverse diameter
transverse diameter
of the distal end
of the distal end
37
18
34
16
min 33 – max 38
–
23
21
–
13
13
min 10 – max 15
15
16
–
14
14
–
19
20
–
Tab. 11: Measurements (mm) of the second phalanx of Cervus elaphus siciliae from Acquedolci compared with the dimensions of the
remains described by Gliozzi et al., 1993.
Number of Remains
Spatial distribution
Minimum Number of Individuals
Age distribution
Skeletal part composition
Skeletal articulation
Degree of fragmentation
Bone modification
Cervus elaphus siciliae
104
random
absence of orientation
absence of concentration
5
predominantly adult, rare juvenile
absence of mandibles and teeth
rare skull, short bones and phalanges
disarticulated bones
very high
cracking, abrasion (not frequently)
Hippopotamus pentlandi
3.016
random
absence of orientation
extreme concentration
33
adult, juvenile, infantile
all skeletal parts represented
anatomical connection
very low
no
Tab. 12: Comparison between taphonomical features of Cervus elaphus siciliae and Hippopotamus pentlandi remains from Acquedolci deposit (taphonomic data about Hippopotamus pentlandi from Bonfiglio, 1995).
68
Bone modifications are observed at about 20 % of
the remains, showing traces of cracking (stage 1,
according to Behrensmeyer, 1978) and/or abrasion.
A comparison between taphonomical features of
Cervus elaphus siciliae and Hippopotamus pentlandi remains from the Acquedolci deposit is shown in
Tab. 12.
From a taphonomical point of view, the small
number of recovered remains of Cervus elaphus
siciliae, with respect to the extension of the deposit
and to the number of the hippo remains, their random distribution over the entire thickness of the
deposit, the lack of skeletal articulation, the presence of selected skeletal elements and the degree of
fragmentation, indicate an allochthonous fossilization, although the slight traces of abrasion and
cracking suggest a minimal transportation and/or a
short period of subaerial exposure.
The taphonomical analysis indicates that the
remains are allochthonous and probably were
deposited in the lacustrine basin as fragments of
carcasses from animals living in the area,
testifying, therefore, a different accumulation
process in comparison with the remains of
Hippopotamus pentlandi, which accumulated and
fossilized “in situ”, in the lacustrine basin where the
hippos have lived (Bonfiglio, 1995).
Conclusion
The morphological and biometrical features of
the remains are in the range of the variability of
Cervus elaphus siciliae POHLIG, the Pleistocene
endemic deer of Sicily which is characterized by a
moderately reduced size compared to the populations of the Italian peninsula.
The small number of specimens belonging to
deer, as well as those belonging to the other associated species, if compared with the very large number of the recovered hippo remains, probably is to
correlate with the different accumulation processes
of the remains and it does not reflect the real composition of the faunal populations living in the area.
Acknowledgments
Work supported by grants CoFin MURST 2003
“Faunal turnover in Sicily during the two last
Glacial cycles”. Thanks to Dr. R. Sardella, for the crit-
ical reading of the manuscript and the precious advises, and to Prof. K. Krainer, for the helpful suggestions in the revision of the English version.
References
Bada, J. L., Belluomini, G., Bonfiglio, L., Branca, M., Burgio,
E., Delitala, L. (1991): Isoleucine epimerization ages of
quaternary mammals of Sicily. – Il Quaternario, vol. 4
(1a): 5-11.
Badgley, C., Behrensmeyer, A. K. (1980): Paleoecology of
Middle Siwalik sediments and faunas. – Palaeogeography, Palaeoclimatology, Palaeoecology, vol. 30:
133-155.
Behrensmeyer, A. K. (1975): The taphonomy and paleoecology of Plio-Pleistocene vertebrate assemblages of
Lake Rudolf, Kenya. – Museum of Comparative
Zoology Bulletin Harvard, vol. 146: 473-578.
Behrensmeyer, A. K. (1978): Taphonomic and ecologic
information from bone weathering. – Paleobiology,
vol. 4(2): 150-162.
Behrensmeyer, A. K., Dechant Boaz, D. E. (1980): The
recent bones of Amboseli National Park, Kenya, in
relation to East African paleoecology. – In:
Behrensmeyer A. K., Hill A. P. (eds.): Fossils in the
making, 72-93. University of Chicago Press,
Chicago.
Bonfiglio, L. (1985): Prima campagna di scavo dei depositi a mammiferi pleistocenici dell’area della grotta di
S. Teodoro (Acquedolci, Messina, Sicilia). – Geologica
Romana, vol. 22: 271-285.
Bonfiglio, L. (1987): Primi elementi di stratigrafia del
talus della grotta di S. Teodoro (Acquedolci, Messina,
Sicilia). – Il Naturalista Siciliano, s. 4, vol. 10 (1-4):
43-57.
Bonfiglio, L. (1989): Distribuzione quantitativa dei resti di
Hippopotamus sp. del deposito di bacino del talus
della grotta di S. Teodoro (Acquedolci, Messina,
Sicilia). – Atti 3° Simposio di Ecologia e Paleoecologia
delle Comunità bentoniche: 299-317.
Bonfiglio, L. (1992): Campagna di scavo 1987 nel deposito pleistocenico a Hippopotamus pentlandi di
Acquedolci (Sicilia nord-orientale). – Bollettino della
Società Paleontologica Italiana, vol. 30 (2): 157-173.
Bonfiglio, L. (1995): Taphonomy and depositional setting
of Pleistocene mammal-bearing deposits from
Acquedolci (North-Eastern Sicily). – Geobios, M. S.,
vol. 18: 57-68.
Bonfiglio, L., Mangano, G., Marra, A. C., Masini, F. (2001):
A new late Pleistocene vertebrate faunal complex
69
from Sicily (S. Teodoro Cave, North-Eastern Sicily,
Italy). – Bollettino della Società Paleontologica
Italiana, vol. 40 (2): 149-158.
Bonfiglio, L., Marra, A. C., Masini, F., Pavia, M., Petruso, D.
(2002): Pleistocene faunas of Sicily: a review. – In:
Waldren W. H., Ensenyat J. A. (eds.): World Islands in
Prehistory, International Insular Insular Investigations,
428-436. BAR International Series, 1095.
Archaeopress, Oxford.
Gliozzi, E., Malatesta, A, Scalone, E. (1993): Revision of
Cervus elaphus siciliae Pohlig, 1893, Late Pleistocene
endemic deer of the Siculo-Maltese district. –
Geologica Romana, vol. 29: 307-354.
70
Mangano G. (2000): Nuovi resti di elefante e revisione di
alcuni resti di mammiferi del Pleistocene superiore
della Sicilia nord-orientale. – Giornale di Geologia,
Supplemento, serie 3a, vol. 62: 103-109.
Pavia, M. (2001): The Middle Pleistocene fossil avifauna
from the “Elephas mnaidriensis Faunal Complex” of
Sicily (Italy): preliminary results. – In: Cavarretta G.,
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Roma.
Revised manuscript accepted: April 8, 2005
Geo.Alp, Vol. 2, S. 71–76, 2005
EXCAVATIONS OF 2003 AT THE S. TEODORO CAVE (NORTH-EASTERN SICILY, ITALY):
PRELIMINARY FAUNISTIC AND STRATIGRAPHIC DATA
Gabriella Mangano1, Laura Bonfiglio1 & Daria Petruso2
1 Dipartimento di Scienze della Terra, Università degli Studi di Messina; e-mail: [email protected]; [email protected]
2 Dipartimento di Geologia e Geodesia, Università degli Studi di Palermoe-mail: [email protected]
Abstract
Systematic excavations have been carried out at the S. Teodoro Cave since 1998. Two trenches have been
excavated on the eastern side of the cave. The “1998 trench”, located between 9 and 13 meters from the
entrance, covers an area of 25 m2. The “2002 trench” was located between 30 and 32 meters from the entrance and covers an area of 9 m2.
The 2003 excavations included the 2002 trench, which has been deepened for 1 m and enlarged by adding new squares. Remains of Cervus elaphus siciliae, Bos primigenius siciliae, Elephas mnaidriensis, Crocuta
crocuta spelaea, Equus hydruntinus, small mammals, birds, reptiles, invertebrates and plant remains have
been recovered. The composition of the faunal assemblage and the lithologic features of the deposit, which
is composed of clayey sands and gravels, are similar to those recognized in the 1998 trench. Fragmentation
of remains, damages on almost all large mammal bones and abundant hyena coprolites testify an intense
hyena activity. Complete and not damaged remains of elephant and deer have also been collected and actually represent a taphonomic novelty. A new sedimentary unit of clayey sands and gravels which does not
contain fossil remains has been detected in the southern part of the trench. The age and the environmental
significance of this new sedimentary unit are to be clarified.
Introduction
The San Teodoro Cave opens in Jurassic limestone at an altitude of 150 m a.s.l.; it has huge
dimensions (about 60 m long, 20 m wide and up to
20 m high) and a total surface of more than 1.000
m2. In previous excavations the authors (Anca,
1860; Vaufrey, 1928, 1929; Tricomi, 1938; Maviglia,
1941; Graziosi, 1943, 1947; Graziosi and Maviglia,
1946) distinguished an upper sedimentary unit,
Late Glacial in age, containing human feeding
remains (mammal bones) associated with late
Upper Palaeolithic (Epigravettian) stone artifacts
(unit A in Bonfiglio et al., 2001), and a lower sedimentary unit (unit B in Bonfiglio et al., 2001) containing late Pleistocene endemic mammals.
The 1998 excavations were devoted to the
reconstruction of the stratigraphy of the cave
deposits and to a better knowledge of the faunal
assemblages, especially the older one. The 1998
trench has been located on the eastern side of the
cave at a square surface of 25 m2, between 9 and
13 meters from the entrance (coordinates 9-13/E-I)
(Fig. 1) and it has been deepened for 1.40 m. The
investigated unit B is composed of clayey sands and
gravels containing a highly diverse assemblage of
vertebrates, invertebrates (molluscs) and plant
remains. The large mammal assemblage which contains elephant (Elephas mnaidriensis), wild ox (Bos
primigenius siciliae), deer (Cervus elaphus siciliae),
wild boar (Sus scrofa), wolf (Canis lupus), hyena
(Crocuta crocuta spelaea), fox (Vulpes vulpes), associated with the equid Equus hydruntinus and the
small mammal taxa Microtus (Terricola) ex gr. savii,
Apodemus cf. sylvaticus, Erinaceus cf. europaeus
and Crocidura cf. sicula, has been attributed to a
71
Fig. 1: Plan of the S. Teodoro Cave with the location of the excavation areas of 1998 and 2002-2003. The black arrow indicates the
cave entrance.
new faunal complex in the Pleistocene of Sicily,
named “S. Teodoro Cave-Pianetti” faunal complex,
which contain some endemic taxa surviving from
the previous faunal complex (“Elephas mnaidriensis
F.C.”) associated with non-endemic taxa (Equus
hydruntinus, Microtus (Terricola) ex gr. savii,
Erinaceus cf. europaeus) (Bonfiglio et al., 2001).
The various evidences of cave frequentation by
spotted hyena populations are the most prominent
taphonomic feature of this deposit. Evidence comes
from the occurrence of several Crocuta skeletal elements (skull, teeth, limb bones), an impressive quantity of coprolites, and from ubiquitous traces of
crushing, gnawing, chewing and digestion that have
been detected on almost all the large mammal
remains (Bonfiglio et al., 1999, 2001). These taphonomic characters have been found so far only in a
few cave deposits of the Italian peninsula (“Grotta
dei Moscerini”, Stiner, 1990-91; “Buca della Iena”,
Pitti and Tozzi, 1971; Stiner, 1990-91; “Grotta
Guattari”, Piperno and Giacobini, 1990-1991; Stiner,
1990-91; “Tana delle iene”, Giaccio and Coppola,
2000) and are actually a novelty for insular environments.
Geochemical and radiometric data are not available for the deposits of the S. Teodoro Cave; the dispersal to Sicily of the ground vole, which has a fossorial habit, and of horses, that prefer open landscapes, might imply that a fully exposed connection
(a temporary land bridge related to an eustatic sea-
72
level lowstand) existed, perhaps more than once
during the last glaciation (Bonfiglio et al., 2002).
Pollen spectra from samples of coprolites from
unit B show the existence of a vegetation which
was mainly dominated by grass with moderate
arboreal taxa (Artemisia, Ephedra) and low percentages of mesophilous pollen taxa (Quercus, Betula,
Abies, Alnus, Pistacia, among others) which depict a
glacial landscape (Yll et al., in press).
During the 2002 excavations a new trench has
been located on the inner eastern side of the cave
at a square surface of 9 m2, between 30 and 32 m
from the entrance (coordinates 30-32/B-D) (Fig. 1),
in order to verify the extension of the evidences of
the frequentation by spotted hyenas in the inner
part of the cave. The 2002 trench has been deepened for about 40 cm. The sediments of unit B are
again composed of clayey sands and gravels and
contain several carbonatic concretion levels often
incorporating fossil remains. Remains of the same
large mammals collected during the 1998 excavations have been found together with small mammals (Microtus (Terricola) ex gr. savii, Crocidura cf.
sicula, Myotis sp.), birds, reptiles and hyena coprolites. The taphonomic features are very similar to
those detected in the 1998 trench and confirm the
extension of the deposit as far as 32 m from the
entrance of the cave, as well as the intense and
extensive frequentation by hyenas (Mangano and
Bonfiglio, 2003).
Coprolites
Bones
2002 trench
1998 trench
2003 excavations 2002 excavations 1998 excavations
1064
291
4271
543
132
2228
Unidentifiable bones
Identifiable bones
Cervus elaphus siciliae
Bos primigenius siciliae
Equus hydruntinus
Elephas mnaidriensis
Crocuta crocuta spelaea
Sus scrofa
Vulpes vulpes
Canis lupus
437
106
94
38
1686
542
75
6
5
11
9
24
1
5
1
3
2
1
1
392
21
41
26
38
14
7
3
Tab. 1: Number of recovered remains during the three excavation surveys at S. Teodoro Cave.
Excavations of 2003 (G. Mangano)
During the 2003 excavation the “2002 trench”
has been deepened for 1 m and enlarged by adding
two new squares on the southern side (coordinates
33/E-F) (Fig. 1).
Stratigraphic data
Besides unit B containing the fossil remains, in
the southern area of the trench (squares 32B/C/D,
and part of the squares 31B and 31C) a new unit of
clayey sands and gravels lacking fossil remains has
been detected. In this unit numerous white-yellowish pisolith-like elements with phosphatic composition, diameters between 1 and 5 cm and lacking
crystalline structure, are scattered. A subvertical,
quite irregular surface separates the fossiliferous
unit B from the sterile deposit and suggests that an
erosional phase cut the sterile deposit unit before
deposition of unit B. Age and precise environmental
significance of this new sedimentary unit are to be
clarified by deepening the trench.
Faunistic data
A total number of 543 large mammal bones and
1064 coprolites have been recovered (Tab.1). Almost
all the skeletal remains are strongly fragmented, not
articulated and horizontally and vertically scattered
without preferential orientation. A very large number of them (437) is represented by unidentifiable
bone splinters. The composition of the skeletal part
is characterized by the abundance of isolated teeth
and antlers (Tab. 2).
Cervus elaphus siciliae Pohlig, 1893. The endemic
red deer of Sicily is the most aundant species: 8
shed antlers, 13 antlers, 2 skull fragments, 3 hemimandibles, 18 teeth, 3 scapulae, 8 anterior limb
bones, 9 metapodials, 5 podials and 6 phalanges
have been recovered. Morphological and biometrical features ascribe them to Cervus elaphus siciliae
(Gliozzi et al., 1994). Particularly, two almost complete right shed antlers, different in size, have been
recovered arranged side by side. They were totally
covered by carbonatic concretions. The largest one
is 1.20 m long and actually is the largest antler fragment belonging to this species so far recovered (Fig.
2, a). Teeth grooves which cannot be ascribed certainly to hyenas are present on the surface of these
antlers.
Elephas mnaidriensis A.L. Adams, 1870. The elephant is represented by a small fragment of a
mandible, 3 teeth, 2 vertebrae, 1 rib, 1 pelvis, 1
anterior limb bone, 1 posterior limb bone and 1
metapodial. Teeth include one large fragment of
incisor and two very worn molar fragments belonging to an adult specimen. A complete and not damaged right tibia, absolutely lacking typical damages
produced by hyenas, is also preserved (Fig. 2, b).
Morphological features and biometrical data allow
to identify these specimens as Elephas mnaidriensis
(Ambrosetti, 1968; Bonfiglio and Berdar, 1979).
Bos primigenius siciliae Pohlig, 1911. The endemic wild ox of Sicily is represented by 2 hemimandibles, 1 femur shaft, 1 tibia, 1 metatarsal bone
73
Bos
Cervus elaphusprimigenius
siciliae
siciliae
shed antlers
antlers
skull
mandible
teeth
axis
girdles
anterior limb
posterior limb
metapodials
podials
phalanges
8
13
2
3
18
Equus
hydruntinus
2
1
4
3
8
9
5
6
Crocuta
crocuta
spelaea
Elephas
mnaidriensis
2
1
1
1
3
3
1
1
1
1
1
4
3
1
Tab. 2: Skeletal element distribution of large mammals recovered in the excavations of 2003 at S. Teodoro Cave.
and 1 scaphoid bone. M/1, M/2 and M/3 are preserved on mandibular fragments. M/3 has a slightly
inclined hypoconulid. The femur shaft belongs to a
juvenile specimen. The proximal end of the left tibia
was totally removed by crunching of the hyenas
(Fig. 2, c). The dimensions of remains are within the
range of the variation of Bos primigenius siciliae
(Brugal, 1987).
Equus hydruntinus Regàlia, 1904. The small equid
is represented by 1 right mandible fragment including the tooth row from M/2 to P/2 (Fig. 2, d), 2
upper molars and 2 deciduous premolars. The upper
molars have a short protocone and a well marked pli
caballin. In the lower cheek teeth the pli caballin is
less evident.
Crocuta crocuta spelaea (Goldfuss, 1832). The
spotted hyena is the only carnivore recovered during the 2003 excavations. One small maxillar bone
fragment, 4 heminandibles, 3 isolated teeth
(canines) and 1 metapodial small fragment are present. Two right hemimandibles include the tooth
row from M/1 to C (Fig. 2, e). Lower premolars are
sturdy and oval in section.
The preliminary study of mammal remains indicates the predominance of the non-endemic species
Microtus (Terricola) ex gr. savii.
The recovered taxa belong to the “S. Teodoro
Cave-Pianetti” faunal complex, late Pleistocene in
age, just recognized for the first time at the S.
Teodoro Cave (Bonfiglio et al., 2001).
Almost all large mammal bones are fragmentary
and show typical damages produced by the activity
of hyenas, such as strong fragmentation, ragged
edges, tooth grooves, tooth pits, digestion traces,
74
scooping out of cancellous bone (Sutcliffe, 1970;
Brain, 1981; Bunn, 1983). Nevertheless, some complete and undamaged bones of elephant (tibia) and
deer (antlers) have also been recovered and actually represent a taphonomic novelty.
Conclusion
Fossil remains collected during the 2003 excavations at the S. Teodoro Cave belong to the same
taxa previously recovered.
Most of the remains are fragmentary and
unequivocally damaged by hyenas, but some complete and undamaged bones are also present.
The cave is confirmed as a very large hyena den
and the spotted hyena is assumed to be the main
collecting agent of the skeletal elements of unit B,
although some new recognized features could indicate the existence of a different accumulation
process of the faunal remains.
A new sterile sedimentary unit has been discovered, but its age and environmental significance are
to be clarified.
Acknowledgments
Work supported by grants CoFin MURST 2003
“Faunal turnover in Sicily during the two last Glacial
cycles” . The excavations have been supported by
University of Messina (2003, extraordinary contribute to L. Bonfiglio) and by Acquedolci
Fig. 2: a) Cervus elaphus siciliae, right antler, external view; b) Elephas mnaidriensis, right tibia, posterior view; c) Bos primigenius
siciliae, left tibia, posterior view; d) Equus hydruntinus, right mandible, occlusal view; e) Crocuta crocuta spelaea, right mandible,
external view. Scale bar = 10 cm (a, b, c); 5 cm (d, e).
75
Commune. Thanks are due to Dr. G.F. Villari,
Superintendent to Archaeological and Cultural
Heritage of Messina and to Dr. U. Spigo, responsible
of the Archaeological Service. A particular acknowledgment to Prof. A. Kotsakis for the critical reading
of the text and for the revision of English version,
and to Prof. K. Krainer for the helpful suggestions in
the revision of the final text.
References
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Romana, vol. 7: 277-398.
Anca, F. (1860): Note sur deux nouvelles grottes ossifères
découvertes en Sicile en 1859. - Bulletin de la Société
Géologique de France, vol. 17: 684-695.
Bonfiglio, L., Berdar, A. (1979): Gli elefanti delle ghiaie
pleistoceniche di Messina. - Quaternaria, vol. 21:
139-177.
Bonfiglio, L., Mangano, G., Marra, A.C. (1999): Late
Pleistocene hyaena den from a large cave deposits of
Sicily (Italy). - INQUA XV International Congress,
Durban 3-11 August 1999, abstract book: 27-28.
Bonfiglio, L., Mangano, G., Marra, A.C., Masini, F. (2001):
A new Late Pleistocene vertebrate faunal complex
from Sicily (S. Teodoro cave, north-eastern Sicily,
Italy). - Bollettino Società Paleontologica Italiana,
vol. 40 (2): 149-158.
Bonfiglio, L., Mangano, G., Marra, A.C., Masini, F., Pavia,
M., Petruso, D. (2002): Pleistocene Calabrian and
Sicilian bioprovinces. – Geobios, M. S., vol. 24: 29-39.
Brain, C. K. (1981): The hunters or the hunted? An introduction to African cave taphonomy. - pp. 1-365,
Chicago University Press, Chicago.
Brugal, J.P. (1987): Cas de „nanisme“ insulaire chez l’aurochs. - In: 112th Congrès National des Sociétés
savants, Lyon, vol. 2: 53-66.
Bunn, H.T. (1983): Comparative analysis of modern
bone assemblages from a San hunter-gatherer camp
in the Kalahari Desert, Botswana, and from a spotted hyena den near Nairobi, Kenya. - Animal and
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143-148.
Giaccio, B., Coppola, D. (2000): Note preliminari sul contesto stratigrafico e paleoecologico del sito “Tana delle
iene” (Ceglie Messapica, Brindisi, SE Italia). - Il
Quaternario, vol. 13 (1/2): 5-20.
Ghiozzi, E., Malatesta, A., Scalone E. (1994): Revision of
Cervus elaphus siciliae Pohlig, 1893, Late Pleistocene
76
endemic deer of the Siculo-Maltese district. –
Geologica Romana, vol. 29: 307-353.
Graziosi, P. (1943): Gli scavi dell’Istituto Italiano di
Paleontologia Umana nella grotta di S. Teodoro
(Messina): nota preliminare. - Atti Società Toscana
Scienze Naturali, Memorie, vol. 52: 82-99.
Graziosi, P. (1947): Gli uomini paleolitici della grotta di S.
Teodoro (Messina). - Rivista di Scienze Preistoriche,
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Graziosi, P., Maviglia, C. (1946): La grotta di S. Teodoro
(Messina). - Rivista di Scienze Preistoriche, vol. 1 (4):
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Mangano, G., Bonfiglio, L. (2003): Campagna di scavo
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S. Teodoro (Acquedolci, Messina – Sicilia nord-orientale). Giornate di Paleontologia 2003, Alessandria
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study of the paleosurface of Guattari Cave (Monte
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143-161.
Pitti, C., Tozzi, C. (1971): La Grotta del Capriolo e la Buca
della Iena presso Mommio (Camaiore, Lucca). - Rivista
di Scienze Preistoriche, vol. 26 (2): 213-258.
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now. - Quaternaria Nova, vol. 1: 163-192.
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Geo.Alp, Vol. 2, S. 77–90, 2005
LOWER PERMIAN PALAEOICHNOLOGY FROM THE OROBIC BASIN (NORTHERN ITALY)
Giuseppe Santi
With 4 figures and 2 plates
Dipartimento di Scienze della Terra, Via Ferrata 1, 27100 Pavia (Italia). e-mail: [email protected]
Abstract
The Lower Permian palaeoichnofauna of the South-Alpine region comes from the Collio Formation only,
and mainly crops out in the Orobic and Trompia basins. It consists of traces of vertebrates (amphibians and
reptiles) and invertebrates (insects, arthropods, burrowing, probable myriapods, gastropods, freshwater jellyfishes, conchostraceans and freshwater bivalves). It is poor in taxa and similar to the coeval ichnoassociation
of Central Europe, N. America and Argentina. Inside the trophic pyramid relevant to the biodiversity of the
lower unit of the Collio Fm., the top carnivore is absent. This role is partially occupied by reptiles (e.g. araeoscelids) having features similar to true lizards. Shifting from the sediments of the lower part of the Collio Formation to the upper part, impoverishment of the ichnocoenosis composition is linked either to a climatic
shift (from more humid towards drier), or to tectonic activity that prevented the persistence of biotope formation. The Upper Permian uplift marks a strong ichnofaunistic change with the introduction of Triassic
components.
Introduction
Brief stratigraphical framework
In the South-Alpine region the continental
Lower Permian is characterized by vertebrate and
invertebrate ichnofossils and by rare floral remains
(macroplants, spores and pollen). They come
almost exclusively from the Collio Fm. cropping
out in the Orobic and Trompia (= Collio) basins,
with the exception of rare fossiliferous remains
from the Tregiovo Basin. The features, problems
and hypotheses relating to the composition, distribution and behavioural features of the trackmakers and the ichnocoenosis variation are also different, moving from the lower “Collio” to the
upper unit of the same formation, and from this
to the Upper Permian when the faunistic change
is profound. These aspects are very clear from
analysing the fossil beds of the Collio Fm. in the
Orobic Basin; this article will review the ichnofaunistic assemblages from this basin and discuss
their significance and the problems inherent to
them.
For a long time it has been known that the
Palaeozoic of the Alps is very poor in vertebrate
remains, with the only exception being the ichnites, which have recently become a great stratigraphical tool (Avanzini et al., 2001). Recent
detailed research on the Permian of Europe (mainly in France and Germany) has enlarged our
knowledge, particularly of the invertebrates, and
of the stratigraphical-chronological role played by
them (Gand et al., 2001 with references therein).
The South-Alpine region is characterized by a
series of basins created from W to E, as inherited
Hercynian structures have produced structural
highs of a metamorphic or igneous nature
(Cassinis & Perotti, 1994; Cassinis et al., 1999 with
references therein; Perotti, 1999). The main basins
are the Orobic Basin and, to the east, the Trompia
Basin (Fig. 1), but other smaller basins are important for their ichnofossil content (Tregiovo Basin,
Tione Basin) (Conti et al., 1997).
77
Fig. 1: Schematic non-palinspastic section of the main Permian basin distribution of the South-Alpine region (Conti et al., 1997,
mod.). 1 – Pre-Permian basement, 2 – Volcanic deposits, 3 – Clastic units of the first cycle of sedimentation (Basal Conglomerate,
Collio Fm, Ponteranica Conglomerate), 4 – Verrucano Lombardo-Val Gardena Sandstone complex, 5 – Bellerophon Fm.
Permian sediments occur in two tectono-sedimentary cycles separated by an uncertain age gap
(between 14 and 25 My, according to the most
recent data in Cassinis et al., 2002a). The first cycle,
of ?Upper Carboniferous–Lower Permian age, is
composed of a continental succession of volcanic
deposits (from intermediate to acid chemistry) and
by alluvial-to-lacustrine sediments that comprise
the Basal Conglomerate, the Collio Fm, the
Tregiovo Fm, the Ponteranica Conglomerate, the
Dosso dei Galli Conglomerate and the Auccia
Volcanics. The second cycle is assigned to the
Upper Permian and is composed of the reddish
clastic deposits of the Verrucano Lombardo-Val
78
Gardena Sandstone complex (Fig. 2). With their
deposition the Palaeozoic ended.
The question of the use of a two- or three-fold
subdivision of the Permian System for dating of
continental successions has been debated for a
long time. A detailed discussion relating to dating
of the Permian continental beds in the SouthAlpine region was recently carried out by Cassinis
(2003), Cassinis and Ronchi (2001) and Cassinis et
al. (2002b). The traditionally adopted Permian subdivision for research in the South-Alpine area is
“Lower Permian” (from about the Asselian to
Kungurian) and “Upper Permian” (from the
Fig. 2: 1 - Chronostratigraphical sketch of the Permian of the Orobic Basin. 2 - Chronostratigraphical sketch of the Permian of the
Trompia Basin.
Ufimian to Tatarian, according to the CisUralian/Russian Standard Scale), and this last rarely
includes the Middle Permian, corresponding
approximately to the Guadalupian Series
(Menning, 2001; Cassinis, 2003, Fig. 1). It is based
on the palaeontological data from macroplants,
palynomorphs, tetrapod footprints, and the radiometric and palaeomagnetic investigations.
Therefore, the stratigraphical resolution is rather
poor compared with the marine equivalents; so the
absence of detailed data and of the wider correlations for the continental beds prevents the use of
the three-fold subdivision of the Permian System
into “Lower”, “Middle” and “Upper”. Only in those
places where the lateral transition between the
continental and marine deposits (i.e. in the
Dolomite region between the Val Gardena
Sandstone and the Bellerophon Formation, togeth-
er referred to the Upper Permian) is evident can
the use of the marine stages be justified. For these
reasons, and in agreement with Cassinis (2003), in
this study the continental Permian “Lower” and
“Upper” subdivisions are used.
Therefore, it is Lucas’s opinion (pers. comm.)
that in this study the term “Upper Permian” should
include the “Middle Permian” (Ufimian and
Kazanian), and only the Tatarian should really be
“Upper Permian”. As such, it may be better to utilize the marine timescale terms (Roadian, Wordian,
Capitanian, Wuchiapingian, etc.) and not the old
Russian terms. The utilized chronostratigraphy
(Cisuralian and Russian stages) for the Early
Permian represents the international subdivision of
the Permian System, but in the dating of the continental beds, to leave out the post-Kungurian
Russian terms that, in Lucas’s opinion (pers.
79
Fig. 3: Permian stratigraphy (SCPS = Sub-Commission of Permian stratigraphy) (Vachard & Argyriadis, 2002. mod.)
comm.), are only the regional stages for the marine
timescale, is more difficult for the reasons
advanced above. Fig. 3 shows the different scales
of the Permian stratigraphy.
In the classic succession of the Trompia Valley
(Collio Basin) the COLLIO FORMATION was deposited on
volcaniclastic rocks (ignimbrites) which do not
crop out with continuity within the Orobic Basin,
but are abundant in other areas (e.g. in the
Acquaduro Valley –Introbio- and in the Cedrino
Pass) (Sciunnach, 2001) and in the mainly “bergamask” sector of the same basin (Jadoul et al.,
2000). Other subdivisions of lithofacies have been
proposed on a petrographical basis by Cassinis et
al. (1988), Cadel et al. (1996), Forcella et al. (2001)
and Sciunnach (2001). The Collio Fm. can be informally subdivided into two units: the lower one is
composed of grey-green and black sandstones and
siltstones, while the upper unit is defined by mainly reddish sandstones and pelites of volcanic elements with quartz, plagioclase and muscovite. It is
well stratified and locally contains some conglomeratic beds. The typical arenaceous zones frequently contain fragments of black clay (clay chips) and
display planar lamination, while in the pelitic
intervals there are different structures such as mud
80
cracks, raindrop imprints, ripple marks and fossil
plant remains, as well as vertebrate and invertebrate ichnites.
This formation is interfingered with the
Ponteranica Conglomerate (Casati & Gnaccolini,
1965, 1967). Utilising the fossils collected in the
Trompia Basin, the Collio Fm. is referred to the
Lower Permian based on chronological data provided by macroflora (Geinitz, 1869; Jongmans,
1960; Remy & Remy, 1978; Kozur, 1981; Visscher et
al., 1999), pollen (Clement-Westerholf et al., 1974;
Cassinis & Doubinger, 1991, 1992) and tetrapod
footprints (Ceoloni et al., 1987; Conti et al., 1991,
1997), and also for its position below the angular
unconformity ascribed to the main post-Saalian
phase (Palatine) of the Hercynian orogenesis.
Vertebrate and invertebrate ichnocoenoses of the
Orobic Basin
In Italy, early knowledge of vertebrate footprints from the Collio Fm. in the Trompia Valley
was advanced by Geinitz (1869) and Curioni
(1870). Later, these fossils were studied by Gümbel
(1880); the same ichnofauna from the Orobic Basin
was analysed by Dozy (1935) and later re-exam-
ined by Haubold (1971). The studies of Berruti
(1968), Haubold (1996, 2000), Haubold & Stapf
(1998), Casati & Gnaccolini (1967), Ceoloni et al.
(1987), Conti et al. (1991, 1997, 1999), Nicosia et
al. (2000) and Santi & Krieger (2001) have
advanced our knowledge of the vertebrate ichnofauna of the Lower Permian. Footprints from both
the Orobic Basin and the Trompia Valley are of
amphibians and reptiles, and they come from different parts of the volcano-sedimentary deposits
of the Collio Formation (Conti et al., 1991; Santi,
2003) relating to main vegetated areas, to other
alluvial zones, to more emergent humid areas, and
others with shallow water.
Together with small- to medium-sized vertebrates, lived insects and arthropods (Bifurculapes
Hitchcock, 1858, Dendroidichnites elegans
Demathieu, Gand & Toutin-Morin, 1992, cfr.
Heteropodichnus variabilis Walter, 1983,
Eisenachichnus sp. (= Secundumichnus), Tambia
spiralis Müller, 1956, Permichnium Guthörl, 1934,
burrowing invertebrates (?Scoyenia White, 1929),
gastropods (Paleobullia sp. vel. ?Cochlea sp.), probably myriapods and some unidentified trails,
bivalves (Anthracosiidae), small crustaceans
(“Estheria”) and freshwater jellyfish (Medusina
limnica Müller, 1978 and Medusina atava (Pohlig,
1892, Walcott, 1898) (Ronchi & Santi, 2003) (Pl. 1).
Up to now, from these former data the composition of the invertebrate ichnocoenosis shows: (a)
imprints are typically of freshwater animals, (b) a
dominance of surface traces and not infaunal burrows, (c) low biodiversity, (d) a lack of monospecifity, and (e) the ichnodiversity and the taxonomic
composition suggest a terrestrial-freshwater origin.
The tetrapod ichnofauna of the Collio Basin
consists of: Batrachichnus sp., Camunipes cassinisi
Ceoloni et al., 1987, Amphisauropus imminutus
Haubold, 1970, Amphisauropus latus Haubold,
1970, Varanopus curvidactylus Moodie, 1929,
Dromopus lacertoides (Geinitz, 1861), Dromopus
didactylus Moodie, 1930 and Ichniotherium cottae
(Pohlig, 1885). That of the Orobic Basin is composed of: “Batrachichnus” salamandroides (Geinitz,
1861), Camunipes cassinisi Ceoloni et al., 1987,
Amphisauropus imminutus Haubold, 1970,
Amphisauropus latus Haubold, 1970, Varanopus
curvidactylus Moodie, 1929 and Dromopus lacertoides (Geinitz, 1861) (Pl. 2). The ichnocoenoses re-
enter in the so-called “red-bed ichnofacies”
(defined as a variety of fluvial, deltaic, lacustrine
and marginal marine environments; Haubold &
Lucas, 1999), typically different from the
“Chelichnus ichnofacies” related to the desert
environment and aeolian facies (Lockley et al.,
1994; Lockley & Meyer, 2000; Lucas, 2002).
A great affinity between the ichnocoenoses of
the two basins is evident, with the only exception
being Ichniotherium cottae and Dromopus didac tylus presenting together inside the Collio Basin,
but lacking in the Orobic Basin. This last ichnospecies is present not only in the highest strata
of the Collio Fm. in the Trompia Valley, but it is also
a monotypic taxon of the Tregiovo Basin (Conti et
al., 1997; Nicosia et al., 2000). At present I. cottae
should be a local taxon of the Trompia Basin.
Besides, there is the problem linked to the validity
of the ichnogenus Camunipes, namely if it effectively should be a true ichnogenus, or should be
considered a synonym of Erpetopus. A discussion
of this taxonomic problem is advanced by Haubold
& Lucas (2001, 2003) and Santi (2004). On the
whole, the Lower Permian ichnocoenosis actually
consists of mostly reptiles and one amphibian
(Batrachichnus); among the former we have a relevant “large” herbivore component, while the others are of smaller size.
The time interval into which the tetrapod ichnofauna is limited is between 286/283 Ma at the
base and 278/273 Ma at the top (Avanzini et al.,
2001). In agreement with the Permian subdivision
effected by Menning (2001), this ichnoassociation
may belong to the Artinskian and Kungurian, but
other scales (i.e. Harland et al., 1990; Odin, 1994;
Gradstein & Ogg, 1996) consider these values to be
Sakmarian and upper Asselian. The South-Alpine
ichnoassociation has a similarity to that of North
America, with strong Wolfcampian affinities showing a great interaction between W-Central Europe
and this continent.
It is a mostly homogeneous association, but
also very poor in taxa, and even more reduced in
the highest strata of the Collio Fm. In the Orobic
Basin, the passage between the lower unit of this
formation and the upper is marked among the
tetrapod palaeoichnofauna by the absence of
Batrachichnus, Camunipes (Erpetopus) and A.
imminutus, and by the presence of only A. latus, D.
lacertoides and V. curvidactylus, and among the
81
invertebrates, Dendroidichnites and Medusina
atava are present. In agreement with the “Global
Permian series of the marine Permian System”, the
above-mentioned ichnoassociation is considered
coeval with the “Lower Permian Cisuralian”
(Cassinis et al., 2002).
On the whole, factors producing the taxonomic
compression of the Lower Permian palaeoichnofauna are different (Lucas, 1998), but regionally, the
“deposition time compression” hypothesis (Nicosia
et al., 2000) can be advanced on the basis of radiometric data presented by Schaltegger & Brack
(1999) in the volcanic beds at the base and at the
top of the Collio Fm. s.s. (= sedimentary “Collio”) in
the Trompia Valley. According to these authors,
about 700 m of sediments were laid down in 4–5
My: a very high rate linked to strong tectonic activity. In my opinion this would prevent the establishment of useful biotopes for the survival of animals.
A clear example is shown near to the Pizzo del
Diavolo (Brembana Valley) neighbouring the
Bocchetta di Poddavista (“Podavit”) where the lower
unit of the Collio Fm. (600 m up) is well exposed. In
its lower portion abundant “signatures” of the tectonic activity are well evident. Repeated pyroclastic
fall intercalations and the soft sediment deformations (seismites), sedimentary dykes, “ball & pillow”
and slumping structures, were probably triggered by
synsedimentary tectonics and frequent volcanoseismic activity. Only in the homogeneous siltymuddy part (last ten of metres) did the tectonic
“peace” allow the development of more firm
biotopes. Only in this position were the taxa of the
“orobic” ichnoassociation identified.
Furthermore, the orogenic activity is not the
cause, but one cause of the taxonomic paucity,
together with climatic change (Santi, 2004).
Partially in agree with the opinion of Lucas
(pers. comm.) that the global paucity in Permian
ichnotaxa reflects the conservative nature of the
footprint structure (Santi, 2004), the ichnoassociation of the South-Alpine region is very similar to
the other European and extra-European countries
(see later): then a priori it is not possible to exclude
the hypothesis that it could accurately reflect the
original vertebrate biodiversity. Overall, local geological events could have played a crucial role for
the original biodiversity composition in this sector
of Palaeoeurope (“deposition time compression”
hypothesis).
82
Paucity in taxa could depend on internal properties and external conditions:
a) linked to niche dimensions for vertebrates and
invertebrates. In fact, the species with the narrowest niches have high probabilities of speciation either because species are unstable and
have patchy populations, or because there are
potential new niches to invade through evolutionary divergences. The “Collio” area was
undoubtedly large and less ecologically diversified, and this should favour extinction rather
than speciation.
b) Species with small and patchy populations tend
to isolate frequently; consequently this pattern
of species has a greater probability of extinction
(Stanley, 2001). The orogenic forces and climatic changes probably operated above a very brittle biodiversity with low numbers, and determined their extinction. Only the ability of some
taxa to disperse and to colonize different
biotopes might have allowed them to survive
(Amphisauropus, Dromopus, Varanopus), but
probably the attempt did not occur completely
within an unstable framework (coeval orogenesis + climatic changes).
In the palaeo-European domain, documented
examples of terrestrial environments with fossiliferous assemblages have been described (e.g.
Debriette & Gand, 1990; Schneider, 1994; Gand et
al., 1997 a, b, c; Eberth et al., 2000). It is noteworthy that in many European Lower Permian basins,
which can represent excellent analogues to those
of the central Southern Alps, the facies distributions and environmental settings record, from base
to top, an evolution from grey-black alluvial-tolacustrine deposits to reddish flood-plain and
playa sediments. Over a large part of Western
Europe, Early Permian times were characterised by
a climatic shift from warm, with alternating wet
and dry seasons, to semi-arid, up to the very warm
and hot conditions of the Late Permian (Ori, 1988;
Dickins, 1993; Parrish, 1993; Golonka & Ford,
2000). Thus, during the mid to late Early Permian
(Artinskian–Kungurian?), a regional and geologically rapid decrease in the rate of precipitation and
the onset of oxidising climatic conditions were
suggested by both lithofacies and biofacies
changes. In the Orobic Basin (at least in its western
sectors), the dominant alluvial-to-lacustrine darkcoloured facies pass quite abruptly, towards the
stratigraphic top of the succession, to reddish fine
sediments. The former dark deposits suggest that a
higher groundwater level produced reducing conditions, while the red fines indicate muddy playa
conditions with high evaporation rates and an oxic
environment. A similar environmental–climatic
transition could also be envisaged in the western
Val Trompia Basin, where the Collio Fm. fluvial and
lacustrine scenario evolves from the proximal to
distal alluvial-fan facies (Dosso dei Galli
Conglomerate) and up-section to the lateral and
bioturbated, purple-red, fine sandstones and siltstones (Pietra Simona Mb.). The consequences
were, at the beginning of the Upper Permian, a
clear change in fauna with more modern features
(Conti et al., 1999); its origin is contained in the
regional temporal gap which divides the first cycle
from the second.
Behavioural features of
the Early Permian tetrapods
It seems opportune to talk about the problem of
the behavioural features of the trackmakers. The
rarity of fossil remains of vertebrates in the continental deposits of the Permian of Central and South
Europe makes a discussion about their behavioural
features rather difficult, but the ichnoassociation
can be considered as a good starting point for this
goal. The Lower Permian ichnoassociation of the
South-Alpine zone reflects the vertebrate association living in this area of Palaeoeurope at the time,
like those of France, Germany and also North
America and Argentina, with only rare exceptions of
elements considered as “local form” (i.e.
Ichniotherium for the South-Alpine region) (Conti
et al., 1999). Within the ichnoassociation of the
South-Alpine region (Orobic and Trompia Basins),
until now typical prints attributed to a top carnivore are absent; either the trackmaker belonged to
a population effectively reduced in number compared with the herbivores, or it was totally absent.
Maybe during the Lower Permian of southern
Europe, its specific role was partially occupied by
other vertebrates. The low number of taxa (common
also in the Lower Permian ichnoassociations from
other countries) suggests that the ichnodiversity
could be, if not real, then the almost complete composition of the vertebrate biodiversity. Then the
prints can be, if not an exact mirror, then at least a
significant indicator of the original vertebrate and
invertebrate biodiversity. This would not explain
why the trophic pyramid should effectively be that
here carried out, but until now the ichnocoenosis
composition and the frequency with which some
footprints are discovered (i.e. Batrachichnus is very
rare compared with the reptiles, and among these
Amphisauropus latus and Dromopus lacertoides are
clearly much frequent in comparison with
Varanopus) allows us to propose the hypotheses
advanced here. This is rather different to Lucas’s
opinion (pers. comm.) referring to the Moenkopi
ichnoassociation from the Triassic of the USA:
“…The tracks are almost all of archosaurs (chirotheres), but the bones from the same formation
are almost all of amphibians…”.
Not withstanding the paucity of taxa of the
tetrapod ichnofauna, the ichnocoenoses have not
been utilised to examine the behavioural features of
the trackmakers. A similar gap is also underlined by
Kramer et al. (1995) referring to the ichnites from
the Coconino Sandstone (North America): “…behavioural aspects of extinct animals cannot be tested “
(Brand, 1978 p. 81) (Kramer et al., 1995 p. 245).
Furthermore, behavioural evidence from trackmakers can be discussed when studying “terminated
trackways” sensu Kramer et al. (1995), or those that
suddenly change direction. From the “orobic” Lower
Permian beds come some data on the reptilian diet.
Among the components of the ichnocoenosis, the
Dromopus trackmaker is commonly ascribed to the
araeoscelid, considered a consumer of small invertebrates with exoskeletons. Figure 4A suggests the
following event sequence, pointing to a lack of
superimposition of walking-trail and footprints. A
trackmaker arthropod (Dendroidichnites elegans)
is moving on a firm silty bed (point A). On its left
side a probable adult araeoscelid reptile, trackmaker of Dromopus, is approaching. At point B the
arthropod abruptly deviates towards its right side,
probably trying an evasive manoeuvre; by this
point the trail impression is not very clear, probably because the trackmaker was alarmed and
progress was disordered. The final trackway-tract
was not preserved by the sediments, but we realise
that the araeoscelid preyed upon the arthropod
without pursuing it. Figure 4B shows a clear “terminated trackway” sensu Kramer et al. (1995) of an
arthropod (Heteropodichnus trackmaker) pursued
by a Dromopus one; traces of its trail abruptly disappear.
83
Fig. 4: A - Interaction between the Dendroidichnites elegans Demathieu, Gand and Toutin-Morin, 1992 trail and Dromopus sp. footprints. Black arrows indicate the arthropod trail directions. B - “Terminated trackway” of cfr. Heteropodichnus variabilis Walter, 1983
with Dromopus sp
As witnessed by the prints upon the slabs in Fig.
4, it is possible that the predator role in the Lower
Permian of the South-Alpine region was played
partially by these reptiles. Rare amphibians and
mainly reptiles compose the tetrapod ichnocoenosis; it is an association with a paucity in taxa and
comprises
herbivores
from
small
size
(Amphisauropus imminutus) to medium-large size
(Amphisauropus latus). At present, large footprints
referred to large vertebrates (i.e. such as the
Middle Permian pareiasaur Pachypes) have not
been found. A top carnivore seems lacking. Thus, in
the Lower Permian of the South-Alpine region the
trophic pyramid was probably like this:
Primar y consumer. Medium-sized herbivore:
cotylosaurs identified as the trackmaker
Amphisauropus latus, a tetrapod of relatively large
dimensions (the true “giant” of the association in
comparison with the sizes of other trackmakers),
with short and stumpy legs, probably strong and
84
adapted to support a relatively great weight. The
frequency with which the A. latus footprints are
found is highest, so it represented the dominant
animal of “Collio” lands. Similar in size or possibly
larger was the Ichniotherium trackmaker (an
edaphosaur pelycosaur), but as seen above, its
presence is very rare, and thus its role inside the
trophic pyramid is much diminished.
Secondary consumer. Carnivores: the ichnological association seems to lack typical footprints
attributed to this consumer.
Mixed diet. Opportunistic consumers: on the
whole these are small reptiles, morphologically and
in their general structure similar to small lizards,
also with autopodia features and with more or less
sharp teeth (Camunipes trackmakers). Their diet
could be similar to that of true lizards of small
dimensions, swallowing and biting anything either
living or dead. In this category should re-enter the
Dromopus trackmaker which, together with the
Amphisauropus, is a common form, and less frequently that of Varanopus. A novel feature of an
araeoscelid trackmaker (Araeoscelis) is the lateral
temporal opening, which could have been closed in
relation to the skull extension as the consequence
of a more massive dentition (Carroll, 1988). Such
araeoscelids could prey upon protein-bearing
organisms and consume some strong parts such as
their exoskeleton (arthropods), or small vertebrates
(amphibians?) also.
Thus, it does not seem that the Lower Permian
association of the South-Alpine area needs to be
balanced. It is possible that the araeoscelids and
the Dromopus trackmaker could have partially
occupied the small predator role.
The author is deeply indebted to S.G. Lucas
(Albuquerque, New Mexico) for his useful advice
and critical review of the text and S. Jones (Cardiff)
for revision to English. This study was carried out
with a grant from FAR.
Conclusions
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1) The ichnocoenosis has a similarity to those
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South America (Argentina; Melchor &
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2) An impoverishment of the ichnofaunistic composition, shifting from the lower unit to the
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by reptiles, some araeoscelids having features
similar to true lizards.
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Manuscript submitted: November 26, 2004
Plate 1
Plate 1
A – Bifurculapes sp. Bocchetta di Poddavista, Orobic Basin. B – cfr. Heteropodichnus variabilis Walter, 1983. Mincucco
Mt. Orobic Basin. C - Paleobullia Götzinger & Becker, 1932 vel. ?Cochlea Hitchcock, 1858. Brembana Valley, Orobic
Basin. D - Secundumichnus sp. Brembana Valley, Orobic Basin. E – Undetermined traces. Brembana Valley, Orobic
Basin. F – Medusina atava (Pohlig, 1982) Walcott, 1898. Inferno Valley, Orobic Basin. G – Anthracosiidae Trompia
Valley Basin. H – Dendroidichnites elegans Demathieu, Gand & Toutin-Morin, 1992, Mincucco Mt. Orobic Basin.
I, J – Medusina limnica Müller, 1978. Trompia Valley Basin .
Plate 2 (continued on next page)
A- Camunipes cassinisi Ceoloni et al., 1987, reverse print left couple manus-pes. Brembana Valley, Orobic Basin.
B – Amphisauropus latus Haubold, 1971, reverse print right pes. Inferno Valley, Orobic Basin. C - Amphisauropus latus
Haubold, 1971, reverse print left manus. Inferno Valley, Orobic Basin. D – Varanopus curvidactylus Moodie, 1929,
reverse print left pes. Inferno Valley, Orobic Basin. E - Varanopus curvidactylus Moodie, 1929, reverse print left couple
89
Plate 2
manus-pes. Inferno Valley, Orobic Basin. F – Dromopus lacertoides (Geinitz, 1861), trackway. Brembana Valley, Orobic
Basin. G - Amphisauropus latus Haubold, 1971, set reverse print manus-pes. Inferno Valley, Orobic Basin.
H - Camunipes cassinisi Ceoloni et al., 1987, set reverse print manus-pes. Scioc Valley, Orobic Basin. I - Amphisauropus latus Haubold, 1971, reverse print manus?-pes?. Inferno Valley, Orobic Basin. J - Varanopus curvidactylus
Moodie, 1929, trackway. Inferno Valley, Orobic Basin.
90
Geo.Alp, Vol. 2, S. 91–98, 2005
PRELIMINARY ANALYSIS OF THE FIRST LOWER MOLAR VARIABILITY
IN LATE PLEISTOCENE AND LIVING POPULATIONS OF TERRICOLA SAVII (ARVICOLIDAE, RODENTIA)
Maria Teresa Curcio1, Longino Contoli2, Emanuele Di Canzio3, Tassos Kotsakis1
1 Dipartimento di Scienze Geologiche, Università Roma Tre, Largo San Leonardo Murialdo 1 - 00146 Roma, Italy
2 Via Arno 38 - 00198 Roma, Italy
3 Dipartimento di Scienze della Terra, Università di Roma “La Sapienza”, Piazzale Aldo Moro 1 - 00185 Roma, Italy
e-mail: [email protected]; [email protected]; [email protected]; [email protected]
Abstract
The main object of this research is the study of the variability degree of the first lower molar in Late
Pleistocene and living populations of Terricola savii in Italy (whose conspecificity has been proved by genetic analyses) and its comparison with that of fossil populations (assigned to T. savii on a morphological basis)
in order to find a way to attribute isolated fossil remains to specific systematic groups. On this basis, we
attempted to establish, through different analyses and direct observations on the occlusal dental surface
morphology, the relationships that exist between fossil and living populations, and to verify the existence of
a temporal and/or geographic cline.
Introduction
Terricola savii (DE SELYS LONGCHAMPS, 1838)
(Arvicolidae, Rodentia) is the most common living
vole on the Italian peninsula (south of the
Apennines) and it is common in northern Italy too
(between the Alps and the Apennines). During the
Late Pleistocene T. savii colonized Sicily (Petruso,
2002). Voles classified as T. gr. T. savii are present in
Early Toringian mammal assemblages (San Giovanni
di Duino, Venezia Giulia - Campani Quarry, Tuscany
– Case Picconetto, Abruzzi) (Bartolomei, 1976;
Marcolini, 2002; Marcolini et al., 2003) belonging
to the Fontana Ranuccio Faunal Unit (Gliozzi et al.,
1997). True Savi’s ground voles are reported since
the beginning of Late Toringian (Bartolomei, 1980),
corresponding to the latest phase of the Middle
Pleistocene. During the Late Pleistocene the geographic range of T. savi expanded or reduced due to
climatic changes. During the temperate-warm
oscillations, this species reached the Alpine region,
whilst during the cool or cold periods, it was
restricted to the southern and central parts of the
Italian peninsula (Kotsakis et al., 2003).
Studies on local populations of T. savii have been
carried out by several authors: De Giuli (1983),
Corridi (1987), Rustioni et al. (1994), Abbassi &
Brunet-Lecomte (1997), Masini & Abbazzi (1997),
Ronchitelli et al. (1998). A general analysis of fossil
and living populations of Italian ground voles has
been already published by Brunet-Lecomte et al.
(1994a). The present study focuses on the comparison of living and fossil populations of T. savii only.
Our target is to examine the relationships of recent
populations of central and southern areas of the
Italian peninsula with the fossil ones of the same
area. A similar work was performed by BrunetLecomte et al. (1994b) for Terricola gerbei (Gerbe,
1879) (= Terricola pyrenaica (de Sélys Longchamps,
1847)) of northern Spain and south-western
France.
Materials and Methods
The studied samples are derived from 11 localities. Five out of this number are fossil populations,
while six belong to living samples. The latter are
91
Fig. 1: Geographical location of the studied populations.
Fig. 2: Chronostratigraphy of the late Middle Pleistocene and
Holocene.
92
from: Cervia (Ravenna, Emilia Romagna), Civitella
del Tronto (Teramo, Abruzzi), Torraccia di San
Gennaro (Rome, Latium), Casarano (Lecce, Apulia),
Sila National Park (Calabria), Noto (Syracuse, Sicily).
The five fossil populations were collected in
Melpignano (Lecce, Apulia), Ingarano (Foggia,
Apulia), Praia a Mare (Cosenza, Calabria), Ostuni
(Foggia, Apulia) and Riparo Salvini (Latina, Latium)
(fig. 1).
As to the fossil localities, in the fossiliferous site of
Melpignano a fauna testyfing warm climatic conditions has been recognized. In particular the macrofauna collected in sediments of karst cavities allowed
its attribution to MIS 5a-5c (Bologna et al., 1994).
Petronio et al. (1996) assigned the assemblage from
Ingarano to the middle part of MIS 3. Capasso
Barbato & Gliozzi (2001) assigned the small mammal
assemblage from Praia a Mare to the final phase of
MIS 3. The Ostuni fossil assemblage is ascribed to
MIS 2 by Angelone et al. (2004). The fauna from Riparo
Salvini has been ascribed to the latest Pleistocene
(Tardiglacial - final phase of MIS 2) (Cassoli &
Guadagnoli, 1987; Alessio et al., 1993) (fig. 2).
The material studied in this research pertains to
public and private collections. In particular the fossil material from Melpignano and Ingarano is stored
in the Dipartimento di Scienze della Terra of the
University of Rome „La Sapienza“, the fossils from
Praia a Mare, Ostuni and Riparo Salvini are stored in
the Laboratory of Palaeontology of the Dipartimento di Scienze Geologiche of the University
Roma Tre. The recent material belongs to the
„Contoli Collection“ and it is stored in the
Dipartimento di Biologia Animale e dell’Uomo of
the University of Rome „La Sapienza“.
The decision to take dental measurements, particulary on the first lower molars (M1) (fig. 3b;
tabs. 2, 3) is necessary because the systematics of
the Arvicolidae is based on the morphology of this
tooth, and because teeth are often the only common fossil elements available. Quantitative and
qualitative analyses have been carried out on the
studied material. The pictures of teeth were taken
by using a digital camera Nikon Coolpix 995 connected to a stereoscopic microscope Nikon SMZ-U.
The measurements were carried out with the graphic program CorelDraw 8. The statistic analyses were
carried out with the program KyPlot ver.2.0 beta 15.
Some illustrations have been produced with the aid
of a Leica L2 camera lucida and of a graphical
tablet.
Fig. 3 : Morphology of M1 of the ground vole Terricola savii:
a-b) morphometry of Terricola M1 using 23 measures; c) M1 showing the characteristic apomorphy of the group, the Pitymyan
rhombus, the length of the tooth and the anterior loop (Brunet-Lecomte & Chaline, 1992).
Qualitative analyses were carried out in order to
recognize the dominant morphotype of each population. In a second step 23 measurements were
taken on the occlusal surface and some indices were
calculated, following the methods described by
Meulen (1973), Brunet-Lecomte (1990) and
Marcolini (2002) (fig. 3a,c): A/L: (var6-var3)
/var6*100; W/L: (var2/var6); W2/L: (var21/var6); RP:
(var4-var3) /var6*100 (fig. 3a,c).
These ratios give the relationship between the
length, the width, the curvature degree of the tooth
and the development stage of the Anteroconid
Complex (ACC) (Meulen, 1973), respectively.
Moreover multivariate statistical analyses,
Principal Component Analysis (PCA) and Canonical
Discriminant Analysis (CDA), were performed on the
measurements indicated by Brunet-Lecomte (1990)
and on the indices proposed by Meulen (1973); anyway it has to be underlined that these last ones
were calculated on measurements taken following
the method of Brunet-Lecomte (1988). Several
comparisons, with the aid of the previously mentioned statistical methods were made in order to
focus on the differences and/or the affinities
between the analyzed populations and the variability within a single population. Both in the Canonical
93
Discriminant Analysis and in the Principal
Component Analysis the populations were analyzed
in a first moment all together. Subsequently, these
same analyses (PCA and CDA) have been repeated
dividing the populations in fossil and recent ones
and all populations have been compared pair by
pair. In all tests an outgroup was present. The outgroup population comes from the lower level of
Gran Dolina (Atapuerca, Burgos, Spain) and it is
composed by Terricola arvalidens (Cuenca - Bescos
et al., 1995). This material has been found in a karst
filling sediment (approximately 18 meters thick)
partly ascribed to Early Pleistocene and partly to
Middle Pleistocene. This population is not temporally or geographically related to ours (both living and
fossil), nevertheless shows similar characteristics to
those of the studied populations and for this reason
has been included in the analyses.
The matrix used for PCA and CDA are available in the
site http://host.uniroma3.it/laboratori/paleontologia.
Results
Three different morphotypes were identified, on
the basis of the number of salient and re-entrant
angles, of the complication and development of the
Fig. 4: Morphotypes of Terricola savii: a,b,c) morphotype 1
(morphotype savii s.s.); d,e,f) morphotype 2; g,h) morphotype 3.
94
Anterior Loop (AL) and of the greater or smaller
confluence of the triangles in the Anteroconid
Complex.
MORPHOTYPE 1( morphotype savii s.s.) is characterized by a simple and wide anterior loop, with a wide
neck and widely confluent with the triangles T7 and
T6. T5 and T4 are broadly confluent. The reentrant
angles are quite marked and slightly more flattened
on the lingual side (fig. 4 a,b,c).
The anterior loop in MORPHOTYPE 2 is more complex than in morphotype 1. T7 and its reentrant
angle are much more evident while T6 and its reentrant angle are only outlined or even absent. T4 and
T5 are not confluent and consequently the
pitymyan rhombus is not clearly visible (fig. 4 d,e,f).
The anterior loop of MORPHOTYPE 3 is as simple as in
morphotype 1 although the triangles are rather
irregular in shape (fig. 4 g,h).
The analysis of the morphotypes shows a clear
dominance of morphotypes 1 and 2 in all the examined populations, both fossil and recent, while morphotype 3 is present only marginally in the recent
populations (tab.1).
As to the variability of the M1’s within the analyzed populations, as it is shown by the qualitative
data, it is clear that M1 follows a mosaic model
composed by the Anteroconid Complex (ACC),
which is more variable and characterizing most of
the morphotypes and by a more conservative
Talonid-Trigonid Complex (TTC). The observed variability is both inter- and intra-populational.
Moreover, it was possible to divide all the analyzed
M1 into two different morphotypes of both the
fossil populations and the living ones (the third
morphotype is present as we have seen only in the
living populations with low percentages) and, in
both cases, the percentages of the morphotypes
are similar. Nearly none of the performed PCA
have brought statistically significant results. In the
plots obtained by statistically significant analyses
there seem to be no differences within the fossil
populations or the recent ones. And there seem to
be no differences between fossil and living populations.
As to the living populations, differences have
been recognized between the populations of Noto
(Siracusa, Sicily) and Cervia (Ravenna, Emilia
Romagna), but this is a rather obvious result, being
geographically the two farthest populations within those considered. Moreover, the population of
Noto, coming from the island of Sicily, introduces
a)
b)
Fig. 5: On the diagram axis are plotted the scores of canonical
variables resulting from the Discriminant Canonical Analysis.
The two selected variables are those with the higher eigenvalues. The percentages reported along each axes are the explained variances of the variable taken into consideration. a)
Projection of the centroids of fossil populatios of T. savii; b)
Projection of the centroids of living populatios of T. savii.
all those problems which are typical of
insular populations (Petruso, 2002).
From the quantitative analyses conducted with CDA, some differences are
evident between the two groups (fossil
and living populations); the affinity and
homogeneity degree within the fossil
populations (heterochronic) (fig. 5a)
turns out to be smaller with respect to
the living populations (homochronic)
(fig. 5b).
From CDA the following observations
can be made:
a) The fossil Apulian populations (Melpignano,
Ingarano, Ostuni) and the living Apulian population
(Casarano) differ in a sensitive way from the other
analyzed populations (fig. 6), particularly from the
Calabrian ones (Praia a Mare and Sila National Park).
The Calabrian fossil (Praia a Mare) and living (Sila
National Park) populations, on the other hand, seem
to be different from the other elements pertaining
to the same group (fig.7). The Apulian populations,
both recent and fossil, show a large affinity, allowing to hypothesize the provenience of present-day
demes phylogenetically connected with palaeodemes of the same geographic area, from MIS 5a-5c
up to the present (fig. 2). Moreover, it is evident that
the population of Melpignano (MIS 5a-5c) is the
farthest from the living populations, followed by
that of Ingarano, confirming consequently the
biochronologic attribution of these fossil populations, obtained by means of the study of the entire
faunal assemblages. The fossil populations of
Ingarano (MIS 3) and Ostuni (MIS 2), and the living
one from Cassarano have a similar position on the
horizontal axis, but the living population is on a distinct position on the vertical axis (fig. 6).
b) There are some limits in the measurement
method proposed by Brunet-Lecomte, since such
measurements do not take in particular account the
anterior loop, neglecting what has turned out to be
the more variable morphologic feature in the qualitative analysis.
The morphologic/morphotypic variability of the
fossil populations fits that of the recent populations
(whose attribution to the same species is certain,
Fig. 6: Projection of the centroids of both fossil and living populations of
T. savii.
95
Fig. 7: Comparison between Apulian and Calabrian populations
of T. savii.
thanks to genetic analyses), therefore confirming
the correct attribution of the fossil populations to
the species T. savii.
eighties for the systematic studies of the family
Arvicolidae), can differentiate populations of different species and, in a more limited way, populations
of the same species.
The analysis of the fossil population from Praia a
Mare and the living one from Sila National Park
does not give any hint about the existence of
Terricola brachycercus (LEHMANN, 1961), an endemic
Calabrian species whose sympatric coexistence with
Terricola savii has been proved by genetic studies
(Galleni, 1995; Galleni et al., 1998 and references
therein). This discrepancy can be probably explained
by the absence of T. brachycercus from the analysed
sample as T. brachycercus has a very restricted distribution area and is sympatric with T. savii.
Nevertheless, Nappi et al. (2003) recognised differences between some Calabrian populations and T.
savii.
T. savii ground voles from Apulia, both fossil and
living ones, are rather homogeneous and differ from
other populations (fossil and living) of the species.
Apulia probably acted as a refuge area during the
cold oscillations of the Late Pleistocene. Moreover
geomorphological landscape (and consequently
environmental) differences between Apulia and the
Tyrrhenian side of the Peninsula influenced the
morphological divergence of the Apulian populations. Pioneers of T. savii from this region re-colonized the Adriatic side of the Italian peninsula during the Holocene.
The populations from the Tyrrhenian side of
Italy, Praia a Mare (MIS 3), Riparo Salvini
(Tardiglacial, latest MIS 2), Sila National Park and
Torraccia are very similar and differences between
fossil and living populations are minimal. On the
western (warmer) side of the Peninsula, T. savii
survived during the later part of the Late
Pleistocene and was almost isolated from the
Apulian populations.
Conclusions
Acknowledgments
PCA is not conclusive as the obtained results are
not statisticaly significant and it is impossible to
distinguish any important difference between the
eleven studied populations. However, this datum
confirms the attribution of all the material to the
same species, because this kind of analysis clearly
separates different species.
The differences obtained from the CDA demonstrate that the variables of the adopted measurements set (used in Europe since the end of the
96
We wish to thank Prof. B. Sala of the University
of Ferrara and Dr. K. Krainer of Innsbruck University
for revision of the manuscript.
References
Abbassi, M., Brunet-Lecomte, P. (1997): Terricola fatio
1867 (Arvicolidae, Rodentia) de cinq séquences du
Morphotype 1 Morphotype 2
Morphotype 3
Total
number
MELPIGNANO
66.67
33.33
0
18
INGARANO
66.67
33.33
0
12
PRAIA A MARE
71.43
28.57
0
14
OSTUNI
68.42
31.58
0
19
R. SALVINI
77.78
22.22
0
16
CERVIA
79.49
20.51
0
39
CIVITELLA
72.41
24.14
3.45
29
TORRACCIA
61.54
30.77
7.69
26
CASARANO
77.42
19.35
3.23
31
P.N.SILA
80.95
14.29
4.76
21
NOTO
59.46
37.94
2.60
37
Tab. 1: Percentages of the morphotypes for each
population of T. savii.
MELPIGNANO
INGARANO
PRAIA A MARE
OSTUNI
R. SALVINI
CERVIA
CIVITELLA
TORRACCIA
CASARANO
N.P.SILA
NOTO
N° of specimens
Minimum
Maximum
Mean
18
12
14
19
16
39
29
26
31
21
37
2.29
2.51
2.32
2.5
2.43
2.28
2.21
2.45
2.53
2.36
2.38
2.92
2.77
2.7
2.79
3.06
2.85
2.87
3.25
2.91
3.02
2.8
2.57
2.66
2.55
2.65
2.6
2.59
2.55
2.74
2.68
2.66
2.54
Standard deviation
v6
0.18
0.08
0.12
0.18
0.27
0.12
0.15
0.19
0.14
0.2
0.1
Tab. 2: Length of M1 of T. savii.
MELPIGNANO
INGARANO
PRAIA A MARE
OSTUNI
R. SALVINI
CERVIA
CIVITELLA
TORRACCIA
CASARANO
N.P.SILA
NOTO
N° of specimens
Minimum
Maximum
Mean
18
12
14
19
16
39
29
26
31
21
37
0.86
0.91
0.82
0.88
0.83
0.7
0.84
0.9
0.87
0.85
0.84
1.03
1.08
0.97
1.01
1.19
1.11
1.06
1.16
1.09
1.16
1.02
0.92
0.99
0.92
0,92
0.99
0.93
0.94
1.01
0.98
0.99
0.95
Standard deviation
v21
0.05
0.05
0.04
0,05
0.09
0.07
0.07
0.08
0.05
0.09
0.04
Tab. 3: Width of M2 of T. savii.
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micromammal association from Praia a Mare
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Cuenca-Bescós, G., Canudo, J.I., Laplana, C. (1995): Los
arvicólidos (Rodentia, Mammalia) de los niveles inferiores de Gran Dolina (Pleistoceno Inferior, Atapuerca,
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Galleni, L. (1995): Speciation in the Savi pine vole,
Microtus savii (de Sélys-Longchamps) (Rodentia,
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Biogeographical and karyological data of the Microtus
savii group (Rodentia, Arvicolidae) in Italy. – Bonn.
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Mezzabotta, C., Palombo, M. R., Petronio, C., Rook, L.,
Biochronology of selected mammals, molluscs and
ostracods from the middle Pliocene to the late
Paleont. Strat., 103: 369-388.
Kotsakis, T., Abbazzi, L., Angelone, C., Argenti, P., Barisone,
G., Fanfani, F., Marcolini, F., Masini, F. (2003): PlioPleistocene biogeography of Italian mainland micromammals. – DeinseA, 10: 313-342.
Marcolini, F. (2002) – Continental Lower Valdarno rodent
biochronology and two new methods for the systematics of Mimomys (Arvicolidae, Rodentia). – Ph.D.
Thesis, Università di Pisa, 165 p.
Marcolini, F., Bigazzi, G., Bonadonna, F.P., Cioni, R.,
Zanchetta, G. (2003): Tephrochronolgy and tephrostratigraphy of two Pleistocene continental fossiliferous successions of Central Italy. – J. Quat. Sci., 18:
545-556.
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Masini, F., Abbazzi, L. (1997) – L’associazione di mammiferi della grotta di Castelcivita. – In Gambassini, P.
(Ed.) – Il Paleolitico di Castelcivita, 33-59.
Meulen, A. van der (1973): Middle Pleistocene small
mammals from the Monte Peglia (Orvieto, Italy) with
special reference to the phylogeny of Microtus
(Arvocolidae, Rodentia). – Quaternaria, 17: 1-144.
Nappi, A., Montuire, S., Brunet-Lecomte, P. (2003) –
Sintesi sulla morfometria del primo molare inferiore
nel gruppo Microtus (Terricola) savii. – Hystrix, n.s.,
suppl., Abstr. IV Congr. Ital. Teriologia, p. 125.
Petronio, C., Billia, E., Capasso Barbato, L., Di Stefano, G.,
Mussi, M., Parry, S., Sardella, R., Voltaggio, M. (1996):
The late Pleistocene fauna of Ingarano (Gargano,
Italy): biocronological, palaeoecological, palaeoethological and geocronological implications. – Boll. Soc.
Paleont. Ital., 34(1995): 333-339.
Petruso, D. (2002): Il contributo dei micromammiferi alla
stratigrafia e paleogeografia del Quaternario continentale siciliano. – Ph.D. Thesis, Università di Napoli,
315 p.
Ronchitelli, A., Abbazzi, L., Accorsi, C.A., Bandini
Mazzanti, M., Bernardi, M., Masini, F., Mercuri, A.,
Mezzabotta, C., Rook, L. (1998) – The Grotta Grande di
Scario (Salerno – southern Italy): stratigraphy, archaeological finds, pollen and mammals. – Proc. 1st Intern.
Congr. “Science and Technology for the Safeguard of
Cultural Heritage in the Mediterranean Basin”, 15291535, Tip. Rustioni, M., Mazza, P., Abbazzi, L., Delfino,
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fauna from Sternatia (Lecce, Apulia, Italy).
Manuscript accepted: May 25, 2005
Geo.Alp, Vol. 2, S. 99–106, 2005
A TEST APPLICATION OF THE SHE METHOD AS A BIOSTRATIGRAPHICAL PARAMETER
Davide Mana
With 6 figures and 1 table
C.so Traiano 24/8, 10135 – Torino, Italia
Abstract:
Biodiversity – the index “expressing the variety and variability of living organisms and of the ecological
systems comprising them” (Ferrari, 2001) – is essential in the characterization and study of past and present
biological systems, and is generally expressed by a number (the “biodiversity index”), statistically derived
from empirical observations.
The SHE indexing method (Buzas & Hayek, 1996, Hayek & Buzas 1997), is expressed by the Shannon Index,
H (a measure of the system’s entropy) as the composition of two factors representing respectively the number of species in the sample (S) and the distribution uniformity (E).
The SHE index does not only describe in a thorough way the system’s biodiversity, but, as a function of
abundance and evenness, can be used to identify biofacies (SHEBI – SHE for Biofaces Identification) or to
characterize the whole structure of the analysed community (SHECSI – SHE for Community Structure
Identification).
SHE analysis, independently of its application purposes, appears to be highly flexible, does not require the
adoption of specific computer packages beyond a common spreadsheet, and is based on a simple graphical
analysis; widely adopted in botanics, SHEBI analysis in particular has been applied with satisfactory results
to the study of benthic foraminiferal faunas from the Atlantic ocean (Buzas & Hayek,1998].
In this work, the SHEBI method has been applied to 87 samples from the Falconara section (Southern
Sicily) – the purpose of the study is to verify the possibility of applying SHE/SHEBI to Messinian planktonic
foraminiferal assemblages.
Our study has to face issues that are typical of planctonic faunas – such as the lower number of species
and the ample variability in single taxa abundances; a further factor to be taken into account in setting up
and executing the analysis is the progressive deterioration of the ecosystem as the peak of the Messinian crisis approaches. Biofacies identification through SHEBI in less than ideal conditions, but on such a widely
studied and described section, offers an excellent opportunity to test the method and its limits, its application range and the reliability of its results.
1. Introduction – SHE and the measure of diversity
This work aims at identifying and evaluating the
limits (if any) of the application of the SHE analysis
to planktonic foraminiferal faunas, in order to simplify the application of this powerful diversitybased technique to the field of planktonic
foraminiferal biostratigraphy.
Diversity is one of the defining factors in any
study of an ecological system. A number of indices
was developed through the years by different
researchers, to quantitatively express diversity as
observed in the field or in laboratory; among the
more widely used indices are Fisher’s α index (a
measure of species richness), Simpson’s λ index,
Equitability (E, a measure of evenness) and
99
logarithmic transformation of the indices, and while
specific software is easily available to calculate the
values of H and E, given standard sample counts, the
whole analysis can be carried out on a simple
spreadsheet software (i.e., Microsoft Excell or
OpenCalc) with a minimum of fuss.
Conceptually, the analysis can be carried out
through time (i.e., vertically, comparing levels along
a geological section) or through space (i.e. laterally,
comparing sectors in a landscape).
Fig. 1: Location map of the Falconara outcrop in Sicily.
Shannon’s H index (a derivation of the information
function) (Smart, 2002).
Species richness itself (expressed as S, total number of species) has been used in the past as a rough
measure of diversity.
The recognition of the mathematical relation
between Species Richness and Taxa Abundance, and
its meaning in terms of Diversity and Dominance is
the basis of the recent SHE approach to the study of
biodiversity (Hayek & Buzas, 1997).
The mathematical expression summing up this
relationship is
(1) H = ln(S) + ln(E)
in which
H is the Shannon Diversity Index
S is the Species Richness
E is the Dominance or Evenness of the distribution
Relation (1) is constant as long as species proportions are constant.
As a change of the proportions of species to each
other is clearly a sign of change in diversity, the SHE
relationship has to be interpreted as an expression
of diversity.
This allows a simple graphical analysis of the
variations of biodiversity: each of the three variables can in fact be plotted against the Abundance
(N) of the sample; changes in proportions (and
therefore in diversity) will be signalled by a change
in the graphic line slope (a “slope break” in the “hollow curve” following Hayek & Buzas’ terminology).
Operationally, the method is not as mathematically intensive as other well-established analysis
procedures (i.e., Cluster Analysis), requiring simply a
100
Introduced in the late 1990s as a way to sidestep
some perceived limits in more popular diversity
indices (Shannon-Weiner in particular), SHE’s field
of application was later extended and redefined,
with the introduction of SHEBI (SHE Analysis for
Biofacies Identification) and SHECSI (SHE Analysis
for Community Structure Identification) (Buzas and
Hayek, 1996, 1998 ; Hayek and Buzas, 1997, 1998).
Examples of applications of the SHE approach to
biodiversity have been published as part of botanical (Hayek and Buzas, 1996, 1997, Small and
McCarthy 2002) and zoological studies (Leponce et
al. 2004); closer to the concerns of this paper, SHE
has been applied to the study of quaternary benthic foraminiferal faunas in what can be defined as a
non-perturbed environmental setting (Buzas and
Hayek, 1998, Osterman et al, 2002).
By all accounts, when applied to current or
recent environments and populations, SHE appears
to be a solid, easily applied method for describing
diversity; in particular, it allows a high-resolution
visualization of changes in diversity through time or
space; the method allows researchers “to examine
evenness separately from richness within a single
multispecies system” (Buzas and Hayek, 1998) and
it does not suffer from some of the limits signalled
for other diversity-based indices (Hayek and Buzas,
1997).
Some doubts might still remain when SHE is to
be applied to situations in which those factors the
method takes into account (population density, specific richness, etc.) are subject to extreme or unpredictable variations – i.e. due to drastic changes in
environmental conditions, or to other external
causes.
To verify the viability of SHE analysis in such critical conditions, this study has been carried out on
planktonic foraminiferal faunas from Messinian
strata of the Mediterranean, which are normally
characterized by lower species richness (S) than ben-
thic faunas. Proximity to the peak of the Messinian
Salinity Crisis further weakens the species richness
signal, due to increased environmental stress.
This paper briefly summarizes the study and its
results.
2. The Falconara Section
and the planktonic samples
The samples used for this study were collected in
the alternating clay/diatomite cycles of the Tripoli
formation (Upper Tortonian-Messinian) with an
exposed thickness of one hundred meters in the
Falconara Section.
Located on the southern face of Monte
Caltagirone, on the southern coast of Sicily between
Gela and Licata (see fig. 1), the Falconara Section
(fig.2) was originally proposed as the type-section
for the Messinian (Colalongo et al., 1979), and has
been the object of continuing studies, criticism and
revisions, due to its paramount importance for the
comprehension of Mediterranean events; in more
than thirty years, studies have shifted from biostratigraphical and chronostratigraphical concerns
and techniques to cyclostratigraphical and astrochronological methods. (summarized in Hilgen et al.,
2000).
The abundance of previous studies and the
detailed description of the Falconara faunas
(Colalongo et al., 1979, Hilgen & Krijgsman, 1999,
Hilgen et al., 2000) by previous authors provides an
excellent background for our test-run of the SHE
approach to planktonic foraminifera biostratigraphy. Our study does not mean to redefine in any
way the stratigraphy of the Falconara section, but
to use a well-studied section and its wealth of
accumulated paleontological and stratigraphical
knowledge as the consensus against which the
results of the SHE test will be compared for validation.
The samples used in this study were collected
from the Falconara Section in 1994 (fig. 3), as part
of a wide-ranging campaign of studies on the
Messinian Salinity Crisis in the Mediterranean; in
the field, both clay and diatomite layers were sampled separately, and were later subjected to standard micropaleontological analysis and quantitative
studies in the laboratories of the Università degli
Studi, Torino.
Fig. 2: View of the Falconara outcrop.
The environmental information provided by the
faunas contained in the sediments was presented
and discussed in the author’s graduation paper
(Mana, 2001) concerning the same samples used in
this study; in that work, a general biozonation
based on a traditional method (Cluster Analysis),
was proposed, identifying seven distinct biofacies,
each connected with the progressive environmental
crisis of the Messinian sea.
That work, and the excellent synthesis by Hilgen
and Krijgsman (1999) will be our two chief references for comparison.
3. SHE Analysis
For the purposes of this study, 87 samples were
observed, and 300 individuals counted according to
standard statistical data-gathering practices; seventeen planktonic taxa were recognized (see below)
and counted; to these, a class labelled “others” was
added to include the few non-planktonic individuals (mostly Bulimina echinata). For the species
Neogloboquadrina acostaensis, sinistral coiling
individuals were counted separately from dextral
coiling individuals.
101
The taxa used in this study are:
Globigerina angustiumbilicata
Globigerina sp
Globigerinoides ruber
Globigerinoides sp
Turborotalita multiloba
Turborotalita sp
Globorotalia conoidea
Globorotalia praemenardi
Globorotalia sp
Neogloboquadrina continuosa
Neogloboquadrina acostaensis sin.
Neogloboquadrina acostaensis dex.
Orbulina universa
Globigerinella obesa
Globigerinella praesiphoniphera
Globigerinella sp
Sphaeroidinellopsis
Other
Fig. 3: Summary sketch of the Falconara Section, with sample
numbers.
102
As described in Buzas and Hayek (1998), from the
species counts, the cumulative values of N (number
of individuals in sample), S (number of species in
sample or Specific Richness), H (Shannon’s Index)
and E (Evenness) were calculated using an Excell
spreadsheet, and the natural logarithms extracted
for each value (Table 1).
Cumulative values (a stepwise addition of values)
were used so that S will be steadily increasing
through the sequence.
Considering now equation
(1) H = ln(S) + ln(E)
as we have already stated, this relation remains constant as long as species proportions remain constant. More to the point, if – as in the case of our
analysis – the value of S increases steadily due to
the cumulative process, two possibilities can
become apparent: if, as S increases, H remains constant, this will mean a progressive decrease in the
value of the samples’ cumulative Evenness; should
instead the value of ln(E) remain constant, this
would mean a progressive variation in the value of
H.
Plotting linear graphics for
ln(S) vs ln(N)
H vs ln(N)
ln(E) vs ln(N)
allows us to pinpoint biofacies changes, represented
by slope breaks on the graphs (fig. 4).
SAMPLE
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
20
22
24
25
26
27
29
31
33
34
35
37
38
39
40
42
43
44
N
307
288
329
302
315
303
300
327
329
396
285
319
286
296
323
312
328
303
305
172
313
293
209
303
289
300
309
304
321
304
291
316
302
300
318
(N)
307
595
924
1226
1541
1844
2144
2471
2800
3196
3481
3800
4086
4382
4705
5017
5345
5648
5953
6125
6438
6731
6940
7243
7532
7832
8141
8445
8766
9070
9361
9677
9979
10279
10597
ln(N)
5,73
6,39
6,83
7,11
7,34
7,52
7,67
7,81
7,94
8,07
8,16
8,24
8,32
8,39
8,46
8,52
8,58
8,64
8,69
8,72
8,77
8,81
8,85
8,89
8,93
8,97
9,00
9,04
9,08
9,11
9,14
9,18
9,21
9,24
9,27
(S)
7
11
20
22
26
31
31
32
34
35
37
37
38
39
45
46
48
48
49
50
50
50
50
51
52
53
53
54
54
54
54
54
54
54
54
ln(S)
1,95
2,40
3,00
3,09
3,26
3,43
3,43
3,47
3,53
3,56
3,61
3,61
3,64
3,66
3,81
3,83
3,87
3,87
3,89
3,91
3,91
3,91
3,91
3,93
3,95
3,97
3,97
3,99
3,99
3,99
3,99
3,99
3,99
3,99
3,99
H
0,97
1,00
0,86
0,62
0,62
0,53
0,43
0,38
0,34
0,40
0,31
0,27
0,29
0,28
0,30
0,26
0,21
0,23
0,22
0,13
0,20
0,19
0,13
0,20
0,19
0,17
0,17
0,16
0,17
0,15
0,14
0,14
0,14
0,13
0,14
(E)
0,38
0,25
0,12
0,08
0,07
0,05
0,05
0,05
0,04
0,04
0,04
0,04
0,04
0,03
0,03
0,03
0,03
0,03
0,03
0,02
0,02
0,02
0,02
0,02
0,02
0,02
0,02
0,02
0,02
0,02
0,02
0,02
0,02
0,02
0,02
LN(E)
-0,97
-1,40
-2,14
-2,47
-2,64
-2,91
-3,01
-3,09
-3,18
-3,16
-3,30
-3,34
-3,34
-3,39
-3,51
-3,57
-3,66
-3,64
-3,67
-3,78
-3,72
-3,72
-3,78
-3,73
-3,77
-3,80
-3,80
-3,83
-3,82
-3,84
-3,85
-3,85
-3,85
-3,85
-3,85
SAMPLE
45
47
48
49
50
51
52
53
54
55
56
57
58
59
61
62
64
65
67
68
70
72
73
74
75
76
77
78
79
80
81
82
84
85
87
N
316
304
341
343
337
396
347
317
354
326
300
307
198
327
331
350
349
287
344
311
314
320
72
347
332
350
304
197
300
370
338
345
201
267
332
(N)
10913
11217
11558
11901
12238
12634
12981
13298
13652
13978
14278
14585
14783
15110
15441
15791
16140
16427
16771
17082
17396
17716
17788
18135
18467
18817
19121
19318
19618
19988
20326
20671
20872
21139
21471
ln(N)
9,30
9,33
9,36
9,38
9,41
9,44
9,47
9,50
9,52
9,55
9,57
9,59
9,60
9,62
9,64
9,67
9,69
9,71
9,73
9,75
9,76
9,78
9,79
9,81
9,82
9,84
9,86
9,87
9,88
9,90
9,92
9,94
9,95
9,96
9,97
(S)
54
54
54
54
54
54
56
56
57
57
57
57
57
57
58
58
60
60
60
60
60
60
60
60
60
60
60
60
60
60
60
60
60
60
60
ln(S)
3,99
3,99
3,99
3,99
3,99
3,99
4,03
4,03
4,04
4,04
4,04
4,04
4,04
4,04
4,06
4,06
4,09
4,09
4,09
4,09
4,09
4,09
4,09
4,09
4,09
4,09
4,09
4,09
4,09
4,09
4,09
4,09
4,09
4,09
4,09
H
0,14
0,11
0,11
0,15
0,10
0,14
0,13
0,11
0,12
0,11
0,11
0,11
0,07
0,12
0,12
0,10
0,11
0,09
0,11
0,10
0,10
0,10
0,03
0,10
0,08
0,08
0,07
0,06
0,08
0,08
0,08
0,09
0,05
0,06
0,07
(E)
0,02
0,02
0,02
0,02
0,02
0,02
0,02
0,02
0,02
0,02
0,02
0,02
0,02
0,02
0,02
0,02
0,02
0,02
0,02
0,02
0,02
0,02
0,02
0,02
0,02
0,02
0,02
0,02
0,02
0,02
0,02
0,02
0,02
0,02
0,02
LN(E)
-3,85
-3,88
-3,87
-3,84
-3,89
-3,85
-3,90
-3,91
-3,93
-3,93
-3,93
-3,93
-3,98
-3,93
-3,94
-3,97
-3,98
-4,00
-3,99
-3,99
-4,00
-3,99
-4,06
-4,00
-4,01
-4,01
-4,03
-4,03
-4,01
-4,01
-4,02
-4,01
-4,04
-4,04
-4,02
Table 1: List of Falconara samples showing calculated values of indexes for SHE analysis. Missing samples were found to be sterile.
Abbreviations: N, counted individuals; (N), cumulated N; (S), cumulated species richness; H, Shannon’s Index; (E), Evenness.
In fig. 4 the three “hollow curves” (Hayek &
Buzas, 1997, 1998) are plotted in a single graph in
the same order in which we introduced them above;
as our fig. 4 shows, a number of breaks are evident,
each of them potentially marking a change in association, and therefore in biofacies.
It is important at this point to notice that matters of scale, and the high number of individuals
projected, might distort the curve plot, causing a
loss of definition and actually masking some significant slope breaks. To avoid this distortion effect,
the suggested practice consists in breaking the
sequence into smaller intervals – which is achieved
in practice by stepwise deleting the samples whose
trend has already been analysed, recalculating all
the values in the system.
The stepwise deletion procedure also corrects
another important distortion which may present
the single-plot SHE model in fig. 4 – the one caused
by the disappearance of certain taxa as the
sequence develops. Cumulative addition of Specific
Richness alone, does account for the appearance of
new species, but not for the loss of those species
that, while present in the earlier levels of the
sequence, disappear later. By stepwise deleting earlier data-points from the plot as the analysis pro-
gresses, and recalculating the values of S, N, E and
H, disappearances are now computed into the
model.
For the purposes of this work, the SHE analysis
procedure was applied six times (fig. 5) in order to
heighten the definition of the hollow curve.
The resulting graphs appear choppy and uneven,
especially when compared to similar plots for benthic faunas (Buzas and Hayek, 1998]; this is an
effect most likely caused by the characters of the
planktonic assemblage (low Specific Richness, sudden disappearances) and the time interval considered (wide and sudden variations in environmental
conditions as the situation evolves towards the crisis). The operator has also to take into account the
very low values of the indices, a product of the generally low Species Richness and of the scarcity of
biological remains in some samples (fifteen of
which lack fossils).
Our biofacies analysis is based chiefly on the joint
observation of both ln(S) and ln(E) plots; the latter
is considered to be most sensitive to specific assemblage changes by most authors, but considering the
scarcity of species represented in the samples, and
the low abundances, using the former as a control
and as a support in the definition of biofacies
103
over forty couples can be observed in the field – and
which were used as a basis for sampling in this
study, and by many other authors (Hilgen and
Krijgsman, 1999).
The definition of the SHEBI method is excellent,
resolving in some cases changes in population balance (and therefore, in diversity) that occur at the
scale of the single clay/diatomite couple; these were
not considered in this work, as each should deserve
a much more detailed analysis and assessment, but
are shown in the plots collected in fig. 5, in which
they appear as brief breaks in the slope of the hollow curve.
4. Conclusions and future developments
Fig. 4: SHE Analysis of Falconara samples, summary graph.
Data-points (samples) have been thinned to improve readability.
breaks appears as an advisable line of conduct.
The analysis leads to the definition of 21 intervals
which can be considered each characterized by stable or near-stable conditions, their assemblages
being therefore distinct biofacies.
Packing so many biofacies in a stretch of about
one hundred meters could be considered embarrassing by someone – especially when compared to the
seven biofacies intervals identified using the same
samples and a more traditional discrimination
approach (Cluster Analysis) in a previous work
(Mana, 2001); and yet the intervals as identified by
the method are undeniably a result of the observed
species and counted abundances. And the fine subdivision of the Falconara sequence also reflects the
rhythmic cycles of clays and diatomites, of which
104
The SHE/SHEBI method is as reliable as more traditional approaches when applied to planctonic
faunas, and does not require any ad hoc modification. In particular, the differences between planctonic and benthic faunas do not seem to hinder the
application of the method, but simply require a
higher degree of attention on the part of the
researcher.
Similarly, conditions of progressive environmental
crisis do not seem to compromise the method’s functionality, and are easily recorded by the “hollow
curve”. By working on Species Richness S and
Evenness E, SHEBI seems to compensate the progressive loss of data due to thinning of the association
through time.
The biozonation obtained from the application of
the SHE method appears to be consistent with previous zonations obtained through different analytical approaches (such as Cluster Analysis), but shows
a higher sensitivity to minute changes in population
balance, and therefore a higher resolution.
Also, the method leaves a higher degree of freedom
to the operator, who is allowed to fine-tune his
interpretation of the graphs based on his knowledge
of local peculiarities.
While probably regionally restricted due to the
probability of sudden changes in planctonic associations, SHEBI zonation still appears to be an excellent correlation tool when used on different sections – and indeed this seems to be one of the more
promising directions in which future investigation
about the applications of SHE to Messinian faunas
might expand; similarly, the possibility of coupling
the biozonation tool offered by SHEBI with
Fig. 5: SHE Analysis of the Falconara faunas; stepwise deletion of samples examined earlier with each new iteration. Vertical lines
show the position of biofacies breaks.
palaeoecological assessing tools such as ordination
methods (PCA, DCA) might hold great promise for
future developments (Mana, 2004].
(University of Marseilles), for allowing the use of the
samples in the first place.
References
5 . Acknowledgments
The author wishes to express his gratitude to
Prof. Donata Violanti (Università degli Studi, Torino)
for the support and the advice concerning the
Falconara samples, and to professor Jean Pierre Suc
Buzas, M.A., Hayek, L.C. (1996): Biodiversity Resolution: an
integrated approach. – Biodiversity Letters, v. 3: 40-43.
Buzas, M.A., Hayek, L.C. (1998): SHE Analysis for Biofacies
Identification. – Journal of Foraminiferal Research, v.
28: 233-239.
105
Fig. 6: Schematic comparison of the biozonation based on Cluster Analysis [Mana, 2001], and the SHE biozonation (this
work). Colors are purely indicative and have no stratigraphical
meaning.
106
Colalongo, M.L., di Grande, A., D’Onofrio, S., Giannelli, L.,
Iaccarino, S., Mazzei, R., Poppi Brigatti, M.F., Romeo,
M., Rossi, A., Salvatorini, G., (1979): A proposal for the
Tortonian/Messinian boundary. – Ann. Géol. Pays
Hellén., Tome hors série, v. 1, pp. 285-294.
Ferrari, C (2001): Biodiversità. – 36 pp, Zanichelli,
Bologna.
Hayek, L.C., Buzas, M.A. (1997): Surveying Natural
Populations. – 563 pp., Columbia University Press, New
York.
Hayek, L. C., Buzas, M. A. (1998): SHE analysis: an integrated approach to the analysis of forest biodiversity.
– In Dallimeier, F., Comkey, J. (eds.) Forest Biodiversity
Research, Monitoring and Modeling, 311-321,
UNESCO and Parthenon Publishing Group, Paris.
Hilgen, F.J., Krijgsman, W. (1999): Cyclostratigraphy ad
astrochronology of the Tripoli diatomite formation. –
Terra Nova, vol. 11, No. 1: 16-22.
Hilgen F.J., Iaccarino S., Krijgsman,W., Villa, G., Langereis
C.G., Zachariasse W.J. (2000): The Global Boundary
Stratotype Section and Point (GSSP) of the Messinian
Stage (uppermost Miocene). – Episodes, Vol. 23,
no. 3:172-178.
Leponce, M., Theunis, L., Delabie, J. H. C. and Roisin, Y.
(2004): Scale dependence of diversity measures in a
leaf-litter ant assemblage. – Ecography v. 27: 253-267.
Mana, D. (2001): Metodi Quantitativi applicati allo studio
dei Foraminiferi del Messiniano di Falconara (Sicilia
Meridionale). – Università degli Studi, Torino, graduation research paper.
Mana, D. (2004): SHE Characterization of the Planktonic
Foraminifera Assemblages from the Falconara and
Capodarso Sections (Messinian), Sicily, Italy. – Oral
presentation to the 32nd International Geological
Congress, Florence, August 20-28, 2004; abstract in
32nd International Geological Congress, Abstracts,
Part 1, IUGS, Florence.
Osterman, L.E., Buzas, M.A., Hayek, L.C. (2002): SHE
Analysis for Biozonation of Benthic Foraminiferal
Assemblages from Western Arctic Ocean. – PALAIOS,
v. 17: 297-303.
Small, C.J., McCarthy, B.C.(2002): Spatial and temporal
variability of herbaceous vegetation in an eastern
deciduous forest. – Plant Ecology, v. 64: 37-48.
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Revised manuscript accepted: June 2, 2005
Geo.Alp, Vol. 2, S. 107–113, 2005
URSUS SPELAEUS ROSENMÜLLER, 1794 FROM THE VENETIAN REGION OF NORTHERN ITALY:
PRELIMINARY NOTES ON ITS EVOLUTIONARY PATH
Cinzia Galli1, Mario Rossi2, Giuseppe Santi3
With 6 figures and 1 plate
1 Museo Civico di Storia Naturale, Parco del Vecchio Passeggio, I-26100 Cremona (Italy)
2 Museo Civico di Storia Naturale, Lungadige Porta Vittoria 9, I-37129 Verona (Italy)
3 Dipartimento di Scienze della Terra, Via Ferrata 1, I-27100 Pavia (Italy), e-mail [email protected] (corresponding author)
Abstract
According to morphometric data, population remains of bears ascribed to the deningeri-spelaeus group have been
identified in caves of the Venetian region of Northern Italy: Cerè Cave, Covoli di Velo, San Donà di Lamon and Veja.
Some initial observations about the evolutionary path of these cave bears are presented.
Introduction and previous work
radiometric data are available for only a few caves in
Northern Italy. The best known is the Grotta Sopra
Fontana Marella (Varese Province, Lombardy). For other
caves (e.g. the Caverna Generosa, Varese Province) data
are still incomplete (Bona, 2004) and are lacking in the
Venetian region. Radiometric data and aminoacid
Numerous caves distributed over the Alpine and preAlpine areas including Cerè Cave, the Covoli di Velo and
Veja caves (Province of Verona), and the San Donà di
Lamon cave (Province of Belluno) (Fig. 1) are of great
importance, both historically and for the
abundance of bear fossils. They provide a
large number of morphometric data
stimulating several interesting considerations on the presence of these fossils in
the Venetian region. The studied deposits
show that these areas were inhabited
either by Ursus spelaeus Rosenmüller,
1794, or Ursus arctos Linnaeus, 1758
albeit in different proportions; in fact,
the cave bear – Ursus spelaeus - represents the most abundant species. Until
now, the presence of Ursus deningeri Von
Reichenau, 1906 has only been assumed
(Zorzin et al., in press). The most recent
studies (i.e Rossi & Santi, 2005) on newly
found fossils from the Cerè Cave, the
most significant results of which will be
presented in this paper, have confirmed
the presence of this species. Currently,
Fig. 1: Geographic position of the main caves of the Venetian region.
107
Fig. 2: Geographic position of the Cerè Cave. A – Lateral wall of the cave in bone breccias. B – Entrance of the Cerè Cave.
racemization of the bear bones from Grotta Sopra
Fontana Marella provide the following ages: FM4 over
26000 years BP, FM2, 22310 ±200 and FM1 21810±200
years BP (Perego et al., 2001).
For many years the Venetian caves have constituted
an important research target. A review of the inventory
of the Pleistocene-Holocene fauna from these caves was
compiled by Bon et al. (1991, cum bibl.) based on fossils
stored at different localities in Northern Italy. More
recently, studies on populations of bears and other fossil
groups from the Cerè Cave, Covoli di Velo, San Donà di
Lamon and Veja caves have been published by Rossi &
Santi (2001 a, b, 2002), Zorzin et. al. (2003, 2004 and in
press) and Rossi et al. (2004).
Brief background on the stratigraphy
of the Cerè Cave
The Cerè Cave, known also as the “Tana dell’Orso” or
the “Tanasela” (Fig. 2), is located at an altitude of about
750 m a.s.l. and is 12 m deep; it opens at the hydrographically right side of the Vajo dell’Anguilla within the
Rosso Ammonitico Formation about 150 m east of
Ceredo (S. Anna di Alfaedo) village. The entrance is near
a distinct fracture of the slope that characterizes the
right side of the Vajo dei Falconi. From bottom to top, the
108
7.50 m thick stratigraphic succession is composed (Zorzin
et al., 2003) as follows:
1. Ferrous-manganesiferrous clay containing concretions (at the karstic bed rock contact).
2. Calcareous concretions, locally very thick.
3. Fine-grained, mixed carbonate-siliciclastic sand with
small amounts of clay filling the bottom of the
depressions and the karst fissures. Locally, a thin layer
of yellow or reddish clay is present below concretion 4.
4. Concretion rich in siliceous and patinated detrital
material.
5. Plastic clay containing pebbles up to 1 cm in size.
6. Horizon with concretions.
7. Plastic red clay containing rare fossil remains and
siliceous detrital fragments with diameters up to 5
cm.
8. Red clay with abundant pebbles of chert and slightly
altered gravel.
9. Dark layer rich in bone remains mostly belonging to
Canis lupus containing concretions and rich in
siliceous and rare chert pebbles with diameters of 1 to
3 cm.
10. Dark layer rich in bone remains predominantly
belonging to Ursus, with calcareous pebbles about 2
cm in size.
11. Strongly cemented bone-breccia, with abundant
remains of Ursus, Canis lupus and Marmota.
Fig. 3: Ratio between “basal length” and “length of dental row” for bear skulls from Italian and other localities.
Fig. 4: Ratio between “absolute length” and “height of vertical branch” for bear mandibles from Italian and other localities.
12. Breccia containing small amounts of sediment composed of strongly cemented large blocks.
13. Breccia with chert pebbles from 1 to 3 cm in size.
14. Calcareous breccia with chert pebbles from 1 to 5 cm
in size.
15. Breccias with chert pebbles from 1 to 5 cm in size.
Distribution of Ursus species
in the Venetian region
Before presenting the main morphometric data, we
believe it is useful to indicate the distribution of the
Ursus species in the following caves: 1) Cerè Cave: Ursus
109
Fig. 5: Ratio between “absolute length” and “transversal width of the diaphysis” for the II metatarsus of bears from the Cerè Cave
and other Italian and other caves.
Fig. 6: Ratio between “absolute length” and “transversal width of the diaphysis” for the III metatarsus of bears from the Cerè Cave
and other Italian and other caves.
110
deningeri, U. spelaeus, U. arctos; 2) Covoli di Velo: Ursus
spelaeus; 3) Veja: Ursus spelaeus; 4) San Donà di Lamon:
Ursus spelaeus (Pl. 1). Considering the rarity of fossils
pertianing to U. deningeri not only in Northern Italy, but
also in the rest of the peninsula, their presence within the
Cerè Cave is of great importance.
a consequence of a limited expansion of these former
groups that were able to colonize only in this limited area
in Northern Italy. The remaining zones could have been
further colonized starting from a supposed initial point,
represented by the Venetian populations originally from
Central Europe, which experienced a rapid and articulated evolution.
Morphometry
Preliminary concluding remarks
Morphometric analysis was carried out on several
hundreds of fossils from a large portion of the skeleton
(except for the vertebrae, ribs and other anatomic parts
whose limited number of specimens prevented an indisputable analysis) stored in the Museo Civico di Storia
Naturale of Verona and compared with other fossils from
Northern Italy (Grotta del Buco dell’Orso – Laglio, Como
Province; Grotta Sopra Fontana Marella - Varese
Province; Grotta delle Streghe – Sambughetto Valstrona,
Verbania Province) including foreign examples, particularly from Spanish caves (Torres, 1988). The findings have
allowed us to advance a number of hypotheses (Figs. 3-6).
a) Cerè Cave: The morphometric data show the presence
of populations from the deningeri-spelaeus group
and the large number of fossils, especially of the
metapodial bones, have confirmed the above mentioned observations.
b) Covoli di Velo: Unlike the Cerè Cave, the fossils are
exclusively from larger-sized bears while those in
medium- to small-size ranges ones appear to be very
poorly represented.
c) San Donà di Lamon and Veja: The morphometric
analysis of the rather limited remains in these localities confirms the presence of relatively medium- to
large-sized populations similar to those that lived in
the Covoli di Velo region.
Hypothesis about the possible
Ursus deningeri “track of ways”
The presence of the deningerian remains in the Cerè
Cave, rarely found in Central Italy and the Alpine and
pre-Alpine sectors of the Western and Central Alps, may
indicate migration paths that initially followed a N-Sdirection, possibly encouraged by the overall mildness of
the climate in the more southern regions, and later also
in an E-W-direction. The lack of the Ursus deningeri
remains in other areas may be due to a gap in the fossil
record linked to inadequate fossil preservation or unsuccessful discovery of the deposits. However, it may also be
Based on the morphometric analysis that shows the
bear fossils belong to the deningeri-spelaeus group, some
preliminary conclusions can be drawn:
a) The main caves of the Venetian region were inhabited by bears of the deningeri-spelaeus group, but in
the Cerè Cave the continuous presence of both
Ursus deningeri and Ursus spelaeus (medium- to
large-sized) from their intermediate to final evolutionary stages is certain. In other regions only large
sized cave bear populations are evident and linked
to the final phase of the evolutionary path of this
species.
b) The presence of the three species in the Cerè Cave
indicates its prolonged inhabitation in ancient times
compared to the other caves. Hence, the Ursus
deningeri population may represent the original
nucleus from which subsequent forms may have
developed with their final examples being discovered
in the other caves examined. These populations are
morphometrically comparable to those from the more
recent beds of the Grotta Sopra Fontana Marella
dated 21810±200 years BP (Perego et al., 2001).
Some data indicate the presence of Ursus deningeri in
the Delle Ossa Cave near Zandobbio village (Bergamo
Province, Lombardy), but further investigations are
required to confirm its occurrence in this area. If future
research confirms the exclusiveness of the findings in the
Cerè Cave, its importance will increase. In fact, on the
basis of this data, this zone could represent an expansion
nucleus for the Venetian region as well for the whole of
Northern Italy.
Acknowledgments
The authors thank Prof. D. Nagel (Vienna) for useful
advice and critical reading of the manuscript and Dr. G.
Papalia (Pavia) for revision of the English.
This study was supported by a FAR grant contribution.
111
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Rosenmüller & Heinroth, 1794 populations from
“Caverna Generosa” (Lombardy-Italy). – Cahiers
Scientifiques, Hors série 2: 87-98.
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from Grotta sopra Fontana Marella, Campo dei Fiori
Massif (Varese, Italy): morphometry e paleoecology. –
Riv. It. Paleont. Strat., 107 (3): 451-462.
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Grotta del Cerè (Verona). 1 – Prime osservazioni sui
resti craniali e mandibolari di ursidi. – Bollettino del
Museo Civico di Storia Naturale di Verona, sez. di
Geologia, Paleontologia e Preistoria, 25: 59-72.
Rossi, M., Santi, G. (2001b): Archaic and recent Ursus
spelaeus forms from Lombardy and Venetia region
(North Italy). – Cadernos Lab. Xeológico de Laxe
Coruña, 26: 317-323.
Rossi, M., Santi, G. (2002): Gli ursidi dei Covoli di Velo
(Verona) e di S. Donà di Lamon (Belluno). I –
Preliminare analisi morfologica e morfometrica dei
resti craniali e mandibolari. – Bollettino del Museo
Civico di Storia Naturale di Verona, sez. di Geologia,
Paleontologia e Preistoria, 26: 33-41.
Rossi, M., Santi, G. (2005): What differences between
Ursus deningeri Von Reichenau and Ursus spelaeus
Rosenmüller-Heinroth? The bear mandibles from
Venetia Region caves (N. Italy). – “V Giornate di
Paleontologia” Urbino 20-22 Maggio 2005, Abstracts
vol., p. 61.
Rossi, M., Santi, G., Zorzin, R. (2004): Distribuzione di
Ursus gr. deningeri-spelaeus nell’Italia Settentrionale
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Torres Pérez Hidalgo T. (1988) - Osos (Mammalia,
Carnivora, Ursidae) del Pleistocene Ibérico (U.
deningeri Von Reichenau, U. spelaeus RosemüllerHeinroth, U. arctos Linneo). – Boletín Geológico y
Minero. I Filogenia, distribution stratigrafica y
geografica. Estudio anatomico y metrico del craneo:
3-46. II Estudio anatomico y metrico de la mandibula,
hioides, atlas y axis: 220-249. III Estudio anatomico y
metrico del miembro toracico, carpo y metacarpo:
359-412. lV Estudio anatomico y metrico del miembro
pelviano, tarso, metatarso y dedos: 516-577. V
Dentiction decidual, formula dentaria y denticion
superior: 660-714. VI Denticion inferior: 886-940.
Zorzin, R., Santi, G., Rossi, M. (2003): I principali mammiferi quaternari della Grotta del Cere’ (Monti Lessini
- VR) conservati presso il Museo Civico di Storia
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Cranium.
Plate 1: Ursus spelaeus Rosenmüller, 1794. A – Skull V160 (Cerè Cave), dorsal view, B – Skull V 162 (Cerè Cave), dorsal view, C – Mandible V 4673 (Cerè Cave), internal lateral view, D – Mandible V 2886 (Veja), external lateral view, E – Mandible V 2887 (Veja), internal
lateral view, F – Mandible V 9889 B (Covoli di Velo), external lateral view, G – Skull V 161 (Cerè Cave), lateral view.
112
113
Geo.Alp, Vol. 2, S. 115–126, 2005
MORPHOMETRY OF THE URSUS SPELAEUS REMAINS FROM VALSTRONA (NORTHERN ITALY)
Alessandro de Carlis1, Enrico Alluvione1, Alessandro Fonte1, Mario Rossi2 & Giuseppe Santi1
With 14 figures and 2 plates
1 Dipartimento di Scienze della Terra, Via Ferrata 1, I-27100 Pavia (Italy); e-mail: [email protected] (corresponding author)
2 Museo Civico di Storia Naturale, Lungadige Porta Vittoria 9, I-37129 Verona (Italy)
Abstract
Morphometric analyses on cave bear fossils of the Valstrona-Valsesia region (Piedmont, Northern Italy) (in
particular from the Delle Streghe Cave), allow the distinction of at least two or three populations of Ursus
spelaeus Rosenmüller, 1794, of different sizes. Elements of smaller size are likely to be found in the Buco
dell’Orso Cave (Laglio, Como province, Lombardy), and in the older strata of the Grotta Sopra Fontana Marella
–GSFM- (Varese province). Differences in size can be linked to the chronological position of the fossils studied: in fact, fossils of smaller dimensions should be chronologically older, but can also be linked to climatic
and thermoregulation factors. The increase of size could represent a response to a cooler climate. An alternative hypothesis associates this reduction of size to the cooler climatic shift. Observations on the Delle
Streghe Cave fossils indicate that they are very similar to those inferred for the GSFM population, linking
this trend to climatic variation. The reason for an increase in size could also be linked to the rapid evolution
of the cave bears and the Delle Streghe fossils should cover a chronological range approximately similar to
the fossils from GSFM.
Introduction
During the Late Pleistocene Ursus spelaeus
Rosenmüller, 1794 (Rosendhal & Kempe, 2004) colonized a large part of Europe, migrating to the
south (central Italy), towards the west (Spain), and
towards the north-west (Great Britain) exhibiting
the most numerous morphological and morphometrical varieties. Several studies concerning the cave
bear group, Ursus spelaeus and its ancestor Ursus
deningeri Von Reichenau, 1906 and U. arctos
species coeval to U. spelaeus (Rabeder, 1999;
Weinstock, 1999; Perego et al., 2001; Rossi & Santi,
2001 a, b;; Santi & Rossi, 2001; Santi et al., 2003 and
others) have allowed several hypotheses to be
advanced about the relationships between cave
bears, U. deningeri and coeval species, and about
possible phyletic lines indicated by the fossiliferous
record and by recent mtDNA examination
(Hofreiter et al., 2002, 2004). Currently, radiometric
data is only available for a few caves in Northern
Italy: the best known example is the Grotta Sopra
Fontana Marella (Varese Province, Lombardy). For
the other caves (i.e. the Caverna Generosa, Varese
Province) the data are still incomplete (Bona, 2004).
Radiometric data and aminoacid racemization of
the bear bones from Grotta Sopra Fontana Marella
provide the following ages: sample FM4 over 26000
years BP, FM2, 22310 ±200 years BP and FM1
21810±200 years BP (Perego et al., 2001). The
wealth of discoveries in various caves in Northern
Italy have shed new light on the distribution of the
115
of bones, deposits were only slightly disturbed and analysed in situ.
Fig. 1: Geographic position of the Delle Streghe Cave (Valstrona, Piedmont,
Northern Italy).
vertebrates in this area. In some zones research has
only just started; one such example is the ValsesiaValstrona (Piedmont) area (Fig. 1). The aim of this
paper is to summarize previous results regarding
these cave bear populations.
Geographical-geological frame of the studied area
Valstrona is a narrow valley with a V-shaped profile in its lower reaches while at its head, near Cima
di Capezzone-Punta del Pizzo (2240 m)-Punta
d’Issola (2146 m), it enlarges into a wide cirque. It
winds for 20 km to Omegna village where it
debouches onto the Orta Lake (Cusio). Near the
Sambughetto village some caves have formed via
karst processes within the lens of the “Marmo
Valstrona” formation; this lenticular body is intercalated between gneisses and micaschists of the “Serie
Kinzigitico-sillimanitica”. Inside the caves the osteological material, accompanied by yellow loessic clay,
collects in the lower parts along the side lanes and
cavities. This sediment is frequently covered by hard
stalagmitic soil (about 15-20 cm thick), and by grey
micaceous and sterile sands interspersed with smaller gravel of more recent age linked to the pluvial
washing away phase. To ensure good preservation
116
Fossils from Valstrona have been
found inside the caves known as
Complesso dell’Intaglio and Caverna
delle Streghe, near the Cava Sambughetto village. The first of these caves
opens out in the upper part of the marble quarry (“Sass Muiè”), it has five
entrances and a subcircular small gallery
complex correlated with an older level of
the water-bearing stratum. The second
cave, called Caverna delle Streghe, is the
widest cave in Verbania Province. It is
composed of a fossil branch presently
foliated by water and by a second active
branch in the marble eroded by the river
(Fig. 2). The water source is from the
Chignolo stream that, after having
crossed the cave and swelled water from
other tributaries, re-emerges in the
Strona River.
The Valsesia fossils are derived from the
Mt. Fenera (Fig. 3) caves and mainly from the
“Ciutarun” and the “Ciota Ciara” cave. The former is
situated at 650 m asl, with a large ogival entrance,
and it is 55 m long and up to 13 m high. The “Ciota
Ciara” is located at an altitude of 685 m asl, it is 57
m long and the difference in levels internally is up
to 18 m. There are two entrances: a southern, natural and a northwestern entrance which was
formed by the collapse of a part of the vault. This
cave rises upward from SE-NW and ends towards
the N (Strobino, 1981).
Materials and methods
About one thousand Ursus remains currently
stored in the Museo Civico di Storia Naturale di
Milano have been analysed. They have been labelled
MCSNM V”, (abbrevation of Museo Civico di Storia
“M
Naturale and Vertebrate), followed by a progressive
number. A substantial portion of the skeletons of
cubs, juveniles and adult elements is represented
(Pls. 1-2). The material is rarely complete, especially
the skull remains, and in particular in the case of
cubs only skull-caps have been preserved.
Preservation is generally good, although some
Fig. 2: A – Planimetric scheme and profiles; B – of the Delle Streghe Cave (Cella, 1993, mod.).
117
Fig. 3: Distribution of the main caves in the Fenera Mt. (Valsesia, Piedmont, Northern Italy). Number 1 is the “Ciutarun”, 2 and 3
refer to the “Ciota Ciara”. (Strobino, 1981, mod.).
Panthera leo spelaea (Goldfuss, 1810) (Fig. 5). Most
of the fossils belong to Ursus spelaeus Rosenmüller,
1794, while others with disputed morphological
features could be classified as Ursus deningeri Von
Reichenau 1906. However we have considered these
remains as U. spelaeus on the basis of the broader
morphological relationships within this species.
Useful morphometric parameters were deduced
from Hue (1908), Von den Driesch (1976) and Torres
(1988).
Morphometry
Fig. 4: A. Pathological Ursus bone (specimen MSNM V 4362,
Delle Streghe Cave). B. Predatory activity traces (specimen
MSNM V 4097, Delle Streghe Cave).
traces of erosion can be found in the proximal and
distal ends of limb bones. In addition, some specimens showed traces of pathologies (e.g. periarthritis and pesudoarthrosis) and generic malformations,
traces of predator activities (Fig. 4). The presence of
predators is indicated by the catlike remains inside
the Delle Streghe fauna with an incomplete right
radius fragment (MSNM V4329) belonging to
118
SKULL – These fossils, although incomplete, have
some morphometric features that seem to be typical of cave bears. They are generally similar in size
to examples of U. spelaeus from caves in Spain and
slightly larger than those from Caverna delle Ossa
(Zandobbio, Bergamo Province, North of Milan).
MANDIBLE – The relationship between the transversal diameter of the condyle and the vertical
diameter (Fig. 6) confirms what has been inferred
regarding skull morphometric analysis. The
Sambughetto specimens are similar in size to the
typical spelaeus (in this paper represented by fossils
from Covoli di Velo Veronese, Verona Province), but
they are larger than those from the Buco dell’Orso
cave, whose small sizes can be linked to climatic
factors (Bergmann’s rule). The dimensions of the
mandibular condyle, but especially the height of
the mandible below P4, provided additional evidence supporting what has been deduced from
skull analysis. Comparison between the fossils studied and samples from some Venetia caves (Grotta
del Cerè whose population appears to be older,
Covoli di Velo Veronese and S. Donà di Lamon) and
from Grotta Sopra Fontana Marella –GSFM- (Varese
Province, Lombardy), allows us to place the Delle
Streghe bears in an intermediate position between
ancient and modern forms. These data are also supported by dental surface features. Data referred to
the M1 and M2 show the greatest range compared
to those of the other specimens considered (Pocala,
Equi, GSFM, Covoli di Velo, Buco dell’Orso, Caverna
delle Fate, Grotta delle Ossa) and a smaller
length/width ratio. This feature could be probably
related to local factors and particularly to food
preferences. But we cannot exclude that this difference in size may be related to sexual dimorphism.
HUMERUS – As shown in diagram Fig. 7, the Delle
Streghe specimens show similar features to those
from GSFM. In fact, the absolute dimensions are
similar. The main difference is evident from the
greater deformation of the diaphyses of the
analysed remains, and particularly in the more
recent forms due to a smaller antero-posterior
diameter.
RADIUS – Data concerning the radius seem to
confirm what is shown by
the humeri. In particular
some morphometric relationships (Fig. 8) allow us
to affirm that: a) the
morphometric characteristics of the specimens
studied are comparable
with those of the GSFM,
b) generally, adult elements can be compared
with those from the older
and intermediate levels
of the GSFM, while the
Fig. 5: Panthera leo spelaea (Goldfuss, 1810). Specimen MSNM
V 4329 (Delle Streghe Cave). Right radius. A: External view,
B – Internal view.
Fig. 6: Relationship between Transversal Diameter of the condyle and Vertical diameter in
mandibles of Ursus spelaeus from c
119
Similar conclusions can be advanced for
the ulnae as well.
Fig. 7: Antero-posterior diameter of the diaphysis (ordinate) and
Transversal diameter of the diaphysis (abscissa) relationship in the humeri
of the Ursus spelaeus from Delle Streghe and Grotta Sopra Fontana
Marella (GSFM) caves. Symbol legend: , Delle Streghe specimens. Grotta
Sopra Fontana Marella specimens: L juveniles from FM2, G juveniles
from FM1, I juveniles from FM4, ∆ adults from FM2, o adults from FM1,
adults from FM4 and FM2 (Perego et al., 2001 mod.).
PISIFORM – Morphometric data referring to
pisiform (Fig. 9) have allowed us to distinguish three clear size ranges: 1) a group with
forms comparable to the U. deningeri and U.
arctos species from caves in Spain; 2) a second
group with elements comparable to the U.
spelaeus (smaller sized) from the Buco
dell’Orso cave (Laglio, Como province,
Lombardy) but more massive, and: 3) a third
group with large elements. The hypothesis
that U. spelaeus corresponded to the smaller
elements is based on the clear speloid morphology (see Torres, 1988) but they could also
be females or juvenile forms, or related to a
cooler climatic phase (Gerhard, 2001). It is
more likely that they would be female specimens because the points are close to those
from the Buco dell’Orso Cave that are indisputably adult forms (Santi et al., 2003). The
presence of one group of adult medium- tosmall sized elements with another group having medium dimensions is very interesting. In
fact, the lack of intermediate forms can be
simply related to the quantity of useful data,
but also to the actual presence of two separate populations.
METACARPUS – The morphometric features of
the studied remains (Fig. 10a) are very similar
to those from the Buco dell’Orso cave (clearly
spelaeus). They are of smaller size than the
typical spelaeus. When compared with the
data from the literature (Di Canzio & Petronio,
2001; Santi et al., 2003), one can conclude
Fig. 8: Antero-posterior diameter of the diaphysis (ordinate) and that a female element is probably present
Transversal diameter of the diaphysis (abscissa) relationship in the radii of among the II° metacarpus specimens. The diathe Ursus spelaeus from Delle Streghe and Grotta Sopra Fontana Marella gram relating to the V° metacarpus (Fig. 10b)
(GSFM) caves. Asterisks represent the Delle Streghe specimens, for the
shows that three elements are more massive
legend of the other symbols see Fig. 7 (Perego et al., 2001 mod.).
than the others used for comparison. These
different morphometric features could depend on
younger elements cover the whole time interval, c)
dimorphic character or different evolutionary phases.
some remains display dimensions similar to the
largest among the more recent GSFM forms. Such
FEMUR AND TIBIAE - Morphometric data (Fig. 11)
show similar features to adult elements from the
an irregular distribution may depend on: 1) sexual
GSFM and the Buco dell’Orso cave. Compared with
dimorphism, 2) the presence of elements related to
the GSFM, the studied remains appear to corredifferent evolutionary stages (the smaller sized
spond to the temporal arch also covered by the
specimens being older, while the larger ones are
compared fossils. It is therefore possible that they
more recent), 3) climatic factors.
120
Fig. 9: Distribution points of the greatest length
and greatest width ratio in the pisiforms of different Ursus species from caves in Italy and Spain
(Santi et al., 2003 mod.).
Fig. 10: a. Distribution points of the greatest length
and the smallest diaphyseal width ratio in the II
metacarpus of different Ursus species from caves in
Italy and Spain . b. Distribution points of the greatest length and the transversal diaphyseal width
ratio in the V metacarpus of different Ursus species
from caves in Italy and Spain (Santi et al., 2003
mod.).
9
may represent different evolutionary steps
within the same population. Fig. 11 also
shows the presence of a juvenile element.
Similar conclusions are also advanced for
the tibiae in comparison with the GSFM
and Buco dell’Orso populations.
ASTRAGALUS, SCAPHOID AND METATARSUS –
Analogous to proposals for other parts of
the skeleton, data concerning the astragalus (Fig. 12) show more deformed bones
than those used for comparison (Buco
dell’Orso). The paucity of data inhibits a
profound analysis of the scaphoids; nevertheless initial analysis seems to confirm
observations also advanced for the astragalus. In addition, morphometric data concerning the III metatarsus (Fig. 13) confirm
that they belong to the U. spelaeus. Their
small size probably indicates the presence
of females.
10a
PHALANGES – Generally, the data show
morphometric features similar to the Buco
dell’Orso bears. The distribution of the
points relating to the II phalanx (Fig. 14)
shows two clear clouds possibly due to
dimorphism.
Concluding remarks
The discovery of an incomplete radius of
Panthera leo spelaea (Goldfuss, 1810) next
to Ursus specimens, widens the faunistic
association of the Delle Streghe cave to
10b
121
Fig. 11: Antero-posterior diameter of the diaphysis
(ordinate) and Transversal diameter of the diaphysis
(abscissa) ratio in the femurs of Ursus spelaeus from
Delle Streghe and Grotta Sopra Fontana Marella
caves. Asterisks indicate the Delle Streghe specimens,
for the legend of the other symbols see Fig. 7
(Perego et al., 2001 mod.).
Fig. 12: Greatest length and the thickness relationship in the astragali of Ursus spelaeus from caves in
Italy.
Fig. 13: Smallest diaphyseal width and the greatest
length ratio in the III metatarsus of Ursus spelaeus
from caves in Italy and Germany (Santi et al., 2003
mod.).
Fig. 14: Greatest length and the diameter transversal
diaphysis relationship in the II phalanx of Ursus
spelaeus from caves in Italy.
122
other nearby caves (Buco dell’Orso Cave, Delle Ossa
Cave – Zandobbio in Bergamo Province).
Pathologies are rare, mainly confined to limbs, and
related to the senescence of the bears.
Morphometric data indicate the presence of at least
two populations of cave bears characterized by different sizes: the small-size bears are comparable to
the Buco dell’Orso cave bears and those specimens
from the older levels to the Grotta Sopra Fontana
Marella. According to Perego et al. (2001), the difference in size is related to a different evolutionary
step of the bear; small size could correspond to
more ancient forms, namely more primitive ones.
The increase in size can be linked to a thermoregulation factor following Bergmann’s rule (1847): the
increase in body size yields an advantage in thermoregulation. Loss of heat in bodies of large size is
lower, causing a smaller surface-to-volume ratio. In
this manner large sized populations can colonize
cool regions. Moreover, in the case of the studied
bears, an increase in dimensions could also represent a response to a shift towards a cooler climate.
In contrast to these authors, Gerhard (2001) and
Rabeder & Nagel (2001) associate a similar reduction in size to the shift toward cooler conditions
although this should be observable in high Alpine
regions. The similarity between the Grotta Sopra
Fontana Marella and Delle Streghe Cave fossils leads
us to link this trend to a climatic change, rather
than to rapid evolution by cave bears.
Acknowledgments
The authors thank D. Nagel (Vienna) for useful
advise and critical reading of the manuscript, and G.
Papalia (Pavia) for revision of the English. This study
was supported by a FAR grant contribution.
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Von den Driesch, A. (1976): A guide to the measurement
of animal bones from archaeological sites. – Peabody
Museum Bullettin, 1: 1-137.
Plate 1: Ursus spelaeus Rosenmüller, 1794. Delle Streghe Cave (Sambughetto Valstrona, Piedmont, North Italy). A – Skull. Specimen
MSNM V 4486, dorsal view. B – Skull. Specimen MSNM V 5043, dorsal view. C – Skull. Specimen MSNM V 5041, dorsal view. D –
Skull-cap of cub. Specimen MSNM V 4614, dorsal view. E – Skull-cap of cub. Specimen MSNM V 4736, dorsal view. F – Skull-cap of
cub. Specimen MSNM V 4721, dorsal view. G - III phalanx. Specimen MSNM V 5028, lateral view. H – Mandible. Specimen MSNM V
5059, internal view. I - Skull. Specimen MSNM V 5043, frontal view. J – I phalanx. Specimen MSNM V 4988, dorsal view. K – Radius.
Specimen MSNM V 4331, external view. L – Scapholunar. Specimen MSNM 4781, lateral view. M – Astragalus. Specimen MSNM 4874,
dorsal view. N – Femur. Specimen MSNM V 4393, caudal view. O – Radius. Specimen MSNM V 4304, dorsal view.
124
125
Plate 2: Ursus spelaeus Rosenmüller, 1794. Delle Streghe Cave (Sambughetto Valstrona, Piedmont, North Italy). A - Calcaneus.
Specimen MSNM V 4904, dorsal view. B - IV° metacarpus. Specimen MSNM V 4827, medial view. C – III° metacarpus. Specimen
MSNM V 4824, medial view. D – IV° metacarpus. Specimen MSNM V 4828, medial view. E – II° metacarpus. – V° metacarpus.
Specimen MSNM V 4845, lateral view.
126
Geo.Alp, Vol. 2, S. 127–129, 2005
THE ACTIVITIES OF THE LIGABUE STUDY RESEARCH CENTRE
ON THE THIRTIETH ANNIVERSARY OF ITS FOUNDATION
Francesco Garofalo1, Fabrizio Bizzarini2, Federica Ferrieri3
With 4 figures
1 Via Monte San Michele 20/A, 30171 Mestre – [email protected]
2 Cannaregio 1269/A, 30121 Venezia
3 Università Ca’ Foscari di Venezia, Dottorato in Studi Iberici, Anglo-Americani e dell’Europa Orientale
Abstract
In 2003 the Ligabue Study Research Centre celebrated its first thirty years of activity with various projects
concerning research and scientific promotion: the opening of a new exhibition area in the Venice Museum
of Natural History and the creation of a multi-themed exhibition in the Palazzo delle Miniere at Fiera di
Primiero (Trento).
The new room in the Venice Museum is dedicated to the scientific expedition which took place in the
Ténéré Desert between 1971 and 1973. It briefly examines the history of the expedition, which contributed
towards the foundation of the Ligabue Study Research Centre.
The exhibition “From Meteorites to Dinosaurs … to Men” has been staged with the collaboration of the
Comprensorio del Primiero (Trento). Theories about biological evolution act as a bond throughout the exhibition: gathering a wide range of exhibits, the exhibition links the evolutionary potentials which can be found
in the history of terrestrial organisms to the global evolution of the solar system and to human cultural evolution.
Riassunto
L’attività del CENTRO STUDI RICERCHE LIGABUE in occasione del trentennale della sua fondazione.
Nel 2003 il Centro Studi Ricerche Ligabue ha celebrato i suoi primi trent’anni di attività con numerose iniziative nel campo della ricerca e della divulgazione scientifica. Questo secondo aspetto è stato caratterizzato da due manifestazioni: l’apertura di un nuovo percorso espositivo al Museo di Storia Naturale di Venezia e
una mostra politematica presso il Palazzo delle Miniere a Fiera di Primiero.
La nuova sala del Museo Veneziano è dedicata alla spedizione scientifica nel deserto del Ténéré, svoltasi
negli anni tra il 1971 e il 1973. Riassume brevemente la storia di quella spedizione che stimolò la nascita
stessa del Centro Studi Ricerche Ligabue.
La mostra “dalle Meteoriti ai Dinosauri…all’Uomo” è stata realizzata in collaborazione con il Comprensorio
del Primiero. Le teorie dell’evoluzione biologica fanno da collante all’intero percorso espositivo, che nel riunire l’ampia varietà di reperti, collega le potenzialità evolutive riscontrabili nella storia degli organismi terrestri, all’evoluzione complessiva del Sistema Solare e alla stessa evoluzione culturale umana.
127
In 2003 the Ligabue Study Research Centre celebrated its first thirty years of scientific-cultural activities with various projects concerning both research and promotion. In particular, the Centre’s
endeavours have been promoted by two exhibitions.
On August 9, 2003, the permanent exhibition entitled “From Meteorites to Dinosaurs … to Men” was
opened in the 14th century Palazzo delle Miniere at
Fiera di Primiero, while on October 25, 2003 the
Venice Museum of Natural History, including a
room called the “Dinosaur Fossil Deposit”, was officially reopened to the public. This room is dedicated
to the scientific expedition conducted in 1973 by
the Ligabue Study Research Centre and the National Museum of Natural History of Paris, and led by
Giancarlo Ligabue and Philippe Taquet. This expedition enabled the study of the dinosaur fossil deposits of Gadoufaoua, in the Ténéré Desert (Niger),
whose sands yielded the skeleton of an Ouranosaurus nigeriensis, now exhibited in Venice. The
sediments of this deposit belong to the Elrhaz formation, upper Aptian (lower Cretaceous), and
formed in a marshy and deltaic environment, which
was rich in vegetation and populated by dinosaurs,
crocodiles, pterosaurs and fish. The exhibition area
of the Venice Museum enables the visitor to retrace
the history of the expedition, its difficulties and the
technologies which were used to save the palaeontologic material. The central part of the exhibition
is dominated by the skeleton of the Ouranosaurus
nigeriensis as well as the sizeable skull and the rest
of the dermic part of the Sarcosuchus imperator,
possibly the largest crocodile found to date. The interactive material and a big central screen for the
projection of footage relating to the expedition
permit the exhibitors to engage the public immersively in the history of the Gadoufaoua deposit and
the discovery of the remains of dinosaurs, crocodiles, turtles, fish and shellfish, as well as vegetable
finds which are now exhibited inside the showcases
that complete the exhibition area. Therefore, not
only the public, particularly young visitors, can admire the richness of the exhibited material, but they
can also experience the main moments of the first
expedition with Italian participants dedicated to
the research and the study of dinosaurs.
The exhibition “From Meteorites to Dinosaurs…
to Men” is the result of a collaboration between the
Ligabue Study Research Centre and the seven towns
of Primiero. It is currently hosted in two rooms of
the 14th century Palazzo delle Miniere of Fiera di
Primiero, already the venue of an ethnographic museum. The exhibited findings represent a part of
those which have been gathered during the activities of the Research Centre. The materials come
from different continents, in addition to various geological eras and historical periods. Their acquisition by the region of Fiera di Primiero represented
the origin of a small but active scientific museum,
which is clearly separated from similar initiatives in
the area, mainly centred on materials of local origin. Therefore, a private collection became a public
heritage and an instrument for the development
and the promotion of scientific culture.
The exhibition includes some fragments of meteorites, which document the origin and the first
phases of the solar system; various fossilised remains of different organisms; and two manufactured exhibits – a female statuette of Olmecan origin and a fragment of cuneiform writing – which
Fig.1: The ceremony of the new exhibition area at the Venice
Museum of Natural History.
Fig. 2: An example of the interactive material in the Venetian
show room.
128
constitute evidence of ancient human civilizations.
The main theme of the exhibition as a whole is the
state of transformation pervading Nature and the
possibility of reconstructing the subsequent phases
of Natural History through the analysis and the interpretation of documents. Man is, at the same
time, both the spectator and interested party of
Natural History; he is the result of biological evolution as well as of the cultural evolution which
emerges in various terms and conditions.
In order to organise the exhibition area, it was
necessary to start with the chronological sequence
of the finds, but we tried to avoid suggesting the
idea of a “project” which – according to some people – could act as a background to the evolution of
living organisms, a progression from initial simplicity towards the ultimate improvement of the organisms. On the contrary, we emphasised the synchronic aspects of evolution, classifying contemporary
events on parallel levels of the exhibition. For example, in the showcase dedicated to invertebrates,
we tried to show the evolutionary potential of “life
without vertebrates” and underline the structural
complexity which has been present since the very
first moments of the Cambrian explosion of life. In
contrast, the evolution of vertebrates is not seen as
a progression towards the colonisation of the emergent lands, but as a contemporary development of
different evolutionary lines, that allowed dinosaurs
– widely present in the two exhibition rooms – to
adapt to a great variety of habitats in the Mesozoic
era. In that period there also appeared the first
mammals and birds: in the exhibition, the latter are
represented by the rare specimen of Cathayornis.
The same period ended with the decline of large
reptiles and the subsequent ascendancy of mammals. However, the link between the skull of the
Cynodont, a small mammal of the Triassic period,
and that of the Miocenic Machairodus giganteus,
the extraordinary sabre-toothed tiger which is the
symbol of the current exhibition, is not at all linear.
Similarly, the subsequent appearance of Man does
not seem automatic. However, it is only in the light
of Darwin’s Theory of Evolution that such phenomena find their explanation, even though the new
discoveries, which came consecutively in the past
hundred and fifty years that separate us from the
publication of The Origin of Species, led to an overall revision of Darwin’s original idea. Unfortunately,
the confirmation of the exact development of evolutionary processes will never come from a labora-
Fig. 3: The Miocenic Machairodus giganteus, the sabre-toothed
tiger, symbol of the exhibition “From Meteorites to Dinosaurus
… to Men”.
tory test, which is conventional for the empirical
sciences. On the contrary, evolutionist biology,
which adopts the method of historical sciences, became the research of those biological traces that
mark the different phases of the history of living
organisms. In this paradigm, palaeontology remains
a field full of potentials, which could provide solutions to some problems concerning the origin and
the extinction of the species. Therefore, fossils contain proof of the transformations that constitute
the history of life on this planet, petrified remains
of organisms that lived in a remote past, and which
now, from a show-glass in a small mountain museum, continue to educate us in the “grandeur in this
view of life”.
Fig. 4: The characteristic skull of Psittacosaurus mongoliensis, a
Cretaceous dinosaur, the latest acquisition of the exhibition of
the Primiero.
129
Geo.Alp, Vol. 2, S. 131, 2005
THE ORIGIN OF THE PALAEONTOLOGICAL FOSSIL CONCEPT
Nicola Dall’Olio
Provincia di Parma, Piazzale Barezzi 3, 43100 Parma; e-mail: n.dall’[email protected]
In the history of science, the interpretation of
fossils as petrified remains of living organisms was
a first decisive step towards both the development
of a dynamic and evolutional conception of geological and biological forms, and the adoption of a
temporal perspective on a scale of billions of years.
In line with an underlying radicalism particularly
widespread within the scientific community, the
current definition of the fossil, and the related
attribution of an organic origin to a particular class
of stone objects, are usually seen as assumptions
that arose almost automatically when, in the modern age, natural scientists set aside their religious
dogma and metaphysical speculation and began to
carefully observe the world around them with an
open and objective mind, in an attempt to work out
„how things really stood“. Today, the ease and
immediacy with which we recognise the vestiges of
what was once a living thing in a spiral object set
in rock, lead us to conclude that a careful and
objective observation, free from prejudice or preconceived ideas based on mere speculation, is
enough to determine the organic origin of fossils
(or at least most of them) and to clearly distinguish
them from other mineral stone objects.
In light of a historical examination of fossil theories developed in Europe between 1500 and 1600,
this intuitive and simplified conception of the origin
of palaeontology would appear to be incorrect and
unfounded. Although the recognition of a research
method based on the careful observation of the
natural world was fundamental in achieving the
system of classification shared today, this nevertheless appears to be insufficient from a historical point
of view. That which is considered an almost logical
consequence of the adoption of an objective point
of view, would appear rather as the result of assuming a vast combination of theories on nature and
the workings of the physical world which act as filters and classifiers of the object being examined.
When the problem of the classification of fossils is
limited to the recognition or denial of their organic
origin through observation, distinctions, classifications and the dividing lines between natural worlds
and beings (such as that between organic and inorganic) are taken for granted. These factors are the
result of a complex theoretical scheme, indeed only
a few centuries ago they neither existed nor could
they even be outlined. In the absence of these dividing lines, the term fossil, coined by Georg Bauer,
better known as Georgius Agricola in the 16th century, was simply used to describe any object in rock
extracted from the subsoil.
The poster, with the help of some illustrations
from that era, aims to represent the decisive epistemological change which, at the beginning of the
17th century, enabled us to conceive the world of
mineral „things“ as distinct from that of organic
„things“, thus providing the essential bases for the
formulation of a more restrictive palaeontological
concept of fossils.
References:
Aldrovandi, U. (1648): Museum Metallicum; Ferroni e
Bernia ed., Bologna.
Dall’Olio, N. (2004): Vedere il Tempo. Fossili e strati nella
Scienza tra 1600 e 1700. _ MUP ed., Parma.
Morello, N. (1979): La nascita della paleontologia nel
Seicento: Colonna, Stenone, Scilla. – Franco Angeli,
Milano
Rossi P. (1979): I segni deI Tempo. Storia della Terra e storia delle nazioni da Hooke a Vico. – Feltrinelli, Milano
Rudwick, M.J.S. (1976): The meaning of fossils. Episodes
in the History of Paleontology. – The University of
Chicago Press, Chicago & London
Stenone, N. (1667) Canis Carchariae dissecturn caput
[Trad. it. a cura di N. Morello in Morello, 1979, op. cit.]
Stenone, N. (1669) De solido intra solidum naturaliter
contento dissertationis prodromus. – Trad. it. A cura di
A. Mottana, Teknos Ed., Roma 1995.
131
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