Revision of a classic site for Permian tetrapod ichnology (Collio Formation, Trompia and Caffaro valleys, N. Italy), new evidences for the radiation of captorhinomorph footprints
Introduction
The renewed interest on Early Permian tetrapod ichnology of Italy has revealed an unexpected potential, for the quality of preservation, relative abundance and ichnodiversity of the material found (Bernardi and Avanzini, 2011, Marchetti et al., 2013a, Marchetti et al., 2013b, Marchetti et al., 2015, Petti et al., 2014). The age of the studied formations, being constrained by radiometric datings (Schaltegger and Brack, 2007), is late Early Permian (Kungurian). This is a key period for the evolution of captorhinomorphs (non-diapsid eureptiles): footprints related to these trackmakers become abundant and diverse in Italy and in the coeval sites of Europe and North America (Haubold and Lucas, 2001, Haubold and Lucas, 2003, Gand and Durand, 2006, Voigt et al., 2013, Marchetti et al., 2014, Marchetti et al., 2015), thus this Italian material has become very significant in the ichnotaxonomic and biostratigraphic discussion.
In this regard, the most important and historically known among the Early Permian continental successions of northern Italy is located in the Upper Trompia Valley (Brescia Province, Lombardy). Here the Collio Formation (Cassinis, 1966a, Cassinis, 1966b) crops out extensively, and yields a rich fossil content (tetrapod and invertebrate trace fossils, plant fossils, sporomorphs, freshwater jellyfishes, bivalves, and conchostracans). Tetrapod footprints from this site have been known since the 19th century (Don G. Bruni in Curioni, 1870). Successive studies were based on scarce material found in 1967 by Berruti together with a few older specimens (Berruti, 1969, Haubold and Katzung, 1975, Ceoloni et al., 1987). In 1987–1989, intensive field work carried out by M.A. Conti, U. Nicosia, N. Mariotti, and P. Mietto, in the Trompia and Caffaro valleys, greatly enlarged the collections with the storage of about 600 specimens. Apart from the preliminary description and biostratigraphic interpretation made by Conti et al. (1991) and Conti et al. (1997), and the preliminary revision of Marchetti et al. (2013b), this material was never analyzed in detail, even if the chronostratigraphic position of the Collio Formation (early Kungurian) is constrained by precise isotopic dating (Schaltegger and Brack, 2007), thus giving important hints for a temporal assignation of the ichnologic association.
The purpose of this revision study is to analyze all the collected material, in order to definitely assess the taxonomic, biostratigraphic and palaeoecological importance of the Trompia–Caffaro site.
Section snippets
Geology and localities
The siliciclastic and volcanic succession of the continental Collio Basin has been studied since the 19th century, and represents the most known Early Permian succession from Italy. The first geological description is from G. Ragazzoni (1856, 1865 in Schirolli, 2010). After that, a number of scientists described the Trompia Valley succession (Curioni, 1865, Curioni, 1870, Suess, 1869, Stache, 1874, Stache, 1884, Lepsius, 1878), until the introduction of the term “Collio Schichten” by Gümbel
Material and methods
Our study is based on the analysis of more than 600 specimens collected in 1870 by Don Bruni, in 1967 by Berruti and in 1987–1989 by M.A. Conti, U. Nicosia, N. Mariotti and P. Mietto in many different localities of the Trompia–Caffaro site (Fig. 1B). About 180 specimens were selected for the taxonomic study, and among these only the best preserved were considered for the determination and the measurement of the ichnologic parameters.
The selection of the material, preserved as concave epireliefs
Former identifications
The first description of vertebrate fossil footprints in the Early Permian of the Southern Alps is from Curioni (1870), in a careful work on the geology of the Trompia Valley. The examined slab was found by Don Bruni and comes from the locality known as “Pulpito”, in the Upper Trompia Valley. It is the first scientific description of vertebrate fossil footprints from Italy and also the first known figured description of the ichnogenus Amphisauropus Haubold, 1970, although any new scientific
Biostratigraphy
The fossil footprints of land vertebrates are a useful tool for stratigraphy, since it is well possible to distinguish characteristic ichnoassociations in different geologic intervals, thus determining their biochronologic attribution/position (Haubold, 1971, Lucas, 2007). If they are strictly linked to the anatomy of the body fossils, they follow the evolution of the trackmakers, although with a minor resolution. However, trace fossils are always in situ, occur in different lithofacies and are
Palaeoecologic inferences
The possibility of placing all the fossiliferous localities of the Trompia–Caffaro site (TR, CF) in a correct stratigraphic position, and evaluating the corresponding lithofacies, enable some stratigraphic and paleoenvironmental considerations. A reduction of the ichnoassociation diversity, from the lower to the upper parts of the succession, was yet hypothesized by Conti et al., 1991, Conti et al., 1997. These authors gave a stratigraphic explanation, and placed FO and LO of four selected
Conclusions and perspectives
The first comprehensive taxonomic revision on the Trompia–Caffaro site confirms the ichnotaxa indicated by Marchetti et al. (2013b) and definitively assesses the taxonomic and biostratigraphic significance of the Collio Formation in a wider global context. The tetrapod ichnoassociation includes: A. kablikae, cf. Batrachichnus isp., D. lacertoides, E. cassinisi, H. bifurcatus, L. heterodactylus, and Varanopus isp. It is characterized by the abundance and diversity of captorhinomorph footprints (
Acknowledgments
We would like to thank Giuseppe Cassinis, Marco Avanzini, Sebastian Voigt, Matteo Belvedere, Paolo Mietto, Umberto Nicosia, and Nino Mariotti for their fruitful discussion, their lab work, and their useful suggestions. We are also grateful to Maria Rita Palombo and Linda Riti (Università “La Sapienza”), for the access to the Rome collection and to two anonymous reviewers who gave very useful suggestions. Dr. Nicoletta Carlo-Stella is particularly thanked for the revision of the manuscript into
References (121)
- et al.
A stratigraphic and tectonic review of the Italian Southern Alpine Permian
Palaeoworld
(2007) In situ pollen from gymnospermous cones from the Upper Permian of the Italian Alps — a preliminary account
Rev. Palaeobot. Palynol.
(1974)- et al.
Palaeoenvironmental and stratigraphic aspects of animal traces and plant remains in Spanish Permian red beds (Peña Sagra, Cantabrian Mountains, Spain)
Geobios
(1997) - et al.
On a moderately diverse continental ichnofauna from the Permian Ikakern Formation (Argana Basin, Western High Atlas, Morocco)
J. Afr. Earth Sci.
(2012) - et al.
Late Early Permian continental ichnofauna from Lake Kemp, north-central Texas, USA
Palaeogeogr. Palaeoclimatol. Palaeoecol.
(2011) - et al.
Permian tetrapod footprints from W. Orobic Basin (Northern Italy). Biochronological and evolutionary remarks
Geobios
(2000) - et al.
Transition between terrestrial-submerged walking and swimming revealed by Early Permian amphibian trackways and a new proposal for the nomenclature of compound trace fossils
Palaeogeogr. Palaeoclimatol. Palaeoecol.
(2014) - et al.
Geologischer Querschnitt durch die Ostalpen vom Allgäu bis zum Gardasse
- et al.
Sulla stratigrafia del massiccio delle tre valli bresciane
Boll. Serv. Geol. Ital.
(1952) - et al.
Early Permian vertebrate ichnoassociations in Scioc Valley — Orobic Basin — (Lombardy — Northern Italy)
Neues Jb. Geol. Paläontol. Monat.
(2003)
Revisione della stratigrafia permotriassica della Val Camonica meridionale (Lombardia)
Riv. Ital. Paleontol. Stratigr.
A new Early Permian ichnocenosis from the “Gruppo vulcanico atesino” (Mt. Luco, Southern Alps, Italy)
Studi Trentini Sci. Nat. Acta Geol.
Ichnosystematics of the Lower Permian invertebrate traces from the Collio and Mt. Luco Basins (North Italy)
Ichnos
Revision of the Pennsylvanian and Permian footprints Limnopus, Allopus and Baropus
J. Paleontol.
Footprints from the Cutler Formation
U. S. Geol. Surv. Prof. Pap.
Geologie der Umgebungen des Iseosees
Geol. Palaeontol. Abh.
Locomotor behavior in early reptiles: insight from an unusual Erpetopus trackway
J. Paleontol.
U–Pb zircon geochronology of volcanic deposits from the Permian basin of the Orobic Alps (Southern Alps, Lombardy): chronostratigraphic and geological implications
Geol. Mag.
Osservazioni biostratigrafiche sulle formazioni continentali pre-quaternarie delle Valli Trompia e Sabbia
Nat. Bresciana Ann. Museo Civico Sci. Nat.
Geologia della regione tra il Sebino e l'Eridio. Parte I: La porzione centrale
Atti Ist. Geol. Univ. Pavia
Crystal-rich mass flow deposits related to the eruption of a sublacustrine silicic cryptodome (Early Permian Collio Basin, Italian Alps)
J. Volcanol. Geotherm. Res.
Morfogenesi delle Prealpi lombarde ed in particolare di quelle in provincia di Brescia
La Formazione di Collio nell'area-tipo dell'alta Val Trompia (Permiano inferiore bresciano)
Riv. Ital. Paleontol. Stratigr.
Rassegna delle formazioni permiane dell'alta Val Trompia (Brescia)
Atti Ist. Geol. Univ. Pavia
Conglomerato del Dosso dei Galli. Studi III
Carta Geol. Ital.
Vulcaniti di Auccia. Studi III
Carta Geol. Ital.
Il Permiano nel gruppo dell'Adamello, alla luce delle ricerche sui coevi terreni delle aree contermini
Mem. Soc. Geol. Ital.
On the geological time of the typical Collio and Tregiovo continental beds in the South Alpine Permian (Italy), and some additional observations
Atti Ticinensi Sci. Terra
Artinskian and Ufimian palynomorph assemblages from the central Southern Alps, Italy, and their stratigraphic regional implications
Interazione strutturale permiana tra la linea delle Giudicarie ed i bacini di Collio, Tione e Tragiovo (Sudalpino centrale, N. Italia)
Boll. Soc. Geol. Ital.
Tectonics and sedimentation in the western sector of the Permian Continental Collio basin, Southern Alps, Italy
Permian chronostratigraphy of the Southern Alps (Italy) — an update
Studio petrografico e mineralogico della Formazione di Collio nel Permiano inferiore dell' alta Val Trompia (Prealpi bresciane)
Mem. Sci. Geol.
Examples of late Hercynian transtensional tectonics in the Southern Alps (Italy). Late Palaeozoic and Early Mesozoic circum-pacific events and their global correlation
World Reg. Geol.
Stratigraphy and facies of the Permian deposits between Eastern Lombardy and the Western Dolomites
Permian to Triassic geodynamic and magmatic evolution of the Brescian Prealps (eastern Lombardy, Italy)
Boll. Soc. Geol. Ital.
Permian continental basins in the Southern Alps (Italy) and peri-mediterranean correlations
Int. J. Earth Sci.
Tetrapod footprints from Collio Formation (Lombardy, Northern Italy)
Mem. Sci. Geol.
New observations on the invertebrate ichnofossils of the Lower Permian basins from Southern Alps (Northern Italy)
Rev. Paléobiol.
Nuove ricerche sugli icnofossili della Formazione di Collio in Val Trompia
Nat. Bresciana Ann. Mus. Civico Storia Nat. Brescia
Succession of selected bioevents in the continental Permian of the Southern Alps (Italy): improvements in intrabasinal and interregional correlations
Di alcuni vegetali dell'epoca carbonifera scoperti nei monti della Val Camonica
Rend. Regio Ist. Lomb. Sci. Lett.
Osservazioni geologiche sulla Val Trompia
Rend. Ist. Lomb. Sci. Lett. Arti Mem.
I depositi permiani delle Prealpi Bresciane. Sedimentologia dei depositi lacustri della Formazione di Collio e rilevamento geologico dell'area tra il M. Crestoso e la Val Dasdana (Collio, Brescia)
Empreintes de pieds de grands quadrupedes dans le Permien Inferieur de l'Herault
Acad. Sci. Lett. Montpellier. Mém. Sect. Sci.
Einige Tierfahrten aus dem unteren Perm der Bergamasker Alpen
Paläontol. Z.
The problem of the transition from the Permian to the Tsiassie Series in southeastern France: comparison with other Peritethyan regions
Les traces de vertébrés tétrapodes du Permien français
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