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J Zool Syst Evol Res. 2021;00:1–46. wileyonlinelibrary.com/journal/jzs
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1© 2021 Wiley-VCH GmbH
Received: 5 March 2021
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Revised: 8 April 2021
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Accepted: 11 April 2021
DOI : 10.1111/jzs.12479
ORIGINAL ARTICLE
Revision of the genus Buccinanops (Mollusca: Neogastropoda:
Nassariidae), an endemic group of gastropods from the
Southwestern Atlantic, including a new genus and accounts on
the Buccinanopsinae classification
Guido Pastorino1 | Luiz Ricardo L. Simone2
Contri buting autho r: Guido Pasto rino (gpasto rino@macn. gov.ar)
Zoobank link: L S I D : h t t p : / / z o o b a n k . o r g / u r n : l s i d : z o o b a n k . o r g : p u b : 9 3 3 8 6 9 D 9 - 2 5 B 2 - 4 2 A 0 - 9 B 5 6 - 1 1 5 C 9 9 2 E D 0 F E O n l i n e I S S N : 1 4 3 9 - 0 4 6 9
1Museo Argentino de Ciencias Naturales
“Bernardino Rivadavia”, Ciudad Autónoma
de Buenos Aires, Argentina
2Museu de Zoologia da Universidade de
São Paulo, São Paulo, SP, Brazil
Correspondence
Luiz Ricardo L. Simone, Museu de Zoologia
da Universidade de São Paulo, São Paulo,
SP Cx. Postal 42494; 04218- 970, Brazil.
Emails: lrsimone@usp.br;
lrlsimone@gmail.com
Abstract
The nominal species referred to the endemic southern South American genus
Buccinanops are revised, clarifying the definition of Buccinanopsinae. All valid Recent
species are redescribed based on type material and other specimens emphasizing
anatomy and illustrated using SEM photographs of radulae, embryos, and shell ultras-
tructure. Only seven of 13 nominal species described are considered valid: Buccinanops
cochlidium (Dillwyn, 1817) (type species); B. latus sp. nov. (formerly B. gradatus or
B. lamarckii from Brazil only); B. monilifer (Kiener, 1834); B. deforme (King, 1832);
B. paytense (Kiener, 1834); B. uruguayense (Pilsbry, 1897) and B. duartei Klappenbach,
1961. A phylogenetic analysis based on morphological characters of these species and
12 other buccinoideans is carried out in order to compare results with recent DNA
sequence approaches. The ingroup cladogram is ((B. monilifer (B. latus- B. cochlidium))
(B. duartei (B. uruguayense (B. deforme- B. paytense)))). The former genus Buccinanops,
supported by 11 synapomorphies, can be divided, and Buccinastrum gen. nov. (3
synapomorphies of support) is introduced for the 4 latter species in the above clad-
ogram; its type species is Buccinastrum deforme. Buccinanops (8 synapomorphies) is
thus restricted to B. cochlidium, B. monilifer, and B. latus, which are large- sized, living in
relative deep waters (subtidal), while Buccinastrum includes small- sized species mostly
restricted to the intertidal zone. The analysis of morphological characters is sugges-
tive of a traditional subfamily Dorsaninae, with 12 synapomorphies, including four
genera— Dorsanum, Bullia, Buccinanops, and the new Buccinastrum, but this result is
only informative, as it has been considered paraphyletic in a DNA sequence approach,
being subdivided into Dorsaninae, Buccinanopsinae, and Buliinae.
KEYWORDS
Anatomy, Argentina, Brazil, Dorsaninae, Mollusks, Nassariidae, Phylogeny, Taxonomy
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1 | INTRODUCTION
The genus Buccinanops d’Orbigny, 1841 presently comprises a
group of the common mostly carnivorous gastropods living in sandy
beaches along the Southwestern Atlantic coast, from southern Brazil
to Tierra del Fuego. Considerable variation of relatively featureless
shell morphology and a broad latitudinal distribution combine to ob-
scure the true number and definition of the species belonging in this
genus. Nevertheless, new collections, a revision of all existing type
materials and museum specimens, and a detailed morphological and
anatomical study, allow us to identify more precisely the status of
each name recorded for this genus throughout its entire distribution
area.
The last comprehensive account of the genus Buccinanops
was in the revision of the Argentine and Uruguayan “Dorsaninae”
by Carcelles and Parodiz (1939), ~80 years ago. In that pioneering
paper, most of the species were included, with good photographs
and radula sketches for the first time. However, no anatomical data
were included, and the range of each species was taken mostly from
old literature, thus veiling the real distribution.
A similar shell morphology probably induced previous authors
(i.e., Allmon, 1990; Carcelles & Parodiz, 1939) to include the genus
Buccinanops with Dorsanum Gray, 1847 and Bullia Gray, 1833, in var-
ious systematic arrangements, always related, despite the fact that
the latter genera are currently living in western and southern Africa,
and in the Indian Ocean. Pastorino (1993) reported some of the ge-
neric and subgeneric combinations of these three genera proposed
by different authors. Also, in a comparative scenario among these
three genera, Simone and Pastorino (2014) described the anatomy
of species of Dorsanum and Bullia from W Africa, concluding that
Buccinanops was recognized as a separate genus. The three gen-
era have been considered related, sometimes as members of the
same subfamily; sometimes as members of monotypic subfamilies—
Buccinanopsinae, Dorsaninae, and Bulliinae (Galindo et al., 2016),
but always within the family Nassariidae.
The analysis by Allmon (1990, text Figure 25) recovered a
monophyletic group (Dorsanum (Bullia – Buccinanops)), supporting
the then new subfamily Bulliinae. However, Haasl (2000, Figure 4)
showed the three taxa in a paraphyletic arrangement, with Bullia-
Buccinanops in one branch, followed by Dorsanum, as sister taxon
of a Photinae/Nassariinae branch. In the molecular genetic study
by Galindo et al. (2016), which used mitochondrial (COI, 16S, 12S)
and nuclear (28S, H3) genes, the three supposedly closely related
genera were retrieved in separated positions in their phylogenetic
trees obtained using maximum likelihood and Bayesian algorithms.
Buccinanops was placed at the base of the Nassariidae, as its first
branch; Bullia appeared in a more internal position, separated from
Buccinanops by samples now considered Photinae and Cylleninae
(formerly buccinids). Samples of Anentominae and a subfamily- indet.
Nassodonta are two branches separating Dorsanum from Bullia, sis-
ter group of remaining nassariids allocated in Nassariinae. The three
genera— Buccinanops, Bullia, and Dorsanum— were thus included in
proper monotypic subfamilies— Buccinanopsinae (described then
as new), Bulliinae Allmon, 1990 and Dorsaninae Cossmann, 1901.
Curiously, Bullia granulosa did not fall within the Bullia branch, but
instead into Nassariine branch of the Naytia group. Only the new
subfamily, that is, Buccinanopsinae had characters defined in that
paper, the set of features is stressed in discussion section herein.
The Galindo et al. (2016) arrangement has been currently ac-
cepted (e.g., WoRMS website, 2020); however, a morphological
definition of these taxa, from subfamily to species level, is still
mostly wanting. Based on a relatively comprehensive nassariid/
buccinid morphological dataset that has been produced by our
team, both in Nassariidae in general (e.g., Abbate & Cavallari, 2013;
Abbate et al., 2018; Couto & Simone, 2019; Gernet et al., 2019;
Simone, 2007, 2011; Abbate & Simone, submitted), and in the taxa
Buccinanops/Bullia/Dorsanum in particular (e.g., Abbate & Simone,
2016; Pastorino, 1993a, b; Simone & Pastorino, 2014), it appears
plausible that these three genera are more closely related based
on morphology, rather than their surprising widespread dispersal
among different branches within Nassariidae, proposed by Galindo
et al. (2016). It is not our intention to contradict the DNA sequence
results (Galindo et al., 2016), as the studied assembly is much smaller
and focused on these three genera. Thus, no taxonomical change
is proposed here in family/subfamily levels. However, as the search
for morphological definitions of the abovementioned taxa has re-
vealed a different scenario, the different result looks interesting to
be exposed, representing a counterpoint, and raising interesting and
well- based discussion.
The present paper includes type material examination, as well
as anatomical features to be added to the conchological and radu-
lar characters traditionally assigned to the genus Buccinanops and
allies. The main objective was a revision of the Recent species that
have been included in Buccinanops s.l.. Furthermore, the relation-
ships between Buccinanops s.l. and other members of the superfam-
ily Buccinoidea were considered, from a phenotypical perspective,
and in comparison with results from other approaches (e.g., DNA
sequence data). The present database is inserted in an already
published phylogenetic approach (Simone, 2011). Additionally, the
buccinoidean representation in that scenario is enriched, with the
intention of a better- based discussion, saving long explanations on
the methodologies.
2 | MATERIALS AND METHODS
Most of the material reviewed for this work belongs to the largest
collections from South American institutions: Museo Argentino de
Ciencias Naturales "Bernardino Rivadavia," Buenos Aires (MACN)
and Museo de La Plata (MLP) from Argentina; Museu de Zoologia
da Universidade de São Paulo (MZSP) and Museu Nacional, Rio de
Janeiro (MNRJ) from Brazil; Museo Nacional de Historia Natural,
Montevideo, Uruguay (MNHNM) and United States National
Museum, Washington D.C., USA (USNM). The type material is
housed in the following European and American museums: The
Natural History Museum, London (NHMUK); Muséum d'Histoire
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naturelle de la Ville de Genève, Switzerland (MHNG); Zoological
Museum - Natural History Museum of Denmark, (ZMUC); Museo
Nacional de Historia Natural, Montevideo, Uruguay (MNHNM);
and Academy of Natural Sciences of Drexel University (formerly
Academy of Natural Sciences of Philadelphia) (ANSP). Fresh mate-
rial was collected from different localities around Brazil, Uruguay,
and Argentina throughout the current geographic distribution of the
Recent species of Buccinanops.
Dissections were performed on ethanol- preserved specimens
for studies of the external morphology and anatomy. Radulae were
prepared according to the method described in Pastorino (2005) and
observed using a Philips XL 30 scanning electron microscope (SEM)
at the MACN. Shell ultrastructure data were procured from freshly
fractured collabral sections taken from the central portion of the lip
on the last whorl of at least, two individuals per taxon.
Most photographs were taken using a Nikon D100 digital camera
with a 60 mm macro lens, except those— usually type specimens—
sent by different curators. All images were digitally processed with
Photoshop CS6 and Corel PhotoPaint X6 software.
Abbreviations used in anatomical drawings: aa: anterior aorta,
ac: siphonal fold anterior to gill, ad: adrectal sinus, ae: anterior
esophagus, af: afferent gill vessel, ag: albumen gland, an: anus, au:
auricle, bg: buccal ganglion, bc: bursa copulatrix, br: subradular
membrane, bv: blood vessel, cc: cerebral commissure, ce: cerebral
ganglion or cerebro- pleural ganglion, cg: capsule gland, cl: cement
gland, cm: columellar muscle, cs: columellar muscle secondary flap,
cv: ctenidial vein, dd: duct to diges t ive gl and, df: dors al fo ld of bu c-
cal mass, dg: digestive gland, di: diaphragmatic septum, ed: epipo-
dial tentacle, ek: efferent renal vessel, ep: posterior esophagus,
es: esophagus, fp: female pore, ft: foot, fs: foot sole/mesopodium,
gd: duct of gland of Leiblein, ge: subesophageal/visceral ganglion,
gi: gill, gl: gland of Leiblein, gm: gill muscle, go: gonopericardial
duct, gp: pleural ganglion, gv: visceral ganglion, he: hemocoel, hg:
hypobranchial gland, ig: ingesting gland, in: intestine, ki: kidney,
kl: kidney lobe, km: kidney membrane with pallial cavity, m1– m11:
odontophore muscles, mb: mantle border, me: mid esophagus, mj:
jaw/peribuccal muscle, mo: mouth, mt: mantle, ne: nephrostome,
ng: nephridial gland, nr: nerve ring, nv: nerve, oa: opercular pad,
oc: odontophoral cartilage, od: odontophore, op: operculum, os:
osphradium, ot: oral tube, ov: pallial oviduct, oy: ovary, pa: pos-
terior aorta, pb: proboscis, pc: pericardium, pd: penial duct, pe:
penis, pg: anterior furrow of pedal glands, pl: pedal ganglion, pn:
penial gland, pp: penial papilla, pr: propodium, pt: prostate, py:
pallial cavity, ra: radula, rn: radular nucleus, rm: retractor muscle
of proboscis, rs: radular sac, rt: rectum, rv: afferent renal vessel,
ry: rhynchostome, sa: salivary gland aperture, sm: stomach mus-
cle, sc: subradular cartilage, sd: salivary duct, se: septum between
odontophore and esophageal portions of buccal mass, si: siphon,
sg: salivary glands, sp: salivary duct swollen region, ss: subterminal
bulged portion of salivary duct, st: stomach, sv: seminal vesicle, te:
tentacle, tg: integument, tm: transverse muscle between esopha-
gus and odontophore, ts: testis, vd, vas deferens, ve: ventricle, vg:
vaginal atrium, vl: valve of Leiblein, vo: visceral oviduct.
In text spm: specimen(s).
The phylogenetic analysis methodology described by Simone
(2011) was applied, which basically used the TNT and Winclada soft-
wares, with no special additional algorithms. A total of 14 new lines
were added to the cladogram published by Simone (2011) and filled
with characters of each species (Table 1) representing the ingroup
Buccinanops and other taxa of Buccinoidea. No character needed
to be added to Simone's list of characters, yet, some states were
essential to improve the resolution of the buccinoideans currently
introduced, as follows:
2.1 | Characters in Simone (2011)
113: added state 5 = pair of posterior epipodial tentacles (Bullia,
Dorsanum).
127: added state 3 = located lateral to short transverse flap
(Buccinanops, Bullia, Dorsanum).
206: added state 4 = right edge convex (B. deforme, B. paytense,
B. uruguayense).
316: added state 3 = pair of lateral, dorso- ventral thin muscles
(Buccinanops, Buccinastrum, Bullia, Dorsanum).
332: added state 5 = laterally positioned, like lateral protractors
(Buccinanops, Buccinastrum).
340: added state 2 = insertion on radular sac, radular nucleus
and posterior end of cartilages (buccinoideans).
341: added state 3 = m2c (along median line – Bullia).
343: added state 3 = as circular thin fibers as opposite of m6
(buccinoideans).
344: a d de d st at e 12 ( C) = m3 l lo ng it u di na l la t er al pa ir ( Buccinanops,
Buccinastrum) (actually 678).
352: added state 2 = multiple origins in outer edge of cartilages
(Buccinanops, Buccinastrum except B. duartei, Bullia, Dorsanum).
359: added state 3 = originated in opposed side of m4 in carti-
lages (buccinoideans).
377: added states 3 = dorsal, Y- shaped (Buccinanops,
Buccinastrum, Bullia, Engoniophos, Dorsanum, Buccinum?); 4 = similar,
but multiple (2 or more) (B. deforme, B. uruguayense); 5 = similar, but
inserting directly in cartilages instead of m3 (Bullia).
384: added state 4 = originated at cartilages posterior end by
side of m5 (buccinoideans).
395: added state 2 = absent but with lateral arrangement
connected to anterior end of cartilages only (buccinoideans,
volutoideans).
418: added states: 4 = arched, with 2– 3 cusps at edge (Pugilina,
Pustulatirus, Hemipolygona); 5 = arched, with ~10 cusps at edge
(Engoniophos, Buccinanops, Buccinastrum, Buccinum); 6 = arched, with
~20 cusps at edge (Bullia, Dorsanum).
424: added state 4 = wide, oblique, with long cusp in each end,
being lateral cusp larger (Pugilina, Bullia, Dorsanum, Engoniophos,
Buccinum, Buccinanops, Buccinastrum).
426: added state 3 = presence of intermediar y small cusps (Bullia
laevissima, Buccinum, Buccinanops).
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450: added state 5 = very narrow and long, running attached to
esophagus by side of aorta branch (buccinoideans).
452: added states 3 = close to median line, ventral (Pugilina,
B. cochlidium, B. latus, B. monilifer); 4 = lateral, close to dorsal folds
(Bullia, Dorsanum, Engoniophos, Pustulatirus, Hemipolygona, B. du-
artei); 5 = lateral, far from dorsal folds (B. deforme, B. paytense);
(B. uruguayense = 0).
478: added state 3 = broad, thick walled (Buccinastrum, Bullia).
485: added state 4 = absent, but valve only vestigial (B. latus,
B. monilifer, B. cochlidium).
493: added state 2 = similar, but with bulged portion towards
posterior (Buccinanops, Buccinastrum, Bullia, Dorsanum) (additive).
554: added state 2 = gonopericardial duct present (Buccinanops,
Buccinastrum, Bullia).
575: added state 3 = in subterminal exposed small papilla
(Buccinanops, Buccinastrum).
660: added state 2 = with buccal commissure imbibed in nerve
ring (Buccinanops, Buccinastrum).
667: added states 2 = concentrated, but with cerebral commis-
sure long, forming small strap- like arc (Buccinanops, Buccinastrum,
Bullia, Dorsanum); 3 = same, but with commissure very long (about
2– 3 times longer) (B. latus, B. monilifer, B. cochlidium).
The resulted consensus cladogram is presented in the Figure 35,
with the synapomorphies of each node presented. A few states pre-
sented ambiguity in the optimization in the cladogram, in these cases
the acctran algorithm was used in that Figure. The characters with
minor ambiguity in some branches are: 9, 180, 331, 377, 418, 424,
426, 478, 498.
3 | RESULTS
3.1 | Systematics
3.1.1 | Subfamily Buccinanopsinae Galindo,
Puillandre, Louzet & Bouchet, 2016
Additional diagnosis: Fusiform shell from obese to relatively elon-
gated. Sculptured mainly with undulating growth lines. Suture well-
marked. Aper ture simple, from 1/3 to 1/2 of shell leng th. Outer lip
simple, non- determinate growth. Siphonal canal simple, wide, non-
protruded. Callus normally thin. Head- foot usually unpigmented,
lacking eyes; head flap- like, with tentacles on both sides. Siphon
long, bearing small fold in its base. Lateral, dorso- ventral muscles
uniting ante rior regi ons of esophagus and odonto phore. Pair of od-
ontophore retractor muscles (m2) with multiple origins. Rachidian
flattened, arched, ~20 cusps in cutting- edge. Stomach with poste-
rior bulge. Penis duct thick- walled, convolute, muscular. Nerve ring
with cerebral commissure long, widely arched.
Included taxa: Genera: Buccinanops (type), Buccinastrum gen. nov.
3.1.2 | Genus Buccinanops d'Orbigny, 1841
Type species: Buccinum cochlidium Chemnitz, 1795 (=B. cochlidium
Dillwyn, 1817) by original designation (d’Orbigny, 1841: 432). Chemnitz's
work was rejected by the ICZN Direction 1 (1954). Cernohorsky (1984:
23) mentioned Herrmannsen (1846) as the source of designation.
TAB LE 1 Species added to the database in Simone (2011) to enlarge the number of samples of Buccinoidea
Species Family Source
Oligohalinophila dorri (Wattebled, 1886) Nassariidae Simone (2007, 2011) (as Nassodonta)
Amphissa acuminata (Smith, 1915) Columbellidae Simone and Leme (2001); Simone (2011)
Amphissa cancellata (Castellanos, 1979) “ “
Pustulatirus ogum (Petuch, 1979) Fasciolariidae Couto et al. (2015)
Hemipolygona beckyae (Snyder, 2000) “ “
Pugilina morio (Linnaeus, 1758) Melongenidae Abbate and Simone (2015)
Pugilina tupiniquim Abbate & Simone, 2015 “ “
Buccinum undatum Linnaeus, 1758 Buccinidae Wilsmann (1942); person. obs.
Engoniophos unicinctus (Say, 1826) Nassariidae Abbate et al. (2018)
Dorsanum miran (Bruguière, 1789) Dorsaninae Simone and Pastorino (2014)
Bullia granulosa (Lamarck, 1822) “ “
Bullia laevissima (Gmelin, 1791) “Abbate and Simone (2016)
Buccinanops monilifer (Kiener, 1834) “Simone (1996, 2011), this study
Buccinanops latus sp. nov. “ “
Buccinanops cochlidium (Dillwyn, 1817) “This study
Buccinastrum duartei (Klappenbach, 1961) “ “
Buccinastrum uruguayense (Pilsbry, 1897) “ “
Buccinastrum deforme (King, 1832) “ “
Buccinastrum paytense (Kiener, 1834) “ “
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However, while on page 127 of Herrmannsen (1846) it is clearly stated
that cochlidium is the type species, it is evident that Herrmannsen was
only following d’Orbigny's original designation and not designating the
type species himself. Allmon (1990: 20) wrongly stated that the type
species was Buccinum globulosum Kiener, 1834. No reference to this
type designation can be found in d’Orbigny (1841). Buccianops Adams
& Adams, 1858: 113 is a typing error pro Buccinanops. The gender is
masculine according to the art. 30.1.4.3 of the ICZN (1999).
Diagnosis: Shell bucciniform, medium or large, smooth, slightly
shining; walls thin. Sometimes with shallow subsutural flat cords
and/or small sharp nodes or spines along keel in last two or three
whorls. Flat cords also covering basal surface of last whorl. First
teleoconch whorls with axial riblets. Spire variable, always less
than 1/3 of maximum shell height. Weak terminal columellar
fold always present. Parietal callus variable, ranging from thin
glazed area to thicker protuberant callus. Operculum large with
smoot h margins and terminal nucleus. Duplicity of pair of ext rinsic
odontophore muscles m1b; aperture of salivary glands ventral,
close to median line; simplified anterior esophagus; papilla- like fe-
male genital pore. Radular central tooth multicuspidate, with 5– 12
cusps increasing in size towards middle of tooth, rachidian base
strongly curved; lateral teeth with inner and outer cusps hook-
shaped, always with 1 to 3 intermediate cusps. Egg capsules al-
ways at tached on the callus area of female shells only. Nur se eggs,
non- planktonic larvae and crawling veliger.
Species included: B. cochlidium, B. latus sp. nov., and B. monilifer.
3.1.3 | Buccinanops cochlidium (Dillwyn, 1817)
(Figures 1– 8)
Buccinum cochlidium Chemnitz, 1795: 275, pl. 209, figs. 2053,
2054 (invalid name); Dillwyn, 1817: 627. Kiener, 1834: 10, pl. 6, fig.
17; Anton, 1838: 91; Deshayes, 1844: 187.
FIGURE 1 Buccinanops cochlidium type shells. (a– b) Buccinum cochlidium specimen figured by Chemnitz, 1795, ZMUC unnumbered (L
77.3 mm). (c) Chemnitz’ (1795) plate 209, Figs. 2053, 2054; (d) Kiener’s (1834) plate 6, Figure 17. (e– f) Buccinum gradatum Deshayes, 1844,
probable specimen figured in Kiener (1834) (L 43.1 mm), MHNG 1296/18. (g– h) Bullia gradata pampeana Ihering, 1907, holotype, MACN- Pal.
855, from Quaternary deposits of Puerto Belgrano, Buenos Aires (L 28.5 mm). (i– j) Buccinum lamarckii Kiener, 1834, probable Kiener’s (1834)
specimen, MHNG 1296/4 (L 64.2 mm). (k) Kiener’s (1834) plate 6, Figure 6. Scale bar = 1 cm for the shells only
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Buccinum Lamarkii (sic) Kiener, 1834: 5, pl. 3, fig. 6.
Buccinanops cochlidium: d'Orbigny, [1839] pl. 61, fig. 25; M. E.
Gray, 1850: 20, 139, pl. 98, fig. 9; Carcelles & Parodiz, 1939: 751,
figs. 2, 6; Carcelles, 1944: 250.
Bullia Cochlidium: Gray, 1839: 126.
Buccinum (Buccinanops) cochlidium: d'Orbigny, [1841]: 434.
Buccinum gradatum Deshayes, 1844: 186.
Bullia gradata: Reeve, 1846: pl. 1, fig. 3; Chenú, 1859: 160; Peile,
1937: 185, fig. 21; Abbott & Dance, 1986: 177, fig. 9.
Bullia cochlidium: Reeve, 1847: pl. 4, fig. 23; Hanley, 1856: 117, pl.
23, fig. 95; Chenú, 1859: 159, fig. 747.
Bullia (Buccianops) (sic) cochlidium: H. & A. Adams, 1858: 113.
Bullia (Buccinanops) cochlidium: H. & A. Adams, 1858: 113; Tryon,
1882: 13, pl. 5, fig. 73, pl. 6, figs. 76, 78.
Buccinanops (Buccinanops) cochlidium: Fischer, 1884: 636.
Bullia (Buccinanops) gradatum: Watson, 1886: 190.
Bullia (Buccinanops) cochlidium: Paetel, 1888: 116.
Bullia cochlidium: Cossmann, 1901: 220.
Bullia gradata pampeana Ihering, 1907: 445; Feruglio, 1933: 236,
239.
Buccinanops (Buccinanops) cochlidium: Thiele, 1929: 32.
Buccinanops gradatum: Carcelles & Parodiz, 1939: 754, pl. 3– 5, fig.
3; Barattini & Ureta, 1960: 114; Castellanos, 1970: 92, pl. 7, fig. 6; Rios,
1970: 93, pl. 27, 1985: 103, pl. 35, fig. 453; Penchaszadeh, 1971a, 1971b:
480, figs. 1e, 2f, 2g, 1973: 15, fig. 1; Scarabino, 1977: 188, pl. 5, fig. 4;
Cernohorsky, 1984: 28; Farinatti, 1985: 219; Calvo, 1987: 143, fig. 110.
Buccinanops cochlidium: Barattini & Ureta, 1960: 113, 114;
Castellanos, 1970: 93, pl. 7, fig. 10; Cernohorsky, 1984: 27, figs.
FIGURE 2 Buccinanops cochlidium shells and operculum. (a– b) MACN- In 20099, Ostende, Buenos Aires province (L 80.4 mm). (c– d) MZSP
80628, from Chubut, Argentina (voucher by Galindo et al., 2016) (L 75.5 mm). (e) MZSP, female, Playa Villarino, Golfo San José, Chubut,
Argentina (L 83.0 mm). (f– g) MACN- In 43749 Mar del Plata (L 37.0 mm); (h– i) MZUSP, male, Playa Villarino, Golfo San José, Chubut, Argentina
(L 72.5 mm). (j– l), MACN- In 43749, Mar del Plata, female (L 67.0 mm). (m– n) operculum of the specimen in ( j– l) (L 34 mm)
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94– 95; Aguirre, 1993: 33, pl. 2, figs. 2 a- b; Pastorino, 1993a: 162, fig.
4– 6; 1993b: 153, figs. 2– 5.
Buccinanops lamarckii: Richards & Craig, 1963: 140; Rios, 1970:
93; Rios, 1975: 96, pl. 27, fig. 401; Scarabino, 1977: 189, pl. 5, fig. 3;
Rios, 1985: 103, pl. 35, fig. 454; Calvo, 1987: 143, fig. 111.
Bullia (Buccinanops) cochlidia: Allmon, 1990: 20, pl. 3, figs. 1, 13,
pl. 6, fig. 34.
Type material: Buccinum cochlidium: holotype ZMUC, figured
by Aguirre (1993a, pl. 2, Figure 2a– b), here in Figure 1a– c; B. gra-
datum: holotype MHNG 1296/18 (Figure 1d– f); B. lamarckii: holo-
type MHNG 1296/4 (Figure 1i– k); Bullia gradata pampeana: holotype
MACN- Pal 855 (Figure 1g– h).
Type localities: Buccinum cochlidium: Islands of South seas (in
error); B. gradatum: New Zealand (in error); B. lamarckii: unknown,
no locality mentioned; Bullia gradata pampeana: “dépôts pampiens
de Puerto Militar” (Quaternary deposits at Puerto Belgrano, Buenos
Aires province).
3.1.4 | Description
Shell: (Figures 1– 2) Large (maximum height 95 mm), somewhat vari-
able, usually fusiform. Protoconch paucispiral of 1.5 to 2 whorls,
1.5– 1.7 mm wide in second whorl; first whorl smooth, followed
by 10– 12 weak axial riblets after second whorl (Figure 3a– b).
Teleoconch of 6– 7 shouldered or globose whorls. Spire rounded,
~1/3 of total shell length. Suture well defined, impressed; blunt sub-
sutural carina in ~50% of specimens; aperture large, usually half of
total shell leng th. Pariet al callus usually weak, stronger in large spec-
imens. Sculpture absent except for growth lines and shallow undula-
tions. Left edge of siphonal canal with pointed anterior beak of ~30°.
Ultrastructure of shell usually with three layers, outermost layer
very thin or absent, of amorphous calcite, intermediate layer thicker,
of crossed- lamellar aragonite, with crystal faces oriented collabrally,
and innermost layer thinner, of crossed- lamellar aragonite oriented
perpendicular to outer lip.
Head- foot: (Figures 3a– c, 4a– c) Head weakly protruded, some-
what socket- like, narrow (~50% of head- foot width); color dark-
brown, almost black (Figure 3c) in exposed areas (except for foot
sole and opercular pad), particularly darker in head and in median-
dorsal area of foot. Tentacles well- separated from each other;
elongated and narrow, eyes absent (Figure 4a– c). Rhynchostome
as longitudinal slit located in middle region of lightly protruded
head ventral surface (Figure 4a– c: ry). Foot broad, of about 3/4
whorl when retracted. Sole rather triangular, edges thick and
rounded (Figure 4a– b: fs). Anterior furrow of pedal glands almost
FIGURE 3 Buccinanops cochlidium anatomical photographs and SEM. (a) foot, transversely sectioned along sagittal plane, hear removed,
scale = 10 mm. (b) posterior region of foot, dorsal view, scale = 10 mm. (c) detail of posterior end of foot, dorsal view, operculum removed,
showing black color absent in opercular pad (MZSP 133346), scale = 5 mm. (d) penis tip, SEM, scale = 1 mm. (e– f) radula, frontal and lateral
views, scales = 200 µm. (g– h) two views of a prehatching young shell, scale = 1 mm
8
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PASTORIN O ANd SIMO NE
imperceptible, marked by narrow furrow in anterior edge (Figures
3a: pg, 4b); pedal gland thick in medial region, immersed in pedal
musculature (Figure 3a: superior pg). Single, small epipodial tenta-
cle (ed) located in dorso- posterior end of foot, originating in dorsal
region of edge posterior to operculum. Opercular pad elliptical,
~50% as wide as dorsal surface of foot; attachment with opercu-
lum occupying ~80% of its area (Figures 3c, 4b: oa). Columellar
muscle thick, spanning, ~1.5 whorl; distal end bifid, right element
wide (cm), left element narrow (cs), encas ed in furrow kept by colu-
mellar fold (Figure 4a- b). Female with small orifice of cement gland
located on median line of anterior sole region (Figure 3a: cl); inner
space wider than narrow duct.
Operculum: (Figures 2m- n, 4a: op) Elliptical, corneous, pale
brown, translucent. Nucleus terminal in inferior corner. Outer edge
straighter than inner edge. Outer surface with concentric growth
lines, forming undulations. Scar oval, occupying about 80% of inner
area, somewhat dislocated closer to inner edge.
Pallial cavity: (Figure 5a, d) Mantle edge simple, thick. Siphon long,
slender, ~10% of mantle edge length, almost as long as 2/3 of pallial
cavity when retracted (Figure 5a: si); edges simple. Osphradium of
~1/2 length, and 1/10 width of pallial cavity; anterior end pointed,
but more rounded than narrow, tail- like posterior third (Figure 5a: os);
osphradium filaments symmetrical, relatively low, isosceles- triangular
(Figure 5d: os). Very narrow area between osphradium and gill.
Ctenidial vein (cv) narrow, with uniform width along its length. Gill (gi)
~80% of pallial cavity length, ~1/2 its width at widest middle region;
anterior end broadly pointed, located at some distance from mantle
edge. Posterior gill end slightly more pointed than anterior end, touch-
ing pericardial area. Afferent gill vessel very narrow, lying at short dis-
tance from right edge of gill (Figure 5d: af). Gill separated from right
edge of pallial cavity by area equivalent to its width. Hypobranchial
glan d thin, lig ht- br own, covering most of area betwe en gill and rect um,
slightly thicker in anterior quarter, vaguely pigmented light- brown.
Rectum broad, with thick walls, running along right edge of pallial
FIGURE 4 Buccinanops cochlidium anatomy. (a) head- foot, male, right- slightly anterior view. (b) same, female, operculum extracted. (c)
head and hemocoel, ventral view, foot and columellar muscle removed, some structures deflected. Scales = 5 mm
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PASTORIN O ANd SIMO NE
cavity (Figure 5d: rt). Anus simple, siphoned (terminally detached),
located near mid length of pallial cavity (Figures 5d, 7c: an). Pallial go-
noducts located between rectum and pallial right edge as described
below.
Visceral mass: (Figure 5a) Anterior quarter of whorl mostly oc-
cupied by kidney (km) and pericardium (pc). Digestive gland light-
brown, located along inferior (siphonal) region of each visceral
whorl, covering middle digestive tubes and two whorls posterior to
stomach. Gonad pale light- brown, lying along superior and columel-
lar surfaces of visceral whorls posterior to stomach. Stomach small,
located half whorl posterior to pallial cavity (Figure 5a: st).
Circulator y and excretory systems: (Figure 5a) Pericardium located
just posterior to gill, along left anterior region of visceral mass (pc),
slightly dislocated anteriorly along right side of posterior 1/10 of gill.
Auricle (au) small, attached to anterior surface of pericardium, with
ctenidial vein entering from left and connected to kidney at its right
end. Auricle connected to anterior surface of ventricle. Ventricle (ve)
large, rounded. Aortas located along postero- left region of ventricle.
Kidney volume equivalent to ~1/4 of pallial cavity volume, 4- times
larger than pericardium; located along middle and right regions of
anterior end of visceral mass. Nephridial gland (ng) triangular in sec-
tion; lying along dorsal- right region of reno- pericardial wall. Renal
FIGURE 5 Buccinanops cochlidium anatomy. (a) pallial cavity and anterior portion of visceral mass, ventral view, digestive gland portion
ventral to stomach removed, kidney opened longitudinally and deflected, ventral pericardial wall removed. (b) foregut, right view, mostly
uncoiled, right transverse muscles (tm) sectioned. (c) detail of insertion of gland of Leiblein, esophagus wall sectioned and rectified (scale as
Figure 17). (d) pallial cavity roof, transverse section in middle level of osphradium (os). (e) region of insertion of gland of Leiblein, outer view.
(f) region of vestigial valve of Leiblein, opened longitudinally. Scales = 5 mm
10
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PASTORIN O ANd SIMO NE
lobe (kl) in three longitudinal, thick flaps, turned to left, transversely
folded, occupying most of kidney's interior volume (Figure 5a: kl);
intestine running along kidney right side. Efferent renal vessel (ek),
well- developed, inserted in dorsal kidney lobe flap.
Digestive system: (Figures 4c, 5b, c, e– f, 6) Proboscis as long as
foot, 1/4 of head- foot width (Figure 4c: pb). Circular mouth at pro-
boscis tip (Figure 5b: mo). Buccal cavity with pair of broad and low
dorsal folds (Figure 6a: df), occupying ~1/3 of dorsal wall, with inter-
val between them slightly broader than each fold. Odontophore oval,
as long as retracted proboscis (Figure 5b: od). Odontophore tube
connecting it with buccal cavity, length equivalent to ~1/5 of odon-
tophore length, possessing mostly longitudinal muscles (Figure 6a, c:
ot). Odontophore muscles (Figure 6a– f ): m1, several small jugal mus-
cles connecting buccal mass to adjacent inner surface of proboscis;
one pair more developed originating in peribuccal lateral region of
proboscis, running posteriorly, inserting in lateral side between mid-
dle and posterior thirds of odontophore (Figure 6b– c: m1l); tm, pair
of transverse muscles connecting lateral walls of odontophore with
adjacent wall of esophagus (Figures 5b, 6a: tm), along anterior third
of odontophore; set of m2, several retractor muscles of odonto-
phore, part of them working also as dorsal tensor muscles of radula,
all originating in ventral surface of hemocoel, in region just posterior
to proboscis, running anteriorly, some median fibers running through
nerve ring, insertions as following: m2m, median m2, single, inserting
FIGURE 6 Buccinanops cochlidium anatomy. (a) buccal mass, right view, mostly opened longitudinally, odontophore only partially shown.
(b) odontophore, dorsal- slightly right view. (c) same, dorsal view, first layer of muscles sectioned and deflected. (d) same, dorsal view,
some more superficial muscles partially removed. (e) same, right- slightly dorsal view, more dorsal muscles deflected to show more internal
structures. (f) odontophore cartilages, dorsal view, most intrinsic muscle partially removed, showing only their origin and some adjacent
muscles. Scales = 2 mm (all same scale)
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PASTORIN O ANd SIMO NE
on radular nucleus together with small branch of aorta (Figure 6b– e);
m2a, broad and thick pair auxiliary of m2 and dorsal tensor mus-
cles of radula, with 4– 5 separated pairs of bundles, inserting along
radular sac in its middle- lateral regions (Figure 6b– e); m2c, lateral
odontophore retractors, inserting at posterior end of each odonto-
phore cartilage (Figure 6b– e); m3c, single circular, constrictor muscle
covering? dorsal surface of odontophore, inserting along outer side
of dorsal edge of cartilages at along ~70% their length (Figure 6b– f);
m3l, pair of narrow, lateral longitudinal muscles, originating in lateral
region of odontophore tube, running posteriorly attached to odon-
tophore lateral surface, inserting in fan- like manner lateral side of
m3c posterior surface (Figure 6b); m4, strong pair of dorsal tensor
radular muscles, originating in odontophore cartilages on posterior-
ventral surface, running toward dorsal surrounding lateral surface
of cartilages, inserting laterally along radular sac (Figure 6b– f); m4a,
4– 5 small pairs auxiliary of m4, of similar attributes, but originating
in bundles along outer- dorsal edge of cartilages, in their posterior
quarter (Figure 6c– f); m5, pair of secondary dorsal tensor muscles
of radula, originating in posterior- dorsal surface of posterior end of
cartilages, running dorsally and medially mixing with m4, inserting
into radular sac alongside and medial to m4 insertion (Figure 6e– f);
m6, thin horizontal muscle, uniting both odontophore cartilages just
posterior to anterior fusion, along ~70% of cartilage length, insert-
ing along ventral and internal edge of cartilages, becoming gradu-
ally wider posteriorly (Figure 6e– f); m8, pair of cartilages shortener
muscles, located along dorsal edge of anterior region of cartilages,
along almost 1/3 their length, thicker anteriorly, gradually narrowing
posteriorly (Figure 6d– f); m9, two pairs of narrow protractor muscles
of radula, one located just anterior from another (posterior slightly
broader), originating on posterior half of dorsal edge of cartilages,
running medially towards anterior, inserting along m3c anterior fi-
bers (Figure 6c– d); m11, pair of ventral tensor muscles of radula,
FIGURE 7 Buccinanops cochlidium anatomy. (a) middle region of male genital structures, ventral view, some adjacent structures also
shown. (b) penis and adjacent region of pallial loor, dorsal view, portion of integument artificially removed in middle level of pallial vas
deferens (vd), penis duct (pd) shown if it was translucent. (c) pallial oviduct and adjacent structures, ventral view, transverse section in middle
level of oviduct artificially done. (d) same, isolated pallial oviduct opened longitudinally and deflected to show inner surface; (e) penis, detail
of apical region, dorsal view. Scales = 2 mm
12
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PASTORIN O ANd SIMO NE
thin, narrow, originating in posterior- ventral end of cartilages, run-
ning anteriorly, and medially covering m6, inserting into ventral edge
of radula and subradular cartilage, and some inner portion preceding
this (Figure 6e). Subradular cartilage expanding in exposed region
of radula into buccal cavity, covering neighboring surface of radula
(Figure 6c: sc); oc, odontophore cartilages, elongate, furrow- like,
flattened, ~20 times longer than wide; fusion between both carti-
lages in anterior- medial end, along ~20% their length (Figure 6e– f);
subradular membrane, covering inner surface of subradular carti-
lage and radula, having insertions of m4, m5 and m11 (Figure 6d– e).
Radula (Figure 3e– f): slightly longer than odontophore, ~50 rows of
teeth; rachidian tooth wide, comb- like, occupying ~half of radular
width; base curved, width ~3 times greater than its length; ~9 long,
sharp pointed cusps of similar size, except for some diminishment
toward lateral edges, lateral cusps very small, sometimes vestigial;
distance between rachidian and lateral tooth equivalent to 1/3 of
rachidian width; lateral tooth with two major and 2– 4 minor long
cusps; base equal in width to rachidian tooth; base deflected pos-
teriorly by ~50 degrees; outer cusp about as long as tooth base,
scythe- shaped; inner cusp shorter, stout, with 2– 4 shorter cusps
of decreasing size emerging from its surface outer cusp and set of
inner cusps separated from each other by smooth area equivalent
to 1/4 tooth's width. Salivary glands enveloping esophagus along
region of valve of Leiblein and nerve ring (Figures 4c, 5b: sg), size
slightly larger than nerve ring; their ducts very narrow, running along
anterior esophagus wall and, more anteriorly, inside dorsal folds of
buccal cavity (Figure 6a: sd); opening with very small pore (Figure 6a:
sa), into anterior- middle region of dorsal folds of buccal cavity; sal-
ivary ducts with enlarged bulges along middle area of dorsal folds
(Figure 6a: ss). Anterior esophagus with somewhat thick walls, twice
length of odontophore; inner surface with 12– 15 narrow, uniform
longitudinal folds; posterior half zigzagging intensely (Figures 4c, 5b:
ae). Valve of Leiblein almost imperceptible externally; inner struc-
ture only marked by transverse wide glandular fold (Figure 5f: vl),
lacking any valve or long cilia; smooth area anterior and posterior to
transverse fold (Figure 5f); oblique furrow (bypass) absent. Middle
esophagus ~5% of anterior esophagus length (Figure 5e: me), walls
thin; inner surface like that of anterior esophagus; aperture of gland
of Leiblein as a minute pore. Gland of Leiblein narrow, elongated
(Figures 4c, 5b: gl), ~70% posterior esophagus length; anterior re-
gion ~double of middle esophagus width, gradually tapering toward
posterior; middle portion surrounding local portion of anterior
aorta (Figures 4c, 5b: aa). Duct of gland of Leiblein short, narrow,
simple (Figure 5b, e: gd). Posterior esophagus (Figures 4c, 5b: ep)
~70% anterior esophagus length, very wide (4– 5 times wider than
wider portion of anterior esophagus), slightly flattened, narrowing
gradually posteriorly (Figure 5a: ep) in sam e width as anterior esoph-
agus; inner surface with 12– 15 narrow, tall, irregular, sometimes
coalescent longitudinal folds. Stomach small blind sac, ~1/4 width
of adjacent visceral whorl, ~1/4 whorl long; located ~half whorl pos-
terior to pallial cavity, approximately in middle position of that whorl
(Figure 5a: st). Esophageal insertion and intestinal origin in anterior
side of stomach relatively well- separated from each other; 2 narrow
ducts to digestive gland, being one in esophageal insertion, turned
posteriorly, another in intestinal origin, turned left (Figure 5a: dd).
Gastric inner surface mostly smooth. Intestine about as wide as ad-
jacent posterior esophagus; weakly sigmoid visceral loop (Figure 5a:
in) immersed initially in digestive gland, afterwards passing though
ventro- left side of kidney, immersed in renal lobe. Rectum and anus
described above (pallial cavity).
Genital system: Male: (Figures 4a, 7a, b, e) Gonad along columel-
lar region of each visceral whorl. Intensely coiled seminal vesicle
(Figure 7a: sv) along ~1/2 whorl in region preceding pallial cavity;
placed slightly obliquely; more posterior region narrower, gradually
widening anteriorly; narrowing in region preceding pallial cavity in
short zigzag in central- left region of visceral anterior border, just pos-
terior to nephrostome (Figure 7a: ne). gonopericardial duct relatively
wide, inserting in vas deferens at end of its coiled portion (Figure 7a:
go). Prostate spanning about 1/3 pallial cavity length, narrow, closed
(tubular), lying ventrally and to right of rectum (Figure 7a: pt). In re-
gion posterior to anus, vas deferens gradually crossing to pallial cavity
floor, narrow; walls relatively thick, running relatively straight up to
penis base (Figure 7b: vd) immersed in pallial floor integument. Penis
slender, curved backwards in base, about ~80% of pallial cavity length,
~15 times longer than its width, dorso- ventrally flattened (Figures 4a,
7b: pe); base narrow, located at some distance from right tentacle
(Figure 4a); middle level twisting, narrowing gradually up to blunt tip,
with small right papilla (Figures 3d, 7b, e: pp). Penial duct running ap-
proximately along penis center, narrow, intensely coiled along its en-
tire length, slightly less coiled and narrow in distal 1/5 (Figure 7b: pd).
Penial aperture in apex of papilla (Figure 7e: pd), very small.
Female: (Figure 7c– d) Visceral oviduct narrow, running along
middle region of columellar surface of last whorl of visceral mass,
closely preceding pallial cavity (Figure 7c: vo). Posterior region of pal-
lial oviduct protruding into kidney. Albumen, whitish, (Figure 7c: ag)
and capsule, pale beige, (cg) glands adjacent, albumen gland spanning
posterior 1/5 of pallial oviduct. Seminal receptacle absent. Capsule
gland with flat lumen, vaginal furrow running along its left edge,
FIGURE 8 Buccinanops cochlidium central nervous system.
(a) dorsal (anterior) view. (b) ventral view. Scale = 2 mm
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PASTORIN O ANd SIMO NE
with surface smooth (Figure 7d: cg). Vaginal duct as tapering region
preceding female pore (Figure 7c– d: vg), wall thin, inner surface
narrow longitudinal folds. Female pore narrow, weakly protruded,
papilla- like, sheltered in shallow cavity (Figure 7c: fp). Bursa copula-
trix absent. Cement gland in pedal sole described above (head- foot)
(Figure 3a: cl).
Central nervous system: (Figure 8a– b) Well- concentrated, most
connectives and commissures indistinct; total nerve ring volume
~1/30 that of hemocoel. No distinction between pleural and cerebral
ganglia (ce). Each cerebral/pleural ganglia ~1/4 of ner ve ring volume.
Cerebral commissure very long, strap- like, extending dorsally ~30%
more than nerve ring size; narrow in middle, thickening on both sides,
without clear distinction with cerebral ganglia (Figure 8a– b: cc). Pedal
ganglia (pl) slightly smaller than cerebro- pleural ganglia; located just
ventral to cerebro- pleural ganglia as almost single mass; deep furrow
separating both pedal ganglia (Figure 8b: pl). Buccal ganglia (Figure 8a:
bg) small, located side by side in region close to left cerebral ganglion.
Visceral ganglion (Figure 8a: vg) ~1/4 of pedal ganglion size, located
3- times nerve ring length posteriorly to it. Subesophageal ganglion
not detected, possibly embedded within nerve ring.
Habitat: Inhabits sandy bottoms, from 6 to ~50 m depth. It feeds
primarily on dead fishes and crustaceans. It is frequently found as-
sociated with the sea anemone Antholoba achates (Drayton in Dana,
1846) (as Phlyctenanthus australis in Pastorino, 1993) in the Peninsula
Valdes area (~42°S), living on the dorsum of the shell. Occasionally it
has been observed feeding on live bivalves.
Distribution: From Rio Grande do Sul, Brazil (~32°S) to Puerto
Madryn, Golfo Nuevo, Chubut, Argentina (~42°S).
Material examined: BRAZIL. Rio Grande do Sul. Rio Grande,
Tramandaí, MZSP 77556, 1 spm, MNRJ 6092, 2 spm; Praia do
Imbe, MNRJ 2923, 2 spm, MNRJ 6062, 2 spm; Cassino, MZSP
61463, 7 spm, MZSP 19197, 3 spm, MZSP 44023, 2 spm, MACN- In
8221, 2 spm, MNR J 690, 2 spm, MNRJ 3467, 3 spm, MNRJ 4172,
1 spm; Albardão, MZSP 16203, 2 spm; MZSP 51731, 1 spm in 13 m,
32°58'S, 52°30'W; MZSP36633, 1 spm; MZSP 53201, 3 spm; 531,
6 spm; MZSP 19178, 3 spm; MZSP 19179, 5 spm; MZSP 19180,
3 spm; MZSP 19181, 1 spm; MZSP 19182, 3 spm; MZSP 19184,
1 spm; MZSP 19186, ~10 spm; 29°43′S 49°55′W, 24 m, MZSP
19187, 1 spm (GEDIP sta. 1715, RV W. Besnard col, 8.iv.1972);
MZSP 19188, 5 spm; MZSP 19189, 1 spm, Albardão- Sarita, 18–
28 m, MNRJ 2872, 3 spm; Pta. Cidreira, MNRJ 2922, 3 spm; São
Jose do Norte, Praia Grande, MNR J 691, 2 spm; Ponta da Cidreira,
MNRJ 2177, 2 spm; No precise loca li ty, MZSP 19190, 1 spm; MZ SP
19192, 2 spm; MZSP 19197, 1 spm; MZSP 32892, 3 spm; MZSP
32902, 3 spm; MZSP 51313; 33°24′S, 52°47′W, MACN- In 15347, in
13 m, 3 spm. URUGUAY. Rocha; MZSP 51679, 2 spm , MZSP 52424,
1 spm; Cabo Santa Maria, MZSP 13394, 1 spm; MACN- In 23569,
6 spm; USNM 331316, 1 spm; La Coronilla, USNM 359259, 1 spm;
La Pal oma , MZ SP 19183 , 2 spm, MZ SP 19196 , 3 spm; MZ S P 33348,
1 spm; MZSP 33400, 2 spm, MNRJ 1236, 3 spm, MNRJ 6546, 2 spm,
MACN- In 15151, 14 spm; MACN- In 30484, 3 m, 3 spm, MACN- In
15314, 6 spm. Maldonado; MZSP 10050, 10 spm, MZSP 51654,
2 spm; Punta del Este, MZSP33353, 1 spm; Piriapolis, MACN- In
18158 , ~30 spm, MZSP 57510 , 34°5′S 53 °30′W, in 20 m, MACN- In
15922, 34°38′S, 52°15′W, 128 m, 1 spm. ARGENTINA. Buenos
Aires; MACN- In 15851, 35°24′S, 53°10′W, 28 m, 1 spm, MACN- In
25200, 35°40′S 56°03′W, 9 m, 22 spm, MACN- In 15980, 35°08′S
52°35′W, 6 spm, MACN- In 24147 36°24′S 55°53′W; MACN- In
24300, 36°44′S 56°34′W, 9 m, 9 spm; off Ostende, MACN- In
21779, 1 spm; Mar Chiquita, MACN- In 3922, 1 spm; Mar del Plata,
MACN- In 10287, 2 spm, MACN- In 19140, 3 spm, MACN- In 19273,
8 spm, MACN- In 10768, 5 spm, MACN- In 12069, 1 spm, MACN- In
11999, 3 spm, MACN- In 9363- 12, MACN- In 19139- 1, 1 spm; Villa
Gesell, MACN- In 28827, 2 spm; 38°08′S 56°58′W, MACN- In
23380, 60 m, 4 spm; Puerto Quequén, MACN- In 23206, 26 spm,
MACN- In 18672, 5 spm; Necochea, MACN- In 9368- 28, 2 spm;
Ostende, MACN- In 20099, 7 spm; Monte Hermoso, MACN- In
6619– 31, 8 spm, MACN- In 11230, 18 spm, MACN- In 9209- 17,
3 spm, MZSP 51650, 10 spm, MZSP 389, 7 spm, 10056, 1 spm;
USNM 219862, 6 spm; USNM 465581, 1 spm; Cabo San Antonio,
MACN- In 16306, ~36 spm; Carmen de Patagones, MZSP10058,
10 spm; Arroyo Parejas, Bahía Blanca, MACN- In 11232, 12 spm;
Puerto Belgrano, MACN- In 11123– 1, many spm; Bahía San Blas
MACN- In 22943, 3 spm, MACN- In 20273– 1, 2 spm; Mouth of
Rio Negro river, MACN- In 20526, 1 spm; 40°56′S 62°28′W, 17 m,
MACN- In 20671, 2 spm. Rio Negro; Puerto San Antonio, USNM
568254, 2 spm; San Antonio, MZSP52101, 1 spm; Faro Punta
Villarino, San Antonio Oeste, MACN- In 13146- 2, 2 spm, MACN- In
9376; MACN- In 9379– 30, 4 spm, MACN- In 9152, MACN- In 9152-
14, 18 spm, MACN- In 9379– 31, ~26 spm, MACN- In 23705, 1 spm.
Chubut; Golfo San Matias, MZSP 51328, 1 spm, MZSP 10057,
2 spm, MACN- In21203, 4 spm, MNRJ 36, 1 spm; Faro Punta
Villarino, MACN- In 13146- 1, 1 spm; Puerto San José, MACN- In
9174, 2 spm; Playa Larralde, MACN- In 37814, 12 spm; Puerto San
José, MACN- In 26435, 128 m, 14 spm; Golfo San José, MACN- In
9174– 15, 2 spm, MACN- In 9441- 6, 9 spm; Playa Villarino, 42°24′S
64°17′W, MACN- In 38162, 10 m, 25 spm; Peninsula Valdes, MNRJ
2927, 2 spm, MNRJ 9399, 2 spm; Puerto Pirámides, MACN- In
26434, 14 spm. MACN- In 11484, 7 spm; Punta Norte, MACN- In
9158- 7, 2 spm; Puerto Madryn, MZSP10054, 2 spm.
Remarks: Buccinum cochlidium Chemnitz, 1795 was rejected by
ICZN Opinion 184 (1944) and Direction 1 (1954), which rejected
and made invalid all new specific names proposed in that work. The
next available name in line was also Buccinum cochlidium, changing
the author to Dillwyn (1817). When Deshayes (1844) reviewed the
2nd edition of Lamarck's Histoire Naturelle des animaux sans vértebrés,
he described Buccinum gradatum based on the specimen housed in
Lamarck's collection and figured by Kiener (1834: 10, pl. 6, Figure 17)
(Figure 1d) as Buccinum cochlidium. The material is now at MHNG
(Figure 1e– f). Tryon (1882) raised some doubts about the validity of
B. gradatum; however, the name was sustained by most later authors.
The probable holotype of Buccinum lamarckii Kiener, 1834, housed
in MNHNG (Figure 1i– k) clearly represents the southern morph of
Buccinanops cochlidium.
A. d'Orbigny (1839, pl. 61, Figure 25) (reillustrated by M.
E. Gray, 1850) depicts an apparently juvenile specimen of
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PASTORIN O ANd SIMO NE
Buccinanops cochlidium. Later, Pilsbry (1897) considered it as Bullia
uruguayensis. The figure is not clear and based on a juvenile speci-
men and could be any one of these species. The Buccinanops mate-
rial from d’Orbigny's collection, housed in NHMUK was examined,
and we conclude that it contains several lots with the following
species: B. deforme: 1854.12.4.469, 4 sp., 1854.12.4.470, 11 sp.;
1854.4.466, 6 sp.; B. uruguayense 1854.12.4.467, 12 sp.; B. monil-
ifer: 1854.21.4.465, 9 sp.; B. paytense: 1854.21.4.463, 1 sp.; B. co-
chlidium: 1854.21.4.461, 4 sp.; 460, 2 sp.; 462, 1 sp.; 464, 3 sp.;
459, 1 sp.; 458, 1 sp.
Ihering (1907) described Bullia gradata pampeana, a fossil sub-
species from Quaternary deposits of Pleistocene age according to
Parodiz (1996). Ihering (1907) also mentioned doubts about the dif-
ferences between his Bullia gradata and B. cochlidium, showing that
the typical form without subsutural cord (B. cochlidium) gradually
change to that of thicker shell, with subquadrate profile and rectan-
gular whorls (B. gradatum). Cernohorsky (1984) also mentioned the
possibility that Buccinanops gradatum could be only a variation of
B. cochlidium. However, several modern authors (e.g., Calvo, 1987;
Rios, 1985) keep both names in use. Later, Allmon (1990) included
B. gradatum in the list of synonyms of B. cochlidium. Allmon (1990)
also subordinated Buccinanops as a subgenus of Bullia which was dis-
cussed by Pastorino (1993a) who reinstated the full generic status.
Buccinum turritum Perry, 1811, was mentioned by Deshayes
(1844) as a dubious synonym of B. cochlidium Chemnitz. Petit (2010)
stated that Perry's name could be nomen dubium unti l “speci alist wi th
all the available types rule out about the validity of this name.” We
have seen all the existent type material related with these names,
and there is none with real similarity to the picture of B. turritum. The
regular axial ribs, particularly on the last whorl, appear as a charac-
teristic feature of B. turritum, which are absolutely abs ent in any spe-
cies of Buccinanops. We agree with Petit's designation of B. turritum
as a nomen dubium and according to our observations, without any
relationship with the genus Buccinanops.
Tryon (1882) and Allmon (1990) synonymized Buccinum labyrin-
thum Gmelin, 1791 from the Netherlands coast with Buccinanops
cochlidium. Pastorino and Kantor (1998) discovered that Gmelin's
species is actually an anomalous specimen of Buccinum undatum L.
with no connection to the South American species.
Penchaszadeh (1971, 1973) described the egg capsules of sev-
eral species of Buccinanops including material of the form "gradatum"
from Mar del Plata, Buenos Aires, Argentina. The morphology and
ornamentation of the capsules are exactly the same of those from
specimens from Puerto Pirámide, off Chubut Province where the
only available form is "cochlidium."
3.2 | Buccinanops latus sp. nov.
(Figures 9– 11)
Buccinanops lamarckii: Rios, 1970: 93; 1975: 96, pl. 27, fig. 401;
1985: 103, pl. 35, fig. 454; 2009: 246 (fig.) (non Kiener, 1834).
Buccinanops gradatum: Rios, 1970: 93, pl. 27; 1985: 103, pl. 35,
fig. 453; 2009: 246 (in syn); Rios & Calvo, 1987: 143, fig. 110 (non
Deshayes, 1844).
Buccinanops gradatus: Simone, 1996: 88– 96, figs. 1– 3, 6, 12– 30;
Galindo et al., 2016: 342, fig. 1 (non Deshayes, 1844).
urn:lsid:zoobank.org:act:52D86791- 0AA4- 4E0D- A258-
F82317E0257C.
Type material: Holotype MZSP 28190 (Figure 9a– c). Paratypes:
Brazil. Rio de Janeiro; São Tomé, off Cabo de São Tomé, MZSP 72606,
2 shells (Figure 9i– j) (Femorale; trawled, 60 m, fishermen, viii/2002).
São Paulo; off Ubatuba, off Flamengo Beach, MZSP 51656, 3 spm
(W Narchi leg, 14/vii/1957) (Figure 9f– h); Guarujá, Praia do Perequê,
MACN- In, 3 spm (Figure 9d– e); Santos, Barra de Santos, 46°19′S
24°04′W, MZSP 28078, 1 shell, MZSP 28079, 1 shell (21.ix.1970);
Cananéia, off Bom Abrigo Island, 25°08′S 47°50′W, 8 m, MZSP
35775, 1 spm (otter trawl, Magenta leg., viii/2002) (Figure 9k– l),
MZSP 47125, 13 spm (BIOTA sta. 83i, 16.x.2001).
Type locality: BRAZIL. São Paulo; Praia Grande, off Boqueirão
Beach, 24°04′S 46°28″W, ~8 m depth (otter trawl, local fishermen
col., i/1994).
Etymology: The new species is named in reference to the broad
profile of the shell, from Latin latus, meaning broad, wide.
Diagnosis: Shell up to 80 mm; usually simple, globose. Spire
angle 65– 70°. Left edge of canal with blunt beak of ~90°. Head-
foot mostly colorless. Kidney tissue with two longitudinal folds.
Single odontophore pair m9; pair m2a reduced. Radular rachid-
ian with 7– 8 uneven cusps. Stomach larger. Anal gland present.
Nerve ring with cerebral commissure slightly shorter than ring
itself.
Description: This supplements the conchological and anatomical
descriptions provided by Simone (1996) (at that time as B. “gradatus”).
Shell: (Figures 9– 10) Size large (to 80 mm), spire short (half
shell length), and wide. Spire angle 65– 70°, obtuse. Protoconch of
~2.8 mm, first two whorls smooth, with axial riblets appearing sub-
sequently, from suture to suture, each riblet with clear subsutural
node (Figure 10a– c); riblets gradually disappearing after two whorls,
leaving only growth lines and shallow spiral cords (Figure 10d– e),
clearer along adapical half of each whorl, ~10 per whorl. Shouldered
shells (Figure 9f– h) more uncommon than in preceding species
(~10%). Left (adaxial) edge of canal with blunt anterior beak of ~90°
(Figure 9a, d, f, i, k).
Head- foot: Color of exposed areas lighter, practically light tan,
except for some sparse pale- brown spots, varying in density among
specimens. Females with well- developed cement gland, similar to
that of preceding species, but shorter and wider (Figure 11b: cl).
Operculum: Similar to B. cochlidium.
Pallial cavity: Features similar to preceding species, except for
more bluntly pointed anterior end of gill (in Simone, 1996: figure 12,
gi).
Circulator y and excretory systems: Heart similar to that of B. co-
chlidium. Kidney also similar, except for renal lobe divided into
two longitudinal folds (instead of three), each fold with different
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15
PASTORIN O ANd SIMO NE
glandular constitution (Simone, 1996: figure 28: gk, vp) (uniform in
B. cochlidium).
Digestive system: Most characters similar to those of B. coch-
lidium. Differences include the following: Odontophore muscles
described in Simone (1996). Table 2 shows a correlation of mus-
cles terminology. Odontophore muscle m9 as single pair, bifurcat-
ing from each other more anteriorly (Simone, 1996: figure 21: m5);
pairs m2a more poorly developed, being narrower fibers (Simone,
1996: figure 21: m4). Radula: rachidian with 7– 8 cusps, each cusp
broader, flatter, and more uneven than those of preceding species;
lateral teeth with inner set of cusps with only three cusps, being
medial cusp ~3 times larger than remaining cusps (Simone, 1996:
figure 6). Transverse muscles between esophagus and odontophore
(Figure 5b: tm) poorly developed. Anterior esophagus much simpler,
practically straight (Simone, 1996: figure 15: ao). Pair of salivary
glands ~twice size of those in preceding species. Valve of Leiblein
much less well developed, only marked by sudden change of inner
esophageal folds (Simo ne, 1996: fig ure 16: vl). Stomach ~50% larger
and internally more complex (Simone, 1996: Figure 18). Inner anal
gland subterminal, close to anus (Simone, 1996: figure 19: ga).
Genital system: Most features similar to preceding species.
Distinctions include the following: Seminal vesicle (sv) slightly
smaller; despite in also having gonopericardial duct (go), addition-
ally bearing aperture to pallial cavity in its posterior pallial portion
(Simone, 1996: figure 29: va). Penis with subterminal papilla slightly
more developed, and penis duct slightly less coiled. Pallial oviduct
FIGURE 9 Buccinanops latus new species, shells of types. (a– c) Holotype MZSP 28190, frontal, right and dorsal views (L 57.8 mm).
(d– e) paratype MACN- In 43748, from Praia do Perequê, Guaruja (SP) Brazil (L 46.6 mm). (f– h) paratype MZSP 51656, from Ubatuba,
SP, frontal, dorsal and right views (L 57.0 mm). (i– j) paratype MZSP 72606, from São Tomé, RJ, frontal and dorsal views (L 82.0 mm).
(k– l) paratype MZSP 35775, from Bom Abrigo, SP, frontal and dorsal views (L 48.1 mm)
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PASTORIN O ANd SIMO NE
similar, including vaginal atrium length and papilla- like female aper-
ture. Cement gland slightly shorter (Figure 11b), well- separated from
hemocoel.
Central nervous system: (Figure 11a, c) Nerve ring similar to that of
preceding species, including concentration of ganglia, indistinction
of some ganglia by fusion, and position of buccal ganglia. Cerebral
commissure (cc) about half length and thicker than that of preceding
species, relatively uniform in width along its length. Visceral gan-
glion (gv) relatively smaller, located closer to nerve ring, distance
equivalent to length of nerve ring.
Development: Egg capsules attached to callus parietal region of
female's shell (Figure 10f). Each egg capsule of ~10 mm, oval, flat-
tened, with thin walls, translucent, containing ~15 embryos.
Habitat: Sandy mud and muddy bottoms, 5– 40 m, normally col-
lected by otter trawl.
Distribution: Espírito Santo (~20°S) to north Rio Grande do Sul
(~31°S), Brazil.
Material examined: BRAZIL. Espírito Santo; Guarapari, MZSP
39453, 1 spm, MZSP 77303, 2 spm. Rio de Janeiro; Cabo de São
Tomé, MZSP 28184, 1 spm, MZSP 34820, 1 spm, MZSP 34870,
1 spm, MZSP 72606, 2 shells, 100– 120 m (Femorale, viii.2002); off
Ilha de Santana, Macaé, MNRJ 8838, 2 spm, MNRJ 1194, 2 spm,
MZSP 73814, 60 m, 1 shell (otter trawl, Femorale, viii/2000); off
Ponta de Guaratiba, 35– 40 m, MNRJ 8836, 3 spm; Cabo Frio, MZSP
51658, 1 spm; Niterói, Itaipu, off Ilha do Pai, 30– 40 m, MNRJ 8837,
2 spm; off Ilha de Guanabara, MNRJ 8835, 2 spm; Rio de Janeiro,
MZSP 33399, 1 spm, Copacabana, MZSP 51651, 1 spm; Ilha Grande,
MZSP 51378, 1 spm, MZSP 51629, 1 spm; Angra dos Reis, MZSP
51320, 2 spm, MZSP 51341, 1 spm, MZSP 51641, 2 spm, MZSP
51647, 1 spm, MZSP 51390, 1 spm; Paraty, MZSP 48307, 1 spm. São
Paulo; Ubatuba MZSP 28080, 6 spm, 28081, 1 spm, 33443, 1 spm,
28186, 2 spm, 37921, 2 spm, 37925, 3 spm,; 38000, 6 spm, 38001,
1 spm, 38330, 19 spm, 38002, 2 spm, 38498, 13 spm, 38499, 8 spm,
38336, 1 spm, 40010, 3 spm, 40020, 5 spm, 45467, 1 spm, 45505,
3 spm,; 47092, 5 spm, 47126, 17 spm, 51314, 7 spm, 51316, 3 spm,
Barra Seca, MZSP 51676, 1 spm, Praia de Maranduba, MZSP 51802,
1 spm, Praia do Flamengo, MZSP 51661, 2 spm, 51656, 1 spm (W.
Narchi col, 14.vii.1957), Praia de Itaguá, MZSP 51645, 3 spm, 51633,
1 spm, 64201, 3 spm; Caraguatatuba, MZSP 40229, 1 spm, 40278,
1 spm, 38332, 8 spm, 37923, 2 spm, 38338, 1 spm, 38340, 2 spm,
45245, 1 spm, 45252, 1 spm, 45253, 1 spm, 45480, 9 spm, 45526,
3 spm, 45647, 4 spm, 45648, 3 spm, 45649, 3 spm, 45664, 67 spm,
45665, 23 spm, 45666, 20 spm, 47086, 2 spm, 47135, 1 spm, 47099,
2 spm, 47125, 13 spm, 54189, 1 sp; São Sebastião, MZSP 37922,
1 spm,; 38003, 1 spm, 38004, 4 spm, 38331, 12 spm (23°52.330′S
45°34.227′W, Biota- Fapesp, sta. 163i, 27/vi.2002), 38333, 2 spm,
FIGURE 11 Buccinanops latus new species, anatomy. (a) central nervous system, ventral view. (b) foot, female, sagittal section in its
anterior region. (c) central nervous system, dorsal view. Scales = 2 mm
FIGURE 10 Buccinanops latus new species, protoconchs and
capsules. (a– c) intracapsular specimen, MZSP 82754, frontal,
dorsal and right views (L 5.1 mm). (d– e) young specimen, sale lot,
dorsal and frontal views (L 10.1 mm). (f) whole frontal view, MZSP
38331, adult female with egg capsules attached to shell (L 60 mm).
Scales = 1 mm
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PASTORIN O ANd SIMO NE
38334, 2 spm, 38337, 2 spm, 38339, 1 spm, 38341, 21 spm, 38343,
1 spm, 38402, 1 spm, 45239, 2 spm, 45539, 1 spm, 45543, 3 sp.;
47131, 3 spm, 10053, 3 spm, Praia do Gato, MZSP 14259, 6 spm;
Ilha dos Buzios, MZSP 15339, 1 spm; Bertioga, MZSP 61464, 40 spm,
61448, 17 spm, 61451, 2 spm, off Boracéia Beach, MZSP 112055, 1
shell (Ex Colella col., i/1971); Guarujá, MZSP 57506, 2 spm, Praia do
Perequê, MZSP 28188, 1 spm, 28192, 20 spm, 43579, 6 spm, 43736,
1 spm, Praia do Góis, 2 m, MZSP 44007, 18 spm, 61160, 1 spm MZSP
33429, 1 spm; Santos, MZSP 33475, 1 spm, 33344, 1 sp, 35397,
2 spm, 51631, 1 spm, 51642, 1 spm, 61443, 4 spm, 36627, 2 spm,
51352, 1 spm, 51639, 1 spm, 32 m, 51804, 1 spm, off Santos, MZSP
36466, 2 spm (Coltro leg, iv/2003), 24°05.704′S 46°20.270′W, 22 m,
MZSP 108269, 2 spm (otter trawl, Instituto de Pesca, 4/vii/2012); be-
tween Ilha da Queimada Grande and Lage de Santos, MZSP 78246,
12 spm, 30– 35 m (otter trawl, xii/1998); São Vicente, MZSP 61151,
1 spm; Praia Grande, off Praia do Boqueirão (type locality) MZSP
27319, 2 spm, 28190, 2 spm, 60047, 10 spm, 61081, 100 spm,
61147, 1 spm, 61148, 3 spm, 61149, 3 spm, 61152, 5 spm, 61153,
7 spm, 61442, 15 spm, 61444, 9 spm, 61445, 9 spm, 61446, 1 spm,
61447, 2 spm, 61449, 2 spm, 1450, 3 spm, 61453, 10 spm, 61466,
35 spm, 61467, 20 spm, 61469, 1 spm, 28079, 1 spm, 51681, 2 spm,
47141, 3 spm, 28082, 18 spm, 28183, 17 spm, 28193, 7 spm, 35392
~30 spm; Mongaguá, MZSP 51385, 1 spm; Itanhaém, MZSP 33398,
6 spm; Peruíbe, MZSP 61150, 1 spm, Praia das Enchovas, MNRJ 2925,
3 spm, Icaparra, MZSP 51649, 4 spm; Iguape, MZSP 51333, 1 spm,
51807, 1 spm. Paraná; Camburiú, MZSP 14768, 1 spm; Paranaguá,
MZSP 34494, 1 spm. Santa Catarina; Penha, 40– 50 m, MZSP 53552,
34 spm; Itajai, MZSP 61465, 40 spm; Porto Belo, MZSP 51329, 1 spm,
51330, 3 spm, 51331, 3 spm, 51655, 3 spm, 32867, 1 spm, 44019,
2 spm, 17908, 1 spm; Florianópolis, MZSP 33442, 1 spm, Praia de
Piçarras, MZSP 51675, 11 spm, Itaporocóia, MZSP 15792, 1 spm;
Barra Velha, MNRJ 2929, 2 spm; Itajubá, MNRJ 3498, 2 spm; Pântano
do Sul da Ilha de Santa Catarina, MNRJ 2932, 2 spm, MZSP 16157,
2 spm.
Remarks: Buccinanops latus sp. nov. is morphologically close to
the type species. It was apparently identified under three names,
that is, B. cochlidium, B. lamarckii, or B. gradatus by different au-
thors. All these nominal species were described without accurate
type locality (i.e., southern seas for B. cochlidium), wrong (i.e., New
Zealand for B. gradatum), or unknown. Buccinanops lamarckii is
a clear synonym of B. cochlidium and B. gradatum was split from
B. cochlidium, probably based only on very few shells. When sev-
eral hundred lots are compared, from a large latitudinal range, a
single variable species could be recognized. Our anatomical stud-
ies, and a DNA sequence treatment produced elsewhere (B. gra-
datus in Galindo et al., 2016, based on sample MZSP 108269, also
included herein), suggested the presence of a new species with a
more restricted range but a comparable morphology. Once we es-
ta bli she d the exis ten ce of a new spe cie s, the use of one of thes e old
names was attempted. However, we decided that with the avail-
able knowledge (i.e., only shells with no locality to backup those
names) no clear proof were enough to use any of them, cochlidium
FIGURE 12 Buccinanops monilifer shells. (a– c) MACN- In 43750 Mar del Plata (L 40.7 mm). (d) MACN- In 43750 other specimen Mar del
Plata (L 39.3 mm). (e– g) MACN- In 43750 other specimen Mar del Plata (L 32.6 mm). (h) Buccinum moniliferum by Kiener’s (1834) plate 3,
Figure 8. Scale = 1 cm
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PASTORIN O ANd SIMO NE
or gradatus, for the new entity. On the contrary, we have abun-
dant evidence, anatomical and DNA sequence, with a considerable
amount of lots, sufficient to describe a new species. Additionally,
there are conchological differences explored in the distinctive de-
scription above and in the discussion below.
The shell in B. latus is smaller (up to 80 mm, against ~120 mm of
B. cochlidium) and has a shorter spire (up to half of total shell length,
while B. cochlidium usually has a longer, slender, spire), and the left
(adaxial) edge of the canal usually has an anterior blunt beak of ~90°
versus ~30° of B. cochlidium. However, some immature specimens
could be dubious. The head- foot color is also diagnostic as B. cochlid-
ium is almost black in exposed areas (Figure 3c) while B. latus mostly
lacks pigmentation or shows only some sparse brown spots. The re-
maining anatomical differences are explored above, drawing attention
to the distinctions in the nerve ring, as B. cochlidium has a much lon-
ger and narrower cerebral commissure (Figure 8a– b: cc) than that of
B. latus (Figure 11a, c). In addition, the valve of Leiblein is much more
reduced in B. latus than in B. cochlidium (Figure 5f). Most specimens of
the new species have shells of uniform light- brown color (Figure 9a– c,
f– l); however, about 30% of the specimens also have axial irregular
bands (Figure 9d– e). The periostracum is thin, velvet- like in freshly col-
lected specimens (Figure 9a– d, f– h). B. cochlidium and B. latus do not
overlap in geographic distribution according to our data. B. latus has as
southernmost limit 31°S, while the northernmost limit of B. cochlidium
is 32°S.
Compared the egg capsules of B. cochlidium illustrated by
Penchaszadeh (1971, 1973) and Averbuj and Penchaszadeh (2010a),
B. latus (Figure 10f) has more rounded and flattened capsules with-
out ornamentation.
3.3 | Buccinanops monilifer (Kiener, 1834)
(Figures 12– 14)
Buccinum moniliferum Valenciennes in Kiener, 1834: pl. 3, fig. 8;
Anton, 1839: 91; Deshayes, 1844:191; d’Orbigny, 1845: 199.
Bullia armata Gray, 1839: 126; Reeve, 1846, pl. 1, fig. 2 a, b;
Kobelt, 1877: 290.
FIGURE 13 Buccinanops monilifer images in SEM. (a– b) radulae, frontal, and slightly lateral views, scale = 200 µm. (c) transverse section of
shell, scale = 200 µm. (d) penis, dorsal view, scale = 1 mm. (e– f) two views of prehatching young shell, scale = 1 mm
TAB LE 2 Correlation of terminology used by Simone (1996) with that used in remaining papers including odontophore descriptions, and
respective functional or topological name
In Simone (1996) Current Name/function/topology
m1 m1 Jugal muscles
m2 m3c Dorsal circular tensor muscle
m3 m2c Retractor muscles of odontophore, car tilage component
m4 m4+m2a Main dorsal tensor muscles of radula
m5 m9 Radular dorsal protractor muscles
m6 m8 Antero- dorsal tensor muscles of cartilages
m7 m9 Radular dorsal protractor muscles (a secondar y branch)
m8 m9 Radular dorsal protractor muscles (a secondar y branch)
m9 m4 Main dorsal tensor muscles of radula (a secondary branch)
m10 m5 Auxiliary dorsal tensor muscles of radula
m11 m6 Horizontal muscle (a ventral circular tensor muscle)
m13 m11 Ventral tensor muscles of radula
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PASTORIN O ANd SIMO NE
Buccinum (Buccinanops) maniliferum [sic]: d’Orbigny, 1846: 434.
Bullia (Buccianops) armata: H. & A. Adams, 1853: 113.
Buccinanops cochlidium (var 3): Gray, 1854: 40 non Chemnitz,
1795 nec Dillwyn, 1817.
Bullia (Buccinanops) moniliferum: Chenu, 1859: 160, fig. 750.
Bullia (Buccinanops) armata: Tryon, 1882: 14, pl. 6, figs. 82, 83;
Paetel, 1888: 116.
Dorsanum armatum: Cossmann, 1901: 218.
Dorsanum moniliferum: Carcelles & Parodiz, 1939: 747, figs. 1, 2;
Carcelles, 1944: 249; Barattini & Ureta, 1960: 113; Rios, 1970: 92, pl.
28; Rios, 1975: 95, pl. 27, fig. 398.
Bullia (Dorsanum) monilifera: Cernohorsky, 1982: 17, fig. 17, 871.
Bullia monilifera: Abbott & Dance, 1983: 177.
Buccinanops moniliferum: Cernohorsky, 1984: 29; Pastorino,
1993a, figs. 1– 3.
Buccinanops (Dorsanum) moniliferum: Rios, 1985: 103, pl. 35, fig.
456.
Buccinanops moniliferum: Calvo, 1987: 143, fig. 112.
Bullia (Buccinanops) monilifera: Allmon, 1990: 25, pl. 2, fig. 12; pl.
6, figs. 1, 2.
Buccinanops moniliferus: Simone, 1996: 96– 100, figs 4– 5, 7– 8,
1 0 – 1 1 , 3 1 – 4 2 .
Type material: Buccinum moniliferum: unknown, never found (not
in MHNP, neither Geneva); Bullia armata: unknown (not in NHMUK).
Type locality: B. monilifer: “Terra Nova”; B. armata: unknown.
Distinctive description: This supplements the conchological and
anatomical description provided by Simone (1996).
Shell: (Figure 12) Large (up to ~55 mm), fusiform, with 6– 7 slightly
convex whorls. Color variable, usually dirty whitish, creamy with two
thick brownish bands, one subsutural and one along middle of last
whorl (Figure 12c, g). Protoconch large, 2.5 whorls, 2.2 mm wide in
2nd whorl, with 8– 10 riblets per whorl (Figure 13e– f). Transition to
teleoconch almost indistinct. Suture well defined, impressed. Spire
high, 1/2 to 1/3 of total shell height; first two whorls with incom-
plete axial riblets, 3rd and subsequent whorls with 10– 14 subsutural
peripheral spines or knobs, somewhat blunt, and of variable size or
development. Sculpture absent, except for growth lines; base of
last whorl with few spiral threads. Parietal callus weak but visible
(Figure 12d). Aperture smaller than half of total length, 60– 70% of
shell width; subcircular, posterior notch usually present, with thicker
area on parietal side of anal angle (Figure 12a, d, e). Outer lip profile
slightly sinuous (Figure 12b, f). Inner lip with strong inflexure at base
of siphon (Figure 12a, d, e).
Ultrastructure of shell of three layers, outermost layer of amor-
phous calcite, intermediate thicker of crossed- lamellar aragonite;
innermost thinner also of crossed- lamellar aragonite (Figure 13c).
Remaining characters similar to preceding species.
Head- foot: Characters similar to those in B. cochlidium. Female
cement gland (Figure 14b: cl) more anteriorly located, shorter and
broader. Dark pigment absent.
Pallial cavity: Characters like those of B. cochlidium, except for
more elongated gill and osphradium, both with more uniform width
throughout their length (Simone, 1996: figure 33: gi, os).
Digestive system: Buccal mass organization similar to preceding
species, differences explored in Simone (1996: figure 34). Salivary
ducts anterior region running along dorsal folds (Figure 14a; sd),
lacking subterminal bulged region, strong curve preceding its ap-
erture towards medial; salivary aperture ventral, at short distance
from median line, approximately in middle level between mouth and
odontophore (Figure 14a: sa). Radula (Figure 13a– b), central tooth
FIGURE 14 Buccinanops monilifer anatomy. (a) foregut, left view, detail of its anterior region, mostly opened longitudinally. (b) foot,
female, sagittal section in its anterior region. (c) central nervous system, dorsal view. Scales = 2 mm
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PASTORIN O ANd SIMO NE
with 5– 13 cusps of variable size, some specimens uniform, others
with central cusp larger. Rachidian base curved. Lateral teeth with
outer cusp hook- shaped, inner cusp thicker than 2– 5 intermediate
cusps, sometimes with different number on each side; lateral tooth
base with blunt inner cusp. Valve and gland of Leiblein, and stomach
similar to those of B. latus, rather than B. cochlidium.
Genital system: Male genital structures mostly similar to those
of preceding species (see Simone, 1996: figs. 39– 41). Penis with
blunt tip, almost bilobed, with papilla in its mid region (Figure 13d).
Females with pore preceded by rather bulged, muscular region.
Central nervous system: (Figure 14c) Closer to that described for
B. latus, except for relatively shorter cerebral commissure (cc), and
clearer pair of buccal ganglia (bg).
Habitat: Sandy mud and muddy bottoms, 5– 25 m, normally col-
lected by otter trawl.
Distribution: From Rio de Janeiro, Brazil (~23°S) to mouth of Rio
Negro area (~41°S), Argentina.
Material examined: BRAZIL. Rio de Janeiro; MZSP 34771,
1 spm; Recreio dos Bandeirantes, MNR J 6059, 3 spm; Cabo Frio,
MNRJ 6462, 2 spm; Cabo de São Tomé, MZSP19591, 1 spm.
São Paulo; MZSP 34443, 10 spm, MZSP 35404, 2 spm, MZSP
51673, 8 spm; Ubatuba, MZSP 33401, 1 spm, MZSP 33406,
1 spm, MZSP 38349, 1 spm, MZSP 48192, 1 spm, MZSP 51287,
1 spm, MZSP 38005, 2 spm, MZSP 51662, 2 spm; São Sebastião,
MZSP 398, 16 spm, MZSP 51664, 3 spm, MZSP 53330, 1 spm;
Caraguatatuba, MZSP 37924, 1 spm; Bertioga, MZSP 61404,
1 spm, MZSP 61426, 14 spm; Guarujá, Praia de Pernambuco,
91.5 m, MZSP51399, 2 spm; Santos, MZSP 51388,1 spm, MZSP
51805, 4 spm, MZSP 51799, 2 spm, MZSP 35403, 1 spm, MZSP
15233, 1 spm, MZSP 33402, 2 spm, MZSP 78817, 2 spm; São
Vicente, Ponte Pensil, MZSP 61423, 4 spm; Praia Grande, MNR J
1523, 2 spm, off Boqueirão, MZSP 26865, 2 spm, MZSP 27320,
3 spm, MZSP 28151, 56 spm, MZSP 28152, 18 spm, MZSP 28153,
18 spm, MZSP 28175, 15 spm, MZSP 28176, 28 spm, MZSP 28177,
FIGURE 15 Buccinastrum deforme shells and operculum. (a– c) Buccinum deforme King, 1832, syntype NHMUK 1854.12.4.470 (L 28.1 mm).
(d– f) NHMUK 1854.12.4.470, syntype (L 28.6 mm). (g– h) B. globulosum holotype, MHNG 1296/17/1 (L 42.7 mm). (i) MACN- In 35384 Caleta
Olivia, Santa Cruz, Argentina (L 53.0 mm). (j– l) MZSP, male, from 42°24′S, 64°17′W, Playa Villarino, Golfo San José, Chubut, Argentina (L
22.7 mm). (m– n) MZUSP, male, from off Montevideo, Uruguay (L 20.5 mm)
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PASTORIN O ANd SIMO NE
9 spm , MZ SP 34444, 14 spm, MZSP 34 445, 19 spm, MZ SP 61398,
3 spm, MZSP 61399, 2 spm, MZSP 61400, 14 spm, MZSP 61401,
1 spm, MZSP 61402, 1 spm, MZSP 61403, 1 spm, MZSP 51332,
1 spm, MZSP 61406, 7 spm, MZSP 61407, 14 spm, MZSP 61409,
3 spm, MZSP 61410, 6 spm, MZSP 61411, 2 spm MZSP 61413,
5 spm, MZSP 61414, 4 spm, MZSP 61415, 6 spm, MZSP 61416,
8 spm MZSP 61417, 6 spm MZSP 61418, 6 spm, MZSP 61420,
15 spm, MZSP 61421, 12 spm, MZSP 61422, 2 spm, MZSP 61424,
2 spm, MZSP 61425, 35 spm, MZSP 61427, 26 spm, MZSP 61428,
20 spm, MZSP 61430, 20 spm, MZSP 61432, 20 spm, MZSP
61435, 36 sp m, MZS P 61436, 60 sp m, MZS P 61437, 80 spm, MZ SP
61438, ~100 spm, MZSP 61439, 70 spm, MZSP 35841, 1 spm,
MZSP 60070, 2 spm, MZSP 61450, 2 spm; Iguape, MNR J 11427,
2 spm; Suarão, MZSP 15208, 1 spm; Itanhaém, MZSP 28180,
1 spm, MZSP 33357, 68 spm, MZSP 51338, 2 spm, MZSP 53589,
5 spm, MZSP 51793, 1 spm; Peruíbe, off Praia Central, MZSP
15127, 2 spm, MZSP 61412, 10 spm, MZSP 61408, 8 spm, MZSP
61431, 12 spm; Iguape, MZSP 10059, 6 spm, MZSP 14145, 2 spm,
MZSP 51657, 3 spm, MZSP 51663, 9 spm, MZSP 51803, 1 spm,
MZSP 47227, 2 spm, MACN- In 1855, 19 spm. Paraná; Ilha do
Mel, MZSP 17855, 2 spm, MZSP 17854, 1 spm; Paranaguá, MZSP
51348, 1 spm; Guaratuba, MZSP 51653, 2 spm. Santa Catarina;
Piçarras, Praia Alegre, MZSP 61419, 6 spm, MNRJ 2931, 2 spm.
Rio Grande do Sul; MZSP 32901, 4 spm, MZSP 19191, 9 spm,
MZSP 19195, 12 spm, MZSP 43088, 1 spm, MNRJ 3868, 1 spm;
off Torres, MZSP 32724, 4 spm; Vitoria do Palmar, 15– 25 m, MZSP
51666, 5 spm; Cassino, MZSP 61405, 2 spm, MNRJ 3470, 2 spm,
MNRJ 7806, 1 spm, MACN- In 8219, 3 spm; Rio Grande, MZSP
40341, 8 spm; Mostardasto São José do Monte, 8– 20 m, MZSP
62976, 3 spm, MZSP 62976, 2 spm; off Chui, MACN- In 15346,
33°24′S 52°47′W, 13 m, 12 spm. URUGUAY. Montevideo; MZSP
10039, 3 spm. Rocha; La Paloma, MZSP 33397, 9 spm, MZSP
60048, 2 spm, MACN- In 35383, 5 spm, MACN- In 15149, 4 spm,
MACN- In 30486, 2 spm, MNRJ 1243, 2 spm, MNRJ 6527, 2 spm;
Cabo Santa Maria, MACN- In 15374, 34°53′S 55°19′W, 20 m,
2 spm, MACN- In 11593 1 spm. Maldonado; Maldonado, MACN- In
9356– 6, 1 spm. ARGENTINA. MZSP 51345, 15 spm, MZSP 51342,
12 spm. Buenos Aires; littoral, MNRJ 34, 35 spm, MZSP 51344,
2 spm, MZSP 51345, 3 spm; Punta Médanos, MACN- In 25197,
35°40′S 56°03′W, 10 m, ~80 spm, MACN- In 23517, 35°55′S
55°19′W, 21 spm, MACN- In 11843, 2 spm; Mar del Plata, MZSP
10037, 3 spm, MACN- In 16544, ~50 spm, MZSP 22389, ~40 spm,
MACN- In 4660– 1, 32 spm, 19274, 38 spm MACN- In 22389–
1, 2 spm; MZSP 19139, 8 spm, MACN- In 9217– 4, 6 spm, MZSP
11372, 4 spm, MZSP 11998, 7 spm, MZSP 10109, 2 spm, MZSP
29459 ~ 200 sp., MACN- In 8587– 6, 2 spm, MZSP 9490, 1 spm,
MZSP 8734, 1 spm, MZSP 12221, 1 spm, MZSP 9361, 2 spm;
Miramar, MACN- In 9247– 6, 6 spm, MACN- In 9252– 2, 2 spm;
Necochea, MZSP 51660, 9 spm, MACN- In 29661, Punta Negra,
1 spm; Monte Hermoso, MZSP 10035, 9 spm, MACN- In 9209–
16, 7 spm, MACN- In 6619– 28, 4 spm, off Faro Recalada, 1 spm;
Punta Carballido, Quequén, MACN- In 26281, 8 spm; Puerto
Quequén, MACN- In 18671, 30 spm; Bahía Blanca, MACN- In
14867, 30 spm; Arroyo Parejas, MACN- In 29663, 2 spm, MACN- In
11232– 2, 1 spm; Puerto Belgrano, MACN- In 11233, 2 spm, 6620–
18, 1 sp.; Bahia Blanca, MACN- In Caleta/Brightman, 6 spm; Isla
Trinidad, MACN- In 19669, 7 spm; Bahía San Blas, MACN 20273,
1 sp m; Ca rmen de Pat ago nes , MZSP 1003 4, 16 spm; Buen os Aires,
MACN- In 30025, 4 spm, MACN- In 11696, 27 spm, MACN- In
20670, 40°47′S 62°15′W, 15 m, 9 spm. Río Negro; MACN- In
20527, 9 spm, MZ SP 51678, 3 spm , E of mou th of Rí o Neg ro, MZSP
51363, ~60 spm.
Remarks: This is probably the most common species of
Buccinanops in Buenos Aires province, Uruguay and southern
Brazil. Usually found subtidally. Buccinanops monilifer is easy to
recognize, because it is the only species of Buccinanops with sub-
sutural spines, despite some rare specimens in which these are
very weak of even obsolete (Figure 12e– g; Simone, 1996: figure
11). The shell is slender, and the coloration is diagnostic in those
cases with short or weak spines. It was cited as belonging to the
genus Dorsanum by Cossmann (1901), followed by local authors
(e.g., Carcelles & Parodiz, 1939; Castellanos, 1970; Rios, 1994).
The attribution is probably because of some shell similarity with
Dorsanum miran (Bruguière , 1789), the West African genotype . The
distinction between Buccinanops and Dorsanum is explored else-
where (Pastorino, 1993a; Simone & Pastorino, 2014), restricting
FIGURE 16 Buccinastrum deforme SEM images. (a– b) two views of radula, scales = 200 µm. (c– d) two views of prehatching young shell.
(e) another prehatching young shell; scale = 1 mm. (f) tip of penis showing short papilla, scale = 200 µm
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PASTORIN O ANd SIMO NE
Dorsanum to the African coast. Penchaszadeh (1971) and Averbuj
and Penchaszadeh (2010b) reviewed the reproductive biology
and the development of imposex in the females of this species
(as Dorsanum moniliferum). Martorelli (1991) pointed out the pres-
ence of cercarian parasites in adult specimens of B. monilifer (as
Dorsanum moniliferum).
3.4 | Genus Buccinastrum gen. nov.
Diagnosis: Shell bucciniform to nassariform, medium to small; outer
surface normally mat; walls thick. Sculpture of axial growth lines only;
whorl profile rounded, rarely shouldered; low irregular subsutural
undulations, sometimes with few faint cords. Last whorl smooth,
usually lacking cords. First teleoconch whorls smooth. Left, adaxial
edge of canal preceded by usually strong transverse fold (Figure 15i:
blue arrow). Parietal callus normally well- developed. Radular rachid-
ian with more uniform cusps. Extrinsic odontophore muscles m1b
single. Aperture of salivary glands lateral. Anterior esophagus with
longitudinal folds. Valve of Leiblein well- developed. Stomach with sin-
gle duct to digestive gland. Female genital pore as wide, simple pore.
urn:lsid:zoobank.org:act:3D715FEE- 89DD- 4FB7- A46F-
1F66D827C6AA.
List of included taxa: Buccinastrum deforme (type species), B. duar-
tei, B. pay tense, B. uruguayense (all new combinations).
Etymology: the suffix astrum is a diminutive, jointed to Buccinum
a genus of similar shell shape; Buccinastrum = little Buccinum. Gender
neuter.
3.5 | Buccinastrum deforme (King, 1832) new
combination
(Figures 16– 19)
Buccinum deforme King, 1832: 349.
Buccinum globulosum Kiener, 1834: 12, pl. 10, Figure 33.
Buccinanops globosum var.: d'Orbigny, [1839] pl. 61, Figure 24.
Buccinum (Buccinanops) globulosum: d'Orbigny, [1841]: 435.
FIGURE 17 Buccinastrum deforme anatomy. (a) head- foot, male, right- slightly anterior view. (b) anterior region of visceral mass, ventral
view, ventral wall of kidney opened transversally and deflected, only topology of pericardium shown. (c) pallial cavity roof and visceral mass,
female, mostly ventral view. (d) anterior region of foot, female, sagittal section. (e) pallial cavity roof, female, transverse section in middle
level of osphradium. Scales = 2 mm
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PASTORIN O ANd SIMO NE
Buccinanops globulosum: d'Orbigny, [1842]: 157; Carcelles &
Parodiz, 1939: 764, Figures 7– 9; Carcelles, 1950: 63; Carcelles &
Williamson, 1951: 299; Barattini & Ureta, 1960: 117; Castellanos,
1970: 90, pl. 7, Figure 5; 1992: 13, pl. 2, Figure 10; Penchaszadeh,
1971a, 1971b: 477, Figure 2 a– b; Scarabino, 1977: 189, pl. 5, Figure 5;
Cernohorsky, 1984: 28; Farinatti, 1985: 219, pl. 1.
Buccinum ampullaceum Deshayes, 1844: 203, nomen novum pro
Buccinum globosum Kiener (in error pro globulosum) non Gmelin,
1791, nec Quoy & Gaimard, 1833.
Bullia deformis: Reeve, 1845, pl. 3, Figure 21; Adams & Adams,
1858:113; Kobelt, 1877: 290; Tryon, 1882: pl. 5, fig. 51; Ihering,
1895: 227; 1907: 425; Feruglio, 1933: 36.
Bullia (Buccinanops) deforme: Paetel, 1888: 116; Allmon, 1990:
16, fig. pl. 2, 13.
Bullia globulosa: Reeve, 1846 pl. 1, Figure 5; H. & A. Adams,
1858: 113; Tryon, 1882: 11, pl. 5, fig. 60; Pilsbry, 1897: 6; 1899:
127; Ameghino, 1906: 279; Ihering, 1907: 405; Feruglio, 1933 pl. 9,
Figure 30 a, b; Abbott & Dance, 1986: 178, Figure 1.
Bullia (Buccinanops) globulosum: Paetel, 1888: 116.
Buccinanops globulosum var. elata Strebel, 1906: 155, pl. 11, figs.
75 a– b.
Buccinanops globulosa: Peile, 1937: 185, Figure 20.
Buccinanops squalidus: Richards & Craig, 1963: 140 (non King,
1832).
Bullia (Buccinanops) globulosa: Allmon, 1990: 20, pl. 2, Figure 14.
Type material: Buccinum deforme: NHMUK 1854.12.4.470, 2 syn-
types (Figures 34– 36). B. globulosum: holotype MHNG 1296/17/1
(Figs. 40– 41).
FIGURE 18 Buccinastrum deforme anatomy. (a) head and hemocoel, ventral view, foot and columellar muscle removed, posterior
structures deflected. (b) odontophore, dorsal view, dorsal superficial layer of membranes and muscles removed, m3 sectioned longitudinally
and deflected. (c) anterior region of buccal mass, left view, mostly opened longitudinally. (d) odontophore, dorsal view, radular ribbon
and its annexed muscles removed, m3 sectioned and deflected. (e) region of valve of Leiblein (vl) and mid esophagus (me), mostly opened
longitudinally. Scales = 2 mm
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PASTORIN O ANd SIMO NE
Type locality: Buccinum deforme; ad flumen Plata (Rio de La Plata),
Gorriti [Island], Uruguay; B. globulosum: unknown; Buccinanops
globulosum var. elata: Puerto Madryn, Argentina; B. ampullaceum:
unknown.
3.5.1 | Distinctive description
Shell: medium size (to 70 mm, with usually 45 mm), globose.
Protoconch (Figure 16c– e) of 2– 2.5 whorls, of ~1.3 to 1.6 mm wide
in second whorl (n=3), smooth or with weak sparse spiral lines
(Figure 16e); whorls convex, globose. Spire short, ~40% of length;
last whorl very globose, of ~60% of total length. Suture well de-
fined. Sculpture wanting, except for growth lines (Figure 16c, h,
k). Aperture large, oval, ~60% of shell length and width. Siphonal
fasciole simple, rising below anterior half of callus. Columellar callus
defined, weak (Figure 16a, j) or strongly pronounced (Figure 16d, g,
m). Columella curved, ending in characteristic terminal plait. Outer
lip barely sigmoid (Figure 16b, e, l, n).
Head- foot and operculum: Similar to those of preceding species
(Figure 17a). Clearly bilobed posterior end of columellar muscle (cm),
left- slender component tall (cs). Females with cement gland in middle
region of anterior third, ending away from hemocoel (Figure 17d: cl).
Operculum (Figure 17o, p) large, suboval, with subterminal nucleus.
FIGURE 19 Buccinastrum deforme anatomy. (a) anterior region of visceral mass, ventral view, ventral wall of pericardium and digestive
gland region ventral to stomach removed. (b) anterior region of visceral mass, male, ventral view. (c) pallial oviduct, ventral view, topology of
some adjacent structures also shown. (d) same, transverse section in middle region of capsule gland (cg) shown. (e) penis and adjacent region
of pallial floor, dorsal view, penis duct (pd) seen by translucency. (f), nerve ring, ventral view, topology of esophagus (es) shown. (g) same,
dorsal view. Scales = 2 mm
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PASTORIN O ANd SIMO NE
Pallial cavity: (Fi gur e 17c, e) Features simil ar to preceding species .
Gill (gi) proportionally broader, occupying ~half of pallial roof; gill fil-
aments relatively lower, with right edge usually convex. Osphradium
slightly asymmetrical, with left filaments ~20% smaller than right fil-
aments. Small fold separating anterior end of gill from osphradium,
perpendicular to mantle border (ac).
Circulator y and excretory systems: (Figures 17b– c, 19a) Most
characters alike those of preceding species. Kidney with two flaps
of renal lobes (kl), both connected to efferent renal vessel (ek) and
to intestine (in). Hollow inner space wider, mainly in posterior region.
Digestive system: (Figures 18a– 19a) Most features similar to
those of preceding species. Exclusivities and notable charac-
ters are lateral transverse muscles (tm) slightly less developed
(Figure 18a), odontophore cartilages slightly broader and shorter,
with pair m8 longer (~45% of cartilage's length); fusion of carti-
lages along ~25% of their length (Figure 18d: oc); pair of muscles
m9 as single, broad bulk in each side (Figure 18b: m9); pair m4
with broader origins (Figure 18d: m4); posterior end of m6 and
m3c closer to posterior end of cartilages (Figure 18d), at ~1/7
of cartilages length from their posterior end. Radula: rachidian
tooth with 6– 12 cusps increasing towards center; rachidian base
strongly curved; lateral teeth with outer cusp hook- shaped, inner
cusp larger than 2– 3 intermediate cusps. Lateral teeth base with
blunt inner cusp. Salivary gland ducts lacking subterminal bulged
region (Figure 18c: sd); salivary aperture close to median line on
ventral wall, at middle level of oral tube (Figure 18c: sa); inner
surface with 5– 6 longitudinal, low folds, separated from each
other (Figure 18e: ae). Anterior esophagus simpler, almost straight
(Figure 18a: ae). Valve of Leiblein well- developed (Figure 18a, e:
vl), wit h an inner transvers e glandular fold and long iri desce nt cilia
turned proximally; no bypass furrow. Middle and posterior esoph-
agus internally smooth (Figure 18e: me, ep); middle esophagus
FIGURE 20 Buccinastrum duartei shell and operculum. (a– c) holotype MNHNM 769, dorsal, frontal and right views (L 28.3 mm).
(d) paratype MNHNM 770, frontal view (L 29.0). (e– g) MACN- In 43751 Mar del Plata, Buenos Aires, Argentina (L 31.8 mm). (h– i), MACN- In
43751 other specimen (L 30.3 mm). (j– k) operculum of the specimen of Fig. e, outer and inner views (L 8.5 mm). (l– n) MACN- In 43751 other
specimen (L 27.9 mm)
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PASTORIN O ANd SIMO NE
ve ry sh o r t; po s t e r ior es o p h a g u s gr adually acquiring 5– 6 in ner, lo n -
gitudinal folds (Figure 19a: ep). Gland of Leiblein narrow, shorter
(Figure 18a: gl), its duct short, simple, hollow; opening as simple
pore (Figure 18e: gd). Pair of ducts to digestive gland (Figure 19a:
dd), being esophageal duct ~half of intestinal duct. Stomach
bulged backwards, with strong circular muscle (Figure 19a: st);
internally with two transverse folds, one in esophageal insertion,
another in intestinal origin.
Genital system: (Figure 19b– e) Most attributes similar to those
of preceding species, distinctions following. Male seminal vesicle
slightly less- developed (Figure 19b: sv), with gonopericardial duct
present (go). Prostate simple, narrow, simply as duct swelling. Penis
broad in base (Figure 17a: pe), tapering gradually and uniformly up
to bluntly pointed tip (Figures 19e, 21a: pe); penis duct smoothly zig-
zagging along penis center (pd); papilla very small, subterminal (pp).
Female pallial oviduct with shallow constriction between albumen
and capsule glands (Figure 19c– d: ag, cg), albumen ~half of capsule
gland length; inner lumen ver y flat. Vaginal atrium ~1/5 of pallial ovi-
duct length, with posterior, dark- colored ingesting gland (ig). Female
pore simple, with elevated edges (Figure 19c: fp).
Central nervous system: (Figure 19f– g) Characters similar to those
of preceding species, being slightly broader. Cerebral commissure
(cc) relatively short, broad, rounded. Pair of buccal ganglia (bg) lo-
cated slightly separated from each other.
Habitat : Sandy and sandy- mud beaches, intertidal to shallow
subtidal levels.
Distribution: Cabo Sant a Maria, Uruguay to southern Santa Cruz,
Argentina.
Material examined: URUGUAY. Rocha; Cabo Santa Maria, USNM
340683, 3 spm. Maldonado; USNM 334528, 2 spm, USNM 270864,
1 spm; Punta del Este, MZSP 32081, 5 spm, MZSP 43098, 1 spm; MZSP
51347, 2 spm, MZSP 51628, 6 spm, MNR J 6163, 2 spm; Ilha Dos Lobos ,
MNRJ 1215, 1 spm, USNM 359242, 2 spm; Solis, MACN- In 9536– 2,
2 spm; Piriapolis, MACN- In 15433, 4 spm; Punta Negra, MNRJ 1772,
2 spm, USNM 359231, 4 spm. Canelones; Montevideo, MZSP 52109,
6 spm, USNM 364197, 7 spm; Isla Flores, MACN- In 28970, 2 spm;
MNRJ 856, 3 spm. ARGENTINA. MZSP 54048, 2 spm. Buenos Aires;
Cabo San Antonio, MACN- In 16310, 10 spm; Mar del Plata, MACN- In
8653– 7, 2 spm, MACN- In 9155– 4, 3 spm; Punta Carballido, Puerto
Quequén, MACN- In 26280, 1 spm; Monte Hermoso, MZSP 51592,
7 spm, MZSP52112, 2 spm, MACN- In 11235, 23 spm, USNM 219861,
1 spm, USNM 465578, 2 spm; Bahía Blanca, MZSP 10045, 3 spm,
MACN- In 14868, 40 spm, MACN- In 11236, 21 spm; Puerto Belgrano
MACN- In 11236– 1, 2 spm, MACN- In 6620– 17, 2 spm; Arroyo Parejas,
MACN- In 22471, 2 spm, MACN- In 11234, 3 spm; Bahia San Blas,
MACN- In 20252, 61 spm, MACN- In 20273– 2, 3 spm; Isla Trinidad,
MACN- In 19678, 1 spm; Carmen de Patagones, MZSP 52104, 15 spm;
MZSP 100 48, 4 spm, MZSP 52102, ~100 spm; mouth of the Río Negro,
MACN- In 20528, 17 spm; Río Negro; Puerto San Antonio, MACN- In
9379, 3 spm, MACN- In 9379– 28, 14 spm, MACN- In 9152– 13, 12 spm,
MACN- In 13359, 15 spm, MACN- In 9379– 24, 2 spm, MACN- In 9379-
26/27, 113 spm, MAC N- I n 937 9, 25 sp m, US NM 346842 , 3 spm; Pu e r to
San Antonio Este, MACN- In 23703, 12 spm, MACN- In 13358, 6 spm,
MACN- In 29138, 2 spm, MZSP 52103, 3 spm, MZSP 10032, 2 spm,
MZSP 78270, 3 spm, MACN- In 29662, 2 spm, USNM 381691, 3 spm;
Punta Villarino, MACN- In 13145, 16 spm, MACN- In 13146– 3, 5 spm;
San Antonio Oeste, USNM 807981, 2 spm Chubut; Golfo San Matias,
MACN- In 33041, 3 spm; Puerto San José, MACN- In 9174– 13, 5 spm;
Puerto Madryn, MACN- In 38163, 42°24′S 64°17′W, 3 spm, MACN- In
9676– 1, 5 spm, MACN- In 9281– 2, 2 spm, MACN- In 22477, 35 spm,
MAC N- In 10953, 41 spm , MACN- In 9171– 20, 25 sp m, MACN- In 9040,
12 spm, MACN- In 9171– 31, 3 spm, MACN- In 9675, 32 spm, MZSP
FIGURE 21 Buccinastrum duartei photo and SEM images. (a) detail of radula, scale = 200 µm. (b) radula, ampler view, scale = 200 µm.
(c) penis, dorsal view, scale = 1 mm. (d) same, detail of apical region, scale = 200 µm. (e) broken shell, arrow showing inferior grove in
penultimate whorl, frontal- apical view (MZSP 77556), scale = 5 mm.
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27
PASTORIN O ANd SIMO NE
52111, 8 spm; Golfo San José, MACN- In 9032– 14, 12 spm; Punta Norte,
MACN- In 11485, 51 spm, MACN- In 9158– 6, 5 spm; Puer to Pirámides,
MACN- I n 26400 , ~100 spm , MACN- In 26402 , 3 spm, MACN- In 9177–
4, 12 spm; Bahía Craker, MACN- In 26437, 20 spm; Bahía Bustamante,
MACN- In 26436, 2 spm, MACN- In 14447, 1 spm, MACN- In 26401,
2 spm, MACN- In 9032– 34, 3 spm; Camarones, MACN- In 26438,
11 spm; Rada Tilly, MACN- In 30881, 28 spm; Comodoro Rivadavia,
MACN- In 6826, 12 spm; Golfo de San Jorge, MNRJ 2069, 2 spm. Santa
Cruz; Caleta Olivia, MACN- In 35384, 3 spm.
Remarks: Buccinastrum deforme (King, 1832) was described in
one of the first molluscan reports from southern South America.
However, the type was never illustrated. Kiener (1834) described
Buccinum globulosum without any reference to King (1832).
Later authors (e.gBarattini & Ureta, 1960; Carcelles & Parodiz,
1939; Castellanos, 1970; Cernohorsky, 1984) used both names
as distinct species. The two syntypes housed at the NHMUK
(Figure 16a– f) clearly show extremes of variation of only one
species (Figure 16g– h). Usually, individuals living in Buenos Aires
province and Uruguay have shorter spire, heavier columellar
callus, and reduced aperture. This is probably related to the se-
lective predation of crabs of the genus Danielethus Thoma, Ng &
Felder, 2012 which fracture the shell using a special denticle on
the master claw. Along the coast of Chubut province and south-
wards, these crabs are absent. Thus, the shells develop a higher
spire, thinner whorls, and larger aperture (Figure 16j– l). Gomez
Espinosa et al. (2018) showed the relationship between the crab
D. crenulatus and B. deforme. Narvarte (2006) studied the popu-
lation ecology of the so- called B. globulosum. It is sympatric with
B. cochlidium particularly around Peninsula Valdes area; however,
both species live at different depths. While B. deforme is found in
intertidal settings— usually never deeper than 6 m— B. cochlidium
never is exposed during low tide, living in considerably deeper en-
vironments. The interaction between the American Oystercatcher
(Haematopus palliatus) and B. deforme (as B. globulosum), which
feeds on the leftovers of the crabs (Cyrtograpsus angulatus), was
studied by Daleo et al. (2005).
In the original description of B. deforme, King (1832: 349) men-
tioned the presence of “eggs” attributed to it. According to his
FIGURE 22 Buccinastrum duartei anatomy. (a) head- foot, male, anterior view. (b) pallial cavity roof, transverse section in middle level
of osphradium. (c) pallial cavity roof and visceral mass, mostly ventral view. (d) reno- pericardial region, ventral view, ventral wall of kidney
sectioned and deflected upwards, ventral pericardial wall removed, some adjacent structures also shown. (e) anterior region of foot, female,
transverse section in cement gland (cl) level. Scales = 2 mm
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PASTORIN O ANd SIMO NE
description, those capsules— “transparent nidus, the size of a turtle's
egg”— actually belong to the volutid Adelomelon brasiliana (Lamarck,
1811), a species fairly common in the area.
The radula of B. deforme is uniform in interpopulational studies.
There is no sexual dimorphism, neither differences between distant
populations nor variation correlated with size in specimens (Avaca
et al., 2010).
3.6 | Buccinastrum duartei (Klappenbach, 1961)
new combination
(Figures 20– 24)
Buccinanops duartei Klappenbach, 1961: 87, Figure 1a, b; Rios,
1970: 92, pl. 27; 1975: 96, pl. 27, fig. 400; 1985: 103, fig. 452;
1994: 130, pl. 41, fig. 559; 2009: 246, fig.; Castellanos, 1970: 94,
pl. 8, Figure 13; Cernohorsky, 1984: 28; Allmon, 1990: 27, pl. 2,
Figure 11.
Type material: holotype MNHNM 769 (Figure 20a– c), Paratypes,
MNHNM 770, 2 spm (Figure 20d), MNHNM 523, 6 spm, MNHNM
893, 1 spm, MNHNM 992, 5 spm.
Type locality: La Coronilla, Rocha Department, Uruguay.
3.6.1 | Distinctive description
Shell: (Figures 20e– i, l– n, 21e) Medium size (up to 35 mm), fusiform.
Protoconch paucispiral of 2 globose, smooth whorls. Teleoconch up
to 6 flat whorls. Growth lines all over shell, otherwise smooth. Most
grown specimens usually with small scars— probably boring sponges
FIGURE 23 Buccinastrum duartei anatomy, (a) head and hemocoel, ventral view, foot and columellar muscle removed; (b) buccal mass,
right view, some distal structures deflected; (c) foregut, dorsal view, posterior structures deflected; (d) buccal mass, right view, buccal cavity
opened longitudinally, salivary ducts seen by translucency; (e) region of valve of Leiblein, opened longitudinally; (f) odontophore, dorsal view,
more superficial structures removed or opened longitudinally, most muscles deflected to show more internal structures, right m4 (left in Fig.)
partially removed. Scales = 1 mm
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PASTORIN O ANd SIMO NE
(Clionaidae) (Figure 20h– i). Color pinkish, purplish, or clear brownish;
irregular subsutural white line usually present. Spire less than half of
total shell length. Suture impressed, sometimes irregular, overgrown,
showing enameling. Aperture large, usually half of total shell length
(Figure 20e, h, l). Parietal callus usually well- developed, reducing ap-
erture. Inner lip concave, siphonal inflexure strong. Outer lip barely
convex, almost straight (Figure 20f, m). Inferior inner groove inter-
nally in each whorl (Figure 21e: arrow).
Shell ultrastructure similar to other Buccinastrum spp.
Head- foot and operculum: (Figures 22a, 23a) Characters similar to
preceding species. Operculum (Figure 20j– k) large, suboval, margin
smooth, subterminal nucleus. Female cement gland close to anterior
edge, depth almost touching propodium (Figure 22e: cl).
Pallial cavity: (Figure 22b– c) Most features similar to those of pre-
ceding species. Those unique to this species are as follows: Siphon (si)
proportionally longer, small pallial fold between anterior ends of gill and
osphradium (ac), perpendicular to mantle border; cavity slightly shallower,
~3/4 whorl; gill (gi) and osphradium (os) proportionally broader; wider
portion of gill close to rectum; gill anterior and posterior ends bluntly
pointed; osphradium with uniform width along its length; osphradium
less sessile, with base smaller than filaments (Figure 22b: os); Gill fila-
ments relatively tall, tip protruded, right edge straight (Figure 22b: gi).
Circulator y and excretory s ystems: (Figure 22c– d) Similar to that of
preceding species, with following differences: relatively small kidney
ventral lobe attached to intestine, forming glandular cover of intes-
tine (Figure 22d: superior kl); presence of dorsal lobe, formed by pair
of zigzagging glandular folds surrounding posteriorly and left lobe
attached to intestine (Figure 22d: left kl); efferent renal vessel large
(rv), connected only to ventral lobe.
Digestive system: (Figures 23a– f, 24a) General organization sim-
ilar to those of preceding species, with following interesting fea-
tures: Proboscis retractor muscles thick posteriorly (Figure 24a:
FIGURE 24 Buccinastrum duartei anatomy. (a) foregut, ventral view, proboscis opened longitudinally, its walls and muscles deflected; (b)
midgut as in situ, ventral view; (c) male genital structures as in situ, ventral view in left, dorsal view in right; (d) right region of pallial cavity
roof, female, ventral view, transverse section in indicated region of pallial oviduct shown; (e) penis and adjacent region of pallial floor, dorsal
view, pallial floor artificially sectioned to show vas deferens (vd); (f) nerve ring, ventral view; (g) same, dorsal view. Scales = 1 mm
30
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PASTORIN O ANd SIMO NE
mr), mainly in ventral branches; sparse lateral transverse muscles
connecting esophageal origin and odontophore (Figure 23b: tm).
Odontophore muscles: pair m1l inserted more posteriorly and me-
dially (Figure 24a); retractor set of muscles m2 (m2m, m2a, m2c)
relatively thinner and slender (Figure 23b, f); m3l as continuous
dorsal cover with longitudinal fibers (Figure 23b); pair m4 as single
bundle originated at posterior end of cartilages (Figure 23f); m11a,
presence of two pairs of auxiliary ventral tensor muscles of radula
(beyond m11), originated on inner surface of m3c, in its posterior
region, running anteriorly covered by m9, inserting at end of radu-
lar ribbon, ventrally to insertions of pair m11. Odontophore carti-
lages slenderer, anterior fusion at ~1/5 of their length (Figures 23f,
24a: oc). Radula (Fig ure 21a– b): central tooth with 7– 10 cusps usu-
ally of equal size, except lateral cusps slightly smaller, central cusps
larger or thicker; rachidian base slightly curved. Lateral teeth outer
cusp hook- shaped, inner cusp larger, thicker than cusps 2 or 3;
sometimes asymmetric intermediate cusps; base with faint blunt
inner cusp. Salivary ducts with small subterminal bulged region
(Figure 23d: sp); salivary apertures in lateral region of dorsal folds,
in posterior level of oral tube (Figure 23d: sa). Anterior esophagus
wide and flattened, smooth inner surface (Figure 23b: ae). Valve of
Leiblein well- developed, including transverse glandular fold and
long cilia turned anteriorly (Figure 23c, e: vl), no bypass groove.
Middle esophagus with clear glandular surface, small longitudinal
folds, length ~15% of anterior esophagus (Figure 23c: me). Gland
of Leiblein slender an d sinuo us, ~ha lf of anter ior esophagu s length
and width (Figure 23a, c: gl); duct small, simple, hollow. Posterior
esophagus very wide, flaccid walls; internally sparse, narrow lon-
gitudinal folds (Figure 23a, c: ep). Stomach as blind sac, ~1/3 whorl
long, ~half local visceral width; posterior end bluntly pointed;
FIGURE 25 Buccinastrum paytense shells. (a– c) Buccinum squalidum King, 1832, syntype, NHMUK 20010205 (L 45.4 mm); (d) MACN- In
37899 Puerto San Julián, Santa Cruz, Argentina (L 41.8 mm); (e– f) operculum of specimen of Figure 26a, outer and inner views (L 13.5 mm);
( g – i ) Buccinum citrinum, holotype NHMUK 1874.12.11.149 (L 35.1 mm); (j– k) Buccinum squalidum King, 1832, syntype, NHMUK 20010205 (L
41.8 mm); (l) Kiener’s (1834) pl. 6, Figure 16
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PASTORIN O ANd SIMO NE
walls thin; inner surface with sparse longitudinal, narrow folds
directed toward intestine (Figure 24b: st); single small duct to di-
gestive gland in esophageal insertion (Figure 24b: dd). Anus wide,
shortly projected (Figures 22c, 24d: an).
Genital system: (Figure 24c– e) Most characters like those of pre-
ceding species. Distinctions following. Male with seminal vesicle
narrow, as simple zigzagging wide tube along ~1 whorl (Figure 24c:
sv). Prostate simple, short (~1/3 length of seminal vesicle), elliptical
(Figure 24c: pt). Penis (Figures 21c– d, 22d, 24c, e) relatively narrow,
weakly narrowing along its length, distal half flattened; tip rounded;
penis duct narrow, weakly zigzagging along penis middle region
(Figure 24e: pd); papilla subterminal, very small (Figure 24e: pp). Pallial
oviduct (Figure 24d) preceded by gonopericardial duct (go) inserting
close to visceral oviduct (vo); capsule gland (cg) as continuation from
albumen gland (ag), separated by loose ingesting gland (ig); albumen
with ~60% of capsule gland lengt h, but broader than it. Vaginal atrium
(Figure 24d: vg) ~1/6 of pallial oviduct length, walls flaccid, hollow,
tapering toward pore. Female pore as simple, longitudinal, small aper-
ture (Figure 24d: fp), located posterior and to right of anus.
Central nervous system: (Figure 24f– g) Most characters similar to
those of preceding species. Cerebral commissure (cc) narrow, short
(slightly shorter than each cerebral ganglion width). Shallow constric-
tions between dorsal and ventral set of ganglia. Small orifice between
both cerebral/pleural set and both pedal ganglia; anteriorly edged by
apparent extra cerebro- pleural commissure (Figure 24f). Buccal gan-
glia (bg) lacking clear commissure between both. Subesophageal/vis-
ceral ganglion (ge) small, spherical, connected by narrow nerve with
~double nerve ring length.
Habitat: Intertidal, in sandy and sandy- mud beaches.
Distribution: Rio de Janeiro (~23°S), Brazil to Miramar, Buenos
Aires province (~38°S), Argentina.
Material examined: BRAZIL. Rio de Janeiro; Recreio Dos
Bandeirantes, MNRJ 9104. Santa Catarina; Florianópolis, MZSP
51771, 2 spm, MZSP 51659, 1 spm, Praia da Armação, MZSP 51322,
2 spm. Rio Grande do Sul; MACN- In 25624, 1 spm; off Rio Grande,
USNM 654 363 , in 25 m, 2 spm; Trama ndaí, MZSP 33413, 4 spm, MZS P
52160, 8 spm, MNR J 6051, 2 spm, USNM 710 317, 4 spm; Torres , MZ S P
16131, 3 spm, MZSP 77556, 2 spm; Pinhal, MZSP 27314, 2 spm; São
Jose do Norte, MNRJ 1156, 2 spm; Tamandaré, MNRJ 2928, 2 spm;
Ponta da Cidreira, MNRJ 9332, 3 spm. URUGUAY. Rocha; USNM
381150, 11 spm; Chuy, barra del arroyo, USNM 875579; La Coronilla,
MZSP 33408, 23 spm, MACN- In 28772, 6 spm, MZSP 51616, 2 spm;
Cabo Santa Maria, MNRJ 1771, 2 spm, MNRJ 6541, 2 spm; La Paloma,
MACN- In 30480, 5 spm, MACN- In 30171, 2 spm. Maldonado;
Punta del Este, MACN- In 15375, 34°55′S 55°19′W, 20 m, 1 spm.
ARGENTINA. Buenos Aires; Villa Gesell, MACN- In 29373, 6 spm; Mar
del Plat a, MAC N- In 9363– 10, 5 spm, MACN- In 10240, 2 spm; Miramar,
MACN- In 8451– 25, 1 spm, MACN- In 8421– 25, 2 spm.
3.7 | Buccinastrum paytense (Kiener, 1834) new
combination
(Figures 25– 29)
Buccinum squalidum King, 1832: 349 (non Gmelin, 1791).
Buccinum Paytense Val. Kiener, 1834: 17, pl. 6, Figure 16.
FIGURE 26 Buccinastrum paytense photos and SEM images. (a) specimen fixed protruded (MZSP 122290) (L 25 mm). (b) same, detail of
posterior end of foot, dorsal view, scale = 5 mm. (c– e) radulae in SEM. (c) detail of central region, scale = 200 µm. (d) ample view, scale =
1 mm. (e) slightly lateral view, scale = 300 µm. (f– g), prehatching young shell, SEM. (f) frontal view. (g) dorsal view, scale = 1 mm. (h) same
detail of apex lateral view, scale = 0.5 mm
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PASTORIN O ANd SIMO NE
Buccinum citrinum Reeve, 1846 pl. 9, fig. 70 (non Wood, 1828);
Strebel, 1906: 153, pl. 11, fig. 74 a, b.
Bullia squalida: Reeve, 1847, pl. 4, Figure 26; H. & A. Adams, 1858:
113; Cunningham, 1870: 476; Kobelt, 1877: 289; Ihering, 1907: 537.
Bullia (Buccinanops) squalida: Paetel, 1888: 117.
Buccinanops squalidum: Carcelles & Parodiz, 1939: 758, Figure 10;
Castellanos, 1970: 95, pl. 7, Figure 7; Castellanos, 1992: 12, pl. 2,
Figure 11.
Buccinanops squalidus: Carcelles, 1950: 62, pl. 2, fig. 40; Carcelles
& Williamson, 1951: 299.
Buccinanops paytense: Cernohorsky, 1984: 30.
Bullia (Buccinanops) paytensis: Allmon, 1990: 25, pl. 2, Figure 9.
Type material: B. squalidum: NHMUK 20010205
(Figure 25a– c); B. pay tense: not found, probably lost; B. citrinum:
NHMUK 1874.12.11.149, illustrated (Figure 25g– i).
Type localities: Buccinum squalidum: unknown; B. paytense: Paita,
Perú (in error); B. citrinum: unknown.
3.7.1 | Distinctive description
Shell: (Figures 25a– d, j– l, 26a) Medium (up to 57 mm), fusiform. Color
reddish brown. Protoconch paucispiral of 1 1/2 to 2 whorls, 1.6 mm
wide in second whorl (Figure 26f– h). Sculpture wanting, except for
growth lines. Teleoconch of 6– 7 slightly convex whorls, spire shorter
than 1/2 of shell length. Suture well defined, impressed; 2– 3 subsu-
tural spiral cords, sometimes vanishing, very weak. Aperture large, sub
elliptic, more than half of total shell length. Parietal callus very weak,
but visible (Figure 25a, d, g, j). Inner lip shallowly concave; siphonal
inflexure weak. Outer lip simply convex (Figure 25b, h).
Head- foot and operculum: (Figures 26a– b, 27a, e) General fea-
tures similar to preceding species. Epipodial tentacle small, preceded
by medial white region (Figure 26b). Females with cement gland
close to anterior end of sole (Figure 27e: cl); relatively short, ending
far from hemocoel (Figure 27f: cl). Operculum more rounded, mainly
close to nucleus (Figure 25e– f).
FIGURE 27 Buccinastrum paytense anatomy. (a) head- foot, male, frontal view. (b) pallial cavity hoot and anterior region of visceral
mass, ventral view, ventral wall of pericardium removed. (c) buccal mass, right view, esophagus (ae) slightly deflected. (d) head and foregut,
proboscis (pb) extended and opened longitudinally, most posterior structures uncoiled. (e) extended foot, female, ventral view. (f) same,
sagittal section in its anterior region. (g) pallial cavity hoof, transverse section in middle level of osphradium. Scales = 5 mm
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PASTORIN O ANd SIMO NE
FIGURE 28 Buccinastrum paytense anatomy. (a) foregut, right view, detail of anterior region, oral tube and part of esophagus opened
longitudinally and deflected, portions of dorsal fold sectioned longitudinally to show salivary ducts. (b) odontophore, dorsal view, most
muscles removed, except for their origins, two transverse sections artificially done in equidistant levels. (c) same, detail of posterior end
of right cartilage, medial view, showing muscles origin. (d) esophagus sectioned longitudinally, inner surface exposed, insertion of salivary
ducts and gland of Leiblein shown. (e) anterior region of visceral mass, ventral view, ventral pericardial wall removed, ventral kidney lobe (kl)
deflected upwards, most digestive gland removed to show midgut. (f) anterior region of visceral mass and right- posterior region of pallial
cavity, male, ventral view, mostly showing genital structures as in situ. Scales = 2 mm
FIGURE 29 Buccinastrum paytense anatomy. (a) penis, dorsal view, topology of head also shown. (b) central nervous system, dorsal view.
(c) same, ventral view. Scales = 2 mm
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PASTORIN O ANd SIMO NE
FIGURE 31 Buccinastrum uruguayense SEM images and photograph. (a) radula, scale = 200 µm. (b) scale = 1 mm. (c) scale = 200 µm. (d)
specimen fixed protruded (MZSP 78248) (L 20 mm)
FIGURE 30 Buccinastrum uruguayense shells and operculum. (a– c) Bullia uruguayensis Pilsbry, 1897, ANSP 70507A1, syntype (L 21.1 mm).
(d– f) ANSP 70504B, syntype (L 15 mm). (g– h) MACN- In 30171 from La Paloma beach, Uruguay (L 27.8 mm); (i– k) MACN- In 30171, other
specimen (L 23.0 mm). (l– n) MACN- In 30171, other specimen (L 23.0 mm). (o– p) ANSP 705042C, syntype (L17 mm). (q– r) operculum of
specimen of Fig. (l- n) (L 12.0 mm), outer and inner views
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PASTORIN O ANd SIMO NE
Pallial cavity: (Figure 27b, g) Most attributes like those of preceding
species, with a few distinctions: Siphon (si) relatively shorter, stubby. Gill (gi)
and osphradium (os) relatively shorter and broader; right edge of gill close
to tectum. Gill filaments low, with right edge highly convex (Figure 27g:
gi). Osphradium with base narrower than filaments (Figure 27g: os).
Circulator y and excretory systems: (Figures 27b, 28e) Characters
similar to those of preceding congeneric species, except for single
kidney lobe, not organized in folds (Figure 28e: kl).
Digestive system: (Figures 27c– d, 28a– e) Most attributes alike
those of preceding congeneric species, distinctive features fol-
lowing. Buccal mass with lateral transverse muscles very devel-
oped (Figure 27c: tm). Odontophore with narrow and thin pair m1l
(Figure 27c), originating in septum between esophagus and odon-
tophore, running posteriorly at short distance from median line,
covering m3c, multiple insertions distributed in postero- dorsal
region of odontophore surface; narrow and thin pair m3l, simi-
lar to m1l, but originated from lateral surface of odontophore in
level of esophageal origin (Figure 27c: m3l); m3c and m6 relatively
shorter, keeping free ~1/3 of cartilage length (Figure 28b– c); pair
m4 separated in 5– 6 separated, narrow bundles (Figure 28b), main
posterior branch (m4), another smaller set antero- lateral (m4a);
pair m11 relatively thick and broad. Radula (Figure 26d– c): cen-
tral tooth with 9 cusps of similar size, last two cusps of each mar-
gin smaller; rachidian base curved. Lateral teeth with outer cusp
large, hook- shaped; inner cusp larger than 3, similar sized, inter-
mediate cusps. Lateral teeth base with blunt inner cusp weakly
developed. Salivary ducts possessing subterminal bulged portion
(Figure 28a: ss); salivary aperture in lateral side of oral tube, in
FIGURE 32 Buccinastrum uruguayense anatomy. (a) pallial cavity and last whorl of visceral mass, ventral view, ventral pericardial wall
and ventral cover of digestive gland removed. (b) head- foot, male, anterior view. (c) head and hemocoel, ventral view, foot removed, inner
structures partially deflected, proboscis (pb) sectioned longitudinally. (d) buccal mass, ventral view, odontophore basis partially sectioned
and deflected to right; salivary ducts seen if wall was transparent. Scales = 2 mm
36
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PASTORIN O ANd SIMO NE
its posterior level (Figure 28a: sa). Anterior esophagus relatively
simple, wide, flattened (Figure 27c, d: ae). Inner surface of en-
tire esophagus with longitudinal, low, narrow folds, slightly more
numerous in posterior esophagus (Figure 28d). Valve of Leiblein
well- developed, complete, with transverse glandular fold and long
cilia (Figures 27d, 28d: vl). Middle esophagus with ~1/3 of ante-
rior esophagus length. Hollow gland of Leiblein relatively long,
transversely folded on one side; aperture simple (Figure 28d: gd).
Posterior esophagus wide, walls thick, ~as long as anterior esoph-
agus (Figure 28e– f: ep). Stomach (st) with slightly bilobed posteri-
orly bulged region, walls thick but weakly muscular, but possessing
strong gastric muscle (Figure 28e: sm) originated in columella
close to end of columellar muscle, running for a short distance up
to ventral wall of stomach. Duct to digestive gland single, wide
(Figure 28e: dd), located close to intestinal origin.
Genital system: Most features similar to preceding species,
except for following: Male with seminal vesicle of medium size
(Figure 28f: sv), very narrow caliber; no detected gonopericardial
duct. Prostate as simple bulged region in middle portion of pallial
vas deferens, preceding its crossing to pallial floor (Figure 28f: pt).
Penis (Figures 27a, 29a: pe) possessing elliptical penial gland in
dors al si de, from its middle level up to it s tip (Figu re 29a: pn) ; pen is
papilla very small, subterminal (Figure 29a: pp). Female with shal-
low constriction between albumen and capsule glands (Figure 27b:
ag, cg), with albumen gland ~half of capsule gland. Genital atrium
with small muscular bursa copulatrix (Figure 27b: bc) in its ventral
side. Female aperture as simple transverse pore (fp).
Central nervous system: (Figure 29b– c) Closely similar to preceding
species, but with cerebral commissure short, broad, and rounded (cc).
Pair of buccal ganglia very small (bg), located close to each other.
Habitat: This is a locally common species that hides buried in
sandy beaches during low tide. It can be found because of a low bulge
where the animal waits for the rising tide. Averbuj and Penchaszadeh
(2016) described the reproductive biology of this species.
Distribution: South of Santa Cruz province, to Tierra del Fuego Is.,
in Argentina to Punta Arenas, Straits of Magellan in Chile.
FIGURE 33 Buccinastrum uruguayense anatomy. (a) foregut, detail of uncoiled posterior region, dorsal view, with detail of transverse
section of anterior region of gland of Leiblein. (b) buccal mass, right view, esophagus deflected. (c) odontophore, dorsal view, first layer of
membrane and muscles removed, m3 sectioned longitudinally and deflected. (d) same, most muscles removed except for their origins. (e)
same, detail of posterior end of right cartilage. Scales = 2 mm
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PASTORIN O ANd SIMO NE
Material examined: ARGENTINA. Santa Cruz; San Julián,
MACN- In 9200– 5, 1 spm, MACN- In 9040– 22, 1 spm. Tierra del
Fuego; USNM 126896, 1 spm; San Sebastián, MACN- In 12576,
6 spm; Punta Maria, Rio Grande, MACN- In 35385, 20 spm, MZSP
37899, 24 spm; Rio del Fuego, MACN- In 12574, ~80 spm; Bahía
San Martín, MACN- In 12578, 1 spm; Punta Sinaia, MACN- In
12577, 3 spm; Ushuaia, MZSP 12574, 1 spm, MZSP 52158, 1 spm;
Cabo Peñas, MACN- In 12575– 1, 7 spm. CHILE. Magallanes; Bahia
Gregorio, USNM 102533, 15 spm, USNM 96528, 1 spm; Punta
Arenas, MACN- In 12135– 1, 3 spm; Isla Dawson, MACN- In 12135,
2 spm, MACN- In 14135, 3 spm; Straits of Magellan, San Gregorio,
MNRJ 1563, 2 spm.
Remarks: King (1832) described this species without illustrat-
ing it. He did not notice the previous usage of the name by Gmelin
(1791), making his name a homonym. It was later figured and de-
scribed as Buccinum paytense by Kiener (1834) and as Buccinum
citrinum by Reeve (1847). Kiener (1834: 17) cited Valenciennes as
author of the species; however, the latter author never published
any mention of B. paytense. It was erroneously recorded from Payta,
Perú, but it was never recorded further north than Punta Arenas, in
the Chilean Pacific.
3.8 | Buccinastrum uruguayense (Pilsbry, 1897) new
combination
(Figures 30– 34)
Bullia Uruguayensis Pilsbry, 1897: 6.
?Buccinanops cochlidium: d'Orbigny, 1839, pl. 61, fig. 25; Pilsbry,
1897: 7 (non Dillwyn, 1817).
Buccinanops uruguayensis: Carcelles & Parodiz, 1939: 760, figs. 5
(radula), 12; Castellanos, 1970: 91, pl. 7, fig. 14; Rios, 1985: 103, fig.
456; 1994: 130, fig. 562.
Buccinanops uruguayense: Cernohorsky, 1984: 30.
Bullia (Buccinanops) uruguayensis: Allmon, 1990: 26, pl. 2, fig. 8.
Type material: ANSP 70507, 8 syntypes (Figure 30a– c).
Type locality: Maldonado Bay, Uruguay.
3.8.1 | Distinctive description
Shell: (Figure 30a– p) small to medium size (to 31 mm), fusiform.
Protoconch paucispiral, of 2 globose, smooth whorls. Teleoconch up
to 7 weakly convex whorls, spire around 1/2 of total shell length.
FIGURE 34 Buccinastrum uruguayense anatomy. (a) penis and adjacent structures, dorsal view. (b) pallial oviduct and adjacent structures,
ventral view. (c) central nervous system, dorsal view. (d) same, ventral view. Scales = 2 mm
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PASTORIN O ANd SIMO NE
TABLE 3 Part of the matrix by Simone (2011) used for present phylogenetic analysis, focusing the insertion of 16 buccinoideans (other
three species were already in the first version)
Abbreviations: Bops. = Buccinanops; Btrum. = Buccinastrum; Eng. = Engoniophos; Hemipol. = Hemipoligona
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39
PASTORIN O ANd SIMO NE
Suture well defined, impressed. Sculpture 3– 4 subsutural spiral lirae
present in most of specimens, vanishing in middle of whorl, reap-
pearing at base of last whorl (Figure 30c, f, h– i, n– o). Aperture large,
sh orte r tha n hal f of tot al sh ell le n g t h. Parie tal cal lus weak (Fig ure 30 a,
d, g, j, l, p); inner lip weakly concave; siphonal inflexure relatively
strong. Outer lip profile almost straight (Figure 30b, e, k, m).
Head- foot and operculum: Most features similar to those of pre-
ceding species (Figure 32b). Colorless, with epipodial tentacle well-
developed (Figure 31d: ed). Operculum elongated, pointed close to
nucleus (Figure 30q– r).
Pallial cavity: (Figure 32a) Attributes alike those of preceding spe-
cies, with following special features: Siphon very larger (si). Gill (gi)
and osphradium (os) with anterior and posterior ends more sharply
pointed. Wide space between gill and rectum. Gill and osphradial
filaments similar to B. deforme.
Circulator y and excretory systems: (Figure 32a) Similar to preced-
ing species. Heart proportionally smaller (ve). Kidney lobe with two
flaps attached to intestine (ki), transversely folded.
Digestive system: (Figures 32c– d, 33) General characters sim-
ilar to those of preceding species. Interesting features follow-
ing. Transverse lateral muscles connecting esophageal origin and
odontophore composed of more sparse fibers (Figure 33b: tm).
Odontop hore muscles: pair m1l ins erted in middle level of odonto-
phore length (Figure 33b: m1l); ventral pair m3l broad (Figure 33b);
pair m4 divided into three portions, larger bundle originated at
posterior end of cartilage, smaller bundle originating preceding
posterior 1/4 of cartilages (Figure 33c– d), another very small
bundle originating in median side of main bundle (Figure 33e:
m4); pair m9 divided into two separated bundles, anterior one in-
serted at middle length of radular sac (Figure 33c). Pair of odon-
tophore cartilages slenderer, fused portion shorter than 1/4 of
length (Figure 33c– d: oc). Radula (Figure 31a– c) central tooth with
7– 10 cusps, usually of same size, except for usually smaller lat-
eral cusps. Rachidian base curved. Lateral teeth with outer cusp
hook- shaped; inner cusp larger, thicker than 2 intermediate cusps;
lateral teeth base with very weak blunt inner cusp. Pair of salivary
ducts possessing bulged subterminal portion (Figure 32d: ss); sal-
ivary aperture on medial edge of dorsal folds, at middle length of
oral tube (Figure 32d: sa). Portions of esophagus similar to those
of B. paytense (Figures 32c, 33a: ae, me, ep); inner surface with
longitudinal, irregular, narrow folds. Valve of Leiblein complete,
lacking bypass (Figures 32c, 33a: vl). Gland of Leiblein small, an-
terior half hollow (Figure 33a: transverse section), with 5– 6 longi-
tudinal, glandular folds; duct very short (gd) simple. Stomach with
non- muscular walls (Figure 32a: st), as deep blind- sac, ~1/3 whorl
long; pair of small ducts to digestive gland located in esophagus-
intestine intersection (Figure 32a: dd).
Genital system: Most structures similar to those of preceding
species, remarkable features following. Male with seminal vesicle
of medium size. Prostate like that of B. pay tense. Penis marked by
FIGURE 35 Portion of cladogram containing the ingroup and other buccinoideans inserted in the scenario by Simone (2011), some nodes
are named with the numbers of that paper, new nodes are named by letters. Each node has shown the synapomorphies that base them, with
symbols as in the left frame; number above symbols means character (as in Simone, 2011); number below means the state in that node. Total
cladogram L= 3131; CI = 51; RI = 94
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PASTORIN O ANd SIMO NE
sudden bottleneck portion, producing slender, elongated distal
region of ~1/3 length of penis (Figures 32b, 34a). Penis duct (pd)
convolute, running closer to right side, except for slender distal
portion, containing duct narrow, straight, running centrally; penis
apex blunt, flattened; papilla small, subterminal (Figure 34a: pp).
Female with clear constriction separating albumen and capsule
glands (Figure 34b: ag, cg), capsule gland ~twice length of albu-
men gland. Vaginal atrium (vg) relatively long, ~1/4 of total pal-
lial oviduct length, tapering toward female pore; brown triangular
portion of ingesting gland in its posterior region, small muscular
concavity in its right- side corresponding to a small bursa copula-
trix. Female pore (fp) simple.
Central nervous system: (Figure 34c– d) As concentrated and with
similar attributes as in preceding species. Cerebral commissure
broad and short (cc). Pair of buccal ganglia apparently fused with
each other (bg). Visceral ganglion located posteriorly at a distance
equivalent to nerve ring length (Figure 34d: vg).
Habitat: rare, usually subtidal up to 30 m depth.
Distribution: Angra dos Reis, Rio de Janeiro, Brazil to Bahía San
Blas, Buenos Aires province, Argentina.
Material examined: BRAZIL. Rio Grande do Sul; Pinhal, MZSP
27314, 2 spm; GEDIP expedition MZSP 33750, 1 spm, MZSP 33898,
2 spm, MZSP 57522, 5 spm; off Mostardas, 31°09′S, 50°43′W, 21 m,
MZSP 51405, 4 spm (sta. 1725). URUGUAY. Rocha; La Paloma, MZSP
51406, 1 spm, MACN- In 28784, 2 spm, MACN- In 30481, 1 spm,
MACN- In 30487, 1 spm, MACN- In 30171– 2, 2 spm, MNRJ 6543,
2 spm, MACN- In 15151, 1 spm, MNRJ 1248, 2 spm, MZSP 78248,
33°54.7′S, 53°17.6′W, 15 m; Cabo Santa Maria, MACN- In 23582,
1 spm. Maldonado; Punta del Este, MZSP 51617, 2 spm. ARGENTINA.
Buenos Aires; Necochea, MACN- In 25199, 35°40S 56°03 W, 9.1 m,
14 spm, MACN- In 24202, 35°27′S 54°11′W, 23 m, MACN- In 24146,
36°24S 55°53 W, 16 m, 6 spm, MACN- In 24295, 37°18′S 56°49′W,
8 m, 2 spm, MZSP 10046, 1 spm; Mar del Plata, MACN- In 25364,
17 spm, MACN- In 4660– 4, 2 spm; Miramar, MACN- In 9247– 14,
12 spm; Monte Hermoso, MACN- In 6619– 53, 5 spm, MACN- In
11230– 1, 1 spm, MZSP 10052, 16 spm; Arroyo Parejas, MACN- In
11232– 1, 3 spm; Puerto Belgrano, MACN- In 11341, 3 spm; Bahía San
Blas, MACN- In 20254, 23 spm, MACN- In 20273– 3, 1 spm.
Remarks: Buccinastrum uruguayense has been scarcely men-
tioned in the local or international literature since its original de-
scription. Probably it was not easily recognized because of the
similarity with young specimens of B. cochlidium and others, one
of the locally most common species of Buccinanops. In addition,
Ihering (1907: 445) stated that it can be only a local variety of B. co-
chlidium, an assumption that can justify its absence in subsequent
surveys. Despite the size, which is clearly smaller, and the slender
spire, the presence of several subsutural lirae in the first whorls
(Figure 30h– i, k, n, o) as well as on the base of the last whorl, are
distinguishing characters. In addition, the shape and ornamentation
of the egg capsules are different from those of B. cochlidium which
has a characteristic and unique sculpture on one side. Averbuj and
Penchaszadeh (2016) provided some data on the reproductive bi-
ology of this species.
4 | PHYLOGENETIC ANALYSIS
As explained above, a sample of 19 buccinoideans (Table 1) were
analyzed using the Maximum Parsimony methodology outlined in
Simone (2011, 2020), including the 7 species studied here. Nine ad-
ditional species of buccinoideans, particularly of columbellids, fas-
ciolariids, melongenids, nassariids, and buccinid, were also included,
based on descriptions published elsewhere (Table 1), although five
species were already treated by Simone (2011): two of Buccinanops,
Oligohalinophila dorri (Wattebled, 1886) (as Nassodonta) and the
two species of Amphissa H. & A . Adams, 1853. The main objective
is to check the monophyly of the Buccinanops/Buccinastrum clade
and to provide inputs for a wide discussion and analysis of its rela-
tionships. Although the Buccinanops/Buccinastrum clade includes all
Recent species, it is acknowledged that the representatives of the
other buccinoideans taxa are far from being well- represented, as the
families are diverse. However, some superficial and preliminary in-
ferences are possible and are explored below.
The matrix resulting from the insertion of the 14 buccinoideans
in the matrix published by Simone (2011) is enormous; thus, only the
lines concerned with the present paper are reproduced in Table 3.
The remaining portions of the matrix were identical to the matrix
by Simone (2011). Anyhow, it is important to emphasize that some
states were inserted into that matrix (see material and methods
above) in order to improve the resolution of the present ingroup.
The resulting cladogram is mostly the same as the one shown by
Simone (2011: figure 20), with an expansion and increased resolu-
tion of the nodes 214 to 217 shown in this paper (Figure 35), using
the same node labels as in Simone (2011: figure 20), adding letters
to newly formed nodes (A– O) that are used in the discussion below.
Figure 35 also shows the synapomorphies of each node displayed by
symbols (i.e., black square = non homoplastic synapomorphy; white
square = reversions; black circle = convergencies – of course these op-
timizations take into account the entire cladogram by Simone, 2011).
A tota l of 12 cladogr ams were obt ained in the anal ysis, th e cla do-
gram portion of the ingroup did not possess any polytomy in the
strict consensus, and it is reproduced here (Figure 35), with 3131
steps, CI = 51, RI = 94.
5 | DISCUSSION
According to the analysis of the material studied, Buccinanops and
Buccinastrum are typical representatives of both malacological prov-
inces: Argentine and Magellanic. There are no species of these gen-
era ranging beyond these biogeographical areas.
All species have anatomical features and shell morphology, in-
cluding reproductive biology traits, that help us in the decision that
Buccinanops and Buccinastrum are valid genera. This has been ro-
bustly suggested phylogenetically, with representatives present at
least since the Neogene in deposits from Southern Patagonia (Griffin
& Pastorino, in prep.). The DNA sequence phylogenetic calibra-
tion has shown an age— for Buccinanops— of at least lower Miocene
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PASTORIN O ANd SIMO NE
(Galindo et al., 2016: 12); and the DNA sequence cladistic arrange-
ment is compatible with the present result, with the single studied
Buccinastrum, B. globulosus, as sister of a monophyletic branch that
included the three Buccinanops here considered (Galindo et al.,
2016, Figure 2).
Interestingly, Galindo et al. (2016) recognized some com-
mon characters uniting the three genera (Buccinanops, Bullia, and
Dorsanum), a branch accepted by Allmon (1990), Brown (1982),
Cernohorsky (1984), Haasl (2000), and Simone and Pastorino (2014),
as “the result of convergence during nassariid evolution.” For exam-
ple, the loss of the planktotrophic mode of larval development and
the presence of multicuspid radular lateral teeth. Allmon (1990) con-
cluded that the Nassariidae were monophyletic, with Dorsaninae-
Bulliinae being the sister group to both Cylleninae and Nassariinae;
he also found a (Dorsanum (Bullia – Buccinanops)) clade, that Haasl
(2000) recovered as paraphyletic (CI = 0.381, RI = 0.639).
5.1 | Taxonomical comparisons
In this discussion, the character numbers are the same as in Simone
(2011). The most important features that can be used for distin-
guishing the species formerly considered Buccinanops are properly
explored above, along the distinctive descriptions. Here, only the
more relevant aspects are discussed.
The morpho- anatomical investigation of the seven valid Recent
species formerly considered Buccinanops (Figure 35, node J) easily
shows two reasonably distinct groups. One of them clusters rela-
tively large species with specimens easily reaching 80– 100 mm that
live in subtidal waters and are mostly sampled by otter trawl or div-
ers from 10 to 50 m depths. Anatomically, there are eight synapo-
morphies supporting this clade (Figure 35: node K), some of them
are the duplicity of the pair of extrinsic odontophore muscle m1b
(character 333, state 1); the ventral aperture of the salivary glands,
close to median line (ch. 452, st. 3); the simplification of the anterior
esophagus (ch. 478, reversion to 0); the papilla- like female genital
pore (ch. 618, st. 2). Nevertheless, the most important synapomor-
phy is the reduction in the valve of Leiblein, which is only vestigial
(ch. 485, st. 4), marked only by a transverse fold in B. monilifer and
in B. cochlidium (Figure 5e– f), and only by a change of the internal
pleating in B. latus (Simone, 1996: figure 16). The length of the ce-
rebral commissure is another interesting feature of this branch, as it
tends to be very long (Figures 8a– b, 11a– c, 14c), particularly in B. co-
chlidium. The node K, which includes the type species B. cochlidium,
represents the revised concept of the genus Buccinanops, grouping
only three living species. Its single internal division clusters B. coch-
lidium with B. latus (Figure 35, node L), in which the synapomorphy
is the elongated rather sinuous anterior esophagus (ch. 478, st. 2),
particularly in B. cochlidium (Figure 5b: ae).
The other group formerly classified as Buccinanops is the node
M (Figure 35), which includes the other four ingroup species. Three
synapomorphies support this node, including the presence of irreg-
ular subsutural nodes (character 9, state 1); a well- developed, folded
callus (ch. 46, st. 1); and the presence of a single duct to the diges-
tive gland in the stomach (ch. 498, st. 1) (e.g., Figure 28e). However,
other additional attributes can be added to this branch: the relatively
smaller size of the shell, usually smaller than 50 mm; the intertidal
habitat, rarely occurring subtidally; the invariably smooth first whorl
vs. the presence of axial riblets, and the more robust, thick- walled
shell, possibly to resist the waves. Based on this set of characters,
the new genus, Buccinastrum gen. nov., is introduced to designate
this node. Its more basal branch is B. duartei, with the other three
species united at node N. Three synapomorphies support this taxon,
including the low gill filaments with convex right edges (character
206, state 4); the multiple pairs of odontophore muscles m9 (ch.
377, st. 4), and the separation of the albumen gland from the cap-
sule gland (ch. 632, st. 0). The following branch separates B. uruguay-
ense from node O, which groups B. deforme and B. paytense. These
synapomorphies support node O, including the lateral positioned
salivar y glands aperture (ch. 452, st. 5); the lack of folds in middle
esophagus (ch. 489, st. 0); and a broad duct to digestive gland (st.
509, st. 3).
A set of 11 synapomorphies support node J (Figure 35), that is,
Buccinanops + Buccinastrum, previously considered Buccinanops. The
most important characters are the single posterior epipodial ten-
tacle (character 113, state 2); the osphradium filaments widely at-
tached to mantle (ch. 192, st. 0); the lack of the odontophore muscle
“ma” (ch. 331, st. 0); the presence of lateral buccal mass protractor
muscles (m1a – ch. 332, st. 5); the longitudinal, lateral pair of odonto-
phore muscle m3 (ch. 344, st. 12/C); the arched rachidian, with ~10
cusps at the edge (ch. 418, st. 5); the small, simple papilla at the tip
of penis (ch. 563, st. 1), located subterminal (ch. 575, st. 3); and the
commissure between buccal ganglia incorporated to the nerve ring
(ch. 660, st. 2).
The node I (Figure 35), which represents the west and south
African genus Bullia, is supported by three synapomorphies, in-
cluding the oral tube muscles forming a circular, sphincter- like band
(character 328, state 4); the presence of the odontophore muscle
pair m2c running along median line (ch. 341, st. 3); and the odon-
tophore pair m9 multiple, inserting directly in cartilages instead of
in m3 (ch. 377, ct. 5). Bullia granulosa, one of the species included
here, resulted in a peculiar allocation using a DNA sequence data:
it fells in the nassariine Nay tia branch together with four other spe-
cies (Galindo et al., 2016) formerly considered Nassarius. All these
four species have small, sculptured shells with a large callus, very
typical of what formerly was considered Nassarius. The inclusion
of B. granulosa in this branch renders difficult a definition based on
non- genetic- molecular characters. Notwithstanding, all species are
now considered Naytia (WoRMS website), a very heterogeneous
taxon so far lacking definition. As no Naytia species is include d in the
present study, no additional comment is possible to be taken from a
morphological viewpoint.
Bullia (Figure 35, node I) is the sister- group of the Buccinanops-
Buccinastrum branch (node J) in node H, supported by six synapo-
morphies, including the loss of the eyes (character 130, reversion to
state 0); the flat form of the gill filaments (ch. 205, st. 2); the presence
42
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PASTORIN O ANd SIMO NE
of intermediate small cusps in radular lateral teeth (ch. 426, st. 3); the
thick- walled anterior esophagus (ch, 478, st. 3), and the presence of
gonopericardial duct (ch. 554, st. 2).
The node G (Figure 35) has a branch representing the subfam-
ily Dorsaninae as defined and Galindo et al. (2016) (only having
Dorsanum), and the node H. The node G is Dorsaninae as defined
by Allmon (1990), with the three genera. It is supported by 12
synapomorphies, reuniting Dorsanum with node H. The main
synapomorphies are the tendency for an unpigmented head- foot
(character 86, state 2); the slightly protruded head, flap- like, with
a tentacle at each lateral end (ch. 127, st. 3); a small fold in the
base of the siphon (ch. 162, st. 8); the presence of a pair of thin
lateral, dorso- ventral muscles uniting esophagus and odonto-
phore (ch. 316, st. 3); the multiple origins in outer edge of carti-
lages of th e odontophore musc le m2 (ch. 352, st. 2); the flat ten ed,
arched radular rachidian, with ~20 cusps in cutting edge (ch. 418,
st. 6); the gastric form bulging posteriorly (ch. 493, st 2); the
thick- walled, convolute, and muscular penis duct (ch. 553, st. 3);
the relatively long cerebral commissure forming small strap- like
arc (ch. 667, st. 2). Another synapomorphy (ch. 113, st. 5) refers
to the presence of a pair of epipodial tentacles, which actually is
shared with most nassariids. Its presence in this node is because
of the lack of epipodial tentacles in the nassariid representative
Engoniophos Woodring, 1928 (see discussion below), but possibly
th e state wo u ld suppor t a di f ferent br anc h if more na ssa rii ds were
included. In any case, the typical nassariid pair of epipodial ten-
tacles is large, while those of Dorsanum and Bullia are very small.
This difference can be indicative of different origins. Additional
discussion on the epipodial tentacles in nassariids is found else-
where (Allmon, 1990; Galindo et al., 2016; Haasl, 2000).
The remaining portion of the shown cladogram (Figure 35) refers
to outgroups, considered only for improving the present discussion.
The character survey is thus not as exhaustive. Contrarily, it was
only sufficient to arrange the closer taxa, to test the ingroup mono-
phyly, and for permitting a better- based argumentation. Considering
this scenario, the ensuing information can be given.
The node F gathers Engoniophos with the dorsanines- bulliines-
buccinanopsines (sensu Galindo et al., 2016) (node G). It is sup-
ported by two synapomorphies, including an elongated, relatively
reduced gland of Leiblein (character 486, state 2); and the loss of
the anterior bursa copulatrix in the pallial oviduct of females (ch.
612, st. 0). Engoniophos unicinctus (Say, 1826) is one of the repre-
sentatives of the family Nassariidae, a position that it has acquired
recently (Abbate et al., 2018; Galindo et al., 2016), as it was previ-
ously assumed to be Buccinidae. The inclusion of this nassariid in
the dorsanids is not a surprise, as the latter has been considered a
subfamily of the former. However, as stated above, it is interpreted
that there are enough differences to support a distinction of both
groups at subfamily rank.
The node E reunites Buccinum undatum with the nassariid br anch
(node F). Buccinum undatum is the type species of the genus that
is the type of Buccinoidea; it was thus included as a good repre-
sentative of Buccinidae, as well as its superficial shell similarity
with Buccinanops (literally a Buccinum lacking eyes). Three synapo-
morphies support this node, including a dorsal, Y- shaped pair m9 in
odontophore (character 377, state 3); and the wide, oblique radular
lateral tooth, with long cusp at each end, the lateral cusp being larger
(ch. 424, st. 4).
The two representatives of the Fasciolariidae, Pustulatirus ogum
(Petuch, 1979) and Hemipolygona beckyae (Snyder, 2000), are united
by the node C (Figure 35). Four synapomorphies support this node,
including osphradium as wide as gill (character 196, state 1); arched
lateral edges of radular rachidian (ch, 420, st. 1); and penis sharply
pointed (ch. 581, st. 1). Both representatives of the Melongenidae,
Pugilina morio (Linnaeus, 1758) and P. tupiniquim Abbate & Simone,
2015, are united by node D, which is supported by 12 synapomor-
phies. The more interesting ones are the shouldered shell (ch. 9, st.
3); the hairy periostracum (ch. 22, st. 1); the head preceded by a
long neck (ch. 92, st. 3), with very reduced tentacles (ch. 126, st. 3);
the osphradium with pointed filaments (ch. 183, st. 6); the salivary
glands aperture in median- ventral region of the oral cavity (ch. 452,
st. 3); the loss of the valve of Leiblein (ch. 484, st. 0) and a simple
papilla on the penis (ch. 536, st. 1).
Both, fasciolariids and melongenids, are joined at node B, sup-
ported by six synapomorphies. The more interesting are elongation
of the siphon (character 50, state 1); multiple pairs of proboscis re-
tractor muscles (ch. 98, st. 4), with muscles surrounding the base of
the proboscis (ch. 300, st. 2); and radular rachidian with 3 cusps at
tip (ch. 416, st. 4).
Node 217 (Figure 35) gathers all buccinoideans excluding the
columbellids (node 216). It is supported by nine synapomorphies,
being the more interesting the shell internal furrow in each whorl
(character 24, state 1); the not (or weakly) protruded head (ch. 92,
st. 7); the osphradium filaments with tips wider than their bases (ch.
192, st . 1); th e loss of the anal glan d (ch. 226, st. 0); the pair of retrac-
tor muscles of buccal mass, m2, narrow, long, inserting posteriorly in
odontophore cartilages (ch. 334, st. 2); the loss of the bypass of the
valve of Leiblein (ch. 485, st. 0); and the loss of the bursa copulatrix
(ch. 624, st. 0).
Node 216 brings together both species of Amphissa, which are
the Columbellidae representatives. The node has four synapo-
morphies supporting it, including the presence of an anal canal
in the aperture (character 41, state 1); a rounded edge of the foot
sole (ch. 112, st. 1); radular rachidian reduced or absent (ch. 416,
st. 6); and radular lateral tooth arched inwards, with bifid tip (ch.
424, st. 3).
Node A includes the columbellids (node 216) and node 217
(Figure 35), with seven synapomorphies supporting it. The more
important ones are a small portion of the pair m2 passing through
the nerve ring (character 337, state 2); the odontophore pair m2
inserting on radular sac, radular nucleus and posterior end of car-
tilages (ch. 340, st. 2); the odontophore muscle m3 as circular thin
fibers opposite to m6 (ch. 343, st. 3); the pair m5 originated in op-
posed side of m4 in cartilages (ch. 359, st., 3); the odontophore
pair of ventral radular tensors, m11, originated at posterior end of
cartilage alongside m5 (ch. 384, st. 4); transverse muscles (mt) as
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43
PASTORIN O ANd SIMO NE
two bands (ch. 396, st. 2); and pair of salivary ducts very narrow
and long, running attached to esophagus alongside aorta branch
(ch. 450, st. 5). The branch with Oligohalinophila dorri interposes
bet ween nodes 215 and A (Figure 35), preceded by eight more syn-
apomorphies. From those, the more interesting are the anal notch
at shell aperture close to suture (ch. 49, st. 2); a socket- like head
(ch. 92, st. 5); a branch of the odontophore pair of muscles m4 con-
nected at posterior vertex of cartilages (ch. 354, st. 1); the pair m8
lying on outer- dorsal edge of odontophore cartilages (ch. 375, st.
3); odontophore cartilages several times longer than wide (ch. 398,
st. 2); and the albumen and capsule gland at pallial oviduct difficult
to separate from each other (ch. 610, st. 3). O. dorri has been con-
sidered a nassariid and supposedly should be allocated after node
F (Figure 35). Its strange allocation may be thus considered an arti-
fact caused by missing data, as the available sample of this species
does not permit a full anatomical investigation. Possibly, if more
details are available, a better resolution of O. dorri, and its reallo-
cation closer to nassariids could be obtained. Under this setting,
the synapomorphies of both nodes 215 and A can be considered
synapomorphies of the Buccinoidea. Interestingly, the allocation of
O. dorri was also problematic following a DNA sequence approach,
with low support (Galindo et al., 2016).
An uncontroversial phylogeny of the Neogastropoda is far
from being well- established, both via morphological and DNA se-
quence approaches (e.g., Colgan et al., 2007; Fedosov et al., 2017,
2019; Hayashi, 2005; Modica et al., 2011; Simone, 2004, 2017;
Yang et al., 2019). Despite that, based on most results in the lit-
erature and in the present one, Buccinoidea appear to have the
Columbellidae at the base (Figure 35, node 216). This node has the
Fasciolariidae- Melongenidae in a branch (Figure 35: node B), and
node E (Figure 35). The node E apparently groups the Buccinidae
together with a branch with Nassariidae including Dorsaninae
(sensu Allmon, 1990) (Figure 35: node F). It is important to em-
phasize that the present phylogeny has the major concern to pro-
vide a better- based dis cussion , wit h no inte ntion of challenging the
current taxonomy of the Neogastropoda, mainly concerned to the
nassariid buccinoideans and allies (Dorsanidae, Buccinanopsidae,
etc.). It is, anyway, a step forwards, basing the discussion of future
additional phylogenies.
6 | CONCLUSIONS
1. The previous concept of the genus Buccinanops has alluded
monophyly, supported by 11 synapomorphies, and was restricted
to SE South American coast, with only one species reaching
the southern- east Pacific (B. paytense).
2. The phylogenetic arrangement of the species that was previously
considered Buccinanops allows its subdivision in two genera,
Buccinanops (8 synapomorphies) restricted to three large species
from relatively deep waters, and the new Buccinastrum gen. nov.
(3 synapomorphies) including four small species— with B. deforme
as type— mostly occurring intertidally.
3. Based on the differences and exclusive morphological charac-
teristics, the nassariid subfamily Dorsaninae appeared to be
valid. It is supported by 12 synapomorphies and includes four
genera— Dorsanum, Bullia, Buccinanops, and Buccinastrum gen.
nov. However, it is recognized that it has emerged as paraphyletic
in DNA sequence studies and the present study does not include
all taxa included in those.
4. The putative subfamily Dorsaninae is part of the Nassariidae,
which is sister of Buccinidae, part of the Buccinoidea, sharing the
15 synapomorphies of the superfamily.
ACKNOWLEDGMENTS
We are grateful to Jose C. Tarasconi (Porto Alegre), who provided
information and specimens from Brazil and the following research-
ers and curators helped in many ways to find type material and send
pictures: A. Pimenta (MNRJ), Yves Finet (MHNG), G. Darrigran (MLP),
A. Tablado (MACN), F. Scarabino, W. Serra, S. Wlodek (MNHNM), P.
Ca llom on (ANSP ); E. Stro ng and J. Haras ewyc h (US NM), V. Hero s, L. A.
Galindo and P. Bouchet (MNHNP). Four anonymous referees and edi-
tor made important comments on the manuscript. Special thanks are
due to M. Griffin (MLP) for his thoughtful suggestions. We acknowl-
edge funding by the Consejo Nacional de Investigaciones Científicas y
Técni cas (CO NIC ET) of Arge ntina, to wh ich G.P. belongs as mem ber of
the “Carrera del Investigador Científico y Técnico.” This work was sup-
ported in part by the Project PICT No. 942 from the National Agency
for Scientific and Technological Promotion, Argentina. A grant from
the FAPESP allowed G.P. to visit the collections of MZUSP in 2006.
DATA AVAIL AB ILI T Y STATE MEN T
The raw data and the data basing this paper are entirely reported
in it and also in giving open- accessed references, mainly Simone
(2011).
ORCID
Guido Pastorino https://orcid.org/0000-0003-3341-777X
Luiz Ricardo L. Simone https://orcid.org/0000-0002-1397-9823
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How to cite this article: Pastorino G, Simone LR. Revision of
the genus Buccinanops (Mollusca: Neogastropoda:
Nassariidae), an endemic group of gastropods from the
Southwestern Atlantic, including a new genus and accounts
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2021;00:1–46. https://doi.org /10.1111/jzs.12479
