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Bolhttino dclk Societl Pahontologica ltaliana
43 (3),2003 rssN0375-7633409-45r
8 pls.
Modena, Dicembre 2004
The xenophorid gastropodsof the Mediterranean I
the record of the Siena Basin
GiuseppeMaNcANELLI
Dipartimento di Scienze Ambientali
Universit)rdi Siena
Valeriano SpRolNl
Via Augusto Toti 6
Lucignano (fuezzo)
Simone CnNTnNELLI
Museo di Storia Naturale
"Le
Sezionezoologice
Specola"
Universith di Firenze
IGY II(/ORDS- Gastropoda,Xenophoridae,Xenophora davolii n. sp., Reuision,Typt d
hnb.
ABSTRACT - The systematics
of fossilEuro-MediterrAnennxenophoridsis still rektiuely t
studying them (uery conseVuatiue
shell itructure, few auaikble shell chiracters, the fact that the ,
amounT,the rarity of certain species
and the dfficulty offndingwellpreseraed spicimens).Mos
sic monographson Tbrtiary molluscsby M. Hiimet F Saccoand M. Cossmann.
SeaTnipecia haue 6een reporte7from the Euro-Meditelrnnean Pliocene;Xenophora
cr
-Glibert,
1847), X. infundibulum (Broichi, Ig14), X. plioiralica Sacco,1896, X. scaldensis
S. testigera(Bronn, I83I). Howeaenall but t6reeC( crispa,X. infundibulum, and S. testig
fb;s paper is deuotedto six speciesfound in the Siena PlioceneBasin, a clnssicltalian Plio1,
lum, X. p1i<iitalicaS. plioexterisa,aid S. testigera)are redesribedand current hnowbdgeat
fom a rtzen specimeni;t dzscribedas new: X. davolii n. sp. In the remarksto the new splcies,
-Euro-Mediteianean
Tbrtiaryxenophoridspecies,i.e. X. de'shaye
si (Micheloni, 1847), X. burc
(D'Orbigny, 1852), are discussed.
A neotypeis designatgdfortwo species(X. crispa and S. testigera)and a proposalof neoty,
X. deshayesi,
X. burdigdensis,and X. grateloupi.
RASSUNTO - [Gasteropodixenoforidi del Pliocenedel Mediterraneo: I'esempiodel
rea Euro-MeditelraneasonoAncorApoco noti A cAusAdelle mobedfficohh cheda sempresi inr
podi
pochi caratteri disponibili; il fiito chegli ogeni inaccati ,
'in (conchigliamoho conseruAtiuai
nurneroe-forma e modificnrek forrna dei giri; k iarith di aliune speciTe Fdifficohh di n
quindi, chek leneraturaj;;t rileainte disponi-bileconsistaancoradelli clnsiche minografie sui
Saccoe M. Cossmann.
Nel PlioceneEuro-Mediterrnneo,si trouanosettespecie:Xenophora crispa (KAnig 1825),'.
lum (Brocchi,I8I4), X. plioitalica SAcco,1896, X. sialdensisGfibert, I95i], Stellarlaplioexte
rnt A parte nt (X. crispa,X. infundibulum e S. testigera),le alne sonopraticamenteiconoscit^
Questokuoro e /ed;cato alle sei specietouate nel Bacino Neogenicodi Siena, un'AreAclzt
crispa,X. infundibulum, X. plioitdiia S. plioextensae S. testigErl sonoridesrine e denag
su iiascunAdi esse;k sesta,uio xenoforidzdi grandi dimensioni,ltoto da una dozaina di eseil
sp. Nell'analisi delh nuoat speciesoio stateesilninate k ualidith e le relazioni di alcuni clnsici
X. deshayesi(Michelotti, 1847), X. burdigdensis (Gratehup, 1847) eX. grareloupi (D'Orbi1
(Jn neotipo2 stato designatqper due lpecie (X. crispa e-S. testigera) enna propostaper k
specie(X. deshayesi,X. builigalensis e X.'grateloupil.
INTRODUCTION
Xenophorids are a small group of small to large,
slightly mobile, sediment feeding, marine caenogastropods, having conical to depressed, trochiform
shells, with a narrow to wide, simple to digitate
peripheral flange separating base from spire and an
aperture overhung by " sort of roof consisting of the
final dorsal portion of the last whorl. They are characterized by some interesting behaviours, the most
peculiar of which is the abiliry to attach foreign bodies (shells, pebbles, coral skeletons, plates and spines
of seaurchins, plates of balanid cirripeds, shark teeth,
man-made objects, etc.) to the dorsal surface of the
peripheral flange or digitations of their shell
(Vignon, 1923; Morton, 1958; Faber, 1978; Zhu
Min-Da, 1984;Feinstein6c Cairns, 1998;Ponder6c
De Ke.:vze\1998; IGeipl S( Alf,, 1999).The adaptive
value of this behaviour is not clear. It may be an
antipredatorystrategy(a visual camouflagein shallow waters;an olfactoryor tactilecamouflagein deep
waters) ot have a functional value (lifting the shell
above the ground to enable the mollusc to grar,e
without moving; increasingstabiliry or weight to
preventoverturningof the shell by currentsand turbulence)(Shank, 1969; Linsley 6( Yochelson,1973;
Berg, 1974, 1975; Feinstein 6( Cairns, I 998).
However, other peculiar behaviours, includitg
410
G, MANGANELLI, V SPADINI, S. CIANFANELLI
stromboidean locomotion, feeding in the area Protected by the shell, below th. p.t"iphgr4 flange .""4
the ,oof .teated by the final poitiori of the lasiwhorl
grd.burying their faeces,suggeststhat attachment of
foreign bodles is part of a stratery for escaping-predator detection (Mortofi, 1958; Linsley 6( Yochelson,
1973; Zhu Min-Da, 1984; Ponder & De Keyzer,
1998;IGeiplscAlf, 1999).
Xenophioridshellsarevery aftractiveand therefore
popularwith shellcollectors(Faber,1978;Lambiotte,
igTg; Goud & Kronenberg,1988-89;Kreipl & Al[
1999).The startingpoint oI modern til(onomy of the
family is Ponder's(1983) extensjvereviewof extant
taxa.Although Ponder'sreviewalsoincluded a report
on anatomicalmorpholory, his revisionwas mainly
in his words the
basedon shell charactersl-because
"anatomy
appearsto be fairly uniform" and only I
small amouni of materialwasavailablefor anatomical
study.The shell charactersutilized for til(onomy afe:
the amount and relative size of foreiqn material
attached to the shell surface ( I ), the *idth of the
peripheralflange (2), the spire angle (3), the dorsal
iculfture and ttrap. of the whorls-(4),tlt. sculptqre
of the b"t. (5), thg shape.andsizeof the
and^shape
umbili.tt (6), the shapeof the basalmargin of the
aperture(7) and colour, particularlyof the base(8)
(Ponder,1983,p. 3).
Ponder (1933) recognized22 extant species,all
belongitg to a single gEnus,
. XenophqraFischervon
\fald[eirir, 1807, in turn, split intb threesubgenera:
Xenophora(t.t.), StellariaMoller, 1832, and Onustus
Swaihsolr,1840.
"shell
with
Xenophora(t.t.) is characterizedby
peripheial flange narroq simple, non-porcellanous
,r.rit"lly; umb"ilicus moderat-eto closed; foreigt
objects attachedto all whorls and occuplitg more
thin I 13 of the dorsalsurface;mantle edgetentaculate,foot rypicallywith remnantof soleonmetaPodi,t-, ctetridhl filamentsof moderatelength; operculum smooth externally,subtriangular to suboval,
rarelystromboid" (Ponder,1983, p. l8). Xenophqra
(r.r.)'includes13 species:the Ameii canX. cgnihyliophora (Born, 1780), the Mediterraneanand eastern
htlantic X. crispa(Konig,182r, and the Indo-Pacific
X. cereA(Reeve, 1845),X. corrugara(Reeve,1842),X.
flindersi (Cotton & Godfrey, 1938), X. granulosa
-Ponder,
1983,X. joPonicaKuroda & Habe, 1971,X.
(Ne*tott, 1905), X. neozelanicaSutet
mekronensis
(Reeve, 1842), X. peroniana
pallidula
X.
1908,
(Iredale, 1929), X. solarioides(Reeve, 1845) and X
tenuisFulton, 1938.Accordingto Ponder(1983, P:
into rwo informll
4), thesespeciescan be group-ed"rypical
speciesof
The former lncludes
subgroups.^
medium sizeshowingheavyto moderateattachment
of foreigt objects,a-narrowpSriphefl flangeand_a
very narrow to closed umbilicus", such as X.
X.
X. cere*,X. colTugat*,X. japonica,
con'chyliophora,
"small
to
pallidulA, etc.,attd the latter,
,to"ikniro, X. -species
showing moderate to heavy
medium-sized
attachment of foreign bodies, a narrow peripheral
flange and a moderite umbilicus", such as X crispa
andX. solarioid,es.
"shell
large, widely
Stellaria is characterizedby
umbilicate,with wide peripheralflangewhich is simple in some speciesor il digitate or divided into
spines;
hrr*.tous lottg, hollow, narrow parallel-sided
little
of
dorcbvering
small,
obiects
foreiin
attached
sal surface;mi'ntle 6dg. mosdy smootli, .t.ttidial filamentsvery long; fooi lacksmetapodialsole;operculum r-ooih .it.tttallv, subtriangular to suboval"
extantspecies,
(Ponder,1983,p. 50) and includesTirre
mainly Indo-Pacific:X. sohris (Linnaeus,1758), X
(Pfril_ippir18_4
1) '
(Bronn,_I 83 l) , X. chinenszs
testigera
-gigantta
Schepman, 1909, and X. lamberti
X.
(SorlvErbie,1871t The five speciesassignedto
Stellariamay be sroupedin threeinformal subgroups
-"large
(Ponder, 1983,i. 4). The first includes a
specieswith little foreign material attached,a wide
umbilicus,strong radial sculptureon the baseand a
into tubular spines"
wide peripheralTlaneedivided
"species
with
(X. sokrii, the ,..orrJ a -attached,
of medium size^,
digitateperiphervery little foreignmaterial
al flan Ee,? very narrow umbilicus" (X testigerai,and
the third three-"larsespecieswith little foreignmaterial attach.J,-"-oJ.t"i. ,o wide umbilicus,ismooth
sculptured base and a wide, simple
to strongly
-flange"
(X. cltinensis,X. gigantea,and X.
peripherali
lamberti).
"shell
thin with wide
Onustusis characterizedby
peripheral, {l?tg. or weakly digitate, .porcellanous
below;umbilicusnarrowto wide, sometimesplugged
with callus;foreign objectsamachedto peripheryon
few to all whorls-,small and inconspicuous,leaving
most of shell surface exposed;foot lacks sole on
metapodium, mande .4g. smooth, ctenidial filaments very long; operculum radially ridged externally'r'or "lmost ,-6oth, b."ring only verv fite radialstri(Ponder, 1983, pp. SE-Sg)and includes four
species:the Indo-PaCificX. indica (Gmelin, l79l)
ahd X exuta(Reeve,1842) and the westernAtlantic
X. caribaeaPetit de la Saussaye,1857 and X. longleyi
Bartsch,1931.
proPosedby Ponder
systematics
The supraspecific
( I 983) hai largelybeeh adoptedwithout objections.Only recent$?id Kreipl etÂl.(1999), Kreipl S(.Alf
(1999) and' Nielsen S. De Vries (2002) adopt
Xenopltora,Stellaria and Onustusas distinct genera'
"differences
between these talra are large
because
enoughto advocategenusstate"(IGeipl et 41.,1999,
p. 17"9).However,*File Xenophoraand Onustusseem
to havea consistenttil(onomic sense,Stelkria seems
composite,making it problematicto seParatefrom
Onustus.This g.ttit/t,tbg.ttus includes,tery different
entities: one lpecies, fiochus solaris,wiih tubular
(hollow) peripheraldigitationswhich has sometimes
a distinct monotyPic
recentlybeen-assigned-to
^ genus
(Steltaiia or UaTlphoebusFischer, 1879i M?-llory
1978;Lambiofte,1979;Abbott & Dance,1981);one
XENOPHORIDSFROM MEDITERRANEANPLIOCENE
species,Pltorus testigerus,with solid blunt peripheral
digitations and unique anatomical characters (penis
structure), always regarded as a peculiar member of
Xenophora and never associated with Stellaria (ot
Haliphoebus) or Onustus (or Tugurium); three species,
Tiochus chinensis, Xenophora gif,antea, and X. lamberti, having a large peripheral Tlinge with non porcellanous lower surface and without peripheral digitations, which Lambiofte (1979) assifinedto Tusuiium
together with the four species Iater assigiled to
Onustus by Ponder (1983). Species of Onustus are
really very'similar to these three speciesdiffering only
in pot..flanous lower surface of p.tipheral t'fange;
moreover all these species except X. gigantea and X.
longleyi (assigned rc-TrochotuguVium S"ico, 1896, by
Mallory, 1978) also share a wavy inner basal margin
of peristome.
Species of Stellaria and Onustus share some
"n"to-ical characters(foot without metapodial sole;
mantle edge mostly smooth or smooth; very long
which distinguish them frori
ctenidial fiI"-..ttr)
speciesof Xenophora (foot with reirnant of metapodial sole; mantle edge tentaculate; ctenidial filaments
of moderate length)l Ve therefore prefer to recognize
rwo genera, rather than three: Xenophora and
Stelkiia. The latter genus may be split into subgenera: Stelkria (r.r.) for" T solaris',Stellaria (n. sube.i. ?)
for P testigerusand Stellaria (Onustus) for T chlnensis,
X. gigantea, X. lamberti, T indicus, P exutus, X. caribAeA and X. longleyi. According to Kreipl et al. (1999)
and Kreipl S( Alf (1999), Xenophora is also divided
into subgenera:Xenophora (r.r.i and the monorypic
Austrophora IGeipl, Alf & Kronenberg, 1999 (ryp.
speciei: Onustusjlind.ersi Cotton & Go-dfrey, I 938i.
Shell characters support easy identification of
some extant species,but the identification of others is
somewhat controversial. Shell morpholory is very
conservative and species-specificcharacters are modest, especially within the nominorypical subgenus.
Some characters (dorsal sculpture, whorl and suture
shape) are often difficult to evaluate becausemost of
the spire is covered with attached objects or their scars
and others (amount and size of foreign material
attached to surface) may depend on the available
material. Norwithstanding this, Ponder's ( l9S3) systematic accounr of *enolphorid species has largbly
(Goud 6c Kronenberg, 1988-89;
been
'sfilsonaccepted
et-a|., 1993; lkeipl & Alf,, 1999) and few
points of taxonomic conflict have been raised, for
example the status of X. robusta(Verrill, 1870) and X.
Fischer, 1873 (see Hutsell, 1994; Kreipl
senegalensis
S(,{f,, 1999) (fo. the list of exranr species,seeThb. l).
The systematics of fossil Euro-Mediterranean
xenophorids is still relatively unknown, which is not
surprisirg considering the problems involved in
studying them. Besidesthose listed for extant species,
thel inilude the rariry of certain fossil *.rophorids
".d the difficulry ol finding well pr.r.*.d
specimens. Most of the relevant literature dates back to the
4rr
classic monographs on Tertiary molluscs by M.
Hornes, F. Saccoand M. Cossmann. The only recent
exhaustive survey is a little known paper by Zhu MinDa ( 19 84); other recent contributions include
Glibert (1949, 1952, I 958), Ertinal-Erentoz ( I 958),
Koiumdgieva 6. Strachimirov (1960), Holzl (1962),
Pelosio (tg67), Baldi (1973), Janssen(1984), Baluk
'Wienrich
(1995), Marquet (1997),
(200 1) and
Landau et al. (2004). A narrative fossil history of
xenophoridsis proposedby Ponder (1983, pp. 10-15,
Fig. 5). The paper of Zhu Min-Da (l 984) is basedon
the srudy of 400 specimens assignedto 27 different
species,but does not make any significant to(onomic
contribution. The other papers discuss one or more
xenophorids in the context of faunal reports.
Seven species are reported from the EuroMediterranean Pliocene (Sacco, I 896; Marquet,
1997), and all but three (X. crispa, X. infundibulum,
and S. testigera)are practically utrknown."The present
paper is de"votedto th. six speciesfound in thf Siena
Neogene Basin, one of which is describedas new. The
Siena Neogene Basin, a classic Italian Pliocene area
(for g...rfl
information see Bossio et dl., 1992,
1993), occupies an almost central portion of the long
graben that extends SE from Valle del Serchio to Val
di Paglia. It is limited to the west by the Montagnola
Seneseand the Murlo-Montalcino ridge, to the north
by the Monteriggioni high, that separatesit from the
Val d'Elsa Basin, and to the east by the Rapolano
ridge, that divides it from the Valdichiana Basin, an
areadirectly connected to the Siena Basin, at least in
the Middle Pliocene,and continues to the south with
the Radicofani Basin, from which it is separated by
the Pienza-SanQuirico d'Orcia high. It contains limited lacustrine sediments of the Upper Miocene and
abundant marine sediments of the Lower-Middle
Pliocene. The marine sedimentary cycle began in the
Lower Pliocene (Sphaeroidinellopsisseminulina
early
'zone)
s.l.
and ended il the late Middt. Pliocene
(Globorotalia aemilianA zone). Detailed surveys with
stratigraphic, sedimentological and palaeoenvironmental analysis are in progress by A. Costantini, A.
Lazzarotto, and F. Sandrelli of the Dipartimento di
Scienzedella Terra, Universit) di Siena.
Species found in the Sienese Pliocene are
redescribedand current knowledge about them summarized. Morphological nomencfature is usually that
"spire"
used by Ponder (1983), including the term
for
the upper parts of the whorls, adapical to the peripheral flange; the aperture terms are illustrated in the
Text-fig. l. Dimensions are measured excludirg
attached objects (unlessotherwise specified) and spire
angle is taken as a rough measure of the mean spire
angle (Cox, 1960), i.e. the angle berween the straight
lines joining the apex to the periph ery of the last
whorl on opposite sidesof the shell in apertural view.
The nomenclature and taxonomy of molluscan
speciesfound attached to the shells are according to
Cavallo & Repetto ( 1992). The material examined is
G. MANGANELLI, V SPADINI, S, CIANFANELLI
412
Thb. I - Extanr xenophorids.Marine zoogeographicregionsaccordingto Briggs (1974).
Species
Geographic distribution
Main references
Remarls
Indo-'West Pacific ('Western Indian
Ocean, Red Sea, Indo-Polynesian
and Northwestern Australia
Provinces) and Japan Regions
Ponder(1983),Goud 6c
Kronenberg( 1988-89),
'Wilson
et il. (1993), lireipl
& Alf (1999)
Conchologically and anatomicdly
very similir to the America n X.
coichyliophora .and to the sympatnc A. perontilnil.
Ponder(1983),Goud &
Kronenberg( I 988-89),
Kreipl S{ dilf (1999)
Concholoeicallv and anatomically
very similir to th. tropical Indo-'
\7est Pacific X. cerea ind X. peroni-
The speciesof Xenopltora:
X. cerea(Reeve,1845)
(Born, 1780) Carolina,'Western
X. conchyliophora
'West Atlantic
(Caribbean,
Indian and
Brazilian Provinces), California
(Cortez and San Diego Provinces)
and Eastern Pacific (Mexican and
Panamanian Provinces) Regions
X. conugara (Reeve, 1842)
Indo-'WestPacificRegion ('WesternPonder(1983), Stewart&
and Kosuge(1983),Goud &
Indian Ocean,Indo-Folynesian
Provinces)
Australia
Northwesrern
f,,2t r\B;'$j [:,';i?'olF"n
(Leee)
X. crispa(Ktinig, 1825)
Mediterranean-Atlantic (Lusitania
Province) and Eastern Atlantic
(V/'est African Province) Regions
X. flindersi (Cotton & Godfrey, Southern Austrdia Reeion
i e38)
(Southeastern Australii Province)
X. granuhsaPonder,1983
(lndoIndo-\7est Pacific Region
Polynesian Province)
X. japonica Kuroda 6c Habe,
Indo-'West Pacific (Indo-Polynesian
Province) and Japan Regions
r97|
Ponder(1983),Goud &
G iannu zzi-Savelliet al.
(1997), Kreipl s( Alf (1999)
Ponder(1983),Goud &
Vilson et il. (1993), Ifteipl
& Alf (1999)
I(ronenbers( I 988-89),
Ifteipl s{ Af (1999)
IGonenberg(1988-89),
Kreipl S{ Alf (1999)
There is much controversy about
its relationship with the #estern
Atlantic X. sinegalensisFischer,
1879.
X. flindersi is assignedto the subseius Austophorilfteipl, Alf &
f(totrenbere,1999,disfineuished
by strombdidoperculumind paucispiralprotoco^nch.Extant sfeci-.ttr assienedto the nomino^rvpical
X
subspeciei.Another subspecies,
Ponder,1983,
flinlersi ludbroohae
and
bccurs in the Lower Pleistocene
'Western
possibleUpper Plioceneof
Austrdia.
V.ry similar to, maybeconspecific
with, X. tenuisFulton, 1838-.
Polynesian and Nortliwestern
Auitralia Provinces)
Ponder(1983),Goud &
Vilson et il. (199il, llreipl
s{ Alf (1999)
Extant specimens assigned to the
subspeciêsX mehrone"nsiskonoi
Habê, 1953. Nominorypicd subspeciesonly occurs in thi Pliocene
oT Mekran'Coast, Iran.
X. neozelanica Suter, 1908
Northern New Zealand Region
(Auckland and Kermadec
Provinces)
Ponder(1983),Goud &
IGonenberg( 1988-89),
Kreipl S{ Alf ( 1999)
Two extant subspecies are recognized. The no-ittorypical one
occurs in New Zealand seas,the
other, X. neozeknica kermadecensis
Ponder, 1983, is restricted to
Kermadec Islands.
X. palliduk (Reeve,1842)
Indian
Indo-\ilV'estPacific ('\tr?'estern
Ocean, Indo-Polynesian and
Northwestern Aistralia Provinces),
Iapan and Southern Australia
iSbuth*estern Australia Province)
Regions
Indo-Vest Pacific (Northeastern
Australia and Hawaiian Provinces),
Southern Australia (Southeastern
Australia Province) and Thsmanian
Regions
Vilson et il. (1993), Boschet
al. (1995), Ifteipl S( Alf
(Newton, 1905) Indo-'West Pacific Region (lndoX. mekronensis
X. peroniana(Iredale, 1929)
(I eee)
Ponder(1983),Goud &
I(ronenbers( I 988-89),
Vilson et il. (1993), llreipl
s{ Alf ( 1999)
Two extant subspecies are recognized. The tto-inorypical one
occurs in eastern Austialia, the
other, X.peroniana kondoi Ponder,
1983, is restricted to Hawaii
Islands.
Continued.
XENOPHORIDSFROM MEDITERMNEA,N PLIOCENE
413
Thb. | - Continued.
Fischer, 1873
X. senegalensis
X. sokrioides(Reeve,1845)
Ponder(1983),Goud &
Ponder(1933) regarded
X. seneIGonenberg(1988-89),Kreipl galemziasnot mo-rethan an ecologicd form of X. crispa(Konig, 1825),
S( Alf (19{9)
but accordingto lGeipl S( Alf
(1999) there are no intermediate
forms betweenthe two. Therefore
,lt.y consider X. senegalensis
as a disunct specres.
(1933), 9oq4.&
Indo-\fest Pacific Region ('Western P_onder
Extant specim.ens.assigned
to the
Indian Ocean, Indo-Folynesianand '\ilfilson
nominotypicd subspeZies.
The
Northwestern Australia Provinces)
et il. (1993),Ifteipl
- & Upper Mibcene or PlioceneX
Alf ( L999)
solirioidesiezleri Cox, 1948, is very
similar rcX. senegalezszi
Fischer,
Eastern Atlantic Region ('W'est
African Province)
r879
X. tmuis Fulton, 1938
Japan Region
lkoqgnberg ( I 988-89), IGeipl
S( Alf (199:9)
The speciesof Stellaria:
S. caribaeaPetit de la Saussaye, Carolina and'WesternAtlantic
(Caribbean,'West
Indian and
r857
Brazilian Provinces)Regions
Ponder(1983),Goud &
IGo4qnberg( I 988-89), lireipl
ScAlf (199:9
S. chinensis(Philippi, 1841)
(1983), Got1d.&
Indo-\7est Pacific (W'esternIndian P_onder
Ocean, Red Sea,Indo-Polynesian '\tr7ilson
and Northwestern Australia
et il. (1993), Boschet
Provinces)and Japan Regions
41.(1995), IGeipl ScAlf
(L999)
S. acuta (Reeve,1842)
Indo-\7est Pacific (V'estern Indian
Ocean, Indo-Polynesianand
Northwestern Aistralia Provinces)
and Japan Regions
Indo-\7est Pacific ('WesternIndian
Northwestern Aristralia Provinces)
and J"p* Regions
Indo-\7est Pacific ('WesternIndian
NorthwesternAristralia Provinces)
Region
(IndoIndo-'WestPacificRegion
PolynesianProvince)
S. gigantea(Schepman,1909)
S. indica (Gmelin, l79l)
S. hrnberti (Souverbie,I 87l)
S. hnglcli Bartsch, l93l
S. sohris (Linneaus,1758)
F,xtantspecimensareassignedto
the nominotypicalsubspecres.
.Prnorhersubifecies,S. ihinensis
dunheri (Martin , 1879), occursin
the Miocene of Java. S. chinensisis
very similar to S. kmberti
(Souverbie,I 87l).
Ponder(1983),Goud &
'\U7ilson
et il. (1993), lireipl Bc
Atf(reee)
Ponder(1983),Goud &
\Tilson et il. (1993), Kreipl &
Alf (reee)
Ponder(1983),Goud &
\Tilson et il. (1993), IGeipl 6c
Alf (reee)
Ponder(1983),Goud &
liroqgnberg ( I 988-89), IGeipl
sc Alf (1999)
The extantxenophoridspecieswith
Ponder(1983), Goud &
Kronenberg(1988-39),Ifteipl the most reduceildisribirdon. S.
ScAlf (1999)
hmberti is very similar rc X. chinenszs(Philippi, l'844).
Indo-'WestPacific Region ('Western Ponder(1983),Goud &
Indian Ocean and Inilo-Polynesian I(ronenberg(1988-89),Bosch
Provinces)
et al. (199r, lGeipl s{ Atf
Carolina and'WesternAtlantic
(Caribbean,'West
Indian and
Brezilian Provinces)Regions
(I eee)
(Bronn, 1831)
S. testigera
EasternAtlandc (Vest African
Province),SouthernAfrica
(SouthwestAfrican Province) and
Indo-'WestPacific ('WesternIndian
Ocean Province) Regions
kept in public institutions and private collectionsfor
wllich rhe followirg- acronyms are used: AFMSN,
Museo di Storia Naturale dell'Accademia dei
Fisiocriticidi Siena;IPUM, Dipardmento del Museo
di Paleobiologiae dell'Orto Botanico, Universiti di
Modena e ReggioEmilia; DSTUT Dipartimento di
Ponder(1983), Goud 6c
Extant specimensassignedto dislftonenberg (1988-39), Kreipl
S. teiigera digitata
- tinct subspecies:
(Marr.elg,1878) occuridong"the
S( Alf (199:9)
west African coastand S. tesiigera
profunda(Ponder,1983) occu-rsin
hoithernmost EastAfrica and Gulf
of Aden.
Scienze della Terra, Universith di Torino; FPC, F,
Pizzolarc collection, Arezzo; GMC, G. Manganelli
collection, Siena; IGE Museo di Storia Naiurale,
Sezione di Geologia e Paleontologia, Universith di
Firenze; MMAS, Museo del Mare Antico, Salsomaggiore; MRSN, Museo Regionale di Scienze
G. MANGANELLI, V SPADINI, S. CIANFANELLI
414
Naturali, Torino; VSC, V. Spadini collection,
Lucienano. Details on the collettion localities are
giveri in Tab. 2.
BASIN
OFTHE PLIOCENE
THEXENOPHOzuDS
OF SIENA
ClassGnsrnoPoDACuvier, 1795
SubclassOntHocASTRoPoDA
Ponder& Lindb€rg, 1995
Cox, 1960
SuperorderCeENocASTRoPoDA
OrdeiSonnr,ocoNcHePonder& Lindberg, 1997
PoDA
Suborder HvpsocASTRo
Ponder& Lindberg, 1997
Infraorder LrrrozuNIMoRPHA
1975
Golikov & Starobogatov,
1852
toschel,
XENopHoRoIDEA
Superfamily
-Family
Thoschel,1852
XENoPHoRIDAE
Genus XENopHoRA Fischervon \Taldheim, 1807
Typt species Xenophora laeuigata Fischer von
Waldheim, 1807, by su6sequentdesignation (Harris,
tggz).
X. laeuigata is a junior
conchyliophorusBorn, I 780.
XENopHoRA cRISPA(Konig, 1825)
Pl. 1, figs 1-10, Text-figt 2-3
KONtc,p. 3, Pl. 5, fig. 58.
crispus
1825Trochus
"Not
kno\Mn" accordi.g to Ponder
Typt materia/ clarify the"identity.of
To
45).
definitively
p.
09{5,
we designate ihe specimen figured by
Trochus'crispu-r
Sacco (189'6, Pl. 2, fiE. 22; 85.066.01.006) as the
neorype (Text-fig. 3).
lithology
Locality
Siena Basin
Barca (Castelnuovo Berardenga, SI)
UTM: 32TPN9802
SI)
Finerri(Asciano,
UTM: 32TQN124879
SI)
quarry(Rapolano,
di Rapolano
Fornace
UTM: 32TQNL092
Guistrigona (Castelnuovo Berardenga, SI)
UTM: 32TQN0I99
- Castelnuovo
SI)
Berardenga,
I Sodiquarry(Asciano
UTM: 32TQN0198,0298
La Querce (Castelnuovo Berardenga, SI)
UTM: 32TQN0399
SI)
Monsindoliquarry(Siena,
UTM: 32TPN8894
Podere Capanna (Rapolano, SI)
UTM: 32TQN096999
Podere Terrarossa (Castelnuovo Berardenga, SI)
UTM: 32TQN0801
I'oggiodarno (Asciano, SI)
UTM: 32TQN065935
Poggio Vangelo (Asciano, SI)
UTM: 32TPN996966
SI)
SanVittorio (Asciano,
UTM: 32TQN976937
Vescona (Asciano, SI)
UTM 32TQN026942
Val di Chiana Basin
PoggiGialli (Sindunga,SI)
UTM: 32TQN2189
PodereCasuccia(Torrita di Siena,SI)
UTM: 32TQN258L
Radicofani Basin
Lucciolabella(Pienza,SI)
UTM: 32TQN2568
Monte Calcinaio(Radicofani,SI)
UTM: 32TQN3055
PoggioRotondo (Sarteano,SI)
UTM: 32TQN3064
synonym of Trochus
Lower Pliocene
(G lo borotalia puncticulata zone)
Middle Pliocene
(G lo borota lia aemi liana zone)
Middle Pliocene
(G lo borotalia aemi liana zone)
Middle Pliocene
Middle Pliocene
Middle Pliocene
Lower Pliocene
Middle Pliocene
Middle Pliocene
(Glo borotalia aemiliana zone)
Middle Pliocene
Middle Pliocene
Middle Pliocene
Middle Pliocene
Middle Pliocene
Middle Pliocene
Lower Pliocene
(G lo borota lia p un cti culata zone)
Lower Pliocene
Middle Pliocene
references
sand and sandstone A. Costantini (pets. comm.)
"shale"
A. Costantini (pets. comm.)
"shale"
Brogi et al. ( 1999)
sand, sandstone and A. Costantini (pers. comm.)
silry sand
"shale"
sand, sandstone and A. Costantini (pers. comm.)
silry sand
"shale"
Costantini (200 4) and pers.
comm.
"shale"
A. Costantini (pers. comm.)
sand, sandstone and A. Costantini (pets. comm.)
silry sand
"shale"
"shale"
"shales"
"shale"
silry sand
silry sand
silry shale and silt
Costantini & Dringoli (2003)
"shale"
Liotta 6c Salvatorini (1994)
silt
Costantini 6c Dringoli (2003)
Thb. 2 - Collectionlocalitiesin the PlioceneBasinsof Siena,Val di Chiana and Radicofani
415
Text-fig. I - Nomenclature of base and aperture. On the left a shell of Xenophora infundib
Xenophora crispa (Konig, 1825), both from rhe Pliocene ofTerie Rosse-(V.Spadini collecrir
Typt locallU -.[6n1g (182,5).reported
.l grisgrls
(c
"cum
"in
praecedentelTrochusagglutinansLam.]", i.e.
calcario cerithico et in argilla prope Grignon Galliae,
& c." Fischer(1879) and Sacco(1896) regardedthis
localiry as mistaken. Sacco ( I 896) clairned that it
probably comes from an Italian Pliocene localiry and
Ponder (1983, p. 45) restricted the rype localiry to
'Asti,
Italy (Astian, Upp.r Pliocene)". However, witho.rl designation of a lectorype. ol^ neorype, these
actions are of no nomenclatural significance.After the
present designation of the neoryi'e, the rype localiry
becomesdefinitively'Astigiana".
Diagnosis - A speciesof Xenophora characterized
by small size (l), robust, variably shapedfrom almost
discoidal to trochiform shell (2), with narrow irregular, roughly round (where attached objects absent)
peripheral flange (3); about 7 -8 whorls having variable profile (0; spire surfacefree of attached objects
or their scars, representing about 30-600/oof entire
surface(5), and with fine wavy meandering subspiral,
sometimes opisthocline, riblets (6); attached objects
(mainly bivalveand gastropod shellsand fragments of
shellsand pebbles) from small to proportionally very
large (7); scarsof attached objects usually deep (8);
base from slightly convex to almost flat (9); basal
sculpture consisting of collabral growth lines and spiral grooves more distinct near umbilicus and having
more or lessgranular appearance(10); basal margin
o.fperistome rgry curved and thickened and with
ir,r,ting outer portion ( I I ); umbilicus small and deep but
often slit-like or hidden by parietal callus (12).
Description- Shell (Pl. I , figs I - 10) dextral, small
in size, robust, variably shaped from almost discoidal
to trochiform, with pointed apex and narrow irregul"r, roughly round (where attached objects absent)
peripheral flange; spire more or less conical (angle
rangesfrom 70" ro 105") with probably 7-S.regulaily
growing whorls (first whorls always erodefl; last
whorl large, sometimes slightly dilated; sutures from
shallow to deep; profile of whorls variable: where
attached objects are absent or small, whorls flat to
very slightly convex, where attached objects are large
and numerous, whorl profile irregular with lower part
inflated and surmounting suture or periph ery; spire
surface free of attached objects or their scars, representing about 30-600/o of entire surface, with very
fine wavy meandering subspiral, sometimes opisthocline, riblets intersected by prosocline growth lines;
attached objects from small to proportionally very
large, mainly (about 90o/o) shells and fragments of
shell of bivalves and gastropodsor pebbles, and secondarily (about l0%) balanid cirriped plates, bryozoansand foraminifers (largestobject is a pebble 19
mm in length and l3 mm in width) (for other details
on attached objects see Thb. 3); scars of attached
objects _usually deep; base from slightly convex to
almost flat with very variable ornamentation consisti.g of collabral growth lines more marked near aperture and spiral groovesmore distinct near umbilicus
and having a more or lessgranular appearance,apparently due to intersection with growth lines (in some
well preserved specimens, very thin subspiral or
opisthocline riblets are present in areasof lower sur-
G. MANGANELLI, V SPADINI, S. CUNFANELLI
416
Localities
Diameter
(mm)
TerreRosse 38.6t 3.8
(31- 45)
n: 62
n:33
Podere
Capanna
n:9
Height
(mm)
x4.0
26.9
(r8- 3l)
n:29
Angle
(degrees)
number of
attached
objectsin
last whorl
Attached objects
number of
attached
objectsin
penultimate
whorl
8 4 . 2 x 7 . 3 7 . 2 x 1 . 0 6.9x 1.4
(5- to)
(5- s)
(70- loo)
n:32
n z3 2
n:34
90 - 105
n:3
Shellsand fragmemtsof shellsof bivalves
(AcanthocardiZ, Arnusium cristatntrt, An atdara
diluuii, Chkmys, Charneleagallina, Corbuk
gibba, Ghns iiterrnedia, Glity*eris,
Nucuh pkcentinA, Pecten,
h[egdrcivtr4s,
PeEcyora,Tbllina sp., Timocha oaAtut,Venus
uenitcosaand others unidentified), shellsand
fragmemtsof shellsof gastropods(Aponhais
'peipelecani,? Bittiurn, eabarata cahirAt*,
Cincelhria cancelkta, Cancilk, Cerithiarn,
Ckuank romana, Ckaatula rusticA,Conus,
Cymathium,Dilorna patulurn, Gqmmuk contigua, Nassaius,Natica tigrina, Strioterebwm,
Si elti a uar i cosa, Thrri cuk- di rni di ata, Thrri telk
and others unidentified), fragmentsof shellsof
scaphopods (Dentalium), pla-tesof cirripeds,
tubes oTserpulid annelids-(DitruPa)and Pebbles.
Testsof foraminifera, shellsand fragmentsof
shells of bivalves (Acanthocardia,?Ana'dara
diluuii, Chkmys, Nucuk, Nuankna, PehEora
and others unidentified) and shellsand Frryments of shells of gastropods(Nassariusend
others unidentifieiD.
Thb. 3 - Shell parametersof Xenophorarispa (Kcinig, 1825).
faceof peripheral flange); aPertulel"{g.. 1rr.doblique
becauseinnermost basll mirgin is behind by almost
one third of whorl; peristomJinterruPte{*ilh uPPgr
and lower vertices united bV e pari-etalcallus; columellar margin thickened-andsfight[yreflected;basal
margin of peristome l-shaped with inner portion
.,r*.,ed, thi&ened and-initidly slightly refleciedand
outer portion usuallyjuttitg ahd sharp;externalmarein iriesular, thin ^{rd shi'rp;
-byumbilicus small and
marked spiral line
f,..p, t5-.times bordered
which forms a sort of keel, but often slit-like or coveredby parietalcallus.
Dimensions Diameter: 3l-45 mm; height:
18-30 mm (for moredetails,seeThb.3).Dimensions
of other Italian PliocenesPecimens:D: uP to 37-38
mm (De Franchis,1895);D: uP to 50 mm' H: uP to
35 mm (sacco, 1896). Dimensions of Lower
Pleistocenespecimens:D: uP- to 58-59 mm (P.
D:
Franchis,I S95).Dimensionsof e*tant sPecimens:
(Ponder,
D:
1983);
mm
to
H:
up
mm,
33.4
42
up to
rlb to 43 mIrI, H: up to 34 mm (Goud &
Ko..nberg, 1988-89;K.ipl ScAlf,, 1999).
Material - SienaBasin: 94 shellsfrom Barca ( 1,
GMC), Guistrigona (7, GMC: 6, VSC), Podere
Capanna (9, VSC) and Terre Rosse(13, FPC; 49,
VSC). Val di Chiana Basin:4 shellsfrom PoggiGialli
(3, FPC) and Poderecasuccia(1, FPC). Radicofani
Basin:5 shellsfrom PoggioRotondo (5, VSC).
Geographic and stratigraPhic distribution
Xenoph-oricrispa occurs in the central and western
Mediterraneanand the westernAtlandc from France
(Gulf of Biscay) to Angola, in the Azores and the
CapoVerdeIslinds (Ponder,1983).It hasbeenextensivilv recordedfrom the Plioceneand the Pleistocene
of the Mediterranean area, especiallyfrom Italian
ourcrops.Judgingfrom publish;d repglls:.thgs.pecies
was very widEspieadin the Lower+rtiddle Plibcene
(Russieri, 1950, 1962; Lecointre, 1952; Caretto,
1963i Glibert, 1963; Moroni S. Paonita, 1964;
, l97l; Malatesta'
Caprotti, 1967, 1970;D'Alessandro
tg74; Marasti 6c Raffi, 1976; Pavia, 1976;
Montefameelio et dl., 1980; Anfossi et 41., 1982;
Aimone 6(" Ferrero Mortara, 1983; Zhu Min-Da,
1984; Gonzdes Delgado, 1988; Cavdlo 6( Repetto,
E)(PI.{I{ATION OF PIATE I
rispa(Ktinig, 1825)from the Middle PlioceneofTerreRosse(Caste
Figsl-10 - Xenophora
XENOPHORIDSFROM MEDITE
V SPADIAil,S. CIATVFATVELLI,
G. MANGATVELLT,
Pl. I
418
G. MANGANELLI, V SPADINI, S. CIANMNELLI
1992; Thbanelli 6( Segurini, 1994; Andreoli 6(
et al., l?96i BgSi.& Cauli,
Marsigli, 1996;Vera-Pel{ez
1998;Landauet aL.,2004)th.. Vpp...t
1998;-Repetto,
Plioceneind the Lower-Middle Pleistocene(Cerulli
Irelli, l9l4; De Fiore, 1937; Ruggieri, 1949, 1950;
Giannini, 195l; Malatesta 6( Nicosia, 1955;
Malatesta,1960; Glibert, 1963; Moshkovitz, 1963;
Papani 6( Pelosio,1963; Pelosio,1964; Rug.gieri*
Greco, 1967; Zaccaria, 1968; Compagnoni et dl.,
1969; Keraudren, 1970-71; Di Geronimo, 1975;
Menesini& Uehi, 1983;ZhuMin-D", 1984;Leoneet
in
dl., 1998;Thviini et al,, 1998).Its apparentabsence
(.f
Fontan
a-Zanco,
malacofaunas
Tyrrhenian
the
1933;' Segre, 1952a, b, 1954; De Stefani, 1953;
Malatestal 1954; Comaschi Caria, 1955; Cuerda
Barcelb, 1957;Ruggieri& Buccheri,1968;Keraudren'
1977;
1970-71;Durante,1975;Durante& Settepassi,
Russieri & Unti, 1988) might be due to the fact that
the"TyrrhenianoutcropsarJof littord facies,an environment where X. cri$a doesnot live.
There are three recordsfrom the Miocene, two
"Helvdtien" Aquitaine (Salles,Largileyre,
of
from the
Cossmann 6. Peyrot,
and Saint-Etienne-d'Orthb;
"Tortonian"
of Stazzano
l9l9) and another,from the
(Sacco,1896).The report from Salles(Cossmann&
Peyrot,1919,Pl. 13, figs5-6) is listedwith a guestion
mark in the synonv-v of Xenophora deshayesi
(Michelotti, l84n bi Baluk (199fi, but the shell
illustrated(Cossmann6c Peyrot,1919,Pl. 13, figs 56) actuallyseemsvery similar to that of X. crilpa;-,o1t
the contrary,the identiry of the shell from Saint-Etienne-d'Orthe(Cossmann& Peyrot,1919,Pl. 18, fig.
9l ) is very uncertain and likewisethat of the shell
from Srazzano(Sacco,I 896), kept in the BellardiSaccocollectionat the Museo Regionaledi Scienze
Naturaliof Turin (85.066.0I .01I /0 I ).
The ecolory and biology of live X crispaare relatively unkno#.,. Adam & Knudsen (1955, as X.
mediterruneu)reported that living specimenswere
variable tt"t.tt. between
dredeed from bbttoms of 'West
Africa. Le Loeuff et
dept6s of 97 and 263 m in
in
al.' (197l, as X. mediterrAnet)stated that it lives
'West
of
(below
co.astj
the
m)
150
along
deep water
Afriga. Poppe 6( Goto (1991) ieported that,livinq
"gravel
bottoms between40 and
speclmensoccur on
300 m" on the continlntal slope.
X. crispais common and widespreadin the Lower
and Middle Plioceneof SienaBasin whence it was
alreadyreportedby De Franchis(1895).
Remarks The taxonomy and nomenclatureof
this small xenophoridhas lons been discussedfrom
the point of ,rie* of relatiotithipt between extant
Mediterraneanand Atlantic forms-,and betweenfossil and extant Mediterraneanforms. Much earlycongive
rroversyarosefrom the fact that Konig did no!."cum
the exact localiry of collection, but wrote
praecedente[Troihusag{utinansf rarus"in the legend
if agglutinansLamar&, 1804, was describedFrom
the Eocene of the Basin of Paris) (Text-fig. 2). Shortly
from the Italian Pliocene (Bronn,
after, specimens
^the
Italian Pleistocene(Philippi, 1836), the
1831),
Mediterranean (\7einkaufl 1873) and the Atlantic
(Martens, 1876) were assignedto this species.Fischer
"[:..]
(ttl9, p. 2ll) wrote:
qu'il est lmpossible de
cotrse*Er le nom de Xenophoia crispa; I'espbce n'est
pas ddcrite, son gisement est faux,' et I'on ne Peut
une coquille docbne,
savoir si la fieuie reprdsent
ând
consequendi ProPosedto
miocbne ou pliocbne"
commutata
replace it with the names Xenophora
Fiicher, 1879, for the Italian Pliocene forms,
Xenophora trinacria Fischer, 1879, for the Sicilian
mediterranea Tiberi,
Pleistocene ones, Xenophora
1863, for the extant Mediterranean ones, and
Xenophora senegalensisFischer, 1873, for the extant
Atlantic ones. Flowever, Sacco ( I 896) observed that
the shell figured by Konig (1825) is probably from
the Italian Pliocene, wh ereX. rispa is very common.
Consequendy, he did not think ii necepary to substitute Kdnigk name as proposed by Fischer (1879). In
line with 5"..o (1896), Ponder (i983) recently designated Asti (Italy) as the rype localiry of the species,
but this has no nomenclatural significance, without
designation of a neotype. To deflnitively clarify the
iden-tity of Trochuscriipus an4 define its _rypelocaliry
we designate the specimen figured by Sacco (1896,
'Astigiana"
as the
Pl^.2,fi;. 22; B5.066.01.006)?oneorypJ (Text-fig. 3).
rTnocnus,
'I'
IIlb.
Ifig. irli.
'f
Lintt.
LAtn.
(Ifolhnca.
(]ctstct'oTtoda: pccliniLranclicc ;)
ttoc ttv s nggltt tirttt tt.s l.ltu.
irr cnlt:ario t:eritlrico r:r in argilla propc Crignorr {iallie, &c.
nocrrus
ct'is2rts. lt.
Orrrn praccdctrte, ranls.
DisiinguiLlll'
& ceteris basi rcticrrlato-crispa.
Text-fig. 2 - Origind description and illustration of Xenophora
crispa(Konig, 182r.
4r9
"Astigiana" (B.
Text-fig. 3 - Neorype of Xenophoramispa (Kcinig, 1825) from Pliocene of
I
85.066.0
.006).
Tirrin,
Ioly,
Naturali,
Scienzê
The prevalent opinion is currently that the Plioand Atlantic
and ."t"tt Medit.tt"t.".
Pleisto.*.
forms belong to the same monotypic species.Scholars
rêcently Ponder
includittg Sfcco ( I 896) and -ot.
(1983) who examined much extant and fossil material strongly supported this conclusion. Only a few
recent authors have assigned the extant specimens to
a different species (X. inediterranen) ot ; subspecies
(X. crispa mediterranea), but they were sceptical about
the possibiliry of distinguishing it from X. crispa
(Adam 6{ Knudsen, 1955) or did not go into the
quesrion (Garavelli, 1969; Parenzan, 1976; Le Loeuff
it al., 1971; Settepassi,l97l; D'Angelo & Gargiulo,
1978; Lambiotte, 1979; Nordsieck & GarciaThlavera, 1979; Marin 6c Ros, 1990).
Another controversial subject is the status of X.
Although Lambiotte (1979) and Ponder
senegalensis.
(1953) listed it as i junior synonym of X. mediterrespectively, Le Loeuff et Al.
runea and X. crispa,
'Krot..b.tg
(1988-89) and Kreipl
(1971), Goud &
S( Alf (1999) consider it a diitinct species.Le Loeuff
et Al. (197l) regarded the Mediterrane an X. mediterrAnet as specifically distinct from the \7est Atlantic X.
senegaleniison th; basis of statistical and ecological
'West
dati. Kreipl 6. Alf (1999) considered the
Atlantic X: senegalensisa different species, partially
sympatric with western Atlantic and Mediterranean
"not
able to find any
X. irispa, because they were
intermediate forms between crispa and senegalensis"
(Kreipl & Alf,, 1999, p. 53).
aringusDe Gregorio,1885 (p.70; type
Xenophora
"Alt"rrilla"(pliocene)t)
was establishedfof a
localiryr'
Hornes
form with shell similar to that fisured bv
'cumulAns
( 18 56, Pl. 44, fig. 3) as Xen1phora
(Bronsniart, 1823),but with highef spire,subgranu"tgl-d whorls and small
lar bale, posteriorlyslightly
"l". 'kno
the status of De
umbili..ri. As far' "r
Gregorio'sspecieshas neverbeen discussed.Ruggieri
by
et aT.(1958; p. 8) wrote that the taxa establishEd
De Gregorio are not available,but this is not true. \7e
looked Tot rype material of this species,but without
success.Th;'small umbilicus a.d subgranular base
described by De Gregorio support X. aringus as a
junior synonym of X. crispa, the only Xenophora
known from the Sicilian Pliocene.
species
^
Ponder (1983) grouped X. uispa and the tropical
Indian and Pacific X. sokrioides (Reeve, 1845) into an
"small
to medium speciesshowing
informal group of
moder"t."to h."w attachment of foreign bodies, I
narrow peripheral'flange and a modera; umbilicus"
but remarked that thoigh the rwo speciesare similar,
they are not closely related.
Sacco ( I 896) considered X. crispa to be derived
from the Eocene-OligoceneX cumukns_,through X.
cumulans var. transiens Sacco, 1896, of the Vienna
Basin, but this hypothesis is unlikely in the light of
current knowledge of Mediterranean palaeogeography. Ponder ( 1983) suggested that it may have
evolved from Miocene X. burdigalensis (Grateloup,
1847), a Lower Miocene species accorditg to
Cossmann & Peyrot ( 19 I 9). The most likely hypothesisis that X. rispa evolved in the Atlantic during the
Upper Miocene lnd entered the Mediterranean at the
stirt of the Pliocene. Support for this hypothesis
might come from the re-eiimination of the material
from the Miocene of the Aquitaine Basin assignedto
this speciesby Cossmantt & Peyrot ( l9l9) "t? from
the Miocene of Florida, assignedto Xenophora delecta
floridana Mansfield, 1930, it entity apparently very
iimila r to X. crispa (Text-fi g. 4).
XENopHoRA DAVOLIIn. sp.
Pl. 2, figr l-4, Pl. 3, figt l-3
Diagnosis - A species of Xenophora characterized
by verf large size (j), robust gradate shell (2), witll
narrow irrEgular shaped, shaiply keeled peripheral
flange (3); "6ottt 7-9^whorls hiving irregular profile
420
G. MANGANELLI, V SPADINI, S. CANFANELLI
(4); spire surfacemosdy free of attachedobjects or
their icars (5), smoothwith very irregularand prominent rugae,opisthoclinein last whorls (6); attached
objectslbinali'e and gastropod shell fragments and
pebbles) proportionilly vbry large (7); scars of
amachedobl.itr usudly deep'(S);6at. slightly concave (9); basal sculpture consistingof distinct collabral srowth lines itO); basalmaigin of peristome
slightl/cutued, thickenedand slight'lyreflected(11);
umbilicusabsent(12).
Description- Shell (Pl. 2, figs l-4; PL.3, figs 1-3)
dextral, f,.ry large in size, roblst and gradate,with
shaped,
pointed apex aitd very narrow irregirlar
-conical
(ipire
iharply kelled peripherd flange; spire
anslb about 70\, *ith probab-ly7--9regularly grbwirrf whorls (first whorls "l*"yt eroded); lasi i'horl
lai e, not dilated; profile of ivhorls variable:where
attlched obiects ati absent,whorls slighdy convex,
but where ih.y are present, whorls iit.g,tl"t with
lower Dart inflated and surmounting suture or
periph.^tytsuturesusuallvshallow;spireirrface mostly fiee of att"ched objicts and thiir scars,smooth
*ith very irregular, prominent, drop-like rugae,
which in iastwh"orls,.tp..ially towardperipheA,Iake
opisthoclinedirection,intersectedby faint prosocline
giowth lines ("t points of intersecdon,growth lines
Jometimes have squamulate appearance);opisthocline riblets absenti attachedobjects proportionally
consistingof shell fragmenisahd pebbles
very large,
-obiect
(laieest '
sdll aftachedis a 14 x 27 mm pebble,
but- scars of larger obiects are visible); siars of
amachedobjects isually'deep; baseslighdy concave
with distinct collabral growth lines (spiral lines not
evident in the only aduft shell having bise freeof sediment, but visible in a juvenile shelfwhich lackscollabral growth lines); aperture large and oblique
becauseinnermostbasalmargin is almostone quarter
of a whorl behind; peristomeinterrupted wiqh upper
and lower verticesunited by large paiietal callus cbvering columellar and innei por;io; of basalmargin;
marsin of peristomewith inner portion sliehtly
basaT
curved, tKick.n.d and initially slightfy reflectedand
outer portion initidly iutting and then coinciding
with ihternal surface'if p.tilheral flange; exterttJ
margin unknown; umbilicus absent.
Dimensions-The largestspecimen(Pl. 2, frg. l) is
d 12 cm in height iicludabout 17 cm in diameter-an
irg attachedobjects.
EXPTANATION
4- i#ru;{ f:ni:x#?ir#,rkrf#il:1#t
rex'-ng
apparently very similar rc Xenophoracrispa (the
rhitt is 36 mrl in diameterand 30 -tt irheighu
from Mansfield, 1930, Pl. 18, figs 5-6).
Tlpt mateial - Elevenspecimensfrom the followirg localitiesof the SienaBasin:TerreRosse(6, VSC;
l,-AFMSN), Guistrigona(1, VSC) and L" Qterce (1,
FPC; l, GMC). Ahother specimenwas found at
PodereCasuccia(1, VSC) in ihe Val di Chiana Basin.
The holoqpe (Pl. 2, fig. l) is kept in the Museo di
StoriaNatuiale dell'Accidemiadei Fisiocriticidi Siena.
Typt locality
Berardenga).
Terre Rosse (Castelnuovo
Etymohg - After Franco Davoli (Modena, Italy),
for hii man' important contributionson the Neogene
marine molluscsof the Euro-Mediterraneanarea.
Geographic and stratigraphic distribution
Xenopliori dauolii n. sp. is [nown from a few speci-.ttr collectedin to--e Middle Plioceneoutcrofs of
the Siena Basin, all of them deposited in shallow
marine environments.
Rernarks Before describitg a new species,w€
carefullycheckedthat it did not correspondto some
nominal til(on of the many createdby past authors
for Neogene xenophorids (T"b. 4). Once this was
done, *J proceededto describeit as new.
The nirrow peripheralflangemakesit possibleto
assign. XeVophoiadnaoli!. r... ip. to Xenophora.Its
grganuc rtr6 immediately distinguishesif _from all
E*i"trt Xenophoraspecies.In fac7,large Xenopltora,
such as X.'palliduk (Reeve, 1842), X. neo{ehnica
Suter, 1908; and X. cotrugata(Reeve, 1842) do not
exceed90 mm in shell fiameter. Only two extant
OF PIATE 2
Figs
v I-4 Xenophoradauolii n. sp.
1) Holog/pe, Museo di Storia Naturde dell'Accademiadei Fisiocritici di Siena.l
Berardenga);
2) G. Manginelli collection. Middle Plioceneof L" Q,t.rce (CastelnuovoBerarde
3) V. Spadini collecdon. Middle PlioceneofTerre Rosse(CastelnuovoBerardenga
4) F. Plzzolarocollection. Middle Plioceneof La Querce (CastelnuovoBerardeng:
G. MANGANELLI,V SPADINI,S. CIANFANELLI,XENOPHORIDSFROM MEDITE
PT.2
422
G, MANGANELLI, V,SPADINI, S, CIANFANELLI
species,Stellaria gigantea Schepmann, 1909, and
StellarialongleyiBirtsch, 1931, havelargershells,but
arevery diffLt.ttt from that of X. dauoli/n. sp. (for .f.
gigantea,
seeGoud & lironenberg,1988-89,Figs35,
46; Kreipl 6. Alf,, 1999, Pl. 20; for S. h"gkyi, .see
Goud & Kronenberg,l98S-89,Figs35,.43;Kreipl &
Al[ 1999, Pl. 28). The coarseopisthoclinerugaeare
vaguelyreminiscentof those of X. granuloy Ponder,
(Newton, 1905) and X. tenuis
l9i3 ,'X. mekronensis
Fulton, 1838 (for X. granulosasee Goud 6(
Kronenberg,1988-89, FiEt I , 27; Kreipl 6( Alf,
1999, Pl. 7; for X. mekrinensis,see Kreipl 6( Alf,
1999,Pl. 10; for X. tenuisseeGoud & Kronenberg,
1938-89,Figs l, 26; Kreipl S( Alf, 1999,Pl.-17).
X. dauolil fr. sp. is easilydistinguishedfrom the
such
species
other congenericMedit.tt"t.an PliJ.cgng
asX. crispZ(Konig, 1825),X. infundibulum (Brocchi,
1824) ^nd X. pli6italica (SaccoltS96). It is distinct
from X cris\abv its largersize,absenceof umbilicus'
differett *horl profile a'-ndsculpture (X. crispais small
with dorsal t.nlpt.tt. consisti"-gof fine *iry meandering subspiral,sometimesoplsthocline,riblets and
umbilicussmallor slit-like;pl. t , fist I - l0). It is also
distinct from X infundibulum and X, plioitalica by its
gradateshell, con-rre*whorls, dorsal surfacesmooth
irifi very irregularand prominentrugae'which in the
last whorls, .ip..ially io*ard the pJtipneA, take an
opisthoclinedirection, its capacityto agglutinatevery
lirge bodiesand the curvedbasalmargin of its peristome (X infundibulumhasflat whorls, minor agglutination capiciry,dorsalsculptureconsistitg of evident
opisthocline iibl.tt, aperture very large-and inner
basalmargin of peristomestraight;Pl. i, .figt l-4; Pl.
6, figr I-"3;X. )tloltolica has fl"t or slightly convex
whoils, dorsalstulpture consistitg of wary vermiculate opisthocline,"p.tt,rre smallaild auriculate;Pl. 7,
figr l-3). Moreovir, X. dauolii is distinct from X.
from the Belgian Pliocene by large size,
sc-aldensis
absenceof umbilicus,wKorl profile and iculplu re (X.
is of medium size,with dorsalsculptureconscaldenszi
"irregular
plicae" and umbilicus small;
sisting of
1997,ip. fi-ll, Pl. l, fig.7).
Marq-'uet,
X. dauoliimai alreadyhavebeeniePortedfrom the
Italian Pliocene by Sacco (1896) as Xenophora
on the basis oF one
deshayesi(Michelotii, 1847) 'Astigiana",
Piedmont
incomplete shell from the
has a gradate
Basin.tnit shell (Pl. 4, fig. 6) is lirge,
-bodies,
[ike X.
i.ry large
shape and agglutinates
-ft
ptopotiionally
attached
more
has
but
daiolii,
gr
objectsand its dorsalsurfht. freeof ittached objec.ts
their scarsdoes not seemsmooth. Unfortunately,it
lacks the last whorl (where the rugae become more
evident and prominent) and the aperture,so that a
There is anothdefinitivedetirminationis impossible.
(Della Campana,
er repoft of PlioceneX. deshayesi
1890),but it hasbeenimpossibleto check.
is a speciesfrequently recordedfrom
X. deshayesi
the Europ6^n Tertiary (Hornês,t6S6; Michelotti,
1881; Koenen,I 882;
1861; Loiard, 1877;Seguenza,
Parona,1887; Della Campana,1890; Sacco,1896;
GignouX, l9l3; Cossmann 6. Peyrot, 1919;
FriEdb€rg,1923; Kautsky, 1925; Venzo, 1935;
Sorsenfr;, 1940, 1958; Beets,1946; Sieber,1947,
1951; Glibert, 1949, 1952, 1963; Ertinal-Erentoz,
1958; Kojumdgieva& Strachimirov,1950; Holzl,
1962; Anders6n, I 965; Janssen, 1972, 1984;
Nordsieck, 1972; Baldi, 1973; Baluk Sc Radwarlski,
1977; Ponder,1983; Zhu Min-Da, 1984; Baluk,
1995; Lozouet et dl., 2001; tVienrich, 200I;
Harzhauseret dl,, 2003). Its status is nevertheless
uncertain and its relationshipsto other Miocene
nominal species,such asX. birdigalensis(Grateloup,
2), arenot at
1847) an[ X. grateloupi(D'Orbigny, 185
"e
all clear.Sacco(1896, p. 2l) claimed una specie
che passa gradualissimamentesia alle forme
.o..triche, toifota, cumulans,Benettiae,siaalle viventi no.chiformii,pallidula, ecc. tanto che dopo ayeT
non iolo le figure,ma numerosiesemplari
esamlnato,
di ogni piano e di molte localit).,un vero carattere
nett; distintivodi questaspecienon sapreiindicarlo".
Although someauthors(Sacco,1896; Cossmann
6(rPeyrot,"I9l9; Zhu Min-Da, 1984) have studied
toporypicalmaterial,the lectotypeor the neoryPehas
redescripThe only exhaustive
rerr.t'beendesignated.
(18 59),
Hornes
of
those
are
X.
delhayesi
of
tions
Cossmann 6{ Peyrot ( 1919) and Zhu Min-Da
are
of X burdigalensis
( 1984).The only redescriptions
thoseof Cossmann& Peyrot(1919),Jansien(1984)
(1984)
and Zhu Min-Da (1984),but that of Janssen
may be basedon misidentified material.The onlr
is that of Cossmann6{
redescriptionof X. gratelguPi
-revision
of these taxa is
Peyrot i t g I 9). Although
paper,
it usefulto
we
thought
this
of
beyondthe aim
dehtre them to discusstheir relatiottJhiptwith the
Pliocenespecies.
Statusa/ Phorus deshayesiMichelotti, 1847, and
allied Mioiene species Michelotti (1847,
PP. 173"P
as [horus]. _testa
174) described-Phorusdeshayesi
conici, corporavariaagglutinante;anfractibusplanis,
superner.tgit laxe ."ttlillati; basi sulcisradiantibus
obiiquis inJtructi; umbilico tecto; margine_simplici"
-de
"La
colline de Turin, les environs Bordeaux,
from
la Touraineet la Polosne".According to Michelotti
benet(1847) somematerials"'misidentified
ai Trochus
Pl. 6, figt 3a-3b;
tiae by Brongniart (1823, -p. 56,"la
"Loignan
moniagne de
prEs Bordeaux" and
Supergue prbs Turin"), Pusch ( l8 37, p. I 10;
"Lond6nthon, xr der Supergabei Turin und bei
"faluns
de
Bord,eaui')and Dujarditr ( 1937, p. 284;
Manthelan")matchedto his species.
Grateloup
- ( 1840, but actually 1847, caption of
toques Pl. 1, fig. 1) introduced TrochuscoyclUliofor a shell from Saubrigues'
phoiusvar. burdigalensis
"
listing as synonyms: Troc. Benettiae.Sow. Miner.
ConJh.t. 9'8.f. i. 2. de Bast.Foss.n.o 1. Al. Brogn.
Vic. pl. 6. f. 3. Grat. Thbl. 189. Grat. Cat. zool.n.o
"l
l8l Thesefive entriesincludethe originaldescrip-
423
Teb. 4 - Nominal ta:<aof the species group introduced for Neogene xenophorids. The lis
( 1983) and includes alf .urr.rr"rly lino*n vdid and invaliJ nominal'taxa established
duced by Sacco (1896) as varieties, have controversial nomenclatural status. In facr
as such are not available (ICZN, 1999: Art. 45.6.4). Their infrasubspecific status
til€ often coexist temporallv and seosraphicallv with other forms intluding the :
"Pi
"Piacenziano
X. crispa var. depressiir.
ed futiario: (cdl tipo,"mi"lq,t"nto'rara)",
"...-Piacenziano:
quente)"; X. testigeravar. ektiuicula:
Colla'forma tipica (poco I
dnderArt. 45.6.4.1, i.e. if a subsequent author adopted them as the valid name of
rbination, complete reference, rype localiry (as 6riginally given) and some remarki are given
Nomind
Ty'pe locality
taxon and references
"France,
St-Paul (Landes)"
lPhorus)AquensisD'Orbigny, 1852: 41, by Miocene;
"Da"
... ) St-Paul, fal.
indication (Grateloup,1847,TioquesPl. (D'Orbigny, 1852);
1, figs 3-4, excl.figs I -2 and synonymy). j a u n ., 2 . ' d t . , s u p . " ( G r a t e l o u p , 1 8 4 7 ) .
Xenophora
aringusDe Gregorio,1885:70.
Pliocene; Sicily,
"Altavilla".
"presso
Pliocene; Italy,
Verduno in
Piemonte".
"Zone
of Aricia humerosA"; Burma,
XenophorabirmanicaNoetling, 1901: 280- Miocen e,
Thayetmyo.
28I, Pl. 18, figs 2l-21a.
"
"Terr.
mioc. Thur.
PhorusBorsoniSismonda,1847: 50, by indi- Miocene; Italy,
"La
colline de Tirrin et
cation (Michelotti, 1847, 175, Pl. 7, fig. (Sismonda, 1847);
Tortone" (Michelotti, 1847).
1).
"Dax
.. . ) Saubrigues,Fal.
var. Burdigalensis Miocene; France,
[Trochusconchyliophorus]
e
material
2.
inf.";
from othir localiPl.
l,
fig.
l.
bl.,
dt.,
Grateloup,I 847, Tioques
ties is cited by bibliographicd references.
"Tortonese".
Miocene; Italy,
TrochuscolligensMichelotti, 1840: 14.
TrocbusBelkrdii Michelotti, I 840: 14.
XenophorA
commutataFischer,1879: 2ll.
Pliocene,
".
(Sicilie)
"Astdsan
... rypique IAltavilla
Remarks
A junior objectivesynonymof Trochus
var.parisiensisGrateloup,
conchyliophorus
1847. Accordingto Cossmann& Peyrot
(1919), ajunior synonymof Xenophora
burdigalenszi(Grateloup, 1847).
A junior synonymof Xenophorarispa
(Konig, 1825) (this paper).
Accordingto Sacco(1896),a junior syn(Bronn, l83l).
onym of Stelkria testigera
Accordingto Ponder& Cooper (1983),a
Xenophora (Xenophora).
Accordingto Ponder& Cooper ( I 983), a
Xenophora(Onustus).
Xenophora burdigalensis(Grateloup, I 847).
Accordingto Sacco(1896),a junior syn(Bronn, l83l).
onym of Stellnriatestigera
A junior synonymof Xenophorauispa
(Konig, 182r.
"cum
praecedente lTrochus Xenophora mispa (Ktinig, 1825).
"in
agguitinans Lam.]", i...
calcario cerithico
et in argilla prope Grignon Galliae, & c."
"Asti,
Italy
Typ. localiry restricted to
(Astian, Upper Pliocene)" by Ponder (1983:
45),is definitively fixed by the present neorype designation.
Trochus
crispusKonig, 182523, PI.5, fig.58. Konig reported it
Accordingto Ponder& Cooper (1983),a
Miocene ('Argilla del miocene inferiore");
Pantanelli,I 887: 125Xenophoradepressa
"Torre
Xenophora (Stelkria) species.
Italy,
della Maina'.
126, Pl. 5, figr 6-7.
"
A junior synonym of Xenophoracrispa
lXenophora) (XTenophoral .) mispa var. depres- Pliocene , PiAcenziano ed Astiano ldel
Piemonte e della Liguria]: (col tipo, ma poco (Konig, 1825).
sior Sacco,1896: 22, Pl. 2, fig. 24.
G*ppy, 1876:529, PI.28,
Phorusdelectus
fig. 10.
Miocene; Dominican Republic, Haiti.
frequent€)", Italy. The shell illustrated is
from Ceriale.
"La
colline de Turin, les environs
PhorusdeshayuiMichelotti, 1847: 173-174. Miocene,
de Bordeaux, la Touraine et la Pologne". If
and when the designation of the proposed
neotype is ruled by the ICZN, the rype
"Colli
Torinesi".
localiry will begin
Xenophora deshayesi(Michelotti, 1847).
XenophoraDunkeri Martin, 1879: 7I-72, Pl. Miocen e; Java (Ponder & Cooper, 1983).
12, fig.7.
Stelkria chinensisdunheri (Martin, 1879)
(seePonder,1983).
"
Miocen e, Ehteziano: Colli torinesi (non
hora].) deshayai van
fXenophora) (XTenop
elntespirata
Sacco,1896: 21, Pl. 2, fig. 21. rara)", Italy.
Accordingto Ponder& Coop.f (J 983), a
junior synonym of Xenophoradeshayesi
(Michelotri, 1847).
"
lXenophora)(XIenophoral.) crispavar. elatior Pliocene , PiAcenziano ed Astiano ldel
Piemonte e della Liguria]: (col tipo, ma
Sacco,1896:22, Pl. 2, fig. 23.
A junior synonym of Xenophoracrispa
(Konig, 1825).
dquanto rara)", Italy. The shell illustrated is
from the Lower Pliocene of Bordighera.
Continued.
G. MANGANELLI, V SPADINI, S. CIANFANELLI
424
Thb. 4 - Continued.
lXenophoral (XIenopboral.) testigeravar.
elaiiuscukSaccolts96:25, Pl. 3, fig. 2.
hora) infundibulum) var.
lXenophora (Xenop
e*fansiorSacc6,I 896:24.
"
A junior synonym of Stellariatestigera
Miocen e, Elueziano: Colli torinesi,
( B r o n n ,1 8 3 1 ) .
Baldissero, S. Raffaele (non rara)", Italy.
"
Pliocene , Piacenziano ldel Piemonte e della"
Liguria] : colla forma tipica (poco frequente)
ItJy.
Pliocene, [del Piemonte e della Liguria],
Italy.
A junior synonymof Xenophorainfundibu'
lum (Brocchi,l8l4).
"Terrains
Accordingto Sacco(1896),a iunior synmarinstertiairesdu Midi de la
marines, onym of Xenophorainfundibu'lum(Brocchi,
France.. . dansles marnessableuses
qui forment de petitescollinessur lesrives 1814).
de la T€t )r Millir et I Neffiach,ou sur les
rivesdu Tech au Boulon et i Banyuls-delsAspre".
"Concretionary
Member of Xenophoralbmingt Beu, 1977.
Xenophora
flemingiBey,1977:233-235,figt Lower Miocene,
Otahu Formation,betweenCliFclenLime2 , 5 , 7 ,8 , 1 0 ,l 3 - 1 7 .
stoneand Calamity Point, south bank'Waiau
River,Clifden, Southland",New Zealand.
nochusFarinesiDe Serres,1831:73-75.
Xenophora dzhcta var. floridana Mansfield,
"Harveys
Creek, half a mile above A Xenophora speciesvery similar to
Miocene,
Xenophora criipa (Konig, 182r.
abandoned mill, L.ott Counry Fla.", US.
figs 5-6.
t93o: l2l-122, P1."18,
"France,
Xmophora grateloupi (D'Orbigny, 1852).
Dan, Saubrigues
Phorussaneloupi [sic] D'Orbigny, I 852: 7, Miocene,
"Da*
... I Saubrigues,
bv in?icadon(Grateloup,1847:Thoques (D'Orbigny,1852);
Fal.bl.,2.', dt., inf." (Grateloup,1847).
Pl. t, fig. 2, excl.figs I ,-3, 4)
"Piacenziano:
plioitalicum
A junior synonymof Xenophora
Rio del Molino di
(Tluguium)
Pliocene,
p
var.
p
in
liatm
.
lio
[sic]
)
lXan horal
(Sacco,1896).
Chieri (rara)",Italy.
presso
Galatea
graiilior Sacco,-l896:2t-26, Pl. 3, Fig.7.
"Vaqueros
Accordingtg_Ponder& Cooper (1983), a
horizon,LowerMioceneonly?
XenophorahawleyLoel 6c Corey, 1932: 269'
Abuhdant in wesrernSantaYnez.Mountains Xenophori (Xenophora).
2io, Pl. 63, ffg. 12.
in the abovenoted zrlne,SantaBarbara
Counry and presentin SanLuis Obispo region
(CorrJ de Piêdra
Creek)', California,US.
XenophorahelueticaDollfus & Dautzenberg, Miocene;France,falunsde la Touraine
"faluns
de
(Dollfus & Dautzenberg,1886);
tdg6: 142,by indication("Trochus.
(Dujardin,ig3z).
Manthelan"
Xenophora
Brong.").
non
Dui.
Benneni
"Mayer"
Kaufiiann, 1872: 504,
heluetica
is a nomennidu-.
Xenophorahoi
fid'.4-6.
'Wang,
1984: 350-351,Pl. 4,
Ponder6r Cooper ( 1983) considered
Xenophoraheluiticaavailablefrom Rutsch
0929); actuallyil ifqyailable sinceDollfus
ied
an
il:'"Tf
i?fl,hnil,T
f
*m,:l;:'"
fiCoop.t
(1983), a Xenophori(Xenophora).
A nominal speciesbasedon very badly PreMiocene;Chiufenerh-shan,KuohsingNantou-hsien,centralThiwan-;uPPer-servedspecimens.
hsiang,
"p"tt
of ShihmemMember of the
most
ShuilikengFormation (late earlyMiocene).
According!qPonder & Cooper (1983),a
Xenophori (Onustus).
Gabb, 1873: 241.
Olnustus).imperforatus
Miocene,SanroDomingo, Dominican
Republic.
TrochusinfundibulumBrocchi, 1814: 352353,Pl. 5, fig. 17.
Xenophorainfundibulum (Brocchi,l8l4).
Pliocene,"... nel Piacentino,a San
Geminiano e a Libiano in Valdera",Italy.
"Dax
Fal. A not availableseniorobjectivesynonymof
... I Saubrigues,
Miocene;France,
e
from othErlocali- PhorusgrateloupiD'Orbigty, 1852.
material
inf,";
bL,2. dt.,
var. italica [sic]
conchyliophorusl
lTrochus
Grateloup,ig4l: TioquesPl. I , fig2.
ties is cited by bibliographical references.
XenophoraiezleriCox, 1948220-21, Pl. 1,
figs I la-l lc.
'W'ang,
19842347-349,
Xenophorakantzulina
Pl: 3, figr. l-4, Pl. 4, figl 8.
Neogene (Upper Miocene or Pliocene,
Ponder, 1983); Sabah, Dent Peninsula.
Xenophora sokrioidcs jezhri Cox, 1948 (see
Ponder,1983)
A nominal taxon in need of revision.
Miocene; Hanging bridge of Kantzulin,
Kuohsine-hsiaie, Nantdu-hsien, central
Thiwan; eh"tteH'r'tkeng Member o[ the
Shuilikeng Foimatiori(late early Miocene).
'Western
"Roe
Australi a at Xenophora flindersi ludbroohae Ponder,
Plain, Eucla Basin,
Xenophora(Xenophora)flindersi ludbrookae
1983.
Tower ...
Repeater
Microwave
Hampton
l7i,
13d,
Figs
Pcinder,1983i 27-29,
i.
Lowei Pleistoceneand possible Upper
Pliocene", south'Westein Australii.
"Todani,
According to Ponder & Cooper (1983)'
Osa-cho, Atetsu-gun,
Tusurium makiyamaiItoieawa 6c Nishikawa, Miocene,
probably-a Xenophora (Stelkiia).
Okayama Pref. ... lowermost part oflthe
7-8.
igle, M7-t49, Pl. 35,-'figs
upp6r member of Bihoku group", Japan.
"Mekran
Xmophora mehranense (Newton, 1905) (see
Beach, offthe Ormara
Pliocene,
Newton, 1905: 301,
Turu.riummehranense
Ponder, 1983).
Baluchistan", Iran.
Headland,
Pt. r 7, figs8-lo.
Continued.
425
Thb. 4 - Continued.
"
Accordingto Ponder& Coopel( 1983),a
fXenophora)(Tlugurium].) borsonivar.pago- Miocen e, Elueziano:Colli torinesi,
Baldissero(non rara)", Italy. The shell illus- junior synonymof Xenopbora(Onustus)bordaeformriSacco,1896:28, Pl. 4, fig. 5.
soni(Sismonda,1847).
trated is from Colli Torinesi.
var. Parisiensis
lTrochusconchyliophorus)
Grateloup,I 847, ThoquesPl. 1, figr 3-4.
Xenophora
paulinae Nielsen & De Vries,
2002:75, Figs12-16.
"Dax
Miocene; France,
... i St-Paul, fal.
j a u n ., 2 . ' d t . , s u p . "
Accordingto Cossmann& Peyrot,(I 9,19), a
junior synonym of Xenophoraburdigalensis
(Grateloup,I 847).
Miocene; Intertidal platform at Punta Perro, Xenophora?aulinae Nielsen & De Vries,
2002
central Chile; Navidad Formation,
Tortonian.
A nominal speciesbasedon very badly preXenopbora
peroniLocard,1877: 109-110, Pl. Miocene; Corse, Bonifacio.
servedspecimens.
l , f i g s1 2 - 1 3 .
"
Pliocene, Pincenziano: Monte Castello pres- Stelkria plioextensa(Sacco,I 896).
lXenophora)(Tugurium)plioextensum[sic]
so Alessandria (poco frequente)", Italy.
S a c c o1, 8 9 6 :2 7 , P L . 4 ,f i g . 3 .
lXenophora) (Tugurium)p lioitalicum lsicl
Sacco,1896:25, Pl. 3, fig. 6.
um [sic]
fXenophora) (Tugurium)p ostextens
S a c c o1, 8 9 6 :2 6 , P L . 3 ,f i g s8 , 9 , p L . 4 ,
fig. 1.
"
Pliocene,, Piacenziano: Albenga (rara)", Italy. Xenophoraplioitalica Sacco,I 896.
"
Miocen e, Aquitaniano: Colli torinesi,
Langhe, Ceva (frequente). Langhiano: Colli
torinesi (poco frequente). Elueziana: Colli
torinesi, Robella, BarbarescopressoAlba,
Cavesana, Langhe, Mon regalese(frequente) .
Tortoniano inferiore ?: Stazzano (non raro)",
Italy.
'Wang,
Xenopborasanyiensis
1984: 349-350, Pliocene;Sanyi, Miaoli-hsien, northern
Pl. 4, figr. l-3.
Thiwan; Shihiliufen Shale (early Pliocene).
"Helvdtien
(Bassinde la Loire)",
Xenophoradeshayesi
scaldemzi
Glibert, I958: Miocene,
'Anversien
(Sablesd'Anvers)",
24.
France and
probablejunior synonymof Xenophora
(Pantanelli,I 887).
(Stellaria)depressa
A nominal speciesbased on very badly preserved specimens.
Xenophora scaldensis Glibert, 1958 (see
Marque r, 1997) .
Belgium.
XenophorasokriformisTesch, 1920: 73-74,
Pl. 133, fig. 2l4a-c.
Pliocene;Timor (Ponder & Coop.r, 1983).
According to Ponder (1983), a junior synonym of Stellaria solaris (Linnaeus, 1758).
Xenophora
Boettger1882: 70, Pl.
subconica
5, figs l4a-14b.
Miocene; Sumatra (Ponder & Cooper,
According to Ponder & Cooper (1983), a
Xenophora(Xenophora)
sueziAbbas, 1977:
127, Pl. 5, fig. 6.
Xenophora(Tugurium) tatei Harris, 1897:
254-255,Pl. 7, figs7a-b.
"Burdigalian,
Miocene,
Agrud Member,
Agrud", Egypr.
"Eocene:
Victoria and South Australia . . .
from Muddy Creek" (Harris, 1897);
"Muddy
Creek, Victoria (Balcombian,
Miocene)" (Ponder, I 9S3).
"
Miocene, Elueziano: Colli torinesi (poco
Frequente)",Italy.
uar. taulXenophora)ffienophora).) testigerA
rotunita Sacco,1896: 25, Pl. 3 fig. 3.
XenophoraterpstraiDey, 196l: 58-59, Pl. 5,
figs9, 10.
Phorustestigerus
Bronn, l83l : 61.
testudinariumBenoist, I 878:
Otffirra
.)
Xlenophora).textilina Dall, 1892: 361.
XenophoraTrinacriaFischer,1879: 2ll.
I983).
Miocene of Quilon, Kerala (lndia).
Xenophora (Xenophora) .
Xenophora tatei Harris, 1897 (see Ponder,
I 983).
Accordingto Ponder& Cooper.(1983),a
,unror synonym of Stelkria testigera(Bronn,
l83r).
Accordingto Ponder& Cooper( I ?8.3,),1
jto
junior synonym of Xenophorasolarioides
leri Cox, 1848.
Stelhria testigera(Bronn, I 83 I ).
"Cq."
"Cq.
Ba. C. 15".
is for
"Ba.",
"8c",
is for
Castell'arquato,
a slip for
"C"
"Jiingerer
is for
Bacedasco,
Grobkalk,
blau, thonig, in den Apenninen". Following
the present neorype designation, the rype
localiry becomes: Bacedasco.
"Tortonien
Miocene,
.. . Largileyre, situd
A nomen nudum.
dans le bourg de Salles",France.
"Older
Miocene of the Chipola beds",
Florida, US.
Xenophora (Xenophora) .
Pleistocene; Sicily, Ficarazzi.
XenophoraTuricensis
Heer, 1862: 34 (fide
Miocene, Switzerland.
Ponder& Cooper,1983,who tentatively
attributedauthorshipto J.R.A.Moussun),
Kaufmann,1872: 504.
cumukns) var. trAn- Miocene, Loibersdorf (Hornes, I 856),
lXenophora(Xenophora)
Austria.
siensSacco,1896:23, by indication
(Hornes,1856,Pl. 44, fig. l3).
A junior synonym of Xenophora rispa
(Konig, 1825).
A nomen nudum.
426
G. MANGANELLI, V SPADINI,S. CIANFANELLI
tion (Sowerby, 1815) and four subsequent reports
(Brongniart, 1823; Basterot, 1825; Grateloup, 1832,
1838)-of Trochusbenettiae,a British Eocene species
regarded as a junior synonym _of Stellaria? schroeteri
(Cmelin, 1891) by Ponder & Cooper (1983)
D'Orbigny ( 18 52, P. 7) introduced Phorus
grateloupi {or'the shell fio- Saubrigues, figured as
Trochui conchyliophorus var. italica in Grateloup's
Atlas (Grateloup,-1847,toques Pl. l, fig 2).
Tlre .ott.tpbndence berween P desiayesi and T
conchyliophorui var. burdigalensiswas aPparently suPported by Sismonda (1847), D'Orbigny ( 18 52),
Hornes (t S 56), Locard (1877), and Du Boucher
( 18 84-85) who listed fig. I of Ti'oques plate I of
"
"
Grateloup's Atlas as T binettiAe" or T conchyliopho-1
rus" in the synonymy of X. deshayesLSacco ( I 8 96)
was convin..d ol their synony-i and first realized
prioriry of Grateloup'sname, but he did
the apparent -necessary
to make any nomenclatural
not fhînt it
"dop
o mezzosecolo che si usa da tutti
change because
I'appEllativo Deshayesisembra piir opportuno di conservarloche non di sostituirlo con un nome ProPosto
come varieti e mai da alcuno accennato".
Cossmann & Peyrot ( 1919) revised T conchyliophorusvar. burdigalensisand discussedits relationship
io X. deshayesi.TheV first clarified the exact date of
publicatiori of Grat'eloup'sAtlas as 1847 not 1840,
b..r.me one of them [A.P.] had the proofs of the atlas
"Bon
I tirer, L7 avril 1847" hand-written by
with
Grateloup. Consequendy, under these conditions
they would not have hesitated to gil e .p.rioriry to
Michelotti's name if the rwo specieshad been synonyms. In fact, they regarded T ,conchyliophorusvar.
burd,igalensisand P.'desbayesias distinit tixa, identi"les
stries concenfied 5y the charactersof ihe base:
triques de X. Deshayesi
.sont assez rdgulibrement
dcaitdes,quoique moins distantes vers le centre, tandis que lei fines stries subobliques de X. burdigalensis
sont partout plus serrdes, ponctudes et beaucoup
moini profondes; les plis d'alcroissement de I'dchancrure du plafond ddcrivent - chezX. Deshayesi- une
courbe bien plus ample et plus creuse,comm€ngant
plus t6t vers-le centie, devtnant moins rapidement
parallbleau bord pdriphdrique, tandis que les plis de
burdigalensisrestent plus long"te-qt,p.tq.."*i:y{ .
laires dans leur rayonnement et font I la pdriphdrie
.
.f
5
5) from Sowerby(1815, Pl. 98); 6) from Brongniart(182:
Text-figs5-6 - Trochusbenettiae.
7) T c. v3r. burdigalensisfrom GratelouP ( 1847, Tra
Te*t-fifis 7-8 - TrochusconchyQophorus.
tl
var.burdisatensis,
,,,.,,f1?:i:y{i?!kT:;,ltrI"i}J? it), conchytiophorus
rhesheus
rl^r.itaiiro ^|rd its junio, o61.J,ivesynonym Pfiorusgratet
in t*r-fie. 8 "ii conchyliopho'rui
figures-6dified with respeltto the original.
rGl".
, t "
XENOPHORIDS I. ROM M EDITERRANEAN PLIOCENE
un arc plus subit pour devenir parallblesau bord festonnd". They ftgured a shell from Ldognan
(Cossmann 6{ Peyrot, 1919, Pl. 17, figr 105-106)
(Text-fig. 9) to which they referred as the "neorype de
"pldsiorype"(p.
cette espbce"(p. 258), or as the
261).
Accordirg to Cossmann S. Peyrot ( 19 I 9), the shell
reported as T benettiaeby Brongniart (1823, 56, Pl.
6, figr 3a-3b) (Text-fig. 6) belonged to this species.
On t-h. contrary, rhe s['ell reporr.J by Hornes (l 856,
442, Pl . 44, fig. 12) is actually considered ro belong
ro X. deshayesi.Cossmann & Peyrot ( 19 I 9) assigneii
very small (25 mm in diameter and 12 mm in
height), low-spired shells rc X. deshayesi
based on the
"Hoernes,
fact that
dans son Atlas (pl. 44, fig. l2a), a
fait reprdsenterun spdcimen presque aussi surbaisse
que ceux de Mancient, et il lui attribue la m6me
ddnomination Deshayesi".It is evident that they misinterpreted Hcirnes'slegend of plate 44. In fact, fig.
l2.a does not repres_ent
a low-spired X. deshayesi,
but
an apertural view of a shell of X. deshayesi,
tlren illustrated in basalview as 12.b. \What they considered a
low-spired X. deshayesi
is fig. 13, i.e. a shell assigned
by Hornes to Xenopltora cumulans (Brongnlart,
1823).
Cossmann & Peyrot (l9l 9, 265-267) also revised
P g!teh.up!; which ,h.y
9oLfirmed as a distinct
species,similar rc X. tinacria Fischer, 1879, from the
Pliocene of Sicily. They realized that this nominal
taxon was first named by Grateloup (1847) as Trochtts
concltyliophontsvar. italica, but thought it inappropriate to reintroduce this specific epithet. Finally, they
figured a shell of X, grateloupi from Saubrigues (Pl.
13, figr 3-4) (Text-figl l0) to which they re"ferredas
"ndorype"
(p. 261).
Subsequent authors accepted the distinction
benveen X. deshayesiand X. burdigalensis(? Mongin,
1952; Ertinal-Erentciz,1958; Glibert, 1963; Janssen,
1984; Zhu Min-Da, 1984; Rocher, 2002), with the
exceptionof Baldi (1973) and Baluk (1995) who questioned the validiry of X. burdigalensis,becausethe differencesin basesculpture used-by Cossmann & Peyrot
( I 9l 9) to distinguish this speciesfrom X. deshayesi
were
"hardly
"of
relevant" (Baldi, 197?,p.266) and
very loy
importance, and it may result from individual variabiliry" (Baluk, 1995, p. l'79). Apart from Ponder (l 983),
no one seemsto have reported X grateloupi again.
Michelotti (1847) indicated th-atthe mat6rial of P
"Mus.
"Cabin€r",
deshayesi
was in
Soc. Holl." and his
the former denoting the Socidtd Hollandaise des
SciencesI Haarlem [now Hollandsche Maatschapij
'Wetenschappen
der
te Haarlem] and the latter h is
own collection. Today, the Hollandsche Maatschapij
der'Wetenschappente Haarlem no longer has a paleontologic.al .ojf..tion (B. Dortland-Eier, perion"l
communrcation). Michelotti's private collection was
at the Istituto di Geologia of the Universit) di Roma
[now Dipartimento di Scienze della Terra], in part of
the,_buildingwhich was damaged by the bom6ing of
l gtn J.tly lg++ (Fabiani & Maria, 1953). In vihrt
427
remains of the collection, no material of P deshayesi
is
present (R. Manni, personal communication).
Michelotti's concept of P deshayesi
is based on mare"La
rials from
.olline de firin,
les environs de
Bordeaux, la Touraine et la Pologne". He certainly
"La
studied specimensfrom
colline de Tirrin", whereas the other materials (Brongniart, 1823; Dujardin,
1837; Pusch, lB37) -"y only have been cited from
literature. Cossmann 6r Peyrot (l 9 I 9) consideredthe
shell from Loignan figured by Brongniart (1823) ro
.un
belong ro X. burdigalensisand used
calque de la
base du rype de Colli Torinesi (in Sacco,Moll . terz.
P i e m . 1 8 9 6 , -of
p a r t eX X , p . 2 0 , p l . I I , F \ g . 2 0 ) " f o r t h e i r
re-definition
the species,rirrealing that they (and
others, e.g. Glibert, 1963, p. 24) regarded Colli
Torinesi as the true rype localiry of Michelotti's
species. The shell illustrated by Sacco (1 8 96)
(BS.066.01.00l) (Pl. 4, fig. l) has no rype srarus
becausethere is no sure evidence that it belonged ro
the material studied by Michelotti (1847), bu"r it is
the first from Colli Torinesi to be figured and it has
been used as reference for interpreting Michelotti's
species.It belongsto a subadult specimenand is one
oT b.rt preseru.J of the many spicimens from Colli
{,."*,-^{
"rr-,..ti**'.*
Text-fig. 9 - Xenophoraburdigalensis
sensuCossman& Peryrot
(1919)from Ldognan(from Cossmann& Peyrot,
1919, Pl. 17, figs 105-106). Magnificationand
orientationmodiTiedwith respecrr-orhe original.
428
G, MANGANELLI, V SPADINI, S, CIANFANELLI
Torinesi. \7e therefore proPoseit be designatedas the
before doing this, it
of P. d,eshAyesi.'However
leorype
ls necessaryto defii-titinelyascertainwhether ihe shell
figured by Brongniart (i823, Pl ._6,,_figt3a-3b) srill
.firtr "td if so, ,"o "pply to the ICZN io set aside its
rype
' ^ status.
Type material of X. burdigalensisconsists of shells
collelied at Saubrigues and figured by GratelogP
(1847, Ti"oquesPl.-l , fig. 1) (Tbxt-fig. 7) and the
ih.llr illustratedby Sow.iby (1815, Pl. 98, larget figures, from Great iJritain) and Brongniart (1823, Pl.
6, fis. 3a-3b, from Loignan prbs Bor-deaux)(Text-figs
5-n: The figures of thJse shells are very fine, but that
of Gratelou[ does not show aperture shape.The shell
figured by Brongniart (1823) corresPolds vgrl well'
eslpeciallyin aperture shape, to that from Ldgglan
iflLstrat.d by Cossmann & Peyrot (1919, Pl. 17, figs
105-106) (Text-fig. 6), to which they referred as the
"neotype
de cette Espbce"(P.-258)
Tvp. materi al of X. graieloupi consists of material
by GratelouP
colletied at Saubrig.t.I and ftgured
(1847, Ti'oques Fl. I , fig. 2) (Text-fig. S).
Unfortunately, this shell is illustrated in a wly that
obscures"peiture shape.A fine shell from Saubrigles
& Peyrot (1919, Pl. 13,|gt
is figured by Cossm_ann
3-4f (Text-fig. 10), but it seemsto have a lower spire
than that illustrated by Grateloup. Again, Cossmann
& Peyrot (1919) referred to this specimen as the
"neorype
de cette espbce"(P. 258).
V. do not know whether rhe synrypes of X. burdigalensis and X. grateloupi still exist. Once their loss
is?efinitively "scJttained, we proPosedesign"ilg the
shell illustrated by Cossmann & Peyrol (1 919).in figs
105-106 of plate'17 as the neorype of X. burdigale7szsand that lllustrated in figt 3-4 of plate 13 as the
neorype of X. gratelouPi.- In fact, Cossmann 6.
"ConchoTogie
ndogdnique de I Aquitaine" is a
P.yrot's
-il.rtone of Eurofean NEogene molluscan p{qontology. It has al*ays been witely consulted atid their
t..'ifibn of the Miocene xenophorids has been a basis
for subsequent understandi"g of these nominal taxa.
Consequendy, when a neoryPe-becomes necessztft
one should consider choosing it from their collections
or at least in line with their species co!c€Pt.
Cossmann & Peyrot ( 1919) referredto some of these
"neotypes",
though not in the sense
specimens as
Nomenclature.
of
Code
the
by
rbquired
'Only
dLsignation of the ryPesJan oPe" tb.-yay to
^ modern ievision of the European Miocene
xenophorids enabling clarifications of th. status of
the nominal taxa deslribed till now, the status of the
many different forms usually assigned to X.. deshayest^
and tn. geographic and stritigt"ihi. distribution of
ngI0 (r;rri'
rext"? ":il:T
h3:::n:1
i:"#{:!::(:"i'#
andorienta1919,Pl. 13,figs-3-4).Masnification
to the
tion of somefifuresmodifiedwith resPect
original.
the valid species.Many 9f the &t-Tt.assigned to X.
deshayesidiffer remarkably in shell shape and size,
sculpiure an4 aperture shape and size. For .IlTPlg'
from Colli Torinesi (Pl. 4,
specrmensof X. d.eshayesi
fier l-3) and those from Vienna Basin (Pl. 4, fig. 6;
r.". also Hornes, 1856, Pl. 44, fig. lz.b) have a different structure of the peristomal Sasalmargin. In the
Pliocene specimens,suth differencesare constant and
distinguish different species.
Conclusiue remarks on Xenophora davolii n. sp.
For the present, accePtitg
- these three nominal
species: X. burdigaleisis, X. d'eshayesi, and X.
irattloupi, in the ab6rresense,X. dauolii n. sP. is again
listincf fro- them by larger size, smooth dorsal sur-
EXPLANATION OF PLATE3
dalolii n. sp. from the Middle Plioceneof TerreRosse (Castelnuovo Berardenga) (FiS. l, G. Manganelli collection;
Figs
v l-3 Xenophora_
Figs'2-3,V. Spadinicollection).
G. MAA.GAAIELI,I, V(SPADIIVI,S. CIAIVFATVELI],XE,NOPHORIDS FROM ME,DITE,
G, MANGANELLI, V SPADINI, S. CIANFANELLI
430
face with very irregular and prominent rqgae and its
DroDortionally larger aperture (at leastwith respectto
'X.
deshayesiind X. bLrdigalensis), horyever itJ relationship-to the others remiins obscure. It may be one
of the l"tt offshoots, together with X. infundibulum
deshayesigroup of forms
and X. plioitalica, of thjX
"rather
sensuPonder (1983), a group chaiacreiizedby
large, evenly conical ihelli, with narrow., simple
p.iipheral flanges and, in most but not all species,
ieduced areasoT agglutination" and includitg several
speciesfrom the Eocene to the Pliocene (X. wemmelensisGlibert, 1938, X. PetroPhoraKoenen, 1892, X.
X. burdigalensis, X.-groteloupi, X. infundibud.eshayesi,
lum, and X. plioitalica).
XENopHoRA INFUNDIBULUM(Brocchi, I 8 14)
Pl. 5, figsl-4, Pl. 6, figt l-3
I8l4 Trochusinfundibulum BnoccHI, pp. 352-353,Pl. 5, fig- 17;
Reprinr .if t 843, pp. 132-133,P[. 5, fig. 17.
Typt material- A syltype from San Geminiano
[sic]-isin the Brocchicollectionat the Museo Civico
di Storia Naturale di Milano, no. 4929 (seeRossi
Roncheffi,1955: 159-161,Fig. 81; Pinna & Spezia,
1978:160,PI.40, fig. 1). RossiRonchetti(1955)and
Pinna 6( Spezia(1178) attributed to this specimen
the statusoTa holorype,but this is mistaken.In fact,
Brocchi ( I 814) basedhis descriptionmore than one
three dilferent localities)and
specimen(at leastfrom"rypr".
any
not
designate
fid
Typt localiry- Brocchi ( 18 14) cited three localities:?iacentino, SanGeminiano [sic] and Libiano.
Diagnosis- A speciesof Xenophgra(t.1.)-charactersize(i), robusttrochiform shell(?), with
izedby"large
narro% sli[htly wag sharplykegledperipheralfl.ange
(3); spire*itlr 7-9 flat or very slighdyconvexwhorls
(4); 90o/oof spire surfacefree of-attachedobjectsor
their scars(5)l with fine closevermiculateopisthocline riblets intersectedby Prosoclinegrowth lines
(6); attachedobjects(mainli bi't"lve she-llsand fragments thereof) alwaysvery small or small excePtin
lastquarterof lastwhorl (7); scarsof attachedobjects
trt,t"ily deep(8); basefunnel-shaped(9); basalsculp-
EXPLANAIION
ture consistingof collabralgrowth lines,weak o1vfY
weak spiral lines and somEtimesvery thin opisthocline riblets (10); basalmargin of peristomestraight
and thickened up to periphetY, then absent and
replacedby peripheralf["ttgi; where basalmargin of
peristomeattachesto perip[eral flange,there is often
^"
(12).
ron of sinulus(l l);l-biticus abse-nt
Descrintion- Shell (Pl. 5, figs l-4, Pl- 6, figs l-3)
dextral,l,ery largein size,robusl and trochiform with
pointed apex ind narrow slightly yary, sharply
keeled peripheral flange; spire conical (spire algle
about 8b'; i"rg.r 70":85o)^withabofi 7-9 regulaily
srowing whorli; last whorl large,dilated only in final
6,r"tt.i(shell outline oval or diop-like in dorsalview:
Pl. 5, fig. l, ll.-6, figs l-3)i ttiwhorls alwayse.roded; lastivhorls flat oivery slightly convex'sometimes
last two whorls very slightli concave before lower
suture or periphery; suttires'shallow;about 90o/oof
spire surfate ft.. of attachedobjects or their scars,
*itn fine closevermiculateopisthoclineribletsintersectedby prosoclinegrow*t lines; attaclredobjects
alwaysy;ty small,.4.."pt in last quarte.rof lastwhorl,
consrsungmainly (about 90o/o)oTsmall bivalveshells
and smafffraemints of bivalveshells,and secondarily
(about l}o/of of balanid cirriped plates, small fragments of gastropodshells,tubesof s.tpulid anneliils
and smalllebbles; foreign objects
-sizealwaysattph.e{ by
of attachedobjects
convexsurface;numbei and
in first whorls
so
that
whorls,
all
on
constant
rather
they are closerand thereis propoglonally lesssurface
free; larsestobiectsconsistineôffragmentsof shells
attached
and *h6le shel'lsof bivalvesind gast-ropods
(large-st
is a
found
quarter
obibct
whorl
last
of
in last
Natica ihefl 18 mm in diameter)(for other detailson
attachedobiectsseeThb. 5); scarsof attachedobjects
usually deep; base funnel-shapedwith collabral
growth lines more marked near aperture'sometimes
ileak spiral lines and very weak opisthocline riblets
crossingof spiral
nearpe'riphery (in somespecimens,
linesind-opisthoclineribl-etsgivesbaseedgea microgranul"t "p'pearance);
aperturevery largea-ttdoblique
margin is behind by about
basal
innermost
b..",rt.
half a whorl; peristomeinterlupted with uPPeran4
lower vertices-unitedbv thick porcellanousparietd
callus; columellar margin of piristome covered by
OF PLATE 4
(Micheloffi, 1847) from Colli Torinesi (Bellardi-Saccocollt
Figs l-3 - pnophora dcshayesi
Tirrin, Italy:
figuredby Sacco(1896: Y\.2, fig..20)and hereinproposed
1) BS.066'.01.001,
srrong spiral groovesin Saccir'sfigttte are a photographicartefact;
.004;
2) 85-.066.001
Figs
- 4-j
Fie.
a 6
3) BS.o66.ol.oo9.
f*o xenophorid-shellsfrom Gainfahren (Austria)..NaturhistorischesMuseum
sitesare usuallyassigned
orher Paratethydean
I Aie I lf,e. Sn.Ur from this and"fu,igiana"
(Beilardi-Saccocollecdon, Mirseo
Tile shell from the Plioceneof
85.066.01.001)assignedby Sacco(18:96)to X. deshayui(Michelotti, 1847).
G, MATVGAIVELLI,V SPADIIVI,S, CIAIVFAIVELLI,XENOPHORIDS FROM ME,DITE,
Pl. 4
432
G. MANGANELLI, V,SPADINI, S, CANFANELLI
parietalcallus;inner basalmargin straight,radialand
thickenedup to periphery;outer basalmargin usually absentand replacedby wa{, knobbed (due to large
objects attachedon dorsal surface)lower surfaceof
peripheralflange (only exceptionallythere is a thin
outer basalmargin); externalmargin thin and sharp;
where inner basal margin peristome attaches to
peripheral flange there Ir oft.n a sort of sinulus;
l-Eiticus abserit(alsoi. juvenileshells).
Dimensions Diameter: 66-116 mm; height:
5I-84 mm; spire angle: 70"-85" (fot more detlils,
seeThb. 5). Dimensionsof other Pliocenespecimens:
D: 54.0-115.0 IDIn, H: 30.0-65.0 mm (Sacco,
1896); D: 50.0-120.0 mm, H: 30.0-70.0 mm
(Caprotti, 1967);D: 50.4-120.0trln, H: 30.0-70.0
mm (Malatesta,1974); D: 65.0-136.0 lrrlrr, H:
46.0-83.0 mm (Zhu Min-Da, 1984).
Material - SienaBasin: 48 shellsfrom Barca ( I ,
GMC; 3, VSC), Guistrigona(1, GMC; 15, VSC)
and TerreRosse(28, VSC).
In addition to Sienesematerialwe studied material from the Plioceneof Colli Astesi,Piedmont (5,
MRSN BS.066.01.012and BS.066.01.013)and
Lucenadel Puerto,Huelva (1, MMAS).
X.
Geographicand stratigraphic distribution
infundlbulum had a widE distriburion around the
Mediterranean in the Miocene and Pliocere,
although the Miocene records,often based,or !:^gmentary specimens,requirerevisionas for identification and stratigraphicsefting.The specieshas been
"Helvdtien"
"Pontildvien"
of
reported from the
or
France(Salles;Cossmann6( Peyrot, l9l9; Glibert,
"Tortonian"
1963), the
of Portugal(Cacella;Dollfus
et al., 1903, Cossmann6( Peyrot, 1919) and Italy
(Benestare,Casa Nova di Calisese,Modenese,
Seguenza,l88l; Sacco,1896; Caprotti,_
Srazzema;
"sahdli;n"
of
1970; Ruggie-ri6c Davoli, 1984), the
Tunisia (SiIhepinsky,1938) and Morocco (Dar-belHamri; Gentil, l9l9; Chavan, 1940; Lecointre,
"Miocene"
1952; Glibert, 1963), the
of Sardinia
(Cagliari;ComaschiCaria, 1959)and the Plioceneof
Spain (Estepona,Huelva, Malaga, Millas; Andrds,
1980; Zhu Min-Da, 1984; Castafioet /t1., 1988;
GonzalesDelgado,1988;Santoset a1.,2003;Landau
(Bir Touta, Oued Nador, Sidiet A1.,2004),"A,lgeria
Moussa; De Lamothe 6c Dautzenberg, I 907;
Cossmann, 1916), France (la Thinitd; Risso, 1826),
Italy (Piedmont, Liguria, Emilia Romagtrx,Tuscany,
Umbria; Brocchi, l8l4; Seguenza,1877; Sacco,
1896, 1904; Ruggieri, 1962; Glibert, 1963; Palla,
1967;Caprotti, 1967, 1970, 1974;Malatesta,1974;
Pavia,1976;Montefameglioet dL.,1980;Giannelli et
dl., l98l; Chirli, 1988; Cavdlo 6c Repetto, 1992;
Andreoli & Marsisli, 1996;Vera-Peliezet al., 1996;
Forli et dl., 2003), Syria and Lebanon (Haffii,
Mandjila, Amerko, Nahr Snobar, Cheikh I(halil,
Quemine, Kherbet el Katrid, Tannechmaniyb;
Roman, 1940; Zhu Min-Da, 1984). Malatesta
(1974) reportedit from the Plioceneof Sicily,probably by error (asfar aswe know, the specieshas never
beenfound from the Sicilian Pliocene).
X. infandibulum is the most common and widespreadienophorid speciesin the Lower and Middle
Plioceneof ihe Sieni Basinwhere it was first reporred by De Stefaniand Pantanelli( 1873) and recendy
by Forli et al. (2003). It has also been found in the
Plioceneof the Val di Chiana (Cittl dellaPieve;Verri,
1886) and Radicofani Basins (Poggio Rotondo;
Malatesta,1974).
Remarks- Despiteits abundanceand diffirsion in
the Italian Pliocene,descriptionsand illustrationsof
X. infundibulum are limited (.f. Sacco,1896, -Rossi
p. 23,
Pl. i, fig. 26; Sacco, 1904, Pl. 26, fig. 9;
Ronchetti,1955,pp. 159-161,Fig. 8 I ; Palla,1967,
pp. 956-957, Pl. 7l, figt 9a-9b;Caprotti, 1967, pp.
189-190; Caprotti, 1970, Pl. 7 , fig. 5; Malatesta,
1974,p.21 5;Pl. 16, figs8a-8b;ZhuMin-Da, 1984,
pp. 22:23, Pl. 3, fig. 1; earralloScR.perro, 1992,Fig.
109;Andreoli & Marsigli, 1996,Pl. l, figs4-6). This
is perhapsdue to the fict that the speciei,one of the
largestgqr,rqpodsof the Mediterr"h."n Pliocene,is
easyto iden:'ify and completespecimensare rare.
The Siena Basin material consistsof very fine
specimens(Pl. 5, figs l-4, Pl. 6, figr l-3) that enabled
detailed study whiJh revealedthai some of the more
peculiarcharactersof this speciesare new: last whorl
dilated in final quarter;orrafor drop-like shelloutline
in dorsalview; largeobjectsattachedto terminal portion of last whorl inner basalmargin of peristome
straight and radial as far as peripliery; ont.r basal
-arfitt of peristome absent and repliced by lower
r,trfi.. of peripheral flange; frequent presenceof a
peristome
sort of sinulus where basal mlrgin
"ovalof ^drop-like
or
attachesto peripherd flange. The
shell outline in dorsalview is evidentin the shell figured by Cavallo 6c Repetto (1992, Fig. 109), large
objects(or their scars)attachedto the terminal portion of the last whorl are evident in the shellsfigured
bv Pall" ( 1967,Pl.7l, fig. 9b), Caprotti (1970,-P1.7,
fis. 5), Chirli (1988, PL 2, fig. 3) and Cavallo 6c
REpetto(1992, Fig. 109) and the sinulusis alsoevident in a specimen from Colli Astesi
EXPIANATION OF PIATE 5
(Castelnuovo
Berardenga).
V. SpadinicolFigs l-4 - {englhora infundibulum (Brocchi, 1814)from theMiddlePliocene
of TerreRosse
lecuon.
XENOPHORIDS FROM ME,DITE,
G. MAIVGAIVELLI, V. SPADIIVI,S. CIAIVFATVELLI,
Pt.5
G. IUI'ANGANELLI,V SPADINI, S, CANFANELLI
434
Attached objects
Localides
Diameter
(mm)
Height
(mm)
Anele
(degiees)
number of
attached
objects in
last whorl
number of
attached
objects in
penultimate
whorl
number of
attached
objectsin
andpenulti-mate whorl
Barca
n:4
88- 112
n:3
60-82
n:3
78-80
n:2
7-15
n:2
6-8
n:2
12-15
n:2
Shellsand frasments of shellsof
bivalves (Acaithocardia, ?
DiphdontA, ?Arca, Chameha
gailina, ? EnsislPhaxas,Gari,
Gkns interrnedia, ? Pandnra,
Timoclzaoaata and others
unidentified), platesof cirripeds
and small pebbles.
64x8
bivdves (Acaithocardia,
Chamehasallina, Chkmys,
Ckusinelli, Corbuk sibba,
Dosinia, Gari, Gknslnturrnedia,
Glycymeris,Nucuk pkcentina,
Nuiukna pelk, Osirea, Timoclea
oaataand bthers unidentified),
fragmentsof shellsof gastropods
(Sniorcrebram and others
unidentified), platesof cirripeds
and smdl pebtiles.
bivdves (Acaithocardia, ?
Amusium cristatiltlt, Anadara
diluuii, ? Diplodonta,
Chameltarillina, Chlamys,
Corbula g;bba, Dosinia, Ghns
intermef,ia, Glyc'ymeris,Nucuk
pkcentina, Nicithna commutntA,
'Ostrea,
Paphia, Pecten,Tbllina,
Timocha iaata and others
unidentified), shellsand fr^gments of shellsof gastropo&
(Nassarius,Nati ca)i fragrirents of
shellsof scaphopods
(DentaliamJ,platesof cirripeds,
tubes of serpulid annelids
(Ditrupa) eid small pebbles.
Guistrigona
n: 16
Terre Rosse
n:28
(67- rr2)
n: 13
(5r - 76)
n: 13
77.8x 2.4
(75- 80)
n: 13
10.9t 2.2
(7 - 13)
n:9
12.7x 3.7
(8- le)
n:9
14.2t 2.9
( 1 1- 1 9 )
n:9
93x14
(66- n6)
n:23
98t10
(5r - 84)
n:23
77.5x 3.4
(70- 85)
n:21
8 . 5t 3 . 1
(l - 16)
n:22
8.4* 3.9
(r - 17)
13.2x 3.2
(7- ts)
n:22
n: 18
89t13
Thb. 5 - Shell parametersof Xenophorainfundibulum (Brocchi, l8 14).
(BS.066.01. 012102). Specimensfrom the Upper
Miocene of Salles (France) (Cossmann 6( Peyrot,
1919, Pl. 13, fig. l) and Cacella(Portugal)(Dollfus
et AI., 1903,PL.32,fig. 1) and the Plioceie of Huelva
(GonzalesDelgado,1988, Pl. 3, fig. 3; Santoset Al.,
2003,PL.2, figs l-2,5) and Estepona(Landauet Al.,
2004, Pl. 19: fig. 2b) differ from those from the
Itdian Pliocene5y the absenceof the sinulus;moreover thosefrom Upper Miocene of Sallesand Cacella
E)(PIANATION
Figs I-3 -
{engphora infandibulum
lecuon.
alsodiffer by havitg an evident thin outer basalmar"Ndogbne
de Karaman,d'un
gin. Specimensfrom the
by
niveau plut6t Helvdtien", assignedto X deshayesi
Eriinal-Erentoz(1958, Pl. 3, figt 24-24a) are indistinguishablefrom the Upper Miocene forms of X.
infandibulum such asthosefrom Salles(Cossmann&
Peyrot,1919,Pl. 13, fig. l).
Sacco(1896, p. 24) claimedthat specimensfrom
"Astigiana"
have a wider shell with a lessstreamthe
OF PIATE 6
(Brocchi, l8l4) from the Middle PlioceneofTerre Ros
XENOPHORIDS FROM MEDITE
G, MATVGAIVELLI,V SPADIIV, S, CIAIVFATVELLI,
P l .6
436
G, MANGANELLI, V SPADINI, S, CIANFANELLI
lined spirethan that figuredby Brocchi ( 1S14, Pl. 5,
fig. 17) and those froilr Tirsciny examined by him.
H? thereforesussestedit be assignedto a distinct
This form was
variery:X. infuntibulumv?r. expan"sior.
"Helvdtien"
or
subsequentlyreported from- the
"Pontildvient'
of Salles(Cossmann6{ Peyrot, l9l9;
Glibert, 1963).The spire angle in the Tuscanspecimens examined ransdsfrorn- 70" to 85" and shells
collection
from Colli futesi kep"tin the Bellardi-Sacco
(BS.066.01
.013101,02)
.012101,02, 03, 85.066.01
havesimilar angles.
X. infundibiltu* is one of the most characterized
xenophtirids.The flat whorls, vev large aperture,
straieht and radial inner basal peristoire ind the
absetceof outer basal^peristomedistineuishit from
species.X. deihayesisensu
all other xenophorid
^Pl.
44, ftg. l2.b) has ap6rtureand
Hcirnes(1856,
peristome reminiscent of.-X. infundibulum, but the
aperturels smallerand a thin outer basalperistomeis
pr.r.trt (see also Pl. 4, fig. 5). Other materials
(Pl. 4, fig. 4; Kautslcy,1925,
issigned to X deshayesi
Pl. 6, fis. 9; Gliberi, 1949, Pl. 12, fig. 20) or to x.
(fansser,1984,Pl. 53, fi{,5) haverather
burd.igalensis
flat wJrorlsand tracesof opisthoclineiiblets but differ
by rather large bodies attached to the sutures.X.
grnuloupi r.ti,r Cossmann6( Peyrot (1919, Pl. 13,
Egr 3-4) (Text-fig. l0) hasflat whorl shape,a modest
and a
aitount of attachtd obiects,opisthocline-riblets
larse aperture like X.' infundibulum, but the inner
margin seemsslig6tly concaveand the outer
bas"'al
basalmarein is evident,though thin.
Pondei (1983) listed X." infundibulum and X.
flemingi Beu, 1977, from ihe New Zealand
Lower-Middle Miocene,in an informal, morpholog"species
ic group includine
[ofl medium to largesize,
wiih medium td tafl spire, straight whorl o-utline,
rather narrow peripheial flange-with only small
objectsattached,ttoh-nmbilicatE".These speciesare
noi at all related:X. flemingiis similar to X. infondibulum in whorl shapeand sJulpturebut it is completely
different in aperture and peristome (for X. fb*ingi,
, i g s2 , 5 , 7 , 8 , 1 0 , 1 3 s e eB e u ,1 9 7 7 ,p p . 2 3 3 - 2 3 5F
17).In fact, Ponder(1983) regaidedX. infundibulum
asa speciesof the X deshayesilroupo{ forms,
Cbssmann& Peyrot(1919) regaidedX i"f"ndibuX.
X. Turdisalensis
and zt.
X.
ourdtgaEnsts ano
n A.
Detween
intermediati
intermediate betwee
between
as lntermedlate
lum
lumI as
"sa
baseplr,rsiqfr+di6uliforme".and
due to
deshayesi
"les
itries d'accroissemeni du plafond qui sont bien
longtemps rectilignes, plus subitement arqudesvers la
pdriphdr'ie", but tritr r.int from these two species and
"sa
base
all Eocene and Olieocene forms due to
creusde". X. gratelnipi sensu Cossmann 6c Peyrot
(1919) is the frott simitar speciesto X. infundiQulum,
indeed so similar that differences warrant further
study. X. {rAtehupi sensu Cossmann 6c Peyrot (1919)
and'X. inTundibitum may have descended from forms
with rather flat whorls' and ffaces of opisthocline
riblets, usually assigned to X. deshayesi.
I 896
XENop"o*ni:t;tffiIcArSacco,
1896 lXenophoral(Tugurium) plioitalicum Stcco, p. 25, Pl. 3,
fig.6.
Tvpematerial- Only the holorype(85.066.02.002)
coliectionat the Museo
whicli is in the Bellardi-Sacco
Regionaledi ScienzeNaturali of Tirrin (Pl. 6, fig. l).
Th; holorypehasalsobeenfiguredby FerreroMortara
et Al. (1984,PL.43, figr 2a-2$.
"
Piacenziano:Albenga" Liguria,
Tvpe localin
Italyl'Accordirig to Ferrero Mortara et-Al. ( t-g84):
Lower Pliocene.
1896 lXenophora) (\uguriuml.)
SRcco, pp. 25-26, Pl. 3, Fii.7.
plioitalicum var. gracilior
specimen
The
original
Material
(85.066.02.003) is in the Bellardilsaccocolftction at
the Museo Regionaledi ScienzeNaturali ofTurin.
Locality-"PiAcenziano:Rio del Molino di Galatea
presso Chieri (rara)", Italy. Accorditg to Ferrero
Mortara et Al. (1984): Lower Pliocene.
Diawtosis- A speciesof XenoDhoracharacterized
by larg8size(l), ,obtrmtrochiforni shell (2), with narrow, ilreeular, rouehlv round (whereattachedobiects
absent)i.tipheral-flange (3); spirewith 6-7 (?)flit or
slighdy tonrrex whorls-(4); most of spire surfacefree
of"attachedobjectsor their scars(5)iwith wary vermiculate opisthocline riblets (6); attached objects
(mainly.bir'.4yeand gasffopodshells)small to medium rn srze(7); scarsof attiched objectsusuallydeep
(S); baseslightly concave(9); basalsculptureconsistine of venifaiirt collabral growth lines, many very
tni"tt spiral lines and .'rr..v"thin wa\ry opistlroclin-e
riblets (t O); apertureauriculatewith basalmargin of
peristomeJ-shaped (l l); umbilicus small and partially coveredblcolumellar margin (12).
EXPTANATION OF PTATE 7
Fiss
v l-4 Xenophoraplioitalica Sacco,1896.
l) Holo*p.r Bellardi-Saccocollection,Museo Regionaledi ScienzeNaturdi, f
Albenga;
2-3) Quarantelli collecdon, Museo del Mare Antico, Salsomaggiore,Italy. Upp,
4) G. M"ttg*elli collection. Two fragments(umbilicus and o,it".t basal-argin .
Lower Plioceneof Monsindoli (Siena).
G. MATVGATVELLI,V SPADIIV, S. CIAIVFAIVELLI,XENOPHORIDS FROM ME,DITE,
P l .7
438
V.SPADINI, S, CUNFANELLI
G. A,T/4NGANELLI,
Description- Shell (Pl. 7 , figs 1-3) dextral, largein
size,robust, trochiform, with pointed apex and-narroughly scallopedand round (where
row irregular,
-ob;ects
abient) peripheralflange;spireconiattached
cal (spireangleabout 85oto 100")with about 6-7 (?)
resulirly er6wine whorls; last whorl large, but not
diLted;'lait whofls flat or very slightly jorrrex with
warty and wrinkled appearance;sutures shallow;
-ori of spiresurfacefreibf "ttached obiectsor their
scars,with numerouswavy vermiculate-opisthocline
ribletsintersectedby faint or more evidenfprosocline
growth lines; opisthocline riblets very small in first
i'horls; attached objects small to rnedium in size,
consistingof shellsand fragmentsof bivalve(Nucuk
and otheis unidentified)antr gastropodshells,naticid
gastropodopercula (Natica),-foraminifersand smdl
[ebbl6s; attiched objectssmall, numerous and very
Llor. to eachother in the first whorls; largerand *.ll
spacedin the lastwhorls (12 in the penultlmatewhorl
ahd 11 in the last whorl of th-e holorype of T
plioitalicum); scarsof attachedobjectsusuallymoderltely deep to deep; baseslightly concavewith very
fine faint collabral growth lines (more evident near
aperture),many very small spiral lines (more evident
near centreof base),and veri thin wary opisthocline
riblets (more evident near periphery);apertureauriculate,largeand oblique becauseinnermostbasalmargin is behind by alrirosta quarter of a whorl; peristome apparentlyalmost continuous,with upper and
lower verticesunited by thick porcellanousparietal
callus; columellar margin reflected and thickened;
basalmarginJ-shaped,ihickenedand initidly reflected; outer basalmargin proportionally large;external
margin slightly thickened (sometimesnon porcellanous bur with collabral growth lines); umbilicus
small and partially covered5y peristomalmargin.
Dimensions- The best specimen(holorype of X.
plioitalica) is ll4 mm in diameter and 63 mm in
height.
Material - Siena Basin: three very fragmentary
from Monsindoli (GMC).
specimens
In addition to Sienesematerial, we studied four
specimens, two from the Pliocene of Albenga
(MnSN 85.066.02.002,holorype of X. plioitalicuil)
and Rio del Molino di Galatea (MRSN
85.066.02.003, original specimenof X. plioitalicum
var.gracilior) and twl froni the Upper Mi6cene of the
Torftnte Stirone (Q,t"rantelli collettion, MIvIAS).
X.
Geographicand stratigraphic distribution
plioitalica'was originally rJcordedfrom two localities
iAlb.tre" and Ri; dei Mulino di Galatea presso
Chieril of the Lower Pliocene of Piedmon-t and
Liguria (Sacco,1896). It was subsequendyreported
"Middle
Miocene" of Moirtebelio (irmilia
frJm the
"[Jpper
Romagna) by Moroni ( 1958) and the
Miocene" of Dar-bel-Hamri (Morocco) by Gentil
(1919), but both reportsmust be consideredwith
prudence.The specimensexaminedby us are from
^th.
Upper Miocine of the Torrente Siirone and the
Plioceneof Siena.
The speciesis new for the SienaBasinwhere it is
only known from the Lower Plioceneof Monsindoli
quarry.
'We
Remarks
assignedsome very fragmentary
and crushed specimensfound in the Monsindoli
The most complete
quarp nearSiena,to this species.
speclmenrs a large shell with an estimateddiameter
oT about l0 cm-(Pl. 7 , fig. 4). Its size and dorsd
X. infundibulum (Brocchi,
opisthoclineriblets recarll
l-814), but it has an umbilicui (umbilicusabsentin
X. infandibulum), the free peripheralflange is rough-X
ly slalloped and round- (sharply keJled in
infundibulum), the outer basal peristome is well
developed (outer basal peristome absent in X.
infunrtbuhm), around unibilicus there are distinct,
nirrow spiral lines (spiral lines at the centre of base
very faini in X. infundibulum) and below the peripheral flanse there are very thin opisthocline-riblets
(thin oplsthocline riblets below peripheral flange
absentor barelyevidentin X. infundibilum).
Cossmann(tg16), Ponder& Cooper(1983)and
Ferrero Mortara et al. (1984) regardedthe names
"plioitalicum
"plioitalicum"
varl gracilior" as origand
bIt
iirally introducedin combinationwiih Tugurit4tn,
Sacco( 1896) regardedTuguriumto be a iubgenusof
XenopltorA.Consequendyrthe original combination
uXen-ophora
(Tugurium)
of thesetwo nominal tora is
"
(Tuflirium)
plioitalicum"
plioltalicum
Xenophora
and
-nar.
gracilioi' despitethe fact thit Saccolisted them
"
equiiocally in the text as Tuguritm plioitalicum" and
" plioitaiicum
var.gracilioi'"and,rr6d thesenamesin
T
thelegendof the pla-te(this equivocaltreatmentis not
exclus-iveto thii genus, but the rule throughout
Sacco'stext wheneverthere are subgenera).
The orieinal
-onlv material of X. plioitalica probably
consistsof
one specimen(whictr .ottsiquendy
has the statusof aholotype), like that of X. plioitaliia
var. gracilior. Although Saccodoes not comment on
this,-inthe caseof thJformer he only gavethe dimensionsof one specimenand in the caseof the latter it
is evident that the description
^"forseis basedon a single
b un esemplare
specimenwhen he writes
-of tt6t
these
iompletamente adulto.. .". Examination
specimens,orieinally in the Audenino collection but
rb* in the Billardi-Saccocollection, enabledus to
ascertainthat thesetwo tilra are synonymsand that
the original specimenof X. plioitalica var. gracilior is
indeeda iuvenilespecimen.
X. plioitalica ii one of the best distinguished
EuropeanNeogenexenophorids(no fossil or extant
hasa siinilar clearlyauriculateaperXenophoraspecles
ture'and ttôlarge-sized fossil or .*t"rrt Xenoplrorn
specieshasan uribilicus), but it is alsoa rarespecies,
sb far known from few specimens. It recalls X.
XENOPHORIDSFROM MEDITERMNHN
PLIOCENE
439
(Sacco,1896).
Text-figsl1-13 - Stellariacf.plioextensa
Italy.
1l) ComplCteshell,Quarantellicollection,Museo del Mare Antico, Salsomaggiore,
Salsomaggiore;
12) Fragmentsof dorsalportion of a shell,G. Manganellicollection.Middle Plioceneof ,
l3) Fragmentof basd portion of a shell,V. Spadinicollection.Middle Plioceneof Forna<
infundibulum in some features (spire shape, few
attached objects and vermiculate opisthocline
riblets), but it is distinct from it in many other characters (aperture; umbilicus; basal margin of peristome; basal sculpture; peripheral flange).
Ponder (1983, p. 14) surmised that X. plioitalica
"perhaps
was an offshoot of the X deshayesigroup,
from a form close to X. grateloupi (D'Orbigny,
1852)" . X. grateloupi (in the sense of Cossmann 6(
Peyrot, 1919) sharesvermiculate opisthocline riblets
with X. plioitalica, but differs in shape and size of the
aperture and absenceof umbilicus. As stated above,
becauseit is very similar to the Mioce ne X. infundibulum it is unlikeiy to be related rc X. plioitalica.
Outside the Mediterranean are" the only material
somewhat reminiscent of X. plioitalica is an incomplete shell from the Miocene of \Tinterswijk-Miste
(The Netherlands), published as X. deshayesiby
Janssen(1984, Pl. 54, fig. 1) on the basisof what
seemedto be a small umbilicus.
Moller, 1832
GenusSrEI-I-nruA
Typ. speciesTiochussokris Linnaeus, 1767, by
monorypy.
(Sacco,1896)
cfl plroEXTENSn
SrEI-uq.ruA
Text-figs1l-13
(Tugurium)
1896 lXenophora)
plioextensum
SRCCo,
p. 27, Pl. 4,
figs3-3a.
No q?e material exists in the
Tlpt mateial
Bellardi-Sacco collection at the Museo Regionale di
ScienzeNaturali ofTirrin (Ferrero Mortara et Al., 1984).
440
G, AIANGANELLI, V.SPADINI, S, CANFANELLI
"
Typelocality - Piacenziano: Monte Castello pres(poco
frequente)", Italy.
to Al6ssandria
Diagnosis- A speciesof Stelkria (Onustus),chg;
acteriz.?by largesize(l), fragileand almostdiscoidal
shell (2), with-large thin pJripheralflange, porcellanousbelow (3); ipire with aSout 7 flatbr Jlighdy
convex whorls (4); most of spire surface free of
attachedobjectsor their scars(5)l with very small delicate wavF vermiculate opisthocline riblets (6)i
attachedobiects(shellsand hagmentsof bivalveand
gastropodshells)smallto medium in size(7); scarsof
Ittachbd objectsvery shallowalong suture,but deep;
er along p.iipheral h*.g. (8); basEconvex(9); baial
sculpt"ie consistinsof iobust collabralgrowth lines
and^verythin spirallines (10); basalm"r-giqof peristome curved ind initially thickened and reflexed
(l l); umbilicusround and deep(12).
dextral,.large
. Description Shell.(Text-figl.12-l?).
with pointed
rn srze,very fragileand almost-discoidal
apex and ittg.-thin
-conicalperipheral flange, porcellanous
depiessed(spire "lgl-. obtuse)
belo*; spire
regularly
with abolt seven
srowing whorls; lastwhorl
large,sometimess[[htly,dil,at.d; #horls flat or slig^htly -onvex; suturesvery shallow;most of spire surface
ft.. of attachedobjecisor their scars,with- vgrysmall,
delicatewavy vermiculateopisthocline riblets intersected by weak prosocline growth lines and some
prosoclinerugae;attachedoFjectsonly along sutures
bf lastwhorls,widely spaced,smallto medium in size
and consistinsof frie-ettts of bivalveshells(a specimen from Pli6ce.. oTCamporedi Salsomaggiorihas
gastropod
"contifua, shells;Suehia.fura.ta,GemmularotAta, G.
Stenodrilla belkrdii, Turricuk dimidiata,
etc.;-Iext-fig. ll) amachedalong peripheralflange;
scarsof attiched objects usually shallow (deeperin
the specimen from Pliocene of Campore di
baserobust,convexwith robust colSalsomaggiore);
labral gr6foth and very thin wary spiral lines; growth
lines Jrigitr"r. from umbilicus'*li.r. they "i. disposed diigonally, then collabrally crossbase,terminating ab"ruptly,slightly raised, at junction with
peripKeralflange;,"pJtt,tieunknown; basalmargin of
peristome curved (iudging from many fragments
*irh marked collabralsio#th lines;Text-fig. b) and
initially thickenedand-wellreflected;umbilicuswide,
round and deep.
Dimensions- Estimateddiameterof largestspecimensaround 10-I 5 cm.
Material - SienaBasin:all the materialconsistsof
crushed and fragmentaryshells from the following
localities: Poggio Vangelo, I Sodi, Vescona, San
Vittorio, Poffidarno," Fornace di Rapolaoo, and
Finerri.
material,we studieda shell
In addition to Sienese
from the Pliocene of Campore di Salsomaggiore
(Q.t"tantelli collection, MV4S) and a _b"4lyPreembedded in
served whole specimen, sdll pardy
'Museo
di Storia
shale,without .oll.ction data in the
Naturale dell'Accademiadei Fisiocritici.
S.
Geographic and stratigraphic distribution
plioexte-nti*asdescribedfiom the Plioceneof Monte
Castello near Alessandria(Piedmont, Italy). Our
recordsfor the Plioceneof Sdsomaggioreand Siena
arethe first reportsof th9 speciessinceits description.
The speciesis new for'the SienaBasin,where it
"shale"
outcroPs
may be locafly common in the
deposited in intermediate-depthmarine environmentsof the Middle Pliocene.
'We
tentatively assign this Stellaria
Remarks
(Onustus) species to Xerlophoraplioex.tensaSacco,
1896. The descriptionof ihis nominal specieswas
sure
basedon such bad materialsthat Saccowas not"non
xenoPhorid:
to
a
belonged
actually
they
whether
parebbero neppure Tugurixttll, ricordando invece
ilcune Pleuroiomaria, se non avesserouna grande
somiglianzacoi giovani di T extensumdel Sowerby''.
AlthJugh Sacgo5diagnosisis defective(" TbstacrAssuAnfractusir;*; et medii laeues,keuiter conla, coni"ca.
uexuli, sublucidi, Torpora aliena non colligentes.
Umbilicusktus, profaidus subkeuis"),it is consistent
with a speciesof Onusttls,especiallyin view of the
large ,rtribilicus.Unfortunately the rype material in
collection at the Museo Regionale
rhJ Bellardi-Sacco
di ScienzeNaturali of Turin no longer exists(Ferrero
Tigures are of no
Mortara et al., 1984) and Sacco's
help becausethey illustrate two inlernal moulds.
the study oft.* materialfrom Monte Castello
Oniy'clarify
the identiry of this nominal tanon.
can
Cossmann(l9l6i p. 195) confirmedit as a valid
: X. (Trochotugurium)
species
-d'Italie plioextensA,occllrritg
'tans
Et dans le Scaldisien
Plaisancien
le
d'Anvers"and figured a fine shell (Cossmlon, 1916,
Pl. I l, figs20-2[), without givingthe localiryo{:o!lection. Thir shell does not belong to a Stelkria
(Onustus)speciesand seemsto have a base,aPerrure
and umbiliius similar to the shellfigured by Marquet
Glibert,
(1997, Pl. l, fig. 7) as Xenophorajcald.ensis
1958,from the?lioceneof Kallo (Belgium).
the relaPonder (1983, p. 15) briefly discusses
with
together
that
st-ating
plioixtensa,
of
S.
tionships
the Itafian Miocene Phorusborsini Sismolda, 1847,
Pantanelli,1887, it probably
and Xenophoradepressa
with Trochusextenbelongstb the lineagethat began
"other
nominal taxa,
sus S6werbv, 182f . Many
describedfrom the Tertiary'of northern Italy, belong
or might belong to Stellaria (Onustut), &ryoph.or*
guguiium) oligoitriatum Sacco, 1896, X. Q jubextensumvar. oriatopnrua Sacco, L896, X. G) borsoni
Sacco, 18-96,^\d F. G) Po.stextenvar.pagodaeformis
sum-Sicco," 1896. Apart from P borsqni;they are
basedon badly preservedmaterial and their status
and relationshipi are thereforedifficult to evaluate.
Also X depressa,
which the fine figure by Pantanelli
(1887) t.tfg.tts it was describedoi " *.ll preserved
shell,wasiZtually basedon a shell in very
con' poor
r
dition (persondunpublisheddata).
Of the four extant species,S. longlai (Bartsch,
l93l) is the most similaf to S. cf. plio/xtensabecause
both have the porcellanouslower surface of the
peripheralflanee and a concaveprofile of the inner
b"r"l peristontE;the other speciis of Stelkria with
porcellanouslower surfaceof peripheralflangehavea
*"ry inner basd margin of perisiome (seeKreipl &
Alf,'1999: Pls 26-28).
sr'*nrua
I83I)
H:T:fiH{l;"""'
183I lPhorus)testigerus
BnoNN, p. 61 .
Tvpematerial - It is very improbable
^below). that wDe
mat;rial sdll exists (seeremirks,
To defi"itively clarify the identity of this nominal taxon, we
designate the specimen from Bacedasco(IPUM
9882) illustrated-inPl. 9, fig. 2 as the neorype.
"Cq.
"Cq."
Type localin
Ba. C. 15".
is
-"Ba."
"Bc"
Cast'-ellarquato, is " rlip for
i.e. Bacedasco,
6(C"
are the,'JtittgererGrobf<alk,blau. thonig, in den
Apenninen .
Diapnosis- A species
of Stellariacharacterizedby
^robust
mediuri size (l),
and trochiform shell (2i,
with slightly obtuse peripheral flange, divided into
long laie blunt disiiations, each bEarinea foreign
obiEctdistally(3); s[ire with 7-9 flat who"rls@);8"0of spire surfacefree of attachedobjectsor their
90lo/o
scars(5); with small wa\rysubspiralthreadsmore evident in the lower half of whorls; in last whorl, some
threadsdivert from their subspiralcourse,becoming
more evidentand irregul"r "rd endins in disitationi
sivins their dorsal ruiface a sliehth #arry *itd wrinHed ippg"Iaq-ce(6); attachedo6j..h (sheilsand fragmenti of shells)small to mediuin in size,presentIn
last whorl only on distal end of dieitationi (7); scars
of attachedobjects from shallowio deep (S); base
slighdy convex(9); basalsculptureconsistingof colla5ral growth lines and sometimesweak spi"rallines
( I 0); 5"r"1 margin of peristome J-shaied ( I I );
umbilicussmdl, Jometimlsslit-like (12).
Qesc7iption Shell,(Pl.9,.$Sr l, 4) dextral,medium in size,robust and trochiform with pointed apex
and slightly obtuseperipheralflange,divided int o 79 long largeblunt digitations(8-13 in extant specimens;Ponder,1983);spireconical(angleabout 90";
75"-85" in extant specimens;Ponder, 1983) with
about 7-9 regularly growing whorls (first whorls
alwayseroded);last whorl usuallynot dilated;whorls
always flat; suturesvery shallow; at least about 8090o/oof spiresurfacefreeoFattachedobjectsor rheir
44r
scars,with smallwavy subspiralthreadsmore evident
in the lower half of whorls, intersectedby faint or
more evidentbut irregularprosoclinegrowth lines;in
last whorl, some threadsdivert from their subspiral
course, becoming more evident and irregular and
ending in digitations, giving dorsal surfaceslightly
yarty appearance;attachedobjects small to large in
size,consistingof fragmentsof bivalveshells(in one
casea claw of a decapodcrustacean),presentin last
whorl only on distal end of digitations (those
attachedalong suturesof first whorls are not generally retainedin adult shells);scarsof attachedobjects
usually shallow; baseslightly convex,with collabral
growth lines and weak spiral lines, more evident and
regularnear umbilicus, closeand wavy on peripheral
flange(in somespecimensspirallinesnearumbilicus
very evident and crossinggrowth lines giving granular appearanceto basesimilar to that of X. rispa; see
Pl. 8, fig. 3 and Sacco,1896,Pl. 3, frg. l.); aperture
large and oblique becauseinnermost basd margin is
behind by almost a quarter of a whorl; peristome
interrupted with upper and lower verticesunited by
weak parietal callus; columellar margin sometimes
thickenedand slighdy reflected;basalmargin of peristome J-shaped, thin and sharp; external margin
irregular,thin and sharp; umbilicus small and deep,
someumesslit-like.
Dimensions- Diameter: up to 72 mm; height: up
rc 47 mm. Dimensionsof other Italian Pliocenespecimens: D: up to 63 rnm, H: up to 40 mm (Sacco,
1896); D: 55.8-64.2 ffiffi, H: 40.0-47.5 mm
(Pelosio,1967);D: up to 56.2 ffiffi, H: up to 32 mm
(Caprotti, 1967). Dimensionsof extant specimens:
D: up to 65.0 rnrn, H: up 80.7 mm (Ponder,1983);
D: up to 83 InID, H: up to 67 mm (Goud 6c
lkonenberg, 1938-89;IGeipl & Alf,, 1999).
Materia/ - SienaBasin:most of the materialconsistsof crushedand fragmentaryshellsfrom the following localities: Monsindoli, I Sodi, Podere
Capanna and Terre Rosse;the best specimensare
from Monsindoli (2, FPC; 2, IGF 5985E and IGF
Sl99E) and from Terre Rosse(1, VSC). In the
RadicofaniBasin,south of the SienaBasin,S. testigerais common at Lucciolabellaand Monte Calcinaio.
In addition to Sienesematerid we studied material from other Pliocene localities: Bacedasco( I ,
IPUM 9882), Priocca(1, DSTUT), Rio Torsero(2,
M R S N 8 5 0 6 6 . 0 1. 0 1 4 a n d 8 5 0 6 6 . 0 1. 0 1 6 ) a n d
Savona(1, MRSN 85066.01.015).
Geographicand snatigraphicdisnibution Live S.
testigeraoccur in two distinct areaswith two subspecies:,S. testigera
digitata Martens, 1878, is wide'West
spreadalong the
African coastsfrom Senegalto
Namibia and X testigera
profundâPonder,
1983,in the
Gulf of Adenand nonheast,\hica (Ponder,I 983).
442
G. AUNGANELLI, V SPADINI, S. CANF'4 \ELLI
The ecolosy and bioloey of extant ,S.testigeraare
relatively .,r1nown. Ad;m 6c Knudsen ll g 55)
reported that living specimensof S. testigeradigitata
*it. dredged from- bottoms of variable nature
betweend.Ipths of 170 and 263 m and Le Lo euff et
Al. (1971) itated that this endry lives on the continental slopeand belongsto the bathyalfauna. Ponder
( 1983) t.iotred thar Sl testigera
Profundawasdredged
betweendepthsof 450 and 730 m.
is recordAccordi"g to Ponder(1983), S. testigera
ed from the" Oligocene to the Pliocene of several
European locditlot however we do not known enY
O[e6cene record and'W. Ponder (personalcommunicition) thinks that the Oligocene indication mey
be bas.d ot "t error. The spJcieshas been reported
from the Miocene of Paratethys(Germany, Austria
and Bulgaria), the Miocene and Pliocene of the
(Miocene: Corsica, southern and
Medit.ti"tt."t
northern Italy; Pliocene: southern Spain, Sardinia,
central-northernItaly and Maritime Apt) (Hcirnes,
1856;Locard, 1877;Seguenza,1877, 1881; Koenen'
1882; Sacco, 1896; Cossmann' 1916; Ruggieri,
1957;Moroni, l95S; Kojumdgieva6c Strachimirov,
1960;Dieni 6c Massari,1966;Caprotti, 1967,1970,
lg74; Pelosio, I 967; csepreghy-vteznerics,1969;
Marasti 6c Ram, 1976, t977;Vontefameglio et A,1,,
1980; Anfossi et dl., 1982; Zhu Min-Da, 1984;
Cavallo 6c Repetto, 1992; Thbanelli 6c Segurini,
1994;Andreoli-8cMarsigli, 1996; Harzhauseret Al.,
statedthat
2003;Landav et d1.,2004).Sacco(1896)
"Astiano"
of
from
the
recorded
is
also
the species
"futigiana", but this is questioned P:losio (1967.).
by
was repoited from the SienaFfif !.y
Sl testigerA,
(1878),
Hrirnes (f856) and De Stefani6c Pantanelli
"shale"
outcroPs
in
where it may be loc"lly common
deposited ill intermediate-depth marine environments of the Lower and Middle Pliocene.
Remarks- Fine fossil shellsof S. testigeraare very
rare. Two very fine specimens are from Priocca
(Montefameglioet al., i980, FiB. l0) and from Rio
torr.ro (Saclo,1896, Pl. 3, Frg.-l.);the latter (Pl. 9,
fig. 3) is interesdngbecauseit-has very .4d.:tt spiral
liiles nearthe umbi'licuswhich crossgrowth lines,giting a granularappearanceto the basesimilar to that
o{X. iris\a. Moit of the material is crushedor incomDlete,lackine the last part of the aperture (Pelosio,
\961 , Pl. i6, figr 2ba-20d, Pl. 37, figt la-ld;
Caprotri, 1970, Pl. 7, fig. 3; Cavallo 6c Repetto,
EXPI,{\AIION
Fies l-4 --o-
1992,Fig.110;Andreoli& Marsigli,1996,l!..1,fig.s
7-, and our specimensare no excePtion.Although
this makes c"rêfulcomparisonsbetween fossil and
extant specimensdifficuit, it is evident that at least
Pliocene specimenshave remarkable variabill.ry.in
spire angle,length of peripheraldigitations and dorsal and basd sculpture.
The earliest Miocene specimens
-conical from northern
than Plioceneor
Italy are smdler and more
extant ones.Saccodistinguishedthem pardy esX. testiqeravar. ektiuscuk and pardy asX. testigeravar. tauriturrita,both of which *. "*"lly regardedasjunior
synonymsof X. testigera(Ponder 6c Cooper, 1981).
Aop.futty, the tt"t,tf of these nominal til€ will be
investigaiedagain, especiallythat of X. testigSyvar.
taurotirri to *Kich i ncludesip ecim ens rather di fferen t
from the Plioceneand extant ones.
was describedby Bronn (1331)
Phorustestigerus
"Jiin-gererGrobkalk, blau- thonig, in den
from the
Apenninen" o'f Castellarquato and Bicedasco.
Bionn's collectionwas acquiredby L. Ag*siz for the
Museum of Comparatlive Zooloy . (Harv"t4,
US) (R. Janssen'Persond communlMassachusetts,
cation) and his fossil molluscs are housed in the
InvertebratePaleontologvDept. No synrypeof P testiqerashas been found i-riwhit remainsof the collecti3n (F. Collier, personalcommunication).
The original iescription-ispogr and doesnot offer
elemen$ s;ppofting sure idenlification of the nomind taxon deslribed-byBronn. Only S. testigeraseems
(Hornes, 1856; Cocconi, 1873;
presentat Bacedasco
Pelosio, 1967), but three species (X. crispa, X.
infundibulum and S. testigera)o_ccurat Castellarqurt:
(Hornes, 1856; Cocconi, 1873; Glibert' 1963;
Caprotti, 1967; Zhu Min-Da, 1984).
The first authors to cite Bronnt specieswere
Hauer (1837, p. 420) in a, list of speciesfrom the
Vienna Basinattd Michelotti (1847,PP. 174-175,Plt
7 , fts.6) on the basisof specimensfrbh Tortona and
However,Miihelotti (1840) illustrated a
Bace?asco.
shell whose identity is not clear: some characters
recall S. testigera ind X. crispa (dorsal subspiral
threads)and Jthett match X irispa (absenceof evident peripheral digitations and last whorl dilated)
(TextJi g. l4). S. tistigerawas_subseque_nilt.p9t_r:.d
by Sisnionda(1847, p. 5O),.llotn (1848,-p.999.,
D'Otbigny (1852,p. 41) and Hornes(1856,p. 444,
PL.44, fie. l().Tha latter is the first author to give a
sufficiently clear figure (albeit of an incomplete shell
OF PIATE 8
Stelkria testigera(Bronn, I 83 I ).
l) F. Pizzoleiocoilection, from the Lower Plioceneof Monsind-oli (S_iena);_
ti M",yp;; M;;;; Ji-p"l.obiologia e dell'Orto Botanico, Modena, Itdy, IPUI\
3i Ri; firr.ro, Bellardi-Sacco.ofl..tion, M-useoRegionaledi ScienzeNaturdi,
4in. Spadini collecdon. Middle PlioceneofTerre R6sse(CastelnuovoBerarder
G. MANGAIVELLI, V SPADIIVI,S. CIANFAIVELLI, XE,NOPHORIDS FROM MEDITE
PI. B
444
Thb. 6 - Synopsisof charactersdistinguishingthe Italian PlioceneXenophoraspecies.
X. mispa
X. daaolii
X. infundibulum
X. plioinlica
S. plioextensa
S. testigera
Size
small
very large
large
Shell
robust, from
almost discoidal
to trochiform
robust and vgredate
robust and trochi- robust and uochi- fragile and di,
form
form
.oidal
Peripherd flange narrow irregular,
roughly rouid
(where attached
objectsabsent)
narrow irregular
shaped,shaiply
keeled (where
attachedobjects
absent)
narrow slightly
wxw, sharilv
^J
keelld
Whorl profile
irregular
flat or very slight- flat or slightly
ly convex
convex
flat
flat
most of surface
about 90o/o
most of surface
most of surface
probably 80-90o/o
variable
Percentage of spire about 30-600/o
surface fue from
attached objects or
their scars
large
large
narrow irregular, large,,thinand
porcellanous
roushly touid
(whireattached below
objectsabsent)
rligltllf obruse,
divided into some
lTg. and.long
Dlunt dlgltauons
Spire sculpture
with fine wary
meandering:,rbsplral, someumes
opisthocline,
riblets
with very irregular with closefine
and prominent,
vermiculate
opisthoclinedrop- opisthocline
fike rugae,inrer-^ tiblets
sectedby faint
prosoclinegrowth
lines
with very small
wary vermiculate
opislhocline
tibl.tt
with very smdl
delicate waw vermiculate opisthocline ribleti
with small wary
subspiral threads
tt oti evident in
the lower half of
whorls; in last
whorl some
threadsbecome
more evident and
irrepulat Sttd end
rn clgrtauons
Aaached objects
mainly bivalve
and gastropod
shelli, fragments
of shellsal"a pebbles from smdl to
proportiondly
very large
very large fragttt.tttt oTshelfsof
bivdves and gasuopods, and pebbles
mainly bivalve
shellsand fragments of sami,
alwaysvery small
to small exceptin
last quarter oTlast
whorl
frasments of
biv"alveshells,
srlall to medium
in size
bivdve shells and
fragmentsof
sarie, smdl to
medium in size
bivalve shells and
fragmentsof
same,smdl to
medium in size,
presentin last
whorl only on distal end ofdigir"tions
Scarsof attached usually deep
objects
usually deep
usually deep
usually deep
shdlow rlorg
sutures,deeper
1!org peripherd
tlange
usudly shallow
Base
slighdy concave
funnel-shaped
slightly convex
convex
slightly convex
Basal margin of
Penstome
sliehdy convex to
alilosi flat
J - shaped,thickened and with
jutting outer portlon
curyed, thickened straisht and thick- I - shaped,thick- curved and inidd- J - shaped
.tteJas far as
and sliehtly
lned and inidally ly thickened and
reflectil
periphery then
reflected
reflected
absent;at attachment to peripheral flanee, thire is
usually-asort of
sinulus
Basal sculpture
collabral srowth disdnct collabral
lines and"spiral growth lines
groovesmore distinct near umbilicus and having a
more or lessgranular sculpture
collabral growth
lines, *eik ot
very weak spird
lin6s and sorrretimes very thin
opisthocline
riblets
very weak colrobust collabrd
hbial srowth lines growth lines and
more &ident near
thin spird
i:o
aperrure,many
llnes
very smdl spiral
[n6s and "..y
small wary vermiculate opisthocline riblets
collabral srowth
lines and-somedmes weak spird
lines
Umbilicus
smdl and deep
but often slitlike
or coveredby
parietal callus
absent
round and d.gp
round and deep
or slit-like and'
coveredby peristomal margln
small, sometimes
slit-like
absent
445
|4#Kr&,'tw+f,frui,,:'':
rex'-ng
1LJ'H:'Jii*n,oJ
whether
bui it is uncertain
species,
tionof Bronn's
to X. testigera.
reallybelongs
theshellillustrat-ed
|5{;w!:xTi:;i#
rex,-ng
ffnrl?*,:"'$r1,11l";::;
from Baden)for recognitionof the speciesasit is currendy understood(l5xt-fig. l5).Thinks to this good
figu1L,subsequentidentifiiations haveprobablybeen
cdnsistent.However, becausedesignationof a neotype is advisableto clarify the ideritiry of l nominal
taxon, we designatethe-specimenfrom Bacedasco
(IPUM 9882) i-llustratedin Pl. 9, fig. 2 as the neotyPe.
-Ponder(1983) statesthat fossiland extant specimens must be assignedto distinct chronosubspecies:
incluGs specimens from the
S. testigerat-estig97a
digitaMiocen-eto the Flioceneof Eurobe,S. testigera
taMartens, 1878,specimensfrom'WestAllica arid .S.
testigeraprofundn Ponder, 1983, extant specimens
frori the Gulf of Aden and northeasi Africa.
However,some of the characterslisted by Ponder to
differentiateextant subspeciesfrom fossil S. testigera
haveonly relativevalue due to wide variabititestigera
fossils.itot example,it is doubtful to distinfi
the
w
from extant
euish the fossilnominoryfii.d subspecies
5n.r on the basisof th6 length oflperipheraldigitations (shorter in fossil specimens,longer in extant
ones).
St. Jean (1982) and Ponder ( l9S3) hypothesize
that S. testigeraoriginated in the western Atlantic
from S. co-nica(Dall, 1892) of the Eocene of
Mississippi.The latter it y9ry similar to S. testigera,
size(fot S. conbeingmainly distinguishedby small_er
icA, i.. St. iean, tl98z, firsf two figures at p. l9).
However, the long gap in stratigrlphic distribudon
betweenthe Eoceie-l.' tonica and,the Miocene S. tesis surprising.
tiqera
o
Arrothei rece"'ntlydescribed species, Stellaria
kriegerbartholdiNielsen 6( De Vries, 2002, is very
simllar to S. testigeraexceptfor its closedumbilicus
(Nielsen6c De Vries, 2002, pp. 75-77, Figs 17-20).
Thir speciesis very interestiridbecauseit Is the first
reportôf an endry of the S. tdstigeragrouP from the
southern hemisphere (Millongue Formation,
PeninsulaArauco, south-centralC[ile), but unfortunately its uncertain stratigraphic age (Eocene or
Miocene)preventsmore detailedcomment.
to the original).
ACKNO\TLEDGEMENTS
'We
thank Andrea Benocci, Antonella Daviddi, Leonardo
Gamberucci, and Saulo Bambi for technical assistance,Helen
Ampt for revisingthe English,Armando Costantini (Siena,Italy)
Franca Campanino (Tirrin, Itdy),
for itratigraphicli assista-nce,
Elisabettl Cioppi (Florence, ttaly), Fred Collier (Harvad,
MassachusettsUS), Franco Davoli (Modena, Italy), Maurizio
Forli (Prato, Italy), Riccardo Manni (Rome, Italy), Daniele
Ormezzano (Tirrin, Italy), FrancescoPizzolato (Arezzo, Italy),
and RaffaeleQuarantelli (Salsomaggiore,Italy) for information
about or loan of material from theii Institute or private collecdons, Fausto Barbagli (Pavia,Italy), FrancescaCagnani (Siena,
Italy), Anita Eschner (Vienna, Austria), Viviana Fiorentino
(Palermo, Itdy),
- Elena Gavetti (Turin, Italy), Jeroen Gould
1-,eiden, The Netherlands), Pierre Lozouet (Paris, France),
Robert Marquet (Annverpen, Belgium), PeterMordan (London,
UK), Ruggiro Noto La Diegi (Berlin, Germany), Marco
Franceico Pezzo (Oxford,_UK), Jot.P
Oliverio fRo-a, Italy),
-(Ciutadella
de Me_norca,Spain), Tom
Quintana Cardona
Sihiotte (Copenhagen, Denmark), Ewa Stworzeuricz(IGakow
Poland)andThomas Vilke (Philadelphia,US), for their valuable
in bibliographic research.
help
^'We
than[ Folco Giusti (Siena, Italy), Robert Marquet
(Anrwerpeh, Belgium), and Giulio Pavia (Tirrin, Italy) who Provided numerous-helpful comments which greatly improved the
accuracyand cualiry of this paper.
di Sienaand Museo di Storia
Resbarchfinanc'edby Univ6rsit)r
"La
Specola",Universiti di Firenze
Naturale, SezioneZoologica
(Italy).
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(manuscript receiuedMarch 9, 2004
accepted September 4, 2004)
Giuseppe MnNcANELLI
Dipartimento di Scienze Ambientali
Universiti di Siena
Valeriano SpnotNI
Via Augusto Toti 6
Lucignano (Arer-zo)
Simone CnNTnNELLI
Museo di Storia Naturale
"La
Specola"
Sezione zoologica
Universit)r di Firenze

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