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

https://doi.org/10.1016/j.palaeo.2015.04.005Get rights and content

Highlights

  • Tetrapod ichnotaxonomy based on well-preserved material

  • New material of Erpetopus cassinisi and Varanopus isp. (new unnamed species)

  • Minimum age of the first appearance of Erpetopus in a global context

  • Late Early Permian radiation of captorhinomorph footprints confirmed

  • Facies-controlled tetrapod ichnofauna

Abstract

The recent studies on Permian captorhinomorph (non-diapsid eureptile) footprints from North America, North Africa, and Europe raised the interest on their taxonomic and biostratigraphic significance. They seem to radiate in the late Early Permian, but the scarcity of absolute dating in the classic sections bearing these trace fossils is not helpful in order to unravel their distribution. The continental Collio Formation cropping out extensively in the central Southern Alps of Italy can cover this gap of knowledge, since it preserves a rich testimony of these trace fossils (Erpetopus, Hyloidichnus, Varanopus) and it is well-constrained from radiometric datings and fossil content. For this and because of the huge amount of stored but unanalyzed material, a revision was necessary, through careful taxonomic assignations. These taxa are here confirmed and/or identified as being: Amphisauropus kablikae, cf. Batrachichnus isp., Dromopus lacertoides, Erpetopus cassinisi, Hyloidichnus bifurcatus, Limnopus heterodactylus and Varanopus isp. The occurrence of Erpetopus at the base of the Collio Formation (Kungurian base) suggests its minimum age of first appearance in a global context: this is a very important presence in order to build a reliable tetrapod footprint biostratigraphy during the Permian. The quantity of material yielded in the different facies of the Collio and Dosso dei Galli Formations also permit new insights on the paleoenvironments and climatic settings.

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

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      The sedimentary units from the continental basins of the Southern Alps (Northern Italy) can potentially cover this gap in our knowledge, because they yield an extensive fossil record with good age constraints by radioisotopic dating (late Cisuralian, Kungurian age; Schaltegger and Brack, 2007; Marocchi et al., 2008; Morelli et al., 2012; Gretter et al., 2013; Berra et al., 2015). The successions have been known since the 19th century for their fossil content, mainly consisting of trace fossils, plant remains, freshwater invertebrates, and rare vertebrate remains (e.g., Leonardi, 1959; Geinitz, 1869; Curioni, 1870; Berruti, 1969; Remy and Remy, 1978; Ceoloni et al., 1987; Conti et al., 1991; Krainer and Spötl, 1998; Nicosia et al., 2000; Santi and Krieger, 2001; Visscher et al., 2001; Ronchi and Santi, 2003; Ronchi, 2008; Avanzini et al., 2008, 2011; Marchetti et al., 2015a, 2015b, 2015c, 2018; Forte et al., 2017, 2018a, 2018b; Santi et al., 2020). Palynological analyses have been carried out as well (Clement-Westerhof et al., 1974; Cassinis and Doubinger, 1992; Barth and Mohr, 1994; Neri et al., 1999; Hartkopf-Fröder et al., 2001; Hartkopf-Fröder, 2019).

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