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Bollettino della Società Paleontologica Italiana, 53 (2), 2014, i-xiv. Modena
Aciculid land snails (Gastropoda: Caenogastropoda, Cyclophoroidea)
from the Zanclean (early Pliocene) of Balze di Caspreno (central Italy)
Giuseppe Manganelli, Simone Cianfanelli, Debora Barbato & Andrea Benocci
G. Manganelli, Dipartimento di Scienze Fisiche, della Terra e dell’Ambiente, Università di Siena, Via Mattioli 4, I-53100 Siena, Italy; [email protected]
S. Cianfanelli, Museo di Storia Naturale (Zoologia), Università di Firenze, Via Romana 17, I-50125 Firenze, Italy
D. Barbato, Dipartimento di Scienze Fisiche, della Terra e dell’Ambiente, Università di Siena, Via Mattioli 4, I-53100, Siena, Italy
A. Benocci, Dipartimento di Scienze Fisiche, della Terra e dell’Ambiente, Università di Siena, Via Mattioli 4, I-53100, Siena, Italy
KEY WORDS - Gastropods, Tuscany, Pliocene, Zanclean, Acicula kadolskyi n. sp.
ABSTRACT - Five species of aciculids are reported herein from the Zanclean (early Pliocene) of Balze di Caspreno (Tuscany, central
Italy): one belongs to the genus Acicula Hartmann, 1821, three to Platyla Moquin-Tandon, 1856 and one to Renea Nevill, 1880. Statistical
analysis (PCA and RDA) was performed on ten shell variables to test the taxonomic assignment of four of the taxa, in order to compare them
with relevant recent species. Caspreno Acicula differs from similar taxa by virtue of its more evident external peristomal varix, smaller size
and lower H/D ratio; RDA confirmed its distinction from other congeners: it is therefore assigned to a new species: Acicula kadolskyi n.
sp. The largest taxon, Caspreno Platyla, represented by only a few very fragmentary shells, is tentatively assigned to P. dupuyi (Paladilhe,
1868) by virtue of shell size and shape and structure of the external peristomal varix. The other two taxa of Caspreno Platyla are assigned
to P. gracilis (Clessin, 1877) and P. similis (Reinhardt, 1880); RDA confirmed that no significant statistical difference exists between recent
and fossil populations of the two species. Caspreno Renea, represented by only very fragmentary shells, is tentatively identified as R. veneta
(Pirona, 1865) due to its conical shape and dense ribbing. This conclusion is not supported by RDA, possibly due to the fragmentary state of
the material which made it impossible to measure some important shell variables. Apart from A. kadolskyi n. sp., the other species are still
extant. Two of these (P. gracilis and R. cf. veneta) are reported for the first time as fossils and the other two (P. cf. dupuyi and P. similis) for
the first time from the Pliocene of central Italy, although one of them (P. cf. dupuyi) had previously been recorded from the Fossil Forest of
Dunarobba, based on misidentified material of P. similis.
RIASSUNTO - [Gli aciculidi (Gastropoda: Caenogastropoda, Cyclophoroidea) del Pliocene inferiore delle Balze di Caspreno (Italia
centrale)] - Il sito pliocenico delle Balze di Caspreno (Siena, Italia centrale) presenta una ricca malacofauna non-marina, comprendente tra
l’altro cinque aciculidi: una specie di Acicula Hartmann, 1821, tre di Platyla Moquin-Tandon, 1856 e una di Renea Nevill, 1880. Al fine di
definire meglio il loro inquadramento tassonomico sono state effettuate, quando possibile, analisi statistiche (PCA e RDA) su dieci variabili
conchiliari, confrontando specie fossili e attuali. L’Acicula di Caspreno differisce da specie simili per un cercine peristomale ben marcato,
minori dimensioni e minor rapporto H/D; poiché anche l’RDA conferma la sua distinzione, è stata assegnata a una nuova specie: Acicula
kadolskyi n. sp.
La più grande delle Platyla, rappresentata soltanto da pochi esemplari molto frammentari, è stata identificata come P. cf. dupuyi (Paladilhe,
1868) sulla base delle dimensioni e della struttura del cercine peristomale. Le altre due Platyla sono state assegnate a P. gracilis (Clessin,
1877) e P. similis (Reinhardt, 1880); per entrambe, l’RDA non ha infatti evidenziato differenze statisticamente significative tra popolazioni
recenti e fossili. L’ultima specie, rappresentata soltanto da materiali frammentari, è stata identificata come R. cf. veneta per la forma conica
e la fitta costolatura. Questa conclusione non è supportata dall’RDA; tuttavia ciò potrebbe essere dovuto al fatto che lo stato del materiale
non ha consentito di includere nelle analisi alcune importanti misure conchiliari. Tranne A. kadolskyi n. sp., tutte le altre specie sono ancora
esistenti. Per due di queste specie (P. gracilis e R. cf. veneta) si tratta della prima segnalazione allo stato fossile, mentre per le altre due (P. cf.
dupuyi e P. similis) del primo ritrovamento per il Pliocene dell’Italia centrale, sebbene P. cf. dupuyi fosse già stata segnalata per la Foresta
Fossile di Dunarobba, sulla base di un’erronea determinazione di conchiglie appartenenti a P. similis.
INTRODUCTION
The non marine malacofauna of Balze di Caspreno is
one of the most important from the Neogene of Apennine
Italy (De Stefani, 1876-80; Manganelli & Giusti, 1997,
2000; Manganelli et al., 2008, 2011). It is very diversified,
especially rich in land snails, and older than all the other
Neogene land and freshwater molluscan assemblages of
central-southern Italy, such as those from Umbria and
Latium (Manganelli et al., 2008). It includes nine species
of land operculate prosobranch gastropods: one species
of cochlostomids (Cochlostoma esuanum Manganelli
& Giusti, 1997), five species of aciculids, which are
the subject of this article, and three other species (a
hydrocoenid, a craspedopomatid and a pomatiid) which
will be the subject of a future paper.
ISSN 0375-7633
The aciculids are a small group of minute, cryptic,
cylindrical-shelled, litter- and soil-dwelling operculate
snails, endemic to the western Palaearctic. Their taxonomy
is based exclusively on shell characters, though their
phylogenetic significance has never been proved. The ca.
70 recent species recognized are assigned to four genera
(Acicula Hartmann, 1821, Menkia Boeters, Gittenberger
& Subai, 1985, Platyla Moquin-Tandon, 1856 and Renea
Nevill, 1880) mainly distinguished on the basis of shell
sculpture. Anatomical studies have only been performed
in a few species and have not provided any character
useful for genus and species taxonomy, because of the
simplification and homogeneity of the genitalia and other
internal organs (Creek, 1954; Jackiewicz, 1967; Bodon &
Boato, 1987; Boeters et al., 1989; Bodon, 1994; Bodon
& Cianfanelli, 2008). The shell characters of taxonomic
doi:10.4435/BSPI.2014.07
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Bollettino della Società Paleontologica Italiana, 53 (2), 2014
interest for species distinction are shape and size, presence
vs. absence of sculpture (collabral grooves and ribs;
spiral grooves), presence vs. absence and structure of the
external peristomal varix and sinulus.
The split at species level is high and most species
are distinguished by very weak characters, scored
by qualitative analysis (no quantitative studies on
intraspecific/interspecific variability are reported).
Acicula includes 26 extant (seven with fossil record) and
nine fossil species ranging from Oligocene to Pliocene
(Boeters et al., 1989; Bodon, 1994; Stworzewicz & Soltys,
1996; Bertrand, 2004; Kadolsky, 2008; Subai, 2009);
Menkia includes three extant species and one from the
Pliocene of Celleneuve (Boeters et al., 1989); Platyla
includes 30 extant (four with fossil record) and six fossil
species ranging from Eocene to Pliocene (Boeters et al.,
1989; Ambrosetti et al., 1995; Cianfanelli et al., 2000;
Ciangherotti et al., 2007; Bodon & Cianfanelli, 2008;
Subai, 2009; Torres Alba, 2012); Renea includes eleven
extant (one with fossil record) and four fossil species
ranging from Oligocene to Pliocene (Boeters et al., 1989;
Stworzewicz & Soltys, 1996; Ciangherotti & Esu, 2005;
Niero et al., 2012).
GEOLOGICAL SETTING
Fossils were collected in the 1980-1990s along the
Arbia River (Montaperti-Pianella area, northern sector
of the Neogene Siena Basin), where continental deposits
were exposed discontinuously in small scattered outcrops
(Fig. 1). Now most of them are covered by recent alluvial
deposits, vegetation or rubble from quarries. The best
sections of continental deposits crop out in natural cliffs
Fig. 1 - Location map of the studied area. a) Structural setting of southern Tuscany. b) Geological sketch of the Montaperti-Pianella area.
Numbers (1-6) indicate sites sampled during the 1990s. Most of these outcrops are now covered by recent alluvial deposits, vegetation or
dumped rubble.
G. Manganelli et alii - Aciculids from Tuscan Zanclean
iii
on the left bank of the Arbia river (Balze di Caspreno,
fossiliferous sites 5 and 6 in Fig. 1b). These sediments
are indicative of a generic fluvial setting, with dominantly
clayey floodplain silts (locally containing peat horizons)
and fine sands, with subordinate lens-shaped bodies of
gravelly fluvial sand (Martini et al., 2011). The age of
these deposits has long been debated (see Manganelli et
al., 2011 for details and references). According to Martini
et al. (2011), integrated palaeontological and stratigraphic
analysis of the continental and marine succession exposed
in the surrounding area enables assignment of these
deposits to the MPl3 Zone of planktonic foraminifera
zonation (zonal scheme of Foresi et al., 2001). Most
specimens studied were collected in site 2 (UTM ref.
ED50: X = 1696279 Y = 4802046), about 200 m from the
right bank of the Arbia River, exposed during quarrying
in the 1980-1990s but now covered by vegetation. A few
additional specimens came from site 4 (UTM ref. ED50:
X = 1696551Y = 4801725) exposed on the right bank
and on the bed of the Arbia River in the early 1990s and
now covered by recent alluvial deposits and vegetation.
MATERIALS AND METHODS
Taxonomy
The monograph by Boeters et al. (1989) is the standard
basis for the taxonomy and systematics of the family
Aciculidae. Subsequent major contributions have been
published by Bodon (1994), Stworzewicz & Soltys (1996),
Cianfanelli et al. (2000), Bertrand (2004), Ciangherotti
& Esu (2005), Bodon & Cianfanelli (2008), Kadolsky
(2008), Subai (2009) and Niero et al. (2012).
Dimensions
Dimensions were measured to the nearest 0.01 mm,
using Wild M5A and Leica M205 C microscopes. Ten
variables were considered: shell height (SH), last antepenultimate whorl height (L-AWH), last - penultimate
whorl height (L-PWH), last whorl height (LWH), aperture
height (AH), shell diameter (SD), aperture width (AD),
last whorl diameter (LWD), penultimate whorl diameter
(PWD), antepenultimate whorl diameter (AWD) (Fig. 2).
Statistical analysis
Redundancy analysis (RDA; ter Braak, 1986) was
used to detect possible multivariate relationships between
shell variables and the taxonomic assignment of four
out of five Caspreno aciculids, comparing them with
relevant recent species. For each fossil species the most
complete specimens available were considered (see Tab.
1); for living species, five populations (each represented
by five specimens) for each taxon were evaluated except
in the case of two Renea species for which only one
population was available (for the list of living populations
examined, see Appendix). The factor “species” was used
as constraint factor. A preliminary Principal Component
Analysis (PCA) was performed in order to detect the
degree of correlation between shell variables and their
role in explaining variability. Data was log-transformed
before performing statistical analysis. An ANOVA-like
permutation test for constrained ordination was used to
assess the significance (p-value) of the “species” constraint
Fig. 2 - Dimensional variables considered for statistical analysis:
shell height (SH), last - antepenultimate whorl height (L-AWH),
last - penultimate whorl height (L-PWH), last whorl height (LWH),
aperture height (AH), shell diameter (SD), aperture diameter (AD),
last whorl diameter (LWD), penultimate whorl diameter (PWD),
antepenultimate whorl diameter (AWH). Shell figure reproduced
from Boeters et al. (1989) with permission.
for the first two RDA axes. RDA was performed using the
software R 3.0.2 (R Development Core Team, 2013), and
the package vegan (Oksanen et al., 2006).
Acronyms
FGC: F. Giusti Collection, Dipartimento di Scienze
Fisiche, della Terra e dell’Ambiente, Università di
Siena (Siena, Italy); GMC: G. Manganelli Collection,
Dipartimento di Scienze Fisiche, della Terra e
dell’Ambiente, Università di Siena (Siena, Italy); MBC:
M. Bodon Collection (Genova, Italy); MZUF GC: Museo
di Storia Naturale, Collezione Gasteropodi Continentali,
Università di Firenze (Firenze, Italy); RMNH.MOL.:
Nationaal Natuurhistorisch Museum (Leiden, The
Netherlands); SCC: S. Cianfanelli Collection (Firenze,
Italy).
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Bollettino della Società Paleontologica Italiana, 53 (2), 2014
Acicula
fossils
fusca (5 pop.)
lineata (5 pop.)
sublineata (5 pop.)
szigethyannae (5 pop.)
SH
2.41 ± 0.11
2.39 ± 0.17
3.13 ± 0.15
3.06 ± 0.18
3.27 ± 0.25
L-AWH
1.94 ± 0.12
1.96 ± 0.14
2.52 ± 0.09
2.40 ± 0.10
2.35 ± 0.13
L-PWH
1.57 ± 0.11
1.65 ± 0.10
2.08 ± 0.07
1.97 ± 0.08
1.89 ± 0.11
LWH
1.17 ± 0.04
1.20 ± 0.07
1.52 ± 0.05
1.43 ± 0.05
1.36 ± 0.09
AH
0.75 ± 0.03
0.73 ± 0.04
0.94 ± 0.04
0.88 ± 0.04
0.85 ± 0.05
SD
0.92 ± 0.02
0.90 ± 0.06
1.13 ± 0.05
0.99 ± 0.22
1.05 ± 0.06
AWD
0.77 ± 0.02
0.71 ± 0.05
0.89 ± 0.04
0.85 ± 0.04
0.91 ± 0.04
PWD
0.83 ± 0.03
0.78 ± 0.04
0.99 ± 0.04
0.92 ± 0.04
0.94 ± 0.04
LWD
0.83 ± 0.02
0.82 ± 0.04
1.00 ± 0.03
0.94 ± 0.04
0.93 ± 0.05
AD
0.55 ± 0.02
0.50 ± 0.04
0.65 ± 0.02
0.57 ± 0.12
0.61 ± 0.06
Platyla
recent-dupuyi (1 pop.)
fossil gracilis
recent gracilis (5 pop.)
fossil similis
recent similis (5 pop.)
SH
2.89 ± 0.05
2.10 ± 0.05
2.56 ± 0.15
1.83 ± 0.07
2.15 ± 0.19
L-AWH
2.39 ± 0.03
1.80 ± 0.07
2.13 ± 0.10
1.55 ± 0.05
1.71 ± 0.13
L-PWH
1.99 ± 0.02
1.52 ± 0.04
1.76 ± 0.08
1.30 ± 0.05
1.43 ± 0.08
LWH
1.46 ± 0.02
1.12 ± 0.05
1.29 ± 0.06
0.95 ± 0.03
1.06 ± 0.06
0.89 ± 0.02
0.66 ± 0.03
0.74 ± 0.03
0.58 ± 0.02
0.65 ± 0.04
SD
1.09 ± 0.03
0.86 ± 0.07
0.97 ± 0.05
0.68 ± 0.02
0.78 ± 0.05
AWD
0.91 ± 0.01
0.74 ± 0.01
0.77 ± 0.04
0.54 ± 0.02
0.59 ± 0.03
PWD
0.88 ± 0.01
0.74 ± 0.04
0.83 ± 0.04
0.61 ± 0.01
0.66 ± 0.03
LWD
0.76 ± 0.02
0.70 ± 0.10
0.84 ± 0.07
0.63 ± 0.01
0.68 ± 0.03
0.61 ± 0.03
0.52 ± 0.07
0.53 ± 0.03
0.38 ± 0.01
0.44 ± 0.02
AH
AD
Renea
fossil-dupuyi
0.91 ± 0.04
0.64 ± 0.02
fossils
berica (1 pop.)
gormonti (1 pop.)
veneta (5 pop.)
SH
4.15 ± 0.15
3.60 ± 0.10
3.86 ± 0.22
L-AWH
2.87 ± 0.04
2.73 ± 0.05
2.87 ± 0.11
L-PWH
2.34 ± 0.05
2.20 ± 0.05
2.37 ± 0.10
LWH
1.56 ± 0.03
1.68 ± 0.03
1.61 ± 0.04
1.71 ± 0.07
AH
1.00 ± 0.05
1.03 ± 0.03
0.97 ± 0.05
1.05 ± 0.05
SD
1.35 ± 0.04
1.35 ± 0.04
1.29 ± 0.03
1.35 ± 0.06
AWD
1.10 ± 0.03
1.07 ± 0.02
1.08 ± 0.05
PWD
1.21 ± 0.04
1.16 ± 0.03
1.19 ± 0.05
LWD
1.17 ± 0.05
1.22 ± 0.04
1.16 ± 0.04
1.22 ± 0.04
AD
0.73 ± 0.03
0.76 ± 0.03
0.72 ± 0.04
0.76 ± 0.04
Tab. 1 - Mean and standard deviation of variables considered for statistical analysis; pop., population.
G. Manganelli et alii - Aciculids from Tuscan Zanclean
SYSTEMATIC PALAEONTOLOGY
Clade Caenogastropoda Cox, 1960
Informal group Architaenioglossa Haller, 1892
Superfamily Cyclophoroidea Gray, 1847
Family Aciculidae Gray, 1850
Genus Acicula Hartmann, 1821
Type species - Bulimus lineatus Draparnaud, 1801
Acicula kadolskyi n. sp.
(P1. 1, figs 2-3)
Diagnosis - A small sized species of Acicula with
robust shell characterized by ovate-cylindrical shape,
teleoconch with rather irregularly spaced collabral grooves
and subsquare aperture with thick and well reflexed
peristome and evident obsolete external peristomal varix.
Description - Shell dextral, very small in size, ovatecylindrical in shape with about six to six and a half slightly
convex whorls separated by deep sutures; last whorl
about half shell height, slightly ascendent; umbilicus
closed; aperture subsquare, less than one third of shell
height, with thick parietal callus and evident angular
tooth; peristome thick and reflexed with evident obsolete
external peristomal varix slightly before peristomal
margin; upper peristomal vertex, slightly re-entrant in
lateral view; protoconch finely malleate; teleoconch with
rather irregularly spaced collabral grooves that are closer
near growth interruptions and on varix, more spaced in
other parts (about 45 in penultimate whorl).
Dimensions (n: four) - Holotype: SH: 2.39 mm, SD:
0.93 mm, AH: 0.72 mm, AD: 0.54 mm. Paratype MZUF
GC/39607: SH: 2.36 mm, SD: 0.92 mm, AH: 0.76 mm,
AD: 0.56 mm. Paratype GMC 41848a: SH: 2.56 mm,
SD: 0.88 mm, AH: 0.75 mm, AD: 0.53 mm. Paratype
v
SCC 16363/3659: SH: 2.31 mm, SD: 0.93 mm, AH: not
available, AD: not available.
Type locality - Site 2, Balze di Caspreno, along
Arbia River (Montaperti-Pianella area, east of Siena),
lacustrine grey clays of early Pliocene age (MP13 Zone
of planktonic foraminifera zonation of Foresi et al., 2001)
(Fig. 1).
Type material - Holotype (Pl. 1, fig. 2, MZUF
GC/39606) and 89 paratypes (82 paratypes: one whole
adult shell, ten juvenile shells, 71 fragmented shells
consisting of 71 last whorls and six apices, GMC 41848;
one paratype: whole adult shell, MZUF GC/39607; five
paratypes: five fragmented shells consisting of last whorls,
MZUF GC/16362; one paratype: whole adult shell, SCC
16363/3659). Paratype 39607 (Pl. 1, fig. 3) was broken
during SEM work.
Derivation of name - The new species is named after
our friend Dietrich Kadolsky, geologist (Sanderstead,
United Kingdom), a fine malacologist specialized in fossil
non-marine molluscs and a leading authority on zoological
nomenclature.
Remarks - The size range (2.30 - 2.52 mm in height)
places the Caspreno species among the small sized
Acicula. This group includes very few species: Oligocene
A. filifera Sandberger, 1862 and A. pseudosturani
Kadolsky, 2008, Miocene A. isseli (Flach, 1889), Miocene
to Recent A. fusca (Montagu, 1803) and A. parcelineata
(Clessin, 1911), Pliocene A. sturani Schlickum & Strauch,
1979 and Recent A. moussoni Boettger, 1879. Other
species are at the lower fringe of its size range: Oligocene
A. praediezi Kadolsky, 2008, Miocene A. crassistoma
Stworzewicz & Soltys, 1996 and A. diezi (Flach, 1889),
or at the upper limit: Miocene and Pliocene A. edlaueri
Schlikum, 1970 and Pliocene A. michaudiana Schlickum,
1975. Most of these species have an aperture with thin,
EXPLANATION OF PLATE 1
Recent and fossil aciculids: species of Acicula and Renea.
Fig.1 - Acicula fusca (Montagu, 1803): apertural (a) and lateral (b) views of a shell from Irun (Guipúzcoa, País Vasco, Spain), M.
Calcagno & S. Cianfanelli leg. 1.9.1995 (MZUF GC/7560).
Figs2-3- Acicula kadolskyi n. sp. from Balze di Caspreno.
2 - Holotype from Site 2 (MZUF GC/39606); apertural (a) and lateral (b) views.
3 - Paratype from Site 2 (MZUF GC/39607; this specimen was broken during SEM); apertural (a) and lateral (b) views.
Fig.4 - Renea veneta (Pirona, 1865): apertural (a) and lateral (b) views of a shell from Fiume Natisone (debris), Paderno (Premariacco,
Udine, Italy), M. Calcagno & S. Cianfanelli leg. 28.12.1990 (MZUF GC/3571).
Figs5-9- Renea cf. veneta (Pirona, 1865) from Balze di Caspreno.
5 - Apertural view of an incomplete shell consisting of apex and first whorls (MZUF GC/39609a).
6 - Apertural view of an incomplete shell consisting of last whorl (MZUF GC/39609b).
7-8- Lateral views of two incomplete shells consisting of last whorl (MZUF GC/16361a-b).
9 - Lateral view of an incomplete shell consisting of last whorl (MZUF GC/39609c).
Scale bar: 1 mm.
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Pl. 1
Bollettino della Società Paleontologica Italiana, 53 (2), 2014
G. Manganelli et alii - Aciculids from Tuscan Zanclean
non reflexed peristome and lack any external peristomal
varix (see Boeters et al., 1989; Stworzewicz & Soltys,
1996; Kadolsky, 2008). Only A. crassistoma and A.
michaudiana have an aperture with thick and slightly
reflexed peristome and A. michaudiana also a marked
external peristomal varix. A. kadolskyi is distinguished
from the former (A. crassistoma) by virtue of larger size,
external almost parallel to columellar peristome and
more numerous collabral grooves, and from the latter (A.
michaudiana) by virtue of smaller size, ovate-cylindrical
shell shape and presence of an evident angular tooth (see
for A. crassistoma and A. michaudiana, Boeters et al.,
1989; Stworzewicz & Soltys, 1996). Finally, another
species with unknown aperture, A. isseli is distinguished
by its very slender shell shape (see Boeters et al., 1989;
Stworzewicz & Soltys, 1996).
The Caspreno Acicula shares the evident obsolete
external peristomal varix with other species: A. benoiti
(Bourguignat, 1864) from Sicily, A. corcyrensis (Boettger,
1883) from Greece, A. hausdorfi Boeters, Gittenberger
& Subai, 1989 from Greece, A. multilineata Boeters,
Gittenberger & Subai, 1989 from Greece and A. vezzani
Bodon, 1994 from Liguria. All have larger size, higher
H/D ratio and three (A. hausdorfi, A. multilineata and
A. vezzani) also have much more numerous collabral
grooves. Other taxa occurring in central southern Europe
have a sligthly developed external peristomal varix: A.
lineata sublineata (Andreae, 1883) from northern Italy,
A. lineolata banki Boeters, Gittenberger & Subai, 1989
from northern Italy, A. lineolata lineolata (Pini, 1884)
from central Europe and A. szigethyannae Subai, 1977
from peninsular Italy and Dalmatia (Boeters et al., 1989;
Bodon et al., 1995). They constitute a group of similar
taxa, from which A. kadolskyi is distinguished by virtue of
its much more evident external peristomal varix, smaller
size and lower H/D ratio.
vii
Taxonomic setting was tested analysing statistically
the relationships between Caspreno Acicula and living
A. fusca, A. lineata lineata, A. lineata sublineata and
A. szigethyannae (for the list of living populations
examined, see Appendix). RDA 1 (P<0.01) separated two
main groups of taxa: A. fusca and A. kadolskyi on one
hand and A. szigethyannae, A. lineata sublineata and A.
lineata lineata on the other, confirming the relationships
previously proposed (Fig. 3). PCA revealed that size was
the first determinant of shell morphological variance
since PC1 (91%) was a positive combination of all
variables: indeed, fusca and kadolskyi are smaller than the
group of similar taxa formed by lineata, sublineata and
szigethyanne. As stated above, the former two are easily
distinguished by the external peristomal varix, absent in
fusca and very marked in kadolskyi.
The present report is the first of an Acicula species
from the Pliocene of peninsular Italy. On the other hand,
species of Acicula have already been reported from the
Pliocene of northern Italy (as A. cf. lineata: Sacco, 1887;
or A. lineata: Ciangherotti et al., 2007), however their
taxonomic setting should be revised because it is possible
that they belong to the Caspreno species.
Genus Platyla Moquin-Tandon, 1856
Type species - Acme dupuyi Paladilhe, 1868
Platyla cf. dupuyi (Paladilhe, 1868)
(Pl. 2, figs 7-9)
Material examined - nine specimens from site 2 (five
specimens: five fragmented shells consisting of five
apertural fragments, GMC 41849; four specimens: two
Fig. 3 - Redundancy analysis (RDA) performed on fossil Acicula from Balze di Caspreno (A. kadolskyi) and living Acicula species (A. fusca,
A. lineata lineata, A. lineata sublineata and A. szigethyannae). The first two RDA axes (RDA 1 and RDA 2) were both significant for P<0.01
and explained 73% and 2% of the variation, respectively.
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Bollettino della Società Paleontologica Italiana, 53 (2), 2014
fragmented shells consisting of the first whorls and two
fragmented shells consisting of the last whorls, MZUF
GC/16359).
Remarks - Platyla includes recent and fossil aciculids
having smooth shells (apart from a few growth lines)
without collabral or spiral sculpture (Boeters et al., 1989).
These species can be roughly distinguished into three
groups, without taxonomic value, on the basis of absence/
presence and structure of external peristomal varix. The
first group is characterized by absence of an external
peristomal varix; the second group is characterized by
an external peristomal varix without clear proximal
delimitation; the third group is characterized by an external
peristomal varix having a marked proximal delimitation.
Two of the three Platyla species from Caspreno belong
to the second group; the other belongs to the third group.
The largest Caspreno Platyla, represented by a few
fragmentary specimens, is a medium-sized species
characterized by a cylindrical-conical shell and a weak
external peristomal varix (Pl. 2, figs 8-9; Tab. 1). Platyla
species sharing these features include the Eocene
P. eocaena (Oppenheim, 1895), Miocene P. callosa
(Boettger, 1870) and Recent P. curtii (Wagner, 1912),
P. dupuyi (Paladilhe, 1868), P. orthostoma (Jackiewicz,
1979), P. peloponnesica Boeters, Gittenberger & Subai,
1989, P. procax Boeters, Gittenberger & Subai, 1989,
P. similis (Reinhardt, 1880) and P. wilhelmi (Wagner,
1910). Excluding Eocene P. eocaena, cylindrical shelled
P. orthostoma, obese shelled P. procax and small sized
P. similis, distinction among the remaining species is not
easy. P. dupuyi seems to match the Caspreno material best
and hence it was tentatively assigned to this entity. The
other species seem to have a more (P. wilhelmi) or less (P.
curtii and P. peloponnesica) evident sinulus or a slightly
marked external peristomal varix (P. callosa).
P. dupuyi has a west European distribution restricted to
France and extreme northern Spain (Boeters et al., 1989;
Bank, 2011). It has been reported (as P. cf. dupuyi) from
the Pliocene of Fossil Forest of Dunarobba (Ambrosetti
et al., 1995), but re-examination of the specimen suggests
that it belongs to P. similis. Other Platyla material reported
from the middle Pliocene of northern Italy (as P. cf.
callosa: Ciangherotti et al., 2007) might belong to the
same species as the specimens from Caspreno.
Platyla gracilis (Clessin, 1877)
(P1. 2, figs 4-6)
Material examined - Sixty specimens from site 2
(52 specimens: 1 whole adult shell, 2 juvenile shells,
49 fragmented shells consisting of 49 last whorls and 4
apices, GMC 41850; 2 specimens: 2 whole adult shells,
MZUF GC/39610; 1 specimen: whole adult shell, MZUF
GC/20123; 4 specimens: 4 fragmented shells consisting
of last whorls, MZUF GC/20122; 1 specimen: fragmented
shell consisting of last whorl, SCC 16365/3659).
Remarks - This small to medium sized Platyla belongs
to the group of species characterized by a cylindrical
or cylindrical-conical shell with an evident external
peristomal varix (in section, very high proximally,
resembling a step, and gently curved distally) which ends
in the umbilical area where it may form a more or less
evident basal auricle. This group includes large species
with slender conical shell, having an oblique aperture,
such as P. banatica (Rossmässler, 1842) and P. pinteri
Boeters, Gittenberger & Subai, 1989; medium to large
species with cylindrical-conical shell having vertical
aperture and often an evident basal auricle, such as P.
elisabethae (Pintér & Szigethy, 1973), P. falkneri Boeters,
Gittenberger & Subai, 1989, P. foliniana (Nevill, 1879),
P. subdiaphana (Bivona, 1839) and P. talentii Bodon &
Cianfanelli, 2008; an array of poorly characterized, small
to medium species with cylindrical or cylindrical-conical
shell and vertical aperture with slightly evident or absent
basal auricle, such as P. alta (Clessin, 1911), P. callostoma
(Clessin, 1911), P. gracilis (Clessin, 1877), P. maasseni
Boeters, Gittenberger & Subai, 1989, P. pezzolii Boeters,
EXPLANATION OF PLATE 2
Recent and fossil aciculids: Platyla species.
Figs1-3- Platyla similis (Reinhardt, 1880).
1 - Apertural (a) and lateral (b) views of a shell from Torrente Turrita Secca (debris) (Castelnuovo in Garfagnana, Lucca, Italy),
M. Calcagno, S. Cianfanelli, G. Manganelli & L. Manganelli leg. 1.11.1998 (SCC 9630/1868).
2-3- Apertural (a) and lateral (b) views of two shells from Site 2 of Balze di Caspreno (MZUF GC/39611a-b).
Figs4-6- Platyla gracilis (Clessin, 1877).
4 -Apertural (a) and lateral (b) views of a shell from Montepaldi (San Casciano Val di Pesa, Florence, Italy), P. Agnelli & S.
Cianfanelli leg. 22.7.1993 (SCC 4386/754).
5-6- Apertural (a) and lateral (b) views of two shells from Site 2 of Balze di Caspreno (MZUF GC/39610a-b).
Fig.7 - Platyla dupuyi (Paladilhe, 1868): apertural (a) and lateral (b) views of a shell from Mount Pons (France), 1882 (Paulucci
Collection, MZUF GC/10976).
Figs8-9- Platyla cf. dupuyi (Paladilhe, 1868) from Balze di Caspreno.
8-9- Apertural (a) and lateral (b) views of two incomplete shells consisting of last whorl (MZUF GC/16359a-b).
Scale bar: 1 mm.
G. Manganelli et alii - Aciculids from Tuscan Zanclean
Pl. ix
2
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Bollettino della Società Paleontologica Italiana, 53 (2), 2014
Gittenberger & Subai, 1989, P. polita polita (Hartmann,
1840), P. polita regina (Subai, 1977) and P. subfusca
(Flach, 1889). The Caspreno Platyla is related to the
latter group, within which species distinction is not easy.
P. gracilis seems to best match the Capreno specimens,
due to its truly cylindrical shape. P. gracilis has a southern
European distribution limited to the central eastern sector:
it is widespread in southern Germany, Austria, Italy,
Slovenia, Bosnia and Herzegovina, Croatia, Albania and
Greece (Boeters et al., 1989; Bodon & Cianfanelli, 2008;
Bank, 2011).
Statistical multivariate analysis was used to test for
significant differences in shell biometric variables between
living and fossil P. gracilis and between living and fossil
P. similis (for the list of living populations examined, see
Appendix). RDA results showed that RDA 2 (1%, P>0.05)
tended to separate living from fossil populations but the
axes were not significant. This means that there were no
statistically significant differences between recent and
fossil populations of the two entities (Fig. 4). Conversely,
RDA 1 (80%, P<0.01) axes showed a statistically
significant separation between the entire set of P. gracilis
populations and those of P. similis. The preliminary
PCA revealed that size was the first determinant of shell
morphological variance (PC1 92%) since PC1 was a
positive combination of all variables: indeed, P. similis is
smaller than P. gracilis.
This is the first report of P. gracilis as a fossil.
Platyla similis (Reinhardt, 1880)
(P1. 2, figs 1-3)
Material examined - Eighty-one specimens from site 2
(70 specimens: 11 whole adult shells, nine juvenile shells,
50 fragmented shells consisting of 47 last whorls, three
apertural fragments and four apices, GMC 41851; three
specimens: three whole adult shells, MZUF GC/39611;
three specimens: one whole adult shell, two fragmented
shells consisting of two last whorls, 1 fragmented shell
consisting of the first whorls, MZUF GC/20124; two
specimens: two whole adult shells, MZUF GC/20125; three
specimens: three whole adult shells, SCC 20125/17226).
Four specimens from site 4 (three whole adult shells
and one fragmented shell consisting of the last whorl,
GMC 41852).
Remarks - The smallest Caspreno Platyla, represented
by many specimens, including 14 whole adult shells, is
a small-sized species characterized by a cylindrical or
cylindrical to slightly conical shell and a weak external
peristomal varix (Pl. 2, figs 2-3; Tab. 1). It is very similar to
P. similis, the smallest Platyla species having an external
peristomal varix without clear proximal delimitation. This
species has a southern European distribution limited to the
central eastern sector: it is widespread in Italy, Croatia,
Serbia, Albania, Greece, Romania and Bulgaria (Boeters
et al., 1989; Bodon & Cianfanelli, 2008; Bank, 2011).
Pleistocene fossils are reported from Great Britain and
Germany (Boeters et al., 1989).
Statistical multivariate analysis (RDA 2; 1%, P>0.05)
showed that no significant difference in shell biometric
variables separated living and fossil P. similis (Fig. 4)
(for the list of living populations of P. similis examined,
see Appendix).
This is the first report of P. similis from the early
Pliocene. However, re-examination of the specimen
reported as P. cf. dupuyi by Ambrosetti et al. (1995) from
the Pliocene of Fossil Forest of Dunarobba revealed that
it belongs to P. similis.
Genus Renea Nevill, 1880
Type species - Renea bourguignatiana Nevill, 1880
Renea cf. veneta (Pirona, 1865)
(P1. 1, figs 5-9)
Material examined - Fourty-four specimens from site 2
(34 specimens: 34 fragmented shells consisting of 34 last
whorls and 11 apices, GMC 41853; three specimens: three
fragmented shells consisting of two last whorls and one
shell lacking last whorl, MZUF GC/39609; six specimens:
six fragmented shells consisting of four last whorls and
two apices, MZUF GC/16361; one specimen: fragmented
shell consisting of last whorl, SCC 39609/17226).
Remarks - The Caspreno material belongs to a
medium-sized Renea species having a conical shell with
regular spaced ribbing, a subsquare aperture with evident
angular denticle, a thickened interrupted peristome slightly
flared at base with a straight (orthocline) or slightly
curved forward outer lip (prosocyrt), bordered by an
evident external peristomal varix and no sinulus. The
Caspreno species is easily distinguished from Oligocene
R. microceras (Braun, 1851) and Miocene R. pretiosa
(Andreae, 1904) by virtue of its much larger size, and
from Recent R. spectabilis (Rossmässler, 1839) by virtue
of its smaller size. It is also distinct, by virtue of apertural
features, from Miocene R. leobersdorfensis (Wenz,
1921), which has a very marked external peristomal
varix, from Pliocene R. saccoi Ciangherotti & Esu, 2005,
which has a very marked external peristomal varix and
an oblique aperture, from late Pleistocene and Recent
R. bourguignatiana Nevill, 1880, and from Recent R.
moutonii (Dupuy, 1849), R. paillona Boeters, Gittenberger
& Subai, 1989, and R. singularis (Pollonera, 1905), which
have a notch-like sinulus that causes variable detachment
of the final portion of the last whorl from the suture.
Finally, it is distinct from Recent R. elegantissima (Pini,
1886) and R. gentilei (Pollonera, 1889) by virtue of its
conical spire and sparser axial ribbing and from Recent
R. kobelti (Wagner, 1910) by virtue of its conical spire
and evident external peristomal varix.
The remaining Renea species, Recent R. berica
Niero, Nardi & Braccia, 2012, R. gormonti Boeters,
Gittenberger & Subai, 1989 and R. veneta (Pirona, 1865),
are characterized by evident spiral microscultpure between
axial ribs (absent or very faint in the other Renea species;
Boeters et al., 1989; Niero et al., 2012); this delicate
microsculpture is also present in Caspreno specimens
although visible only in small areas, probably obscured by
recrystallization during fossilization. Among these Renea
species, the one that best matches Caspreno specimens is
R. veneta due to its conical shell and dense ribbing (R.
G. Manganelli et alii - Aciculids from Tuscan Zanclean
xi
Fig. 4 - Redundancy analysis (RDA) performed on living and fossil P. gracilis and P. similis. RDA 1 was significant for P<0.01 and explained
80% of the variation while RDA 2 was not statistically significant (P>0.05) and explained 1% of the variation.
berica has sparse ribbing and R. gormonti a cylindric
shell). They are therefore tentatively assigned to this
entity, although there are minor differences in the aperture
(peristome slightly flared at base and outer lip straight
in apertural view in Caspreno specimens; peristome not
flared at base and outer lip curved in apertural view in
R. veneta).
Unfortunately, the fragmentary state of the fossil
specimens did not allow us to test the taxonomic setting
by analysing statistically relationships between Caspreno
Renea and living R. berica, R. gormonti and R. veneta (for
the list of living populations examined, see Appendix).
Indeed, RDA analysis was only performed on variables
that it was possibile to measure on fossil specimens (five
out of ten: LWH, AH, SD, AD, LWD) (Fig. 5). The first
two axes of RDA accounted for 26% and 2% of variation,
respectively. RDA 1 was significant for P<0.01. Caspreno
fossil specimens were separate from all the others. Living
R. veneta specimens were widely dispersed, with those of
R. gormonti at one extreme and those of R. berica widely
overlapping. These results do not support identification
of the fossil specimens as R. veneta, but this may be due
Fig. 5 - Redundancy analysis (RDA) performed on fossil Renea from Balze di Caspreno (R. cf. veneta) and living Renea species (R. berica, R.
gormonti and R. veneta). RDA 1 was significant for P<0.01 and explained 26% of the variation while RDA 2 was not statistically significant
(P>0.05) and explained 2% of the variation.
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Bollettino della Società Paleontologica Italiana, 53 (2), 2014
to the fact that important shell variables such as PWD
and AWD were omitted (the Renea species examined
were distinguished by shell shape and these two variables
contribute to describing this feature).
This is the first report of R. veneta as a fossil.
ACKNOWLEDGEMENTS
Maurizio Ulivi (Centro Servizio Microscopia Elettronica e
Microanalisi, Università di Firenze, Italy) gave technical assistance
during SEM work. Helen Ampt revised the English. Mikaela
Bernardoni and Florence Conti (Biblioteca di Area ScientificaTecnologica, Università di Siena, Italy) helped with bibliographic
research. Bram van der Bijl loaned material from the Nationaal
Natuurhistorisch Museum (Leiden, The Netherlands); Marco Bodon
(Genova, Italy) and Gianbattista Nardi (Brescia, Italy) loaned
material from their private collections. Daniela Esu (Università
di Roma, Italy) and Piero Giuntelli (Università di Torino, Italy)
provided information on Italian fossil aciculids and Ivan Martini
(Università di Siena, Italy) on stratigraphical setting of the outcrop.
Edmund Gittenberger (Leiden, The Netherlands) gave permission
to reproduce figure 210 from the monograph on the aciculids by
Boeters et al. (1989). Folco Giusti (Siena, Italy) offered discussion
and criticism.
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Manuscript received 25 March 2014
Revised manuscript accepted 2 August 2014
Zoobank registration number urn:lsid:zoobank.org:pub:6D82E278F6DA-471D-9109-7DBE5E159AE0
Zoobank registration date 24 September 2014
Published online 30 September 2014
Editor Annalisa Ferretti
xiv
Bollettino della Società Paleontologica Italiana, 53 (2), 2014
Appendix
Material of Recent taxa considered for statistical analysis or used for comparison.
Acicula fusca (Montagu, 1803)
1) Belgium, Namur, Parc National de Furtooz; A.J.
de Winter leg. 25.6.1983 (RMNH.MOL. 161956). 2)
France, Pyrénées Atlantiques, Itxassou; H.J. Hoenselaar
& J. Hoenselaar leg. 26.9.2000 (ZMA.MOLL. 2282567).
3) Germany, Nordrhein-Westfalen, Neandertal; H. Schütt
& P. Subai leg. 22.6.1975 (RMNH.MOL.161965). 4)
Spain, Guipuzcoa/Navarro, 13 km SE Beasain, Alto de
Lizarrusti; A.J. de Winter leg. 8.4.1985 (RMNH.MOL.
161962). 5) Spain, Santander, entrance Cueva de Tijeras,
SE of Haces near Comillas; A.J. de Winter leg. 17.4.1984
(RMNH.MOL. 161961).
Acicula lineata lineata (Draparnaud, 1801)
1) France, Ain, Serrière sur Ain - Bolozon; H.J.
Hoenselaar & J. Hoenselaar leg. 14.9.1999 (ZMA.MOLL.
2284450). 2) France, Jura, S of Lac de Chalain; E. Subai
& P. Subai leg. 18.7.1977 (ZMA.MOLL. 321439). 3)
Germany, Baden-Württemberg, Isteiner Kloz; A.C.
Gittenberger & E. Gittenberger leg. 20.5.1971 (RMNH.
MOL. 162081). 4) Germany, Baden-Württemberg,
Wutachschlucht; A.J. de Winter leg. 30.6.1982 (RMNH.
MOL. 162072). 5) Switzerland, Bern, Spiezberg; A.J. de
Winter leg. 17.5.1980 (RMNH.MOL. 162088).
Acicula lineata sublineata (Andreae, 1883)
1) Italy, Lombardia, San Pellegrino Terme (Bergamo);
P. Subai leg. 22.7.1976 (RMNH.MOL. 162096). 2) Italy,
Lombardia, 5 km S of Serina (Bergamo); E. Gittenberger
leg. 6.9.1974 (RMNH.MOL. 162099). 3) Switzerland,
Tessin, Lugano, Monte San Salvatore; H.D. Boeters leg.
(RMNH.MOL. 162097). 4) Switzerland, Tessin, Tremona;
E. Gittenberger leg. 26.5.1989 (RMNH.MOL. 45456). 5)
Switzerland, Tessin, S of Meride S of Lugano; B. Hauser
leg. 13.7.1977 (RMNH.MOL. 45459)
Acicula szigethyannae Subai, 1977
1) Italy, Apulia: Foresta Umbra (Monte Sant’Angelo,
Foggia), M. Calcagno & S. Cianfanelli leg. 04/01/1996
(SCC 6556/1142). 2) Italy, Sicily: debris of Fiume
Anapo (Cassaro, Siracusa), S. Cianfanelli & E. Talenti
leg. 24/05/2008 (MZUF GC/26742). 3) Italy, Campania:
Boschetiello Mountain (Calabritto, Avellino), S.
Cianfanelli & E. Talenti leg. 30/04/1995 (MZUF
GC/10874). 4) Italy, Campania: near Grotta dell’Angelo
(Olevano sul Tusciano, Salerno), S. Cianfanelli &
E. Talenti leg. 16/10/1994 (MZUF GC/10798). 5)
Tuscany (Pietrasanta, Lucca): Porta, debris of Fiume
Versilia, M. Calcagno & S. Cianfanelli leg. 25/04/2010
(SSC/28380/6314).
Platyla gracilis (Clessin, 1877)
1) Italy, Marche: Favalanciata, debris of Fiume Tronto
(Acquasanta Terme, Ascoli Piceno), M. Calcagno & S.
Cianfanelli leg. 3.6.1995 (SCC 24703/1122). 2) Italy,
Tuscany: debris of Fiume Sieve (Vicchio, Firenze),
M. Calcagno & S. Cianfanelli leg. 4.2.2007 (SCC
23934/5246). 3) Italy, Tuscany: Colognole, debris of
Torrente Uscioli (Pontassieve, Firenze), A. Arrighi,
M. Calcagno, S. Cianfanelli & C. Volpi leg. 11.4.1999
(SCC 8967/2019). 4) Italy, Tuscany: Porto Santo
Stefano, near Sorgente Appetito (Monte Argentario,
Grosseto), M. Calcagno & S. Cianfanelli leg. 22.9.2012
(SCC 42112/18456). 5) Italy, Tuscany: Porta, debris of
Fiume Versilia (Pietrasanta, Lucca), M. Calcagno & S.
Cianfanelli leg. 25.4.2010 (SCC 28378/6314).
Platyla similis (Reinhardt, 1880)
1) Italy, Tuscany: Porta, debris of Fiume Versilia
(Pietrasanta, Lucca), M. Calcagno & S. Cianfanelli leg.
25.4.2010 (SCC 28379/6314). 2) Italy, Tuscany: Risolaia
near Grezzano, debris of Torrente Rampolli (Borgo
San Lorenzo, Firenze), M. Calcagno & S. Cianfanelli
leg. 15.9.1998 (SCC 10410/1873). 3) Italy, Tuscany:
Isola, debris of Fiume Magra (Aulla, Massa Carrara),
S. Cianfanelli leg. 6.4.1997 (SCC 14533/1565). 4)
Italy, Tuscany: Colognole, debris of Torrente Uscioli
(Pontassieve, Firenze), A. Arrighi, M. Calcagno, S.
Cianfanelli & C. Volpi leg. 11.4.1999 (SCC 8965/2019).
5) Italy, Tuscany: debris of Fiume Sieve (Vicchio,
Firenze); M. Calcagno & S. Cianfanelli leg. 4.2.2007
(SCC 23932/5246).
Renea berica Niero, Nardi & Braccia, 2012
1) Italy, Veneto: Monti Berici, northern slope of Monte
Cengia (Barbarano Vicentino,Vicenza), A. Braccia, P.
Greotti & G. Nardi leg. 19.6.2011 (FGC 40682, MZUF
GC/41614, SCC 41614/13309).
Renea gormonti Boeters, Gittenberger & Subai, 1989
1) France, Maritime Alps, Mentone, N of Monti,
M. Bodon leg. 3.1.1999 (MBC, FGC 41854, SCC
40481/17688).
Renea veneta (Pirona, 1865)
Italy, Liguria: banks of Rio Pennavaira near Ponte
Carpe (Castelbianco, Savona), M. Calcagno & S.
Cianfanelli leg. 24.6.1990 (SCC 1056/3). 2) Italy, Friuli
Venezia Giulia: Campone, debris of Torrente Chiarzo
(Tramonti di Sotto, Pordenone), S. Cianfanelli leg.
2.4.1999 (SCC 8875/2014). 3) Italy, Friuli Venezia Giulia:
debris of Fiume Natisone, Paderno (Premariacco, Udine),
M. Calcagno & S. Cianfanelli leg. 28.12.1990 (SCC
1358/10). 4) Italy, Friuli Venezia Giulia: Azzida, debris
of Torrente Alberone, 500 m upstream from confluence
with Torrente Cosizza (San Pietro al Natisone, Udine); M.
Calcagno & S. Cianfanelli leg. 28.12.1990 (SCC 1558/11).
5) Italy, Friuli Venezia Giulia: Invillino, Cascata della
Plera (Villa Santina, Udine), M. Calcagno & S. Cianfanelli
leg. 25.3.1989 (SCC 22974/470).

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