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Geobios 37 (2004) 631–641 A new large barn owl (Aves, Strigiformes, Tytonidae) from the Middle Pleistocene of Sicily, Italy, and its taphonomical significance Une nouvelle espèce d'effraie géante (Aves, Strigiformes, Tytonidae) du Pléistocène moyen de Sicile, Italie, et son importance taphonomique Dipartimento di Scienze della Terra, University of Torino, Via Accademia delle Scienze 5, 10123 Torino, Italy Received 3 March 2003; accepted 19 May 2003 A new species of Tytonidae, Tyto mourerchauvireae, is described from the Sicilian cave deposits of Spinagallo, Luparello and Marasà, which have yielded a common vertebrate fossil assemblages referred to the early Middle Pleistocene. T. mourerchauvireae nov. sp. shows apronounced increase in body size compared to other congeneric taxa. It is larger than the extant Tyto alba and the extinct Tyto balearica andTyto sanctialbani and comparable in size among the Mediterranean taxa only with the extinct Tyto robusta, which differs in somemorphological characteristics of the long bones. The insular adaptations of T. mourerchauvireae nov. sp. and the differences from the otherforms of the genus Tyto which spread through the Mediterranean area during Neogene and Pleistocene are discussed. The fossil assemblagesof the early Middle Pleistocene of Sicily are dominated by extinct giant Gliridae of the genus Leithia and Maltamys; these taxa are regardedas the primary prey items of T. mourerchauvireae nov. sp. The Gliridae remains were analysed microscopically to detect possible modificationsof bones and teeth caused by ingestion and digestion processes. This type of analysis, together with the qualitative study of the fossilassemblages, allows to show the taphonomical importance of T. mourerchauvireae nov. sp. as agent of accumulation in such Sicilian fossilassemblages. Incidentally the taxonomic validity of T. robusta is discussed and confirmed.
2004 Elsevier SAS. All rights reserved.
Une nouvelle espèce de Tytonidae, Tyto mourerchauvirae, est décrite dans trois gisements de Sicile, les Grottes de Spinagallo et de Luparello, et la Grotte Marasà, qui ont livré d'abondants restes de vertébrés datant du début du Pléistocène moyen. Tyto mourerchauvirae nov.
sp. montre une augmentation de taille prononcée par rapport à d'autres espèces du même genre. Elle est plus grande que l'espèce actuelle Tytoalba et que les espèces éteintes Tyto balearica et Tyto sanctialbani, et parmi les autres espèces méditerranéennes elle peut être comparée entaille uniquement avec l'espèce éteinte Tyto robusta, mais elle s'en distingue par des caractères morphologiques des os post-crâniens. Lesadaptations insulaires de Tyto mourerchauvirae nov. sp. et ses différences par rapport aux autres espèces du genre Tyto qui existaient dans lesrégions méditerranéennes durant le Néogène et le Pléistocène sont discutées. En Sicile les associations de vertébrés du début du Pléistocènemoyen sont dominées par les Gliridae géants éteints des genres Leithia et Maltamys. Ces taxons sont considérés comme la principale ressourcealimentaire de cette nouvelle effraie géante. Les restes de Gliridae ont été analysés au microscope pour vérifier la présence de traces, sur les oset sur les dents, causées par l'ingestion et la digestion. Ce type d'analyse, en même temps que l'étude qualitative des assemblages fossiles, apermis de montrer l'importance taphonomique de Tyto mourerchauvirae en tant qu'agent d'accumulation de ce matériel fossile sicilien. Par lamême occasion la validité taxonomique de l'espèce Tyto robusta a été examinée et confirmée.
2004 Elsevier SAS. All rights reserved.
Keywords: Tytonidae; Tyto mourerchauvireae nov. sp.; Middle Pleistocene; Italy; Insularity; Taphonomy Mots clés : Tytonidae ; Tyto mourerchauvireae nov. sp. ; Pléistocène moyen ; Italie ; Insularité ; Taphonomie E-mail address: (M. Pavia).
0016-6995/$ - see front matter 2004 Elsevier SAS. All rights reserved.

M. Pavia / Geobios 37 (2004) 631–641 The fossil avifauna of Sicily is little known, except for a few studies on bird remains from archaeological sites, whereonly extant species and continental-like bird associations arelisted and for preliminary analyses ofMiddle Pleistocene birds from Contrada Fusco and Spina-gallo Cave (Siracusa, southeastern Sicily) In the recent revisionof the Pleistocene avifaunas of Mediterranean islands, excluded the bird assemblages of Sicilybecause it was supposed that the island and mainland wereconnected, though it was known to sustain endemic verte-brate taxa, due to the isolation of Sicily during Middle and Fig. 1. Map of Sicily (Italy) showing the position of the fossil localities with early Late Pleistocene. In the last two centuries, many Sicil- remains of Tyto mourerchauvireae nov. sp. cited in the text.
Fig. 1. Carte de Sicile (Italie) montrant les localités fossilifères avec les ian localities with fossil vertebrate assemblages have been restes de Tyto mourerchauvireae nov. sp. mentionnés dans cet article.
found and excavated Recentpalaeontological analyses and deposited in the "Museo di Paleontologia" of the Univer- arranged the Pleistocene ver- sity "La Sapienza" of Roma, Italy (MPUR). Six bones and tebrates into five Faunal Complexes (FC). Four of these one bone have instead been found in Luparello and Marasà mainly include endemic fossil mammals and reptiles, while Cave respectively, they are all kept in the Museo Geologico the fifth, dating from the latest Pleistocene, contains extant "G.G. Gemmellaro" of the University of Palermo, Italy continental species together with Palaeolithic artefacts. Fos- sil bird remains were found in each FC Comparisons have been made with recent skeletal mate- rial stored in the "Dipartimento di Scienze della Terra" of the oldest one, the "Monte Pellegrino FC" which contains only University of Torino, Italy (Marco Pavia Osteological Col- endemic small mammals and reptiles lection (MPOC)), in the "Museo Civico di Storia Naturale" The recent analysis of some Sicilian fossil bird as- of Carmagnola, Italy and in the University Claude Bernard semblages furnished detailed information on Lyon-1, Villeurbanne, France. Comparisons have also been the avifaunas of the Middle Pleistocene "Elephas falconeri made with fossil remains of Tyto robusta and Tyto gigantea, FC" and "Elephas mnaidriensis FC", i.e. the 2nd and the 3rd from the Late Miocene of Chirò and Pizzicoli quarries near FC of These data are now included in Apricena, Gargano, Italy stored in a revision of the fossil bird associations of the Mediterranean the National Museum of Natural History (Naturalis) of islands isolated during Middle and Late Pleistocene Leiden, The Netherlands (RGM), the "Dipartimento di Sci- Some endemic forms are enze della Terra" of the University of Firenze, Italy, and in the present in these fossil bird assemblages, particularly in the "Museo di Geologia e Paleontologia" of the University of "Elephas falconeri FC". The most important locality of this Torino (PU); comparisons with bones of Tyto sanctialbani FC, that yielded hundreds of bird bones together with a huge from the Miocene of La Grive-St. Alban, France, and Tyto amount of mammal and reptile bones, is the Spinagallo Cave, balearica from the Pliocene of the Balearic Islands and the near Siracusa (southeastern Sicily) Middle Pleistocene of Corsica This paper describes fossil remains from Spinagallo Cave were made in the University Claude Bernard and other Sicilian localities of the same age (Luparello Cave Lyon-1, Villeurbanne. The microscopic analysis of the bone and Marasà Cave) from which a new species of Tyto has been surfaces was made using a stereomicroscope with variable found. Besides it is clear that the widespread presence of the 6.3× to 50× magnification, backed up with a scanning elec- new Tytonidae in the Sicilian Middle Pleistocene vertebrate tronic microscope (SEM).
assemblages is the consequence of a certain synecological Measurements are in millimetres, in accordance with the role of this strigiform in the middle to small size vertebrate indications proposed by The osteo- communities. Hence, the importance of the new Tyto species logical terminology is from in the formation of those fossil assemblages needs a detaileddiscussion.
3. Systematic palaeontology
2. Material and methods
Class AVES Linnaeus, 1758Order STRIGIFORMES (Wagler, 1830) Sixteen bones referable to the new species of Tyto have Family TYTONIDAE Ridgway, 1914 been found in the material collected in the Spinagallo Cave Genus Tyto Billberg, 1828
M. Pavia / Geobios 37 (2004) 631–641 Remarks: According to the family
T. gigantea and T. robusta were described from the Mio- Tytonidae contains only two living genera: Tyto and Phodi- cene of the Gargano peninsula these lus. The first one is widespread in the world with at least 14 two species, with definite endemic characteristics, were re- species with only one, Tyto alba, occurring in the Western cently synonymized by as T. gigantea, Palearctic Phodilus instead is known to have according to the opinion that they represent diacronous steps two species: Phodilus badius from Asia and Phodilus pri- of a single phyletic lineage. The effective coexistence of the goginei from Africa The genus Tyto two forms in the same karst fissures differs from Phodilus in some morphological characteristics, and some morphological differences pointed out by the as also pointed out by in particular, analysis of new material from Gargano, such as the different in Tyto the processes supracondylaris dorsalis at the distal robustness of the tarsometatarsus, allow to exclude this hypo- humerus is more robust than Phodilus, moreover, the ridge of thesis (author's personal observation), so I am considering the condylus lateralis of the distal tibiotarsus in Phodilus is here T. robusta as a valid species.
straight on the diaphysis more like in the Strigidae than in theTytonidae, the tarsometatarsus of Phodilus is stout, while in Tyto mourerchauvireae, nov. sp.
Tyto it is slender.
The Tytonidae also comprises several fossil genera, such 1998. Tyto nov. sp.—Tyrberg, p. 547.
as Nocturnavis, Necrobyas, Palaeobyas, and Palaeotyto 1999. Tyto undescribed species—Pavia, p. 125–126.
from the Middle Eocene to the Upper Oligocene of the 2000. Tyto nov. sp.—Pavia, p. 50, Pl. 4, Fig. 4, 5.
Phosphorites du Quercy while the Holotype: Spinagallo Cave, MPUR 5218, right femur.
genus Palaeoglaux, found in the Phosphorites du Quercy and Paratypes: Spinagallo Cave: MPUR 5640, left humerus,
in the Middle Eocene of Messel, was formerly referred to the distal part; MPUR 5641, right ulna; MPUR 5220, left femur; monotypic subfamily Palaeoglaucinae MPUR 5700, right femur; MPUR 5756, right tibiotarsus, distal which has been later elected to family rank by part; MPUR 5642, right tarsometatarsus, proximal part.
Referred material: Spinagallo Cave: MPUR 5305 in-
The genus Basityto, described by complete skull; MPUR 5615, right radius, distal part; MPUR with the species B. rummeli and placed in the subfamily 5757, left ulna, distal part; MPUR 5758, left ulna, proximal Tytoninae, family Strigidae, contrary to the most widespread part; MPUR 5760, left ulna, proximal part; MPUR 5617, opinion of the family identity of Tytonidae right ulna, proximal part; MPUR 5423, incomplete synsa- has been recently synony- crum; MPUR 5759, left femur, proximal part; MPUR 5219, mized with the genus Balearica, family Gruidae, by right femur. Marasà Cave: MGUP-MA 229/86, incomplete right scapula. Luparello Cave: MGUP-GL 425/3, incomplete The genus Tyto is known since the Upper Miocene with right scapula; MGUP-GL 434/4, right radius, proximal part; the species T. sanctialbani reported from various European MGUP-GL 238, right ulna, proximal part; MGUP-GL 361, left femur, distal part; MGUP-GL 410, right tibiotarsus, species Tyto ignota from the Middle Miocene of Sansan was distal part; MGUP-GL 345, right tarsometatarsus, proximal long regarded as the oldest form of the genus but has been recently moved by Etymology: This species is dedicated to Cecile Mourer-
into the Strigidae, as Asio (?) ignotus. The Chauviré, of the French CNRS, who has studied and descri- fossil record also comprises a rich variety of insular species, bed many endemic birds, especially Strigiformes. It is a like the ones described from the Neogene and Pleistocene of honour to regard her as my particular friend and teacher from the Mediterranean Islands and from the Pleistocene of the whom I learned all I know on fossil birds.
Western Indies In the Type locality: Spinagallo Cave, near Siracusa, south-
Mediterranean area four extinct species of Tyto are known so far: Tyto melitensis, T. balearica, T. gigantea and T. robusta, Additional localities: Luparello Cave and Marasà Cave,
the latter three with a marked increasing of body-size com- near Palermo, north-western Sicily pared to the living T. alba.
Age: Early Middle Pleistocene, "Elephas falconeri FC"
Tyto melitensis, an extinct form described by from the Pleistocene of Malta, was synonymized with Measurements: See Tables 1,2.
the extant T. alba Diagnosis: A large species of the genus Tyto, larger than
and will not further considered here. T. balearica is an the extant T. alba and the extinct T. balearica and T. sanc- extinct species described by tialbani. The cranium shows a shallow depressio frontalis as an insular form endemic of the Plio-Pleistocene of the with respect to T. alba. The femur shows a small tubercular Balearic Islands and subsequently found in several Neogene prominence in the proximal part of the linea intermuscularis and Pleistocene localities of the Western Mediterranean ba- cranialis, which is absent in T. alba and in the extinct forms sin also with the recently described of the genus; the distal epiphysis of the femur is proportio- subspecies T. balearica cyrneichnusae endemic of Corsica nally wider than the other species, due to the lateral develo- pment of the condylus medialis.

M. Pavia / Geobios 37 (2004) 631–641 Fig. 2. 1–16. Various skeletal element of Tyto mourerchauvireae nov. sp. (1, 2, 3, 6, 11, 14, 15) compared with the extinct Tyto robusta (10) and the extant
Tyto alba (4, 5, 12, 13, 16). Tyto mourerchauvireae nov. sp. from Spinagallo Cave: 1, right femur (holotype, MPUR 5218), caudal view; 2, right femur (holotype,
MPUR 5218), cranial view; 3, right femur (paratype, MPUR 5700), cranial view; 6, right ulna (paratype, MPUR 5641), ventral view; 11, proximal end of right
tarsometatarsus (paratype, MPUR 5642), dorsal view; 14, distal end of left humerus (paratype, MPUR 5640), cranial view; 15, distal end of right tibiotarsus
M. Pavia / Geobios 37 (2004) 631–641 (paratype, MPUR 5756), cranial view. Tyto mourerchauvireae nov. sp. from Luparello Cave: 7, distal part of right tibiotarsus (MGUP-GL 410), lateral view; 8,
distal part of right tibiotarsus (MGUP-GL 410), cranial view; 9, proximal end of right ulna (MGUP-GL 238), cranial view. Tyto robusta Ballmann, 1973 from
Pizzicoli quarry: 10, proximal end of right tarsometatarsus (RGM 425479), dorsal view. Tyto alba (Scopoli, 1769), recent (MPOC 37): 4, right femur, cranial
view; 5, right ulna, ventral view; 12, right tarsometatarsus, dorsal view; 13, left humerus, cranial view; 16, right tibiotarsus, cranial view. Specimens coated with
ammonium chloride to enhance details. The scale bars represent 10 mm.
Fig. 2. : 1–16. Tyto mourerchauvireae nov. sp. de la Grotte de Spinagallo: 1, fémur droit (holotype, MPUR 5218), vue caudale; 2, fémur droit (holotype, MPUR
5218), vue crâniale; 3, fémur droit (paratype, MPUR 5700), vue crâniale; 6, ulna droite (paratype, MPUR 5641), vue ventrale; 11, tarsométatarse droit, partie
proximale (paratype, MPUR 5642), vue dorsale; 14, humérus gauche, partie distale (paratype, MPUR 5640), vue crâniale; 15, tibiotarse droit, partie distale
(paratype, MPUR 5756), vue crâniale. Tyto mourerchauvireae nov. sp. de la Grotte de Luparello: 7, tibiotarse droit, partie distale (MGUP-GL 410), vue latérale;
8, tibiotarse droit, partie distale (MGUP-GL 410), vue crâniale; 9, ulna droite, partie proximale (MGUP-GL 238), vue crâniale. Tyto robusta Ballmann, 1973 de
la carrière Pizzicoli: 10, tarsométatarse droit, partie proximale (RGM 425479), vue dorsale. Tyto alba (Scopoli, 1769), récent (MPOC 37): 4, fémur droit, vue
crâniale; 5, ulna droite, vue ventrale; 12 tarsométatarse droit, vue dorsale; 13 humérus gauche, vue crâniale; 16, tibiotarse droit, vue crâniale. Tous les
exemplaires ont été blanchis au chlorure d'ammonium. Échelle graphique 10 mm.
Description: The fossil bones show morphological char-
also in caudal view the condylus medialis is acteristics which fit exactly those of Tytonidae, in particular narrow. In the proximal part of the tarsometatarsus the arcus those of the genus Tyto. The fossil remains exhibit the fol- extensorius is absent. The femur groups into two different lowing morphological features: at the cranium the os meseth- dimensional classes of bones, even at the same locality, as moidale is wide and pneumatic. The fossa musculi brachialis indicated in Table 2; this can be interpreted as the expression of the distal humerus is well defined and deep; at the proxi- of sexual dimorphism. The same fact has been found by mal epiphysis of the ulna, the tuberculum ligamenti collat- concerning the endemic eralis ventralis forms a little ridge and the trochlea humeralis Tyto neddi of the Pleistocene of the West Indies.
ulnaris is deep and shows a pneumatic surface; the cotyla Comparison: The bones of T. mourerchauvireae nov. sp.
humeralis of the proximal radius is rounded and the depres- have been compared with the extant species T. alba and all sio ligamentosa at the distal part of the bone is deep. The the extinct forms of the Neogene of the Mediterranean basin crista trochanteris of the proximal femur is perpendicular to known so far: T. sanctialbani, T. balearica, T. robusta, and the longitudinal axis of the diaphysis and at the distal part of T. gigantea. The fossil remains are clearly larger than those the bones the proximal portion of the crus condylus lateralis of T. alba, T. sanctialbani, and T. balearica and thus do not is rather pointed in posterior view; the sulcus extensorius on require any comparison; the fossil remains are also definiti- the distal tibiotarsus is not as deep as in the Strigidae and, in vely smaller than those of T. gigantea, so they are not lateral view, the condylus lateralis extends caudally compared further. It is instead necessary to compare the Table 1Measurements of the forelimb bones of Tyto mourerchauvireae nov. sp., compared with the extinct Tyto robusta, Tyto balearica balearica, T. balearicacyrneichnusae and the extant Tyto alba. (Data from (1); (4); (2) ; (3))Dimensions des éléments du membre antérieur de Tyto mourerchauvireae nov. sp., comparées avec celles des espèces éteintes Tyto robusta, Tyto balearicabalearica, T. balearica cyrneichnusae et de l'espèce actuelle Tyto alba. (Données d'après (1); (4); (2);(3)).
Tyto mourerchauvireae nov. sp. MPUR 5640 paratype Tyto robusta (1) Tyto balearica balearica (3) Tyto balearica cyrneichnusae (2) Tyto alba (4) Tyto mourerchauvireae nov. sp. MPUR 5167 Tyto mourerchauvireae nov. sp. MPUR 5641 paratype Tyto mourerchauvireae nov. sp. MPUR 5757 Tyto mourerchauvireae nov. sp. MPUR 5758 Tyto mourerchauvireae nov. sp. MPUR 5760 Tyto mourerchauvireae nov. sp. MGUP–GL 238 Tyto robusta (1) Tyto balearica cyrneichnusae (2) Tyto alba (4) M. Pavia / Geobios 37 (2004) 631–641 Table 2Measurements of the hindlimb bones of Tyto mourerchauvireae nov. sp., compared with the extinct Tyto robusta, Tyto balearica balearica, T. balearicacyrneichnusae and the extant Tyto alba. (Data from (1); (4); (2); (3))Dimensions des éléments du membre postérieur de Tyto mourerchauvireae nov. sp., comparées avec celles des espèces éteintes Tyto robusta, Tyto balearicabalearica, T. balearica cyrneichnusae et de l'espèce actuelle Tyto alba. (Données d'après (1); (4); (2);(3)).
Tyto mourerchauvireae nov. sp. MPUR 5218 holotype Tyto mourerchauvireae nov. sp. MPUR 5219 Tyto mourerchauvireae nov. sp. MPUR 5220 paratype Tyto mourerchauvireae nov. sp. MPUR 5700 paratype Tyto mourerchauvireae nov. sp. MPUR 5759 Tyto mourerchauvireae nov. sp. MGUP-GL 361 Tyto robusta (1) Tyto balearica balearica (3) Tyto balearica cyrneichnusae (2) Tyto alba (4) Tyto mourerchauvireae nov. sp. MPUR 5756 paratype Tyto mourerchauvireae nov. sp. MGUP-GL 410 Tyto robusta (1) Tyto balearica cyrneichnusae (2) Tyto alba (4) Tyto mourerchauvireae nov. sp. MPUR 5642 paratype Tyto robusta (1) Tyto balearica balearica (3) Tyto balearica cyrneichnusae (2) Tyto alba (4) bones of the new species with the corresponding ones of At the distal tibiotarsus of T. mourerchauvireae nov. sp. the T. robusta, because of their similar dimensions in some incisura intercondylaris is wider and the condylus medialis is larger and thinner. The tarsometatarsus of T. robusta is more At the distal part of the humerus of T. mourerchauvireae slender than the one of T. mourerchauvireae nov. sp., which nov. sp. the tuberculum supracondylare ventrale is more also differs in the more protruding eminentia intercondylaris strongly developed than in T. robusta, which also shows a more slender humerus. At the distal part of the ulna, thecondylus ventralis ulnaris is more developed in T. mourer-chauvireae nov. sp. The proximal epiphysis of the femur of the new species is characterized by a small tubercular promi-nence on the linea intermuscularis cranialis close to the facies 4.1. Evolution and synecology articularis antitrochanterica, which is absent in T. robusta; inthe latter species the crista trochanteris extends distally and The few specimens of T. mourerchauvireae nov. sp.
forms a continuous ridge, while in T. mourerchauvireae nov.
known so far do not allow an analysis of the phylogenetic sp. it is interrupted by the impressiones iliotrochantericae relationships among this species and the other Tyto species and thus constitutes of two distinct cristae. At the distal part described in the Mediterranean area, both continental and of the femur of T. mourerchauvireae nov. sp. the fossa insular forms. Among the five FCs in which the Sicilian fossil poplitea is deep; moreover the condylus medialis and the vertebrate assemblages have been divided crista supracondylaris medialis are laterally more developed.
the "Elephas falconeri FC", of early Middle M. Pavia / Geobios 37 (2004) 631–641 Pleistocene, is the oldest one in which bird remains have powerful Bubo bubo, which arrived from the mainland, than been found. Thus the direct ancestors of T. mourerchauvireae to the loss of food source; probably it took place in the middle nov. sp. are unknown since no bird remains occur in the Early part of the Middle Pleistocene, in a time confined between Pleistocene "Monte Pellegrino FC" the "Elephas falconeri FC" and the "Elephas mnaidriensis Nevertheless it seems likely that the new species evolved autochthonously after an early colonization of Sicily The analysis of fossil vertebrate assemblages of the "Ele- by a smaller form like T. balearica, the only Tytonidae phas falconeri FC" found in the karst cavities allows an recorded in the Early Pleistocene of the Mediterranean area estimate of the food range of T. mourerchauvireae nov. sp. It It is well known that insular predatory birds, could have included the giant Leithia melitensis, the other both diurnal and nocturnal, tend to enlarge their size Gliridae and the endemic Crocidura esuae, which were all of and this is particularly true in the genus Tyto, in average abundance. In the food range of the new species a which the trend to gigantism in insular environments is well rich variety of birds probably played an important role: a demonstrated by different species from the Pleistocene of the large number of species is usually present in the fossil assem- West Indies and the Mi- blages and they are interpreted as the product ocene of Gargano This fact seems to of predation; these birds are closely related to different envi- be closely related to prey size, mostly of small mammals, ronments, such as open habitat, as is the case with Calan- which on islands tend to increase their body size, favoured by drella brachydactyla, or dense woods, e.g. Dendrocopos the absence of small terrestrial carnivores.
leucotos, while other species are typical of aquatic environ- In Sicily the presence of endemic Gliridae of the genus ment, like Larus minutus. In fact, according to the abundance Leithia and Maltamys has been reported during the "Elephas of remains, it is possible to conclude that the food range of falconeri FC" One of them, Leithia T. mourerchauvireae nov. sp. was mostly characterized by melitensis, is a gigantic Gliridae, while the other species Leithia melitensis, while other animals like small mammals Leithia cartei and Maltamys wiedincitensis show a slight and several bird species of various dimensions played a increase in body size compared to Glis glis, the largest extant secondary role. In the fossil assemblages, the prevalence of European Gliridae. During the early Middle Pleistocene no one or few species together with the presence of a good terrestrial carnivores inhabited the Sicilian country except for variety of other species is similar to that observed in the the endemic otter Lutra trinacriae pellets of the extant T. alba; while the possibility to take Thus, the presence of various large big rodents might animals of different sizes, from a small Crocidura to a big have favoured the local evolution of the endemic species Anser, is also observed in extant species with body size T. mourerchauvireae nov. sp., which represented the top of similar to T. mourerchauvireae nov. sp., such as Bubo bubo the food chain. In fact, birds are the most common predators and Nyctea scandiaca of small vertebrates on islands, and often they predate onmiddle- and large-sized vertebrates, especially if they have 4.2. Taphonomical importance of T. mourerchauvireae nov. increased their body size The extinction on islands of various endemic large species of Tyto probably was related most often to the loss of their Several authors have pointed out the role of predators as favourite prey species accumulators of small fossil vertebrate remains. It is particu- This is not the case concerning T. mourerchauvireae nov. sp., larly true in the case of the Strigiformes, in which the hun- as in the younger "Elephas mnaidriensis FC" the new species ting, ingestive and digestive processes caused no or scarce has never been recorded and was possibly replaced by the modifications on the skeletal elements found in their pellets extant Bubo bubo, while the endemic giant glirids Leithia melitensis and Maltamys wiedincitensis still survive After the analysis of a big The deep faunal renewal sample of pellets and scats of both nocturnal and diurnal occurred at the end of the Middle Pleistocene, as testified by raptors and mammal carnivores, identified the onset of the "Elephas mnaidriensis FC", involved all the different kinds of modifications of the small mammal bones.
macro-mammals, with the extinction of the dwarf Elephas Five categories were defined and the predator species were falconeri and the arrival of many continental forms grouped according to the degree of modification of the bones During the "Elephas mnaidriensis FC" the of their prey. These five predator groups range from the least endemic bird species became all-extinct and were partially destructive, with very little modifications and no or light replaced by continental forms This digestion traces, to the most destructive with massive modi- renewal, though so deep to involve such sedentary birds as fication and digestion of bones.
the Strigiformes, did not modify the endemic micromammal The first category includes only Strigiformes, such as Tyto communities at all, which disappeared in the more recent alba, Nyctea scandiaca, Asio otus and Asio flammeus; the "Pianetti-S. Teodoro FC" In conclu- second one also includes only Strigiformes, such as Bubo sion, the extinction of T. mourerchauvireae nov. sp. seems to bubo and Strix aluco, but shows evident modifications and be more related to the direct competition with the more breakage both in teeth enamel and in the long bones. From M. Pavia / Geobios 37 (2004) 631–641 the third to the fifth groups the modifications of bones and bed by in the food remains of the extant teeth become more and more evident and they include only T. alba. Following the conclusions of diurnal raptor and terrestrial carnivores. In the same paper, applied those categories to the Late Pleis- species of Strigiformes found as fossils in the "Elephas tocene small-mammal assemblage of Westbury sub-Mendip falconeri FC", e.g. Asio otus and Athene trinacriae, had to (England), to understand the taphonomy of this fossil accu- produce much more marks on the bones, of which there is no mulation. This method has been successfully used in some evidence in the studied material.
other localities The mean dimensions of the presumed prey species also correlate to a large predator, such as T. mourerchauvireae detailed taphonomical analyses carried out in order to im- nov. sp. The giant Gliridae Leithia melitensis was as large as prove the palaeoecological reconstruction. In fact the pa- a rabbit and the other Gliridae were larger than the extant Glis laeoecological inferences obtained from fossil predators are glis In fact also some open land or aquatic not as precise as the ecological studies on living species bird species, probably taken into the assemblages as preys, This is because the various predators can are of medium or large size, such as Anser erythropus and introduce species from different habitats and thus produce an Recurvirostra avosetta and this can be corre- overrepresentation of the favourite prey lated with a wide range of predation.
thus possibly masking the real species composition and dis- In conclusion, the degree of modification and digestion of torting the palaeoenvironmental reconstruction. Taphonomi- the bones, typical of the genus Tyto, together with the mean cal analyses may correct the bias due to the hunting prefe- dimensions of the animals found in the assemblages, indicate rence since the predator is detected and identified T. mourerchauvireae nov. sp. as the main species responsible of the accumulation of small and medium sized vertebrates in Small vertebrate remains, particularly bones of Leithia, Sicily during the "Elephas falconeri FC". These results con- dominate the Sicilian karst fossil localities referred to the firm the importance of predatory birds in fossil vertebrate "Elephas falconeri FC", like the Spinagallo Cave, where accumulations, even on islands, as previously suggested by thousand of Leithia remains have been found and described other studies in continental environments together with many bones of reptiles, bats and the endemic Crocidura esuae Important collections of smallvertebrate bones have been found in other caves, such asPoggio Schinaldo Luparello and Marasà (author's personal observation).
In all these localities the bones of small vertebrates are very T. mourerchauvireae nov. sp. is described from fossil well preserved and mostly complete. The fossil assemblages material of three karst localities of Sicily: Spinagallo Cave, mainly comprise vertebrate species with no troglophilous Luparello Cave and Marasà Cave, whose fossil vertebrate habits, such as Gliridae and aquatic or woodland birds. This assemblages have been attributed to the early Middle Pleis- fact, together with the good preservation of the fossil re- tocene "Elephas falconeri FC" mains, suggests that the fossil assemblages have been gene- T. mourerchauvireae nov. sp. shows a marked in- rated by the hunting activities of owls, such as T. mourer- crease in body size towards gigantism; this characteristic has chauvireae nov. sp., Otus scops, Athene trinacriae, Asio been previously recognized in several other endemic species otus, and cf. Surnia ulula, known so far in the "Elephas of the genus Tyto as the result of an insular evolution: Tyto falconeri FC" neddi, T. noeli, T. ostologa, T. pollens and T. riveroi, from the Western Indies T. robusta In the "Museo di Geologia e Paleontologia" of the Univer- and T. gigantea from the Late Miocene of Gargano sity of Torino, a large sample of Gliridae remains from The presence of T. mourerchauvireae Marasà Cave, collected by Fabiani in 1929, is present to- nov. sp. in the early Middle Pleistocene of Sicily confirms the gether with some bones of Leithia melitensis, Maltamys tendency of the genus Tyto to generate endemic forms in wiedincitensis and Crocidura esuae from Spinagallo Cave insular environments, even after a short colonization period, that I collected during a recent survey. Macroscopic analyses as also demonstrated in other insular areas of this material and of other specimens from Spinagallo Cave and Poggio Schinaldo Cave The analysis of other vertebrate remains in the various show that the skulls and mandibles are in cave localities of the "Elephas falconeri FC" allows to esti- fact often almost complete, with the incisors still in place, mate the food range of T. mourerchauvireae nov. sp. and its and the percentages of complete long bones are very high role in the genesis of the fossil assemblages, which contain a The microscopic analyses made on the bone number of species with scarce or no troglophilous habits, and and teeth surfaces reveal no or very little signs of digestion thus were probably taken into the cave as preys. The width of those features are in accordance to that descri- the food range is similar to that observed in the extant T. alba,

M. Pavia / Geobios 37 (2004) 631–641 Fig. 3. 1–4. Gliridae remains showing no breakage or traces of digestion. Leithia melitensis (Adams, 1863) from Marasà Cave. 1, SEM photograph of fragment
of right maxilla (PU 100046), occlusal view; 3, right mandible (PU 100047), lingual view; 4, left humerus (PU 100048), anterior view. Maltamys wiedincitensis
(Zammit Maempel and De Bruijn, 1982) from Spinagallo Cave. 2, SEM photograph of the distal part of right tibia (PU 100049), anterior view. Specimens 3 and
4 coated with ammonium chloride to enhance details. The scale bars 1 and 2 represent 2 mm; the scale bars 3 and 4 represent 10 mm.
Fig. 3.1–4. Restes de Gliridae ne montrant pas des traces de digestion. Leithia melitensis (Adams, 1863) de la Grotte Marasà. 1, maxillaire droite (PU 100046),
vue occlusale; 3, mandibule droite (PU 100047), vue linguale; 4, humérus gauche (PU 100048), vue antérieure. Maltamys wiedincitensis (Zammit Maempel et
De Bruijn, 1982) de la Grotte de Spinagallo. 2, tibia droit, partie distale (PU 100049), vue antérieure. Les exemplaires 3 et 4 ont été blanchis au chlorure
d'ammonium. L'échelle graphique 1 et 2 représente 2 mm; l'échelle graphique 3 et 4 représente 10 mm.
while the dimensional range and the mean dimensions of the rmed also in insular condition, such as the Sicilian cave prey species are comparable to those observed in other Stri- deposits. On islands, the strigiforms often represent the only giformes with a body size similar to T. mourerchauvireae predators at the top of the food chain nov. sp., such as Bubo bubo and Nyctea scandiaca and thus are probably the most important agent of bone The macro- and microscopic analyses, aimed at detec- accumulation. An understanding of the predator and its role ting what kind of modifications due to predation occurred on in the fossil assemblages is useful for palaeoecological and skeletal remains, revealed that the long bones and cranial palaeoenvironmental reconstructions, even if the endemic remains found in the Sicilian cave deposits are often almost animals sometimes are synecologically different from their complete and their surfaces show very few traces of modifi- continental relatives. For this reason, this model could be cation and digestion. According to the pat- transferred to other fossil assemblages, such as the late Mio- tern of modifications observed in the Sicilian fossil as- cene vertebrates found in several fissure fillings of the Gar- semblages fits well with the one of the genus Tyto thus gano area in which the high number of indicating, together with the dimension of the preys, the main nocturnal raptors, some of big body size role of T. mourerchauvireae nov. sp. as accumulator of small may have played an important role in the formation of and medium size vertebrate remains in the Sicilian karst the very rich small mammal fossil assemblages.
localities of the "Elephas falconeri FC".
Such evidence confirms the importance of taphonomic analyses aimed at selecting, from fossil assemblages, infor- mation useful for reconstructing the palaeoenvironmentalconditions of the fossil locality. The validity of the model I would like to thank Prof. U. Nicosia and Dr. R. Manni for proposed by and successfully applied in the possibility to study the Spinagallo material; I am very some other continental localities by different authors grateful to the late Dr. E. Burgio for the loan of the material from the Luparello and Marasà caves and for his invaluable suggestions. Prof. T. Kotsakis is thanked for his comments on M. Pavia / Geobios 37 (2004) 631–641 the manuscript and for the long discussions on the Sicilian Bonfiglio, L., Mangano, G., Marra, A.C., Masini, F., Pavia, M., Petruso, D., palaeontology, Dr. N. Minciotti for the revision of the En- 2002. Pleistocene Calabrian and Sicilian bioprovinces. In: Monegatti, P.,Cecca, F., Raffi, S. (Eds.), International Conference "Paleobiogeography glish version, and Dr. G. Mayr and Dr. D.W. Steadman for and Paleoecology 2001", Piacenza and Casetll'Arquato 2001. Geobios their useful suggestions. A special thank to Prof. G. Pavia for MS 24, pp. 29–39.
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