Doi:10.1016/j.geobios.2003.05.007
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 and
Tyto 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: [email protected] (M. Pavia).
0016-6995/$ - see front matter 2004 Elsevier SAS. All rights reserved.
doi:10.1016/j.geobios.2003.05.007
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
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Check List 8(2): 264-266, 2012© 2012 Check List and Authors ISSN 1809-127X (available at www.checklist.org.br) Journal of species lists and distribution Mammalia, Rodentia, Cricetidae, Calomys laucha (Fischer, 1814): Distribution extension in southern Brazil Caroline Badzinski 1*, Daniel Galiano 2 and Jorge R. Marinho 1 1 Universidade Regional Integrada do Alto Uruguai e das Missões – Campus de Erechim, Departamento de Ciências Biológicas. Avenida Sete de