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Bollettino della Società Paleontologica Italiana, 48 (1), 2009, 51-57. Modena, 15 maggio 2009
Two new species of Rissoina (Gastropoda: Rissoidae) in the Lower
Miocene of Valle Ceppi (Torino, NW Italy)
Marta ZUNINO & Giulio PAVIA
M. Zunino, Dipartimento di Scienze della Terra, Università degli Studi di Torino, Via Valperga Caluso 35, I-10125 Torino, Italy; [email protected]
G. Pavia, Dipartimento di Scienze della Terra, Università degli Studi di Torino, Via Valperga Caluso 35, I-10125 Torino, Italy; [email protected]
KEY WORDS - Rissoidae, Rissoina, Lower Miocene, Valle Ceppi, Torino Hills, Tertiary Piedmont Basin.
ABSTRACT - Two new species of Rissoina from the Miocene of Torino Hills, Rissoina (R.) giuntellii and Rissoina (R.) sturanii, are
described in this paper. Rissoina (R.) giuntellii n. sp. is frequent in the Lower Miocene deposits of Valle Ceppi and in coeval and more recent
layers of Torino Hills; it is characterized by a small shell with convex whorls, the sculpture is composed by prominent axial ribs and cords
visible at the base of the last whorl. Rissoina (R.) sturanii n. sp. is only known from the Valle Ceppi section; it is characterized by medium-sized
shells with flat whorls and very fine axial and spiral ornamentation. Because of the reduced number of specimens and their state of conservation,
scanning electron microscope photographs were used to study shell micro-architecture where possible. Comparison with the coeval Rissoininae
known from the deposits of Paratethys, Aquitaine Basin and Loire Basin confirmed the election of Rissoina (R.) giuntellii n. sp. and Rissoina
(R.) sturanii n. sp., currently known only in the Miocene deposits of the Tertiary Piedmont Basin.
RIASSUNTO - [Due nuove specie di Rissoina (Gastropoda: Rissoidae) nel Miocene inferiore di Valle Ceppi (Torino, NW Italia)] - Recentemente
è stata avviata la revisione delle associazioni a molluschi del Miocene inferiore della Collina di Torino. I livelli fossiliferi presenti nella
località di Valle Ceppi hanno restituito un’abbondante associazione a micro-molluschi, in particolare gasteropodi, ancora mai studiata nel
dettaglio. In questa prima nota sistematica vengono descritte due nuove specie appartenenti alla famiglia Rissoidae: Rissoina (R.) giuntellii e
Rissoina (R.) sturanii. La prima specie è frequente nei depositi di Valle Ceppi ed è stato possibile riconoscerne la presenza anche nei depositi
del Miocene medio di altre località fossilifere della Collina di Torino. Al contrario la seconda specie, rappresentata da un numero esiguo di
esemplari, è finora stata riconosciuta solo nel Miocene inferiore di Valle Ceppi. Le analisi morfologiche permettono di riconoscere le due
specie da altre congeneriche del Bacino Terziario Piemontese, in base ai seguenti caratteri distintivi: Rissoina (R.) giuntellii n. sp. presenta una
conchiglia di piccole dimensioni con giri molto convessi e un’ornamentazione composta da spesse coste assiali e ornamentazione spirale
limitata alla base dell’ultimo giro; Rissoina (R.) sturanii n. sp. è caratterizzata da giri piatti con un’ornamentazione formata da tenui coste
assiali che si intersecano con i sottili cordoni spirali formando piccole celle rettangolari. Le differenze morfologiche evidenziate rispetto ai
Rissoininae di altre località del Miocene europeo, come la Paratetide, il Bacino di Aquitania e il Bacino della Loira, hanno confermato la
validità dell’istituzione di Rissoina (R.) giuntellii n. sp. e Rissoina (R.) sturanii n. sp. Questi taxa sono al momento conosciuti solo nei depositi
miocenici del Bacino Terziario Piemontese.
INTRODUCTION
Rissoids are small to minute prosobranchs found in
seas throughout the world. They usually prefer shallow
water and the greatest diversity of species occurs in the
lowest intertidal and in shallow sublittoral zone where
algae, stones, corals or other objects provide shelter. The
oldest fossil rissoids are recorded from the European
Middle Jurassic; few taxa are cited from Cretaceous,
whereas most records pertain to the Cainozoic, especially
of Europe, Indonesia, Japan, Australia, and New Zealand
(Ponder, 1984).
Taxonomic definitions within the family Rissoidae are
complicated by the small size of shells and by their
homoeomorphism due to convergence in architecture and
ornamentation, which are often repeated in different
subfamilies and genera. The supraspecific classification
of Rissoidae was revised by Ponder (1984) and was based
on a combination of diagnostic characters found in adult
shells, embryonic whorls, radula, organic operculum and
the internal anatomy of living specimens. In fact, such a
classification hardly applies to fossil rissoids, at any
systematic level, due to the loss of anatomic and organic
elements and because of the influence of shell
ISSN 0375-7633
preservation. Nevertheless, since Ponder’s is the only and
most complete classification available, it is adopted here.
Our task is facilitated by the shell peculiarity of the
genus Rissoina d’Orbigny, 1840, whose representatives
from the Lower Miocene of Valle Ceppi are treated in
this paper. Taxa assigned to Rissoina show relative large
and turret-like shells, D-shaped apertures usually with
varix on the outer lip and with anterior and posterior
channels; furthermore, the protoconch is pauci- to
multispiral, dome shaped, smooth or weakly sculptured.
The taxonomy of European Mio-Pliocene Rissoina
dates back to the classic monographs on Tertiary molluscs
by Sacco (1895), Friedberg (1911-1928), Cossmann &
Peyrot (1919). Recent contributions with short
descriptions of some Rissoina include those of Glibert
(1949, 1962), Kojumdgieva & Strachimirov (1960),
Baluk (1975; 2006), Landau et al. (2004) and Chirli
(2006). The literature records 27 species from the EuroMediterranean Miocene. Five of these are listed by Sacco
(1895) from the Lower Miocene deposits of the Tertiary
Piedmont Basin (TPB): Rissoina (R.) pusilla, R. (R.)
decussata, R. (R.) obsoleta, R. (R.) moravica, Rissoina
(Rissolina) lamellosa. In the Lower Miocene deposits
of Valle Ceppi, just three of these species were
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Bollettino della Società Paleontologica Italiana, 48 (1), 2009
Fig. 1 - a) Structural sketch map of northwestern Italy (modified after Clari et al., 1994). b) Location of scheme c. c) Schematic map of the
sedimentary succession of Torino Hills; dotted areas indicate conglomeratic bodies.
recognized (R. pusilla, R. decussata and R. lamellosa);
in addition, within the examined material, we identified
two new taxa that are the object of the present paper.
Only the new species are described here in detail,
whereas those already described in the literature are
discussed for comparative purposes.
GEOLOGICAL AND PALEONTOLOGICAL DATA
Valle Ceppi is a deep valley cut by the Rio Civera and
located 3 km northeast of the village of Pino Torinese on
the southern slope of the so-called “Colli Torinesi” (Torino
Hills; Fig. 1). In its middle part, just upstream of Tetti
Civera, a set of fossiliferous beds crops out. They match
one of the best known localities described by Sacco (18891890) in the Miocene deposits of the Torino Hills, which
attracted the interest of many palaeontologists since the
beginning of the nineteenth century (e.g. Borson, 18201825). These include Bellardi (1872-1890) and Sacco
(1890-1904), whose studies on mollusc assemblages led
to the compilation of the namesake collection, formerly
held at the “Museo di Geologia e Paleontologia” of the
Torino University and now transferred to the “Museo
Regionale di Scienze Naturali di Torino”.
The Torino Hills are a structural domain of TPB that
rises south of the city of Torino and the Po River (Polino
et al., 1991). The fossiliferous layers of Valle Ceppi
pertain to the Termofourà Formation, within the Eocene
to Miocene molasse-type succession (Festa et al., in
press a, b) and are attributed to the Early Miocene, in
particular to the Burdigalian Biozone N7a (Zunino,
2007). Lithologically, the Termofourà Fm. is composed
by marls, fine to coarse sandstones and conglomerates
with structures and textures testifying gravity deposition
on the outer slope of large submerged fan deltas (Clari
et al., 1994).
The invertebrate assemblages of Valle Ceppi are
dominated by molluscs (Zunino, 2007) and gastropods
are the best represented fossils. Common taxa belong to
Naticidae (Euspira catena, Neverita josephinia),
Nassariidae (Nassarius badensis, N. tessellatus),
Olividae (Oliva cylindracea, Ancilla glandiformis),
Conidae (Conus antiquus, C. brocchii), Turridae
(Gemmula subcoronata, Bathythoma cataphracta). The
microgastropods are nearly unknown: the commonest
groups are Rissoidae (several species of Alvania,
Rissoina decussata, R. pusilla), Diastomidae
(Sanbergeria perpusilla), Litiopidae (Alaba costellata),
Cerithiidae (Bittium reticulatum).
The state of shell preservation is good but traces of
erosion and fractures are common on the teleoconch, and
are present as a rule on the protoconch. Fossils are totally
allochthonous: the gravity processes that led to the
deposition of the fossiliferous beds involved different
biocoenoses, resulting in a mixing of specimens
transported from different environments, ranging from
the littoral to the epibathyal zones.
M. Zunino, G. Pavia - New species of Rissoina in the Lower Miocene
MATERIAL AND METHODS
Molluscs from the Burdigalian of Valle Ceppi derive
from six large samples (weight: 2-5 kg) taken from four
different fossiliferous layers, which have delivered
homologous assemblages (Zunino, 2007). The specimens
of Rissoina described in this paper were extracted from
sediment after washing, fractioning with sieves and
picking out all the mollusc components. Rissoid shells
were mounted on stubs, coated with gold and observed
using a “Cambridge S-360°” scanning electron
microscope. Morphological features of the protoconch
and teleoconch were examined with special attention to
shape, sculpture, size and number of whorls. The whorls
of protoconch were counted following Jablonki & Lutz
(1980: fide Silva, 2001)
Comparative material derives from the “Bellardi &
Sacco” collection, which assembles the original
specimens (Bellardi, 1872-1890; Sacco, 1890-1904)
including types, figured specimens and supplementary
specimens (Ferrero Mortara et al., 1981, 1984; Merlino,
2007). Additional material was furnished by Piero
Giuntelli; his collection includes several specimens of
Rissoina sampled at Valle Ceppi, as well as some others
sites of the TPB Middle and Upper Miocene such as the
Langhian site of Albugnano and the Tortonian to
lowermost Messinian sites of Sant’Agata Fossili and
Borelli (Ghibaudo et al., 1985; Pavia, 1991; Festa et al.,
in press b).
Besides those held in the “Bellardi & Sacco”
collection, all specimens here mentioned are held at the
Geo-palaeontological Museum of the Torino University.
They are inventoried with codes composed by the
acronym PU (Paleontologia Università) and a progressive
number according to the scheme proposed by Pavia &
Pavia (2005).
53
Topotypes - Zunino’s collection: 66 specimens
(PU108598-PU108664), Giuntelli’s collection: 37
specimens (PU108665-PU108702).
SYSTEMATIC PALAEONTOLOGY
Class GASTROPODA
Subclass PROSOBRANCHIA
Order CAENOGASTROPODA Cox, 1959
Suborder MESOGASTROPODA Thiele, 1929
Superfamily RISSOACEA Gray, 1847
Family RISSOIDAE Gray, 1847
Subfamily RISSOININAE Stimpson, 1865
Genus Rissoina d’Orbigny, 1840
Rissoina (Rissoina) giuntellii n. sp.
Figs. 2 a-f; Pl. 1, figs. 1-2
Derivation of the name - Dedicated to Piero
Giuntelli, expert collector of Miocene fossils.
Type horizon and locality - Valle Ceppi (Pino
Torinese), Piedmont, Lower Miocene.
Type material - Holotype (PU108591); 6 paratypes
(PU108592-PU108597).
Fig. 2 - Rissoina (R.) giuntellii n. sp. at scanning electron
microscope, upper Burdigalian, Valle Ceppi. a) Holotype PU108591.
b) Paratype PU108593. c) Paratype PU108592. d) Particular of shell
ornamentation of paratype PU108592. e) Paratype PU108594,
particular of protoconch, oblique view. f) Paratype PU108595,
particular of protoconch, apical view.
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Bollettino della Società Paleontologica Italiana, 48 (1), 2009
Additional material - 31 from Albugnano
(PU108703-PU108734), 1 from Bocca d’Asino
(PU108735), 3 from Borelli (PU108736-PU108756).
Dimensions - Holotype: H: 3.52 mm; L: 1.4 mm.
Diagnosis - Small shell with convex whorls separated
by crenulate sutures; sculpture composed by prominent
axial ribs, very fine spiral grooving between inner-rib
spaces and prominent cords visible at the base of the last
whorl; multispiral protoconch with small nucleus.
Description - Shell small, solid, conic-turret-like. The
protoconch consists of 3-3.5 smooth and convex whorls,
diameter 360-370 µm, with a small nucleus, diameter
50-56 µm. The teleoconch has five convex whorls with
deeply impressed sutures. The last whorl, approximately
a half of the height of entire shell, is strongly convex
and roundly angled at the base. Ornaments include many
prominent, sharp, straight or slightly opisthocline axial
ribs, which on the last whorl do not reach the abapical
edge. They number 12 to 16 on the first whorl of the
teleoconch, and 18 to 21 on the last one. In addition,
thin, thread-like spiral grooves, best visible on an SEM
microscope, are present in inner-rib spaces. Twelve to
19 prominent cords are visible at the base of the last
whorl. The apertures narrow, oval to D-shaped, obliquely
oriented and contracted in a narrow anal sinus and distinct
siphonal furrow. The absence of varix clearly results
from transport of specimens before fossilization.
Remarks - The large number of specimens allow us
to describe variability in Rissoina (R.) giuntellii n. sp.
Intraspecific variability consists mainly in the convexity
of whorls and in the number and thickness of axial ribs
and spiral cords; this is clear in Valle Ceppi samples as
well as in those coming from Middle and Upper Miocene
localities of Torino Hills. In particular, the early
teleoconch whorls are flat to rather convex, whereas the
last whorl is usually strongly globular with a roundly
angular abapical edge. Axial ribs vary in number, spacing
and inclination: they may be straight but they can also be
clearly opisthocline. Spiral cords at the base of the last
whorl may be prominent to weak.
Discussion - The architecture and morphology of
Rissoina (R.) giuntellii n. sp. are easily distinguishable
from those of other species that occur in the terrigenous
deposits of the Miocene Piedmont Basin, in particular:
Rissoina (R.) pusilla (Brocchi): this is the most
common species in the Miocene deposits of the Torino
Hills and is recorded in the European Miocene-Pliocene
assemblages (Korytnica Basin, Badenian; Bulgaria,
Tortonian; Pliocene of North Italy, Spain, Portugal and
France). Compared to Rissoina (R.) giuntellii n. sp., the
teleoconch of Rissoina (R.) pusilla is larger and
composed of six less-convex whorls; the last whorl
displays an evenly rounded profile. Ornamentation is
similar in both species; nevertheless Rissoina (R.) pusilla
shows 20-33 axial ribs, sharp in cross section, and thin
spiral cords at the base of the last whorl; the aperture is
wider (Pl. 1, figs. 3a-b).
Rissoina (R.) moravica (Hörnes): the specimens
described by Sacco (1895) fit sufficiently with the
Paratethyan taxon Rissoina (R.) moravica (Hörnes). This
species differs from Rissoina (R.) giuntellii n. sp. in the
flat profile of whorls and in the axial ornamentation
reduced to large and depressed ribs limited to the first
whorls of the shell. Moreover, Rissoina moravica has a
wide, D-shaped aperture (Pl. 1, fig. 4). The absence of
protoconch on any specimen of Rissoina moravica from
Torino Hills hinders complementary comparison.
With regards to taxa described by Zilch (1934) and
Baluk (1975, 2006) from the Paratethys Middle
Miocene, Rissoina (R.) giuntellii n. sp. is similar to
Rissoina extranea (Eichwald) and Rissoina eleonorae
Boettger in the shape of axial ribs, whereas it differs in
the more convex whorls and the smaller size of aperture.
As far as the French Miocene specimens described
by Cossmann & Peyrot (1919) and Glibert (1949; 1962)
are concerned, only Rissoina (R.) exdecussata Sacco is
similar to Rissoina (R.) giuntellii n. sp. in the general
shape of teleoconch however Rissoina (R.) exdecussata
shows larger axial ribs and spiral cords. The lack of
taxonomic revisions of small taxa coming from Miocene
France Basin does not permit extensive comparisons.
Stratigraphic distribution - Currently, the taxon is
known only from the TPB Miocene. More specifically,
EXPLANATION OF PLATE 1
figs. 1-2
fig. 3
fig. 4
fig. 5a, b
fig. 6
figs. 7-9
- Rissoina (R.) giuntellii n. sp., Lower Miocene (upper Burdigalian) of Valle Ceppi.
1 - Specimen PU108596.
2 - Specimen PU108597.
- Rissoina (R.) pusilla (Brocchi). Giuntelli’s Collection, Middle Miocene (Langhian) of Albugnano (PU108598).
- Rissoina (R.) moravica (Hörnes). Bellardi & Sacco Collection, Miocene of Sciolze (BS.053.030.006).
- Rissoina (R.) decussata (Montagu). Lower Miocene (upper Burdigalian), Valle Ceppi (PU108599).
- Rissoina (R.) obsoleta (Partsch). Bellardi & Sacco Collection, Miocene of Torino Hills (BS.053.030.005).
- Rissoina (R.) sturanii n. sp. Lower Miocene (upper Burdigalian), Valle Ceppi (PU108765).
7 - Holotype (PU108765).
8 - Paratype (PU108762).
9 - Paratype (PU108757).
M. Zunino, G. Pavia - New species of Rissoina in the Lower Miocene
Pl.551
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Bollettino della Società Paleontologica Italiana, 48 (1), 2009
it has been collected in the Burdigalian of Valle Ceppi,
Langhian of Albugnano, Tortonian of Bocca d’Asino and
S. Agata Fossili, lowest Messinian of Borelli.
Rissoina (Rissoina) sturanii n. sp.
Pl. 1, figs. 7-9
Derivation of the name - Dedicated to the late
Professor Carlo Sturani, the paleontologist from Torino
University who first contributed to modern knowledge
of the Miocene stratigraphy of Torino Hills.
Type horizon and locality - Valle Ceppi (Pino
Torinese), Piedmont, Lower Miocene.
Type material - Holotype (PU108765); 2 paratype
(PU108757-PU108762).
Topotypes - Zunino’s collection: 4 specimens
(PU108758-PU108761). Giuntelli’s collection: 1
specimen (PU108763).
Dimensions - Holotype: H: 3.99 mm; L. 1.70 mm.
Diagnosis - Medium-sized shell with flat whorls
separated by straight sutures; reticulate sculpture
composed of thin axial ribs and spiral cords irregular in
spacing and dimension; median-spiral protoconch with
small nucleus.
Description - Shell medium-sized, solid, turret-like
with an elongated spire. The protoconch is conical with
2-2.5 convex and smooth whorls, diameter 415 µm,
and small nucleus, diameter 80 µm. The teleoconch
consists of four or five nearly flat whorls, separated
by straight and impressed sutures. The last whorl, about
half of the total height, is slightly convex and roundly
angled at the base. The axial sculpture consists of thin,
close-set, rounded and curved ribs. Ribs are irregularly
spaced, usually becoming more frequent at the last
whorl; they number 12 to 16 on the first whorls, to 47
to 52 on the last one. In addition, thin spiral cords run
over the surface of the shell. In the early whorls spiral
cords are visible under great magnification between
axial ribs; they become stronger in the last whorl, wide
as axial ribs, and outline a reticulate sculpture. The
aperture is oval, elongated and posterior contracted;
both lips are thin. Only one specimen shows an outer
varix.
Remarks - The specimens from Valle Ceppi are too
few to assess variability in this species; morphological
features are homogeneous for the whorl profile as well
as for the number, shape and dimension of ribs and spiral
cords.
Discussion - Rissoina (R.) sturanii n. sp. has
morphological features that could be confused with those
of coeval species described by Sacco (1895) and Ferrero
Mortara et al. (1984) in the Lower Miocene fossiliferous
beds of the Torino Hills. In particular the new species is
in some ways similar to Rissoina (R.) decussata and
Rissoina (R.) obsoleta; it can however be distinguished
from both on the basis of the following features:
Rissoina (R.) decussata (Montagu): this species
differs from Rissoina (R.) sturanii n. sp. in the general
shape of the shell, i.e. larger size, flatter whorl profile
and in ornaments. Axial ribs are thinner and denser mainly
in the last whorl; spiral ornamentation similarly consists
of numerous and irregular cords clearly recognizable
within ribs, more evident at the base of the last whorl (Pl.
1, figs. 5 a-b).
Rissoina (R.) obsoleta (Partsch): though similar in
the shape of whorls, this taxon differs from Rissoina (R.)
sturanii n. sp. in the less “conical” profile of the last
whorl. Moreover the ornamentation consists of spiral
cords that are further coupled with a narrow spiral groove,
whereas axial ribs are weak, strongly opisthocline and
evanescent near the adapical suture. Also, the sculpture
is more reticulate with pits formed between sculptural
elements and a larger aperture. (Pl. 1, fig. 6).
With regards to Miocene species of Rissoina from
Paratethys, the morphology of Rissoina (R.) sturanii n.
sp. recalls some taxa recorded from the Middle Miocene
of Korytnica by Baluk (1975; 2006) and Kostej by Zilch
(1934), and particularly, in terms of ornamentation,
Rissoina (R.) varicosa Boettger and Rissoina (R.)
sororcula Boettger. Nevertheless, both species show
narrower apertures and more convex whorls.
Furthermore, Rissoina (R.) varicosa is larger and shows
varix on the shell surface.
Another species similar to Rissoina (R.) sturanii n.
sp. is Rissoina (R.) loueli Deshayes, from the Middle
Miocene of Paratethys and from the Lower Miocene of
Aquitaine Basin (Cossmann & Peyrot, 1919) and the
Loire Basin (Glibert, 1962); Deshayes’ taxon has similar
ornamentation but strongly differs in the less convex
whorls, and in aperture size.
In comparison with Rissoina (R.) giuntellii n. sp.,
Rissoina (R.) sturanii n. sp. is bigger and shows a flatter
whorl profile. Actually, the main differences between the
two species lie in ornaments: Rissoina (R.) sturanii has
weak ornamentation with thin axial ribs and spiral cords;
Rissoina (R.) giuntellii shows predominance of axial
ribbing whereas spiral ornaments consist of narrows
grooves in the inner-rib spaces, which appear thicker at
the base of the last whorl. The aperture is similar in the
two species: oval and narrow.
Stratigraphic distribution - As the supplementary
material of the Piedmont Tertiary Basin has shown, the
species is presently known only from the Burdigalian
deposits of Valle Ceppi.
CONCLUDING REMARKS
The recent revision of the Miocene molluscs from
the Torino Hills (Zunino, 2007) has highlighted the
existence of numerous undescribed taxa, especially small
gastropods, which will be dealt with in future systematic
notes.
With regards to the genus Rissoina, fossils from the
Torino Hills Lower Miocene exceptionally preserve the
protoconch and literature does not supply any detailed
M. Zunino, G. Pavia - New species of Rissoina in the Lower Miocene
study on this morphologic argument. On the other hand,
as far as we know the protoconch of Rissoina is very
simple, being paucispiral or multispiral, and composed
by almost smooth whorls. We do not expect any definite
answers from larval shell morphology, i.e. the characters
of the teleoconch seem to be sufficient to distinguish
taxa and thus to propose new species.
Rissoina (R.) giuntellii n. sp. and Rissoina (R.) sturanii
n. sp. seem to be endemic to the Torino Hills, at least on
the basis of the published and figured specimens from
other classic European sites, particularly those of
Aquitaine and Loire Basins, which pertain to the other
species discussed in this paper. In this respect, special
attention should be devoted to mollusc assemblages of
Paratethys Middle Miocene (Badenian), in particular to
those from Vienna Basin, whose expected revision could
confirm their affinity with Mediterranean taxa, as already
pointed out by several authors (e.g. Pavia, 1991; Zunino,
2007)
ACKNOWLEDGEMENTS
We are grateful to Piero Giuntelli (Nole Canavese), who provided
the specimens analyzed in this paper, and to Daniele Ormezzano
(Torino) for facilitating access to “Bellardi & Sacco” collection. We
would also like to thank Simona Cavagna (Torino) for assistance
with SEM.
We are indebted to the reviewers Elio Robba (Milano) and
Bernard Landau (Albufeira, Portugal) whose comments significantly
improved the paper. Ottavio Janni improved the English of the
manuscript.
The research was financially supported by 2007 funds of the
Torino University (PG6020007), resp. G. Pavia.
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Manuscript received 15 July 2008
Revised manuscript accepted 29 April 2009