Original articleNew discoveries on the giant hedgehog Deinogalerix from the Miocene of Gargano (Apulia, Italy)☆
Introduction
Among the most spectacular insular adaptations of terrestrial fauna, the dwarfism of large continental species (large mammals, dinosaurs; Sander et al., 2006) and gigantism of the small ones (micromammals, birds, reptiles) are probably the most characteristic (Sondaar, 1977). This phenomenon is known as the “Island Rule” (Van Valen, 1973, Lomolino, 1985, Lomolino, 2005). Today, most of the species with extreme insular features are extinct. The most spectacular cases are observed in the fossil record (Van Der Geer et al., 2010, Sondaar, 1991). The insular extinct genus we are interested in here is the giant gymnure Deinogalerix, the largest galericine ever (Fig. 1).
Deinogalerix is part of an endemic ecosystem of the Late Miocene in the Abruzzo-Apulia palaeobioprovince, now part of the east coast of southern Italy. This bioprovince was isolated from the continent during the Miocene and formed a large palaeo-island (Rook et al., 2006). Deinogalerix has been mostly recovered among the Late Miocene “Terre Rosse” faunal complex (also known as the Mikrotia fauna) in the Gargano area (Freudenthal, 1971, Masini et al., 2010), but it is also present in the Tortonian limestone deposits of Scontrone in Abruzzo, Southern Italy (Mazza and Rustioni, 2008).
We report in the present paper new discoveries of the genus Deinogalerix from the “Terre Rosse” sediments of a karstic fissure named “Mikrotia 013” (M013), with special emphasis on the cranial remains.
Section snippets
Deinogalerix record in the “Terre Rosse” faunal complex
Rich vertebrate-bearing karst fissure fillings, outcropping as a result of intense quarrying activity in the area between Apricena and Poggio Imperiale in the western part of the Gargano, have been well known in literature since the late 1970's (Freudenthal, 1971, Abbazzi et al., 1996, Masini et al., 2010). The vertebrate fossil-bearing infill of the extensive karst system network (Abbazzi et al., 1996, Pavia et al., 2010) document two different population phases: a late Miocene endemic faunal
Biochronology of M013 fissure
As the Gargano mammals stem from fissure fillings, there is no lithostratigraphical control of their relative ages. The sequence of localities, indispensable for the reconstruction of evolutionary lineages, was determined on the basis of the evolutionary stage of Mikrotia in the various fissures (Freudenthal, 1976, De Giuli et al., 1986). Prolagus provides also a biochronological marker based on its progressive size increase and the successive development of endemic autapomorphic dental
Abbreviations
PU: Museo di Geologia e Paleontologia Università di Torino (2000–2010 field work by Giulio Pavia's team). RGM: Rijksmuseum van Geologie en Mineralogie, Leiden (now Netherlands Centre of Biodiversity, Naturalis; collections of 1970's field work by Mathijs Freudenthal's team). TRF: “Terre Rosse” collection of the Dipartimento di Scienze della Terra Università di Firenze (collections of 1980's field work by the team directed by the late Claudio De Giuli). SCT: Scontone collections at the
Systematic paleontology
Family ERINACEIDAE Fischer, 1814
Subfamily GALERICINAE Pomel, 1848
Tribe GALERICINI Pomel, 1848
Genus Deinogalerix Freudenthal, 1972
Original diagnosis: see Freudenthal (1972).
Revised diagnosis (Butler, 1980): Galericinae which differs from Echinosorex as follows: size greater (linear dimensions of adults about 1.5 times to more than twice those of Echinosorex); I1 much larger than I2 and I3, i3 absent; upper canine premolariform, with two roots like P1, P2 and p2; c–p3 and C–P3 spaced in the
Discussion
With gigantism occurring in various lineages, and a series of collections, which represent a decent amount of time, Gargano offers an ideal theatre for studying the effects of insular evolution in small mammals. In order to get a grip on the evolutionary processes, we need a number of preconditions. First, an understanding of the time involved, and more importantly of the correct sequence of localities is needed. The age of the faunas is still hotly debated, as well as the mode of colonisation.
Conclusion
More than 30 years after its last description, the new find among the oldest “Mikrotia fauna” from Gargano allows us to update the knowledge about the giant hedgehog Deinogalerix. The new species, D. masinii nov. sp., possesses the characteristic traits of the more derived forms of Deinogalerix in association with primitive features, as one would expect from the oldest representative of the genus. This new find at M013 shows that a primitive species of Deinogalerix already occupied the
Acknowledgements
We thank the people in charge of the collections in the University of Turin (M. Pavia); Centre de Conservation du Musée des Confluences in Lyon (D. Berthet and F. Vigouroux); Naturalis in Leiden (R. van Zelst); the University of Palermo (F. Masini). We specially thank Maria Adelaide Rossi of the Soprintendenza per i Beni Archeologici dell’Abruzzo who kindly provided us the access to the material from Scontrone; Giorgio Carnevale for being kind and patient during the drafting of the manuscript
References (66)
- et al.
L’industrie lithique du site Pléistocène inférieur de Pirro Nord (Apricena, Italie du sud) : une occupation humaine entre 1,3 et 1,7 Ma
L’Anthropologie
(2009) - et al.
The age of immigration of the vertebrate faunas found at Gargano (Apulia, Italy) and Scontrone (l’Aquila, Italy)
Comptes Rendus Palevol
(2010) - et al.
Origination and extinction patterns of mammals in three central Western Mediterranean islands from the Late Miocene to Quaternary
Quaternary International
(2008) - et al.
Processes of island colonization by Oligo-Miocene land mammals in the central Mediterranean: new data from Scontrone (Abruzzo, Central Italy) and Gargano (Apulia, Southern Italy)
Palaeogeography Palaeoclimatology Palaeoecology
(2008) - et al.
Galerix iberica sp. nov. (Erinaceidae, Insectivora, Mammalia) from the Late Miocene and Early Pliocene of the Iberian Peninsula
Geobios
(1993) - et al.
Stratigraphical and palaeontological data from the Early Pleistocene Pirro 10 site of Pirro Nord (Puglia, south eastern Italy)
Quaternary International
(2012) - et al.
Lands and endemic mammals in the late Miocene of Italy: constraints for paleogeographic outlines of Tyrrhenian area
Palaeogeography Palaeoclimatology Palaeoecology
(2006) A revised generic classification of the Galericini (Insectivora, Mammalia) with some remarks on their palaeobiogeography and phylogeny
Geobios
(2001)- et al.
A bridge too far, reply to “Processes of island colonization by Oligo-Miocene land mammals in the central Mediterranean: New data from Scontrone (Abruzzo, Central Italy) and Gargano (Apulia, Southern Italy)” (by Mazza, P., Rustioni, M., 2008. Palaeogeography Palaeoclimatology Palaeoecology 267, 208–215)
Palaeogeography Palaeoclimatology Palaeoecology
(2009) The effect of insularity on the Eastern Mediterranean early cervoid Hoplitomeryx: the study of the forelimb
Quaternary International
(2008)
Revision of the Neogene and Pleistocene of the Gargano region (Apulia, Italy). The marine and continental successions and the mammal faunal assemblages in an area between Apricena and Poggio Imperiale (Foggia)
Memorie della Società Geologica Italiana
Optima for Animals
Evidence of earliest human occupation in Europe: the site of Pirro Nord (Southern Italy)
Naturwissenschaften
The ontogeny of molar pattern
Biological Reviews
The giant erinaceid insectivore, Deinogalerix Freudenthal, from the upper Miocene of Gargano, Italy
Scripta Geologica
Insectivores pliocènes du sud de la France (Languedoc-Roussillon) et du nord-est de l’Espagne
Palaeovertebrata
Stertomys laticrestatus, a new glirid (dormice, Rodentia) from the insular fauna of Gargano (Prov. of Foggia, Italy)
Scripta Geologica
Small mammals from the Early Miocene of Sabuncubeli (Manisa, S.W. Anatolia, Turkey)
Beiträge zur Paläontologie
Effetto arcipelago: un esempio nelle faune fossili del Gargano
Bollettino della Società Paleontologica Italiana
Island endemism in the eastern mediterranean mammalian paleofaunas: radiation patterns in the Gargano paleo-archipelago
Atti dei Convegni della Accademia Nazionale dei Lincei
Insectivora from the Upper Aragonian and Lower Vallesian of the Daroca-Villafeliche area in the Calatayud-Teruel Basin (Spain)
Scripta Geologica, Special Issue
The endemic Pleistocene deer of Crete
Major patterns and processes in biodiversity: taxonomic diversity on islands explained in terms of sympatric speciation
The mammals of the Lower Miocene of Aliveri (Island of Evia, Greece). Part 5. The insectivores
Insectivores (Erinaceomorpha, Soricomorpha; Mammalia) from Karydia and Komotini (Thrace, Greece; MN 4/5)
Beiträge zur Paläontologie
Insectivora und Chiroptera (Mammalia) aus dem Neogen der Türkei
Schweizerischen Paläontologischen Abhandlungen
Neogene vertebrates from the Gargano Peninsula, Italy
Scripta Geologica
Deinogalerix koenigswaldi nov. gen., nov. sp., a giant insectivore from the Neogene of Italy
Scripta Geologica
Rodent stratigraphy of some Miocene fissure fillings in Gargano (prov. Foggia, Italy)
Scripta Geologica
Cricetidae (Rodentia) from the Neogene of Gargano (Prov. of Foggia, Italy)
Scripta Geologica
Gliridae (Rodentia, Mammalia) from the Late Miocene fissure filling Biancone 1 (Gargano, Province of Foggia, Italy)
Palaeontologia Electronica
When and how did the Mikrotia fauna reach Gargano (Apulia, Italy)?
Geobios
Insectivora (Mammalia) from the Lower Miocene of the Aktau Mountains, Southeastern Kazakhstan
Senckenbergiana lethaea
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2014, Quaternary Science ReviewsCitation Excerpt :A remarkable example of hypsodonty and other trends in dental morphometrics is found in the endemic extinct murid Mikrotia, from the Gargano palaeo-archipelago (southern Italy) (Freudenthal, 1971, 1976). The attention of the scientific community has recently been focused anew on various aspects of the Messinian endemic vertebrates from the Gargano: several new papers dealing with glirids, insectivores, cricetids, as well as on biochronological, palaeogeographic and biogeographic aspects have been published recently (Freudenthal and Martín-Suárez, 2006; Martín-Suárez and Freudenthal, 2007; Mazza and Rustioni, 2008; Rinaldi and Masini, 2009; Freudenthal and Martín-Suárez, 2010; Masini et al., 2010; Mazza and Rustioni, 2011; Masini et al., 2013; Mazza, 2013; Savorelli, 2013; Villier et al., 2013). Among these, the contribution by Masini et al. (2013) presents a revision of the faunal succession for the early phase of Mikrotia evolution in the palaeo-archipelago, bringing fresh data for resolving the tangled biochronology of this older phase.
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2014, Comptes Rendus - PalevolDietary divergence in space and time - Lessons from the dwarf-goat Myotragus balearicus (Pleisto-Holocene, Mallorca, Spain)
2013, Mammalian BiologyCitation Excerpt :Adaptive radiations, however, are a rare event and require abundant resources and free niches to be exploited by specialised populations, which can then evolve to distinct species. Most examples in the Mammalia concern small species, such as the murids from the Indonesian island of Flores (Locatelli et al. 2012), or the various rodent and insectivore radiations on the palaeoisland Gargano (e.g. Freudenthal 1976; Butler 1980; Freudenthal and Martín-Suarez 2006; Villier et al. 2013). Insular radiations in larger Mediterranean mammals are much rarer; amongst them the peculiar five-horned ruminant Hoplitomeryx from Monte Gargano (Leinders 1984; Van der Geer 2005; Van der Geer 2008; Mazza and Rustioni 2011), the Pleistocene deer Candiacervus from Crete with up to eight species or ‘morphotypes’ (De Vos 1979, 1984; De Vos and Van der Geer 2002; Van der Geer et al. 2006) and at least two sympatric species of the endemic bovid Nesogoral on Sardinia (Palombo et al. 2006, 2013; Rozzi and Palombo 2013).
A new small mammal assemblage from the M013 Terre Rosse fissure filling (Gargano, South-Eastern Italy)
2013, GeobiosCitation Excerpt :M013 needs first of all to be located within the Terre Rosse biochronological succession. The occurrence of a very primitive species of Deinogalerix (Villier et al., 2013), of a murid related to Mikrotia but much more primitive, and of an endemic Cricetodontinae probably derived from a primitive genus, combined with the absence of Apodemus and Prolagus, which are ubiquitous taxa in the Terre Rosse assemblages, indicates M013 as the succession's oldest fissure known to date. Furthermore, the occurrence of Stertomys lyrifer and Stertomys simplex links M013 to Rinascita 1, which Freudenthal (1976) considered the second oldest fissure after Biancone 1 on the basis of Mikrotia's biochronologic ordering.
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Corresponding editor: Giorgio Carnevale.