Table Of ContentBoll. Malacol., 44(I-4): 15-19(2008)
The identity of Nucula perminima Monterosato, 1875
and Yoldia striolata Brugnone, 1876 (BivaSvia: Protobranchia)
Rafael La Perna*
* Dipartimento Abstract
di GeologiaeGeofisica, Two enigmatic protobranchs, Nucula perminima Monterosato, 1875 and Yoldia striolata Brugnone, 1876,
Universitàdi Bari, are revised basing on material from the Monterosato collection. Nucula perminima, described from off
via Orabona4, Sciacca, southwestern Sicily, is a paedomorphic nuculid of which Nucula recondita Gofas & Salas, 1996 is
70125 Bari, Italy, a junior synonym. Yoldia striolata, described as a Pleistocene fossil from Ficarazzi, Palermo, has a complex
[email protected]
systematic history: its synonym list includes Yoldia producía Monterosato, 1880 and Yoldiella seguenzae
Bonfitto & Sabelli, 1995. The newcombination is Yoldiella striolata (Brugnone, 1876).
Riassunto
È stata effettuata la revisione di due specie enigmatiche di protobranchi, Nucula perminima Monterosato,
1875 e Yoldia striolata Brugnone, 1876, sulla base di materiale presente nella collezione Monterosato. Nu-
culaperminima, descritta dai fondi coralligeni al largo di Sciacca, è una specie pedomorfica e non uno sta-
dio giovanile di nuculide come erroneamente ritenuto da Monterosato. Nucula recondita Gofas & Salas,
1996 ne è un sinonimo più giovane. È una specie criptica, capace di vivere anche in ambiente di grotta.
Yoldiastriolata, descritta per il Pleistocene di Ficarazzi presso Palermo, ha una storia sistematica complessa:
la lista dei suoi sinonimi comprende Yoldia producta Monterosato, 1880 e Yoldiella seguenzae Bonfitto &
Sabelli, 1995. La nuova combinazione applicata a questa specie è Yoldiella striolata (Brugnone, 1876). È
una specie di acque profonde, presente in Mediterraneo e nel vicinoAtlantico.
Key words
Systematics, Nucula, Yoldiella, Monterosato, Brugnone, Mediterranean.
Introduction asa synonymofN. perminima. Inthe lastnote, Montero-
sato (1881) definitely regarded N. perminima as thejuve-
Nucida perminima Monterosato, 1875 and Yoldia striolata nile stage ofN. aegeensis.
Brugnone, 1876 are two enigmatic protobranchs descri- The type material ofNucida perminima (MZR 14379, Fig.
bed in the early literature, the former from off Sciacca, 1 D) consists of 12 valves and few complete shells (Fig.
southwestern Sicily, the latter as a Pleistocene fossil 1 A-C). This is a valid species, not a juvenile stage,
from Sicily. Nuculaperminima remained totally forgotten clearly conspecific with the paedomorphic nuculid de-
in the malacological literature, but the recent descrip- scribed from the Mediterranean as Nucula recondita by
tion ofa new species, possibly a synonym ofN. permini- Gofas & Salas (1996: p. 430, figs. 14-22). The synonymy
ma, led theidentity ofthis species tobe debated. Thehi- between N. perminima and N. recóndita was first repor-
story of Yoldia striolata is a long sequence ofsynonymies ted by Giannuzzi Savelli et al. (2001: p. 331, figs. 11-13)
and interpretations. In the present work, the identity of and Palazzi & Villari (2001: p. 24, figs. 91-96), but wi-
both species is discussed basing on material from the thout the supportoftype material.
Monterosatocollection (MuseoCivicodiZoologia, Roma). Nucida perminima actuallyis notably similar to the early
stages of normal sized nuculid, as well documented by
Nucula perminima Monterosato, 1875 Gofas & Salas (1996) and this explains Monterosato's
doubts about the taxonomic status. Up to the size of N.
Monterosato (1875a) described Nucida perminima from perminima (about 1 mm), all the juvenile nuculids have
the coralligenous bottoms off Sciacca, Sicily Channel. It a smooth margin, but this species has well distinct,
was said tobe very small ("La pili piccola delle nostreNii- functional primary teeth whereas they are strongly re-
culae e forse anche di quante se ne conoscono viventi e duced or totally lost in the other species (Gofas & Salas,
fossili"), but apparently consisting of mature specimens. 1996). The larval shell is flatfish, with a central depres-
It was compared with young stages and growth series sion (Gofas & Salas, 1996).
of other nuculids and found specifically distinct, even This species exhibits a remarkablevariability inthe shell
from Nucida tenuis (non Nucida tenuis Montagu, 1803, outline. The type material is less elongate and morecon-
but N. aegeensis Forbes, 1844, now in Ennucuta Iredale, vex ventrally than the specimens illustrated by Gofas &
1931) which shares a smooth margin with N. perminima. Salas (1996). Shell elongation and ventral convexity are
Afew years later, Monterosato (1878) listed some varie- also variable in the shells illustrated by Giannuzzi
ties of Nucida nucleus (L., 1758), including a var. minima Savelli et al. (2001) and Palazzi & Villari (2001). The
Rafael
La
Perna
Fig. 1. Nuculapermìnima Monterosato, 1875. A-C. Syntypes, SicilyChannel, offSclacca (MZR 14379): A, B. 1.02 mm, c. 1.03 mm. D. Monterosato's
label. E-J. Grottadell'Accademia, Ustica Island, southernTyrrhenian (author'scollection): E, F. 1.13 mm; G. 1.24mm; H. 1.05 mm; I,J. 1.23 mm.
Fig. 1.Nuculapermìnima Monterosato, 1875 A-C. Sintipi, Canaledi Sicilia, al largodi Sciacca(MZR 14379): A, B. 1,02 mm; C. 1,03 mm. D. Etichetta
di Monterosato. E-J. Grotta dell'Accademia, Isola di Ustica, MarTirreno meridionale(collezionedell'autore): E, F. 1,13 mm; G. 1,24mm; H. 1,05 mm;
I,J. 1,23 mm.
shells from a shallow water cave (Di Geronimo et al., Yoldin striolata Brugnone, 1876
1997) havea slightly moreobliqueoutline (Fig. 1 E-J).
The lifehabits ofN. permìnima mustbenotablydifferent Brugnone (1876: p. 9, fig. 9) described Yoldia striolata
from those of the normal sized nuculids, inhabiting soft from Ficarazzi (Palermo), an Early Pleistocene (Sicilian)
bottoms as shallow burrowers. The original records of locality, frequently mentioned in the early literature on
N. permìnima and N. recondita are from hard substrates, fossil molluscs (see Greco, 1986). The original illustra-
the former from cavities and crevices in volcanic boul- tion is poor, only showing the external surface of an
ders colonized by Corailum rubrum, the latter from ovate, moderately inequilateral bivalve with a weak
rocky bottoms with Laminaria and from Posidonia beds. commarginai sculpture and a relatively prominent um-
Di Geronimo et al. (1997) reported this species from ca- bo. No type material of Y. striolata was foundin the Bru-
ve assemblages obtained from sediment samples. Also gnone collection (Museo "G.G. Gemmellaro", Univer-
the record by Palazzi & Villari (2001) is from cave as- sityofPalermo) nor intheMonterosatocollection.
semblages. Nucula perminima is evidently a cryptic spe- The history of this species starts with Monterosato
cies, as suggested by Di Geronimo et al. (1997: p. 24). (1875b), who introduced Leda (Yoldia) producía as a no-
Thanks to miniaturization, implying a lowering of men nudum for an extant species from off Palermo, quo-
energy requirement, it is able to cope with the oligotro- ting Yoldia abyssicola Torell, 1859 as a dubious synonym.
phic conditions in the caves (Hayami & Kase, 1993). It was then reported (Monterosato, 1877) from the
Similar considerations could be also applied to Nucula Pleistocene of Ficarazzi, as a synonym of Y. abyssicola
bicornis, anotherpaedomorphic species from the Canary and Y. striolata and again as a synonym of Y. abyssicola
Islands, also describedbyGofas & Salas (1996). in a checklist of living Mediterranean molluscs (Monte-
According to Hanken & Wake (1993), miniaturization rosato, 1878). Finally, Monterosato (1880) made availa-
may imply important phylogenetic perspectives. The ble the name Yoldia producía by giving a reference to the
suggestionby Gofas & Salas (1996) about the need for a published description of Y. striolata Brugnone, mentio-
new genus for these paedomorphic nuculids is herein ning "type given by the author" and Y. abyssicola (non
strongly supported. Concerning this, other cases of mi- Torell) sensu Monterosato, 1878 as a misidentification of
niaturized molluscs, such as the mytilid Dacrydium the same species. Therefore, Yoldia producía is an objecti-
Torell, 1859 (Ockelmann, 1983; Salas & Gofas, 1997) and ve synonym of Y. striolata. It was said to be "different
the nuculanidMicrogloma Sanders &Allen, 1973 (Ockel- from Y. lenticula, Moll. (- Y. abyssicola Torell), which is
,
16 mann & Warén, 1989), constitute systematicprecedents. from Norwayand all theArctic region".
Seguenza (1877: p. 1180, pi. 5, fig. 28) reported Yoldia to pusio Philippi, although these species were kept as
abyssicola Torell, 1859 from the Plio-Pleistocene of distinct by Jeffreys (1879). Since Leda pusio sensu Jeffreys
Southern Italy, with Y. producto and Y. striolata as syno- was mostly based on his var. latior, misidentified as var.
nyms. The illustration shows a bivalve somewhat simi- salicensis Seguenza, 1877 (Jeffreys, 1876, 1879), Warén
lar to Yoldiella philippiana (Nyst, 1845), with a narrower (1989) proposed the synonymystriolata = latior = salicen-
posterior side and a commarginai striation. The same sis and applied the combination Neilonella striolata (in
synonyms were listed for Yoldia abyssicola in a later family Neilonellidae Schileyko, 1989) to Leda pusio var.
work (Seguenza, 1879: p. 284). latiorJeffreys, 1876.
Jeffreys (1879: p. 54) reported Leda striolata Brugnone Allen & Sanders (1996), basing on the illustration of
from some deep water stations in the Northeast Atlan- Yoldia abyssicola by Seguenza (1877), concluded that pro-
tic, with Yoldia abyssicola sensu Seguenza and Yoldia pro- ducía and striolata were the same species, without men-
ducía Monterosato as synonyms. It was compared with tioning that they are objective synonyms, and different
The
Leda lenticula Moller, 1842 as follows: "flatter, more from Leda pusio var. latior to which they applied the
sharply pointed or wedge-shaped at the anterior [sic] combinationNeilonella salicensis. identity
end, and concentrically striated; the striae are regular More recently, Jeffreys' var. latior was moved to the ge-
of
and sometimes numerous, but usually distant and co- nus Pseudoneilonella Laghi, 1986 (Neilonellidae) as P. la-
vering the front only". Jeffreys added: "I had provisio- tior (Jeffreys, 1876) by La Perna (2007). In the same Nucula
nallynamed this speciesacutalis". work, Pseudoneilonella pusio (Philippi, 1844) and P. sali-
Warén (1989) gave an identity to Yoldia striolata, relying censis (Seguenza, 1877) from the Mediterranean Plio-
on material from the Jeffreys collection. For unknown Pleistocene, were kept asdistinct species. perminima
reasons (mislabelling?), he found no difference between Warén's (1989) interpretation of Yoldia striolata as Leda
the material referred therein to striolata Brugnone and pusio var. latiorJeffreys was questioned by Di Geronimo
Monterosato,
1875
and
Yoldia
striolata
Brugnone,
1876
(Bivalvia:
Protobranchia)
Fig. 2. A-C, E-H. Yoldiellastriolata(Brugnone, 1876). A, B. Syntypeof Yoldiaproducía Monterosato, 1880, Palermo, 210 m, 4.68 mm (MZR 14425).
C. Marseille, 3.20 mm (MZR 14425) D. Monterosato's label [Leda (Yoldia)abyssicola Torell= L producía Monterosato Pal. 210ml}. E-G. Archi (sou-
thern Calabria), Early-Middle Pleistocene: E, F. 4.06 mm (author'scollection): G. 3.51 mm (author'scollection). H. SouthernTyrrhenian, Eocumm95 st.
14, 38°20'08" N, 14°10'47" E, 1139 m, 3.20 mm (author's collection). I-K. Yoldiella philippiana (Nyst, 1845): I. Villafranca Tirrena (Messina), Early
Pleistocene, 5.20 mm(author'scollection). J, K. SyntypeofNuculatenuisPhilippi, 1844, 3.60mm(HumboldtMuseum, Berlin; photoA. Warén).
Fig. 2. A-C, E-H. Yoldiellastriolata(Brugnone, 1876). A, B. Sintipodi Yoldiaproducía Monterosato, 1880, Palermo, 210 m, 4,68 mm (MZR 14425). C.
Marsiglia, 3,20 mm (MZR 14425) D. Etichetta di Monterosato [Leda (Yoldia) abyssicola Torell = L. producía Monterosato Pai. 210 mi). E-G. Archi
(Calabria meridionale), Pleistocene inferiore-medio E, F. 4,06 mm(collezionedell'autore): G. 3,51 mm (collezionedell'autore). H. MarTirreno meridio-
nale, Eocumm95 st. 14, 38°20'08" N, 14°10'47" E, 1139 m, 3,20 mm (collezione dell'autore). I-K. Yoldiella philippiana (Nyst, 1845): I. Villafranca
Tirrena (Messina), Pleistocene inferiore, 5,20 mm (collezione dell'autore). J, K. Sintipo di Nucula tenuis Philippi, 1844, 3,60 mm (Museo Humboldt,
Berlino;fotoA. Warén). 17
a of this species: he had the possibility to compare his
Recent material of producía side by side with the fossil
Testaminuta,ovata,subaecjuilatera
+ type of striolata, given by Brugnone (see above), and
Shellsmall,ovate,subaequilateral
realized that they were the same species. The material
transversimtenuiterstriata of Yoldia producía (MCZ 14425, Fig. 2 D) consists of two
+
transversallyfinelystriated valves and a few fragments from off Palermo (Fig. 2 A,
striaein mediotestaeremotiores, B) and of a single, smaller valve from Marseille (Fig. 2
in umbonibusobsoletae + C). This material is conspecific with Yoldiella seguenzae
striaemoredistantinthemiddleofshell, Bonfitto & Sabelli, 1995, described from the Tyrrhenian
lostonumbo Sea (Bonfitto & Sabelli, 1995). Yoldiella striolata
lateribusrotundatis posticosubattenuato, (Brugnone, 1876) is then the new combination for Yoldia
;
vixalteroproductiore striolata.
+
Rafael bothsidesrounded;posteriorlynarrower There is a close match between the description of Yoldia
La andslightlymoreelongate striolata and the characters of Yoldiella seguenzae (Tab. 1
A). Also the differences remarked by Brugnone (1876)
Perna margineintegro
marginsmooth + between Yoldia striolata and Yoldia tenuis (Philippi) cor-
respond to the differences between Yoldiella seguenzae
b
and Y. philippiana (Tab. 1 B). The only apparent discre-
Differta Yoldia tenui(Nucida)Phil.,cujusformam pancy lays in size: Y. striolata was said to be larger than
quadammodosimulat, y. philippiana, but the two species attain a similar maxi-
ItdiffersfromYoldia tenuis,towhichitissomewhat mum shell size, about 5 mm in length, according to the
similarinshape. morphometric data by Bonfitto & Sabelli (1995). Bru-
mm
testa majore gnone's specimen was described as being 6 long,
mm mm
byalarger 4.5 high and 3 wide: it was then particularly
large, but the length to height ratio is 1.33, close to 1.38
ventricosiore mm
moreconvexshell + calculated for the largestvalves (5.0 in length) of Y.
seguenzaefrom the data ofBonfitto & Sabelli (1995).
umbonibus tumidioribusetmagisincurvis + The sculpture of Yoldiella striolata is somewhat variable,
umbomoreinflatedandcurved from a pattern of well defined commarginai striae,
ánguloapicaliacutiore mainly near the ventral margin, to almost consisting of
+
umbonal anglesharper only growth lines, as seen in the present illustrations
(Fig. 2 A-C, E-H) and in other illustrated material
latereposticoretuso
posteriorsideobtuse + (Bonfitto & Sabelli, 1996: fig. 5, 6; Di Geronimo & La
Perna, 1997: pi. 7, figs. 5, 6; Giannuzzi Savelli et al.,
externasuperfcicieargutestriata + 2001: figs. 58, 59). The same illustrations also cover the
outersurfacedistinctlystriated
variability range in shell shape of this species. For com-
Tab. 1. Original description of Yoldia striolata Brugnone, 1876. Crosses parison, material of Yoldiella philippiana is here illustra-
indicate full correspondence with the characters of Yoldiella seguenzae ted (Figs. 2 I-K), including a syntype of Nucida tenuis
Bonfitto& Sabelli, 1995 (a)andwiththedifferencesbetween Y.seguen-
zaeand Y.philippiana(b). Philippi, 1844.
Yoldiella striolata is a deep water species, also occurring
Tab. 1. Descrizione originale di Yoldiastriolata Brugnone, 1876. Le cro-
zceateteBoinnfdiitctoan&o Spaibeenlali,co1r9r9i5sp(oan)deenczoancloendiifcfaerraetntzeerifrdaiYY.osldeigeulleanzsaeeguaennd- min t(Bhoenafidtjtaoce&ntSaAbtellalnit,ic1,99w5i;thSalaasd,ep1t9h96;raLnagePeorfna5,002-020030)0.
Y.philippiana(b). The Ficarazzi stratigraphic sequence includes shallow
water calcarenites and fine grained deposits of outer
& La Perna (1997) and La Perna (2003),but withoutpro- shelf-upper slope deposition (Buccheri, 1983, 1984).
posing an alternative interpretation. Brugnone descri- Some bathyal species were reported from this locality
bed his species using the word striata and compared it by Monterosato (1872).
with Nucida tenuis Philippi, 1844, i.e. Yoldiella philippiana
(Nyst, 1845) (see Warén, 1989 for the nomenclatural hi-
story of Nucida tenuis Philippi, 1844). Yoldia striolata Acknowledgements
must be then a protobranch with a sculpture of corn-
marginal striae, different from the coarser sculpture of Aspecial thank to Lionello Tringali (Rome), for his kind
Pseudoneilonella latiorand similar, in the gross shell mor- assistance at the Monterosato collection and to Anders
phology, to Yoldiella philippiana. On the other hand, To- Warén (Naturhistoriska riksmuseet, Stockholm), for
rell's Yoldia abyssicola is a junior synonym of Yoldiella help at various stages. Serge Gofas (Universidad de
lenticula (Moller, 1842), an Arctic species somewhat si- Màlaga) and John Alien (University Marine Biological
milar to Y. philippiana (Warén, 1989: p. 239, figs. 8c, d; Station, Millport) are acknowledged for their careful re-
lOe, f). view.
The proper understanding of Brugnone's species should Work supported by Fondi di Ricerca d'Ateneo 2006
18 be based on the concept which Monterosato (1880) had (resp. La Perna).
;;; ; ;; ;
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