Spheniopsidae Gardner, 1928 (Bivalvia): Conchological

Characters of Two New Species from off Brazil,

Southwestern Atlantic

Authors: Machado, Fabrizio Marcondes, and Passos, Flávio Dias

Source: American Malacological Bulletin, 33(2) : 212-220
Published By: American Malacological Society
URL: https://doi.org/10.4003/006.033.0207

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 Amer. Malac. Bull. 33(2): 212–220 (2016)

Spheniopsidae Gardner, 1928 (Bivalvia): Conchological characters of two new species

from off Brazil, Southwestern Atlantic

Fabrizio Marcondes Machado1 and Flávio Dias Passos2

1
Programas de Pós-Graduação em Ecologia e Biologia Animal, Universidade Estadual de Campinas (UNICAMP) CEP 13083-970,
Campinas, SP, Brazil
2
Depto. de Biologia Animal, Inst. de Biologia, Universidade Estadual de Campinas (UNICAMP), Caixa Postal 6109, CEP 13083-970,
Campinas, SP, Brazil
Correspondence, Flávio Dias Passos: flaviodp@unicamp.br

Abstract: The Spheniopsidae Gardner, 1928 comprises four living species of Spheniopsis Sandberger, 1861 and eight of Grippina Dall, 1912,
the latter restricted to the Pacific Ocean, the former also occurring in the Atlantic. Spheniopsis brasiliensis new species and Grippina coronata
new species are described from the Brazilian southeastern coast, based on a detailed study of their shells. They are distinguished from already
described species mainly by differences in the outline of the valves, external sculpture and right hinge dentition. Shallow micro-pits present
all over the outer surface of the dissoconch are viewed as similar to those already described for cuspidariids. They may, thus, represent further
evidence for the relatedness of spheniopsids and cuspidarioideans. The new species from Brazil represent the first finding of these rare minute
bivalves of this family from the southwestern Atlantic.

Key words: taxonomy, microbivalves, Anomalodesmata, Grippina, Spheniopsis

Members of the Spheniopsidae Gardner, 1928 are
among the most mysterious bivalve groups, there being
many citations of uncertainties about their phylogenetic
affinities (Mikkelsen and Bieler 2008, Bieler et al. 2010).
Formerly (Gardner 1928, Keen 1969, Coan 1990, Cosel
1995, Coan et al. 2000), they were believed to be related to
the Corbulidae (for example, the type-species of Spheniopsis
Sandberger, 1861 is Corbula scalaris Braun, 1851, a fossil
from the Middle Oligocene) but, more recently, they have
been mostly considered among the Anomalodesmata
(Marshall 2002, Mikkelsen and Bieler 2008). Their tiny
shells and presumed rarity have hampered the finding of
specimens with well-preserved internal tissues for anatomi-
cal and/or molecular studies and, as so, there are only a few,
somewhat speculative, notes about their biology, character-
izing them as infaunal, brooding bivalves (Coan 1990,
Mikkelsen and Bieler 2008).
Apart from Spheniopsis, known from the Pacific and
Atlantic Oceans, there is only one another recognized genus
belonging to the Spheniopsidae (Coan 1990), i.e., Grippina
Dall, 1912, which is restricted to the Pacific. Spheniopsis tri-
quetra (Verrill and Bush, 1898) was the first described
Recent species of this family (originally as a Montacuta
Turton, 1822), from North Carolina, USA (Verrill and Bush
1898). Recently, the taxonomy of both fossil and extant
Spheniopsis spp. were discussed by Coan (1990), who
described Spheniopsis frankbernardi Coan, 1990 from the
northeast Pacific (Baja California, Mexico, to Costa Rica).
212
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More recently, Spheniopsis senegalensis Cosel, 1995 and
Spheniopsis sculpturata Coan and Valentich Scott, 2012 were
also discovered, the first recorded as an endemic species
from Senegal, West Africa (Cosel 1995), and the second
only known from the Islas Galápagos, Ecuador (Coan and
Valentich Scott 2012). The first described species of Grippina
was Grippina californica Dall, 1912, its type-species occur-
ring from Santa Barbara, California (USA), up to Costa
Rica. The Californian Grippina berryana Keen, 1971 was
considered a synonym of G. californica (Coan 1990, Coan
et al. 2000, Coan and Valentich Scott 2012). Marshall (2002)
identified seven species restricted to New Zealand, one of
them being transferred from Mysella Angas, 1877 [Grippina
aupouria (Powell, 1937)] and the other six as new to science.
Apart from all these species, the Spheniopsidae also has
extinct (Oligocene) representatives, all belonging to
Spheniopsis (Coan 1990).
As a result of intense oil exploration activities in the
Campos Basin, southeast Brazil, studies of the sea bottom
have been conducted off the coast of the States of Rio de
Janeiro and Espírito Santo, mostly by the “Habitats Project –
Campos Basin Environmental Heterogeneity”. These research
activities are revealing molluscan taxa not yet recorded from
Brazil (Passos and Machado 2014, Corrêa et al. 2014). From
samples collected in this area, two new species of
Spheniopsidae were obtained and their shell characters are
herein described in detail and compared with the living spe-
cies already described in the literature.
 TWO NEW SPECIES OF SPHENIOPSIDAE (BIVALVIA) FROM OFF BRAZIL
MATERIAL AND METHODS
Hundreds of bottom samples were collected by the
“Habitats Project” using a box corer during the years of 2008
and 2009, both from the shelf and continental slope, and
sieved through a 0.5 mm mesh. Seventy-three of these sam-
ples from 17 to 148 m depths contained empty shells and liv-
ing individuals of spheniopsids, which were initially fixed in
4% formalin and then later preserved in 70% alcohol. Well-
preserved specimens were selected and mounted on alumi-
num stubs for examination under a scanning electron
microscopy (SEM). This set of stubs is deposited in the
Museum of Zoology “Prof. Adão José Cardoso” of the State
University of Campinas (ZUEC-BIV). The holotype, para-
types and non-type specimens are in the same museum; other
paratypes are in the Museum of Zoology of the University of
São Paulo (MZSP) and in the National Museum of Rio de
Janeiro (MNRJ). In the materials’ lists below all the lots are
ZUEC-BIV, unless stated.
RESULTS AND DISCUSSION
SYSTEMATIC
Class Bivalvia Linnaeus, 1758
Family Spheniopsidae Gardner, 1928
Remarks: The shell characters of this family were origi-
nally described by Gardner (1928), and subsequently by Coan
(1990), Coan et al. (2000), Marshall (2002), Mikkelsen and
Bieler (2008), and Coan and Valentich Scott (2012). The lat-
ter authors modified and improved upon the shell character
descriptions. Spheniopsids are generally characterized as
small, < 5.1 mm in length. The valves are either equal or sub-
equal, with a subtrigonal to ovate-trigonal outline, bearing
umbones positioned at the midline or posterior to it. The
anterior margin is rounded, while the posterior is more vari-
able in shape; the valves can be blunt posteriorly, or modestly
pronounced. The lunule and escutcheon are bounded by
radial ridges or ribs. Externally, the surface is either smooth
or sculptured with commarginal ribs. The right hinge plate
has two teeth placed on either side of the internal ligament; in
some species it seems that each tooth is a proximal thickening
of a sub-dorsal lamella (as described by Coan 1990) that
sometimes is better developed, but mostly either reduced or
absent in others. These teeth have also been called “cardinals”
(Dall 1912, Coan and Valentich Scott 2012) or “laterals”
(Coan et al. 2000); the anterior is generally more robust or
sub-equal to the posterior, which can be laminar, reduced or
almost absent. The left valve is edentulous and its dorso-anterior
and dorso-posterior margins fit into submarginal grooves
of the right hinge. The ligament, attached to subumbonal
resilifers, has fibrous (organic) and calcified (= lithodesma)
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213
parts. Dimyarian, with subequal adductor muscle scars; a
shallow, broad pallial sinus is present.
Genus Spheniopsis Sandberger, 1861. Type-species: Corbula
scalaris Braun, in Walchner (1851), by monotypy.
Remarks: Species of Spheniopsis are characterized by
shell valves with a subtrigonal outline, the posterior end being
produced to a variable degree during development and trun-
cated. As noted by Coan (1990), the external sculpture is also
variable, some species bearing fine concentric ribs or more
conspicuous, spaced, coarse ribs. On the dorso-posterior
slope, one radial rib is generally present bordering the
escutcheon, but a second rib can also be present. Coan (1990)
also pointed out that the posterior tooth of the right valve can
be either elongate or peg-like.
Spheniopsis brasiliensis new species (Figs. 1, 2)
Type-material
Holotype: ZUEC BIV 6173, station HAB16 F3 R2,
22°7′43.330″S, 40°18′48.056″W, Brazil -Campos Basin, off Rio
de Janeiro, “Habitats Project” coll, 6/vii/2009, 71 m [1 shell
with disarticulated valves]. Dimensions: Length: 1.78 mm;
Height: 1.29 mm; Width: 0.8 mm.
Paratypes: All collected in Campos Basin, off Rio de
Janeiro and Espírito Santo States, Brazil, during the “Habitats
Project” station HAB16 F3 R3, 22°7′43.380″S, 40°18′46.336″W,
73 m [6 pairs, ZUEC-BIV 6174]; HAB11 E3 R3, 22°8′9.289″S,
40°27′27.441″W, 65 m [18 pairs + 5 isolated valves, 6175];
HAB16 F4 R1, 22°12′38.570″S, 40°13′19.647″W, 99 m [25 pairs
+ 1 valve, 6176]; HAB16 F3 R2, 22°7′43.330″S, 40°18′48.056″W,
71 m [16 pairs, 6177]; HAB11 F2 R2, 22°3′41.580″S,
40°24′8.917″W, 56 m [6 pairs, MNRJ 26281]; HAB16 F2 R1,
22°3′41.420″S, 40°24′9.883″W, 56 m [2 pairs, MNRJ 26282];
HAB11 E4 R2, 22°17′42.207″ S, 40°26′59.691″W, 104 m [4
pairs, MNRJ 26283]; HAB16 F4 R2, 22°12′38.200″S,
40°13′18.927″W, 99 m [14 pairs, MNRJ 26284]; HAB17 D4 R3,
22°23′21.874″S, 40°34′57.133″W, 110 m [6 pairs, MZSP
117134]; HAB16 F3 R1, 22°7′43.550″S, 40°18′47.006″W, 72 m
[8 pairs, MZSP 117135]; HAB11 F3 R3, 22°7′43.147″S,
40°18′46.307″W, 73 m [4 pairs, MZSP 117136]; HAB11 F3 R1,
22°7′43.309″S, 40°18′46.483″W, 73 m [1 pair, MZSP 117137].
Mounted on stubs: Stub A. (ind.1) HAB11 F3 R1, 22°7′43.309″S,
40°18′46.483″W, 73 m [1 pair, ZUEC-BIV 6184]; (ind. 2)
HAB11 E3 R3, 22°8′9.289″S, 40°27′27.441″W, 65 m [1 shell
with disarticulated valves, 6185]; (ind. 3) HAB11 F4 R3,
22°12′37.347″S, 40°13′18.731″W, 99 m [1 shell with disarticu-
lated valves, 6186]; (inds. 4, 5, 6) HAB13 H5 R3, 21°42′37.911″S,
40°8′58.911″W, 147 m [3 pairs, 6187]; (ind. 7) HAB11 E3 R2,
22°8′9.091″S, 40°27′27.679″W, 65 m [1 shell with disarticulated
valves, 6188]; Stub B. (ind. 1) HAB11 C5 R3, 22°57′28.722″S,
40°50′30.265″W, 142 m [1 shell with disarticulated valves,
6189]; (inds. 2, 3) HAB13 H5 R3, 21°42′37.911″S, 40°8′58.911″W,
 214 AMERICAN MALACOLOGICAL BULLETIN
147 m [1 pair + 1 shell with disarticulated valves, 6190]; (ind.
4) HAB11 F2 R2, 22°3′41.580″S, 40°24′8.917″W, 56 m [1 shell
with disarticulated valves, 6191]; Stub C. HAB16 A5 R1,
23°36′14.972″S, 41°21′30.073″W, 145 m [internal tissues, 6192];
Stub D. HAB11 F5 R2, 22°17′ 25.519″S, 40°6′36.262″W, 143 m
[1 pair, 6193].
Non-type material: (See supplemental document at: doi:
10.4003/006.033.0207.s1).
Type-locality
Campos Basin, off Rio de Janeiro (22°7′43.330″S,
40°18′48.056″W), Brazil.
Geographic distribution
Up to now, known only from the Rio de Janeiro and
Espírito Santo States, Brazil. Bathymetric range (based on only
two samples containing specimens with internal tissues):
17–148 meters.
Diagnosis
Shell thin and delicate, small (maximum length of
2.56 mm); valves with a subtrigonal outline, rounded anteri-
orly and produced posteriorly. Umbones nearly central, slightly
prosogyrate and produced. Lunule wide, short, weakly demar-
cated; escutcheon narrow, long, well bounded by a radial rib in
each valve. Prodissoconch small, smooth, circular in shape.
Dissoconch with growth lines, a weak commarginal sculp-
ture and randomly distributed, shallow micro-pits. Right
hinge plate short and narrow, with two diverging peg-like
teeth: an anterior one continuous with a weak subdorsal
lamella and a posterior one which can be lamellar in more pos-
teriorly pointed shells. Left valve edentulous; anterior and pos-
terior margins slightly thickened near the umbones. Ligament
deeply located in an oblique resilifer placed below the umbo-
nes; its fibrous part forms the antero-ventral convex portion,
and the calcified part (lithodesma) forms the dorso-posterior
concave portion. Anterior adductor muscle scar elongated;
posterior adductor muscle scar elliptical-oval, smaller. Siphonal
sinus broad.
Description
Shell small (up to 2.56 mm in length), compressed
(average dimensions of ten largest specimens: Length =
2.05 ± 0.23 mm, Height = 1.3 ± 0.3 mm, Width = 0.7 ± 0.1 mm),
subtrigonal, posteriorly produced and truncated; slightly
inequivalve, with the dorsal margin of left valve fitted just
below the dorsal margin of right valve (Figs. 1A–D).
Umbones nearly central, slightly prosogyrate and pro-
duced. Valves thin and delicate, whitish, translucent,
not gaping, strongly inequilateral. Anterior dorsal margin
straight, with slight concavity next to the umbones;
anterior margin broad, uniformly rounded, smoothly
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· ·
33 2 2016
confluent with the ventral margin. Posterior dorsal margin
straight, longer and steeper than the anterior; posterior
margin slightly convex, vertical and short. Ventral margin
uniformly rounded anteriorly and nearly straight posteri-
orly, with a slight concavity just in front of the posterior
margin. An intraspecific variation was observed in the con-
tour of the valves (although more easily observed inter-
nally): some specimens are broader anteriorly (Figs. 2C,
G), and others are more pointed posteriorly (Figs. 2D, H).
Lunule smooth, wide, short, weakly demarcated; escutch-
eon narrow, long, smooth, bounded by a radial rib running
from the umbones up to the posterior margin dorsal end
(Fig. 1D); a second rib below this is nearly absent.
Prodissoconch I small (173 ± 16 μm, n = 10), circular,
smooth, generally well preserved in adult animals; limits of
prodissoconch II not visible (Figs. 1D–F). Periostracum
thin, transparent, colorless. Apart from the presence of
growth lines and a weak, irregular, commarginal sculpture,
at the external surface of the valves there are randomly dis-
tributed, shallow micro-pits, not crossing the shell; absent
in the prodissoconch (Figs. 1E–H). Inner margins smooth
(Figs. 2A–D, G, H). Right hinge plate short and narrow,
with two diverging teeth separated by the resilifer: an ante-
rior robust, curved, peg-like one continuous with a weak
subdorsal lamella extending up to the dorsal edge of the
anterior adductor muscle scar; and a curved, also peg-like,
posterior one, which can be lamellar in those more poste-
riorly pointed shells (Figs. 2A, C–F). Deep grooves are
present between the anterior and posterior dorsal margins
and their corresponding teeth in the right valve (Figs. 2E,
F). Left valve edentulous, its dorsal anterior and posterior
margins being slightly thickened proximally, near the
umbones (Figs. 2B, G, H); when the valves are united, these
margins fit into the right valve hinge submarginal grooves.
Ligament stout, deeply attached onto sunken oblique resil-
ifers placed below the umbones (Fig. 2I); fibrous part
forming its antero-ventral convex portion; trigonal-
shaped, calcified part (the lithodesma) forming its dorso-
posterior concave portion (Fig. 2J). Anterior adductor
muscle scar oval, elongated; posterior adductor muscle
scar elliptical-oval, smaller than the anterior. Retractor
muscles scars not visible. Pallial line nearly visible; sipho-
nal sinus broad, almost reaching the umbonal line.
Etymology
Relative to its original collection place - Brazilian waters.
Conchological remarks
This new Brazilian bivalve species can be distinguished
from all other extant members of Spheniopsis based on shell
characteristics only. It is distinct from the Pacific S. frankber-
nardi and S. sculpturata, and from S. triquetra and S. senegalensis
 TWO NEW SPECIES OF SPHENIOPSIDAE (BIVALVIA) FROM OFF BRAZIL 215

those of Spheniopsis brasiliensis n. sp.,

in which the valves are delicate, the
right one bearing a small hinge denti-
tion. Apart from these evident differ-
ences in external sculpture, S. sculpturata
can be distinguished by the contours of
the valves, as they present a subtrigonal-
elongated outline, only slightly pro-
duced posteriorly (Coan and Valentich
Scott 2012), different from the subtri-
gonal, posteriorly truncated, valves of
Spheniopsis brasiliensis n. sp.
The shell of Spheniopsis senega-
lensis is longer, with the posterior
end more pronounced than that of
Spheniopsis brasiliensis n. sp., which is
comparably taller and less elongated.
The hinge dentition of the former was
characterized and illustrated as repre-
sented by a “very small anterior cardi-
nal and a narrow but stronger posterior
tooth (more a lateral than a cardinal)
in form of a long laminar ridge parallel
to the postero-dorsal margin” (Cosel
1995), so contrasting with the distinct
peg-like teeth of Spheniopsis brasiliensis
n. sp.
The outline of the valves and the
hinge characters of Spheniopsis brasil-
Figure 1. Spheniopsis brasiliensis n. sp. (Outer view of shell). A–B, photomicrography of ho- iensis n. sp. are similar to those of
lotype (ZUEC BIV 6173), left and right valve, respectively. C–H, SEM views of paratypes; S. triquetra, which was originally
C, outer surface, left valve (ZUEC BIV 6184); D, dorsal view (ZUEC BIV 6185); E, detail of described in detail by Verrill and Bush
prodissoconch and dissoconch (ZUEC BIV 6186); F, detail of the edge between the pro- and
(1898), but with a poor illustration of
dissoconch, showing micro-pits restricted to the surface of the latter (ZUEC BIV 6184); G,
outer surface of dissoconch showing the growth lines, the weak commarginal sculpture and the
its shells. According to Coan and
micro-pits (ZUEC BIV 6184); H, profile view of a broken valve showing micro-pits not cross- Valentich Scott (2012), this species was
ing the shell (ZUEC BIV 6189). Abbreviations: di, dissoconch; es, escutcheon; lu, lunule; mp, cited as “Grippina sp. A” by Mikkelsen
micro-pits; pr, prodissoconch. Scale bars. A–B = 1000 μm; C, D, G = 100 μm; E, F = 20 μm; and Bieler (2008), with detailed SEM
H = 10 μm. (Color shown in electronic version only). views of the external and internal sur-
faces. Examination of these pictures
and of the type of S. triquetra deposited in the United States
described from the Atlantic. Good illustrations of all these National Museum, Smithsonian Institution, Washington
species are available in the literature, permitting comparisons D.C. (USNM 77627), permitted us to distinguish this species
and sound conclusions about the identity of Spheniopsis from Spheniopsis brasiliensis n. sp. mainly by the external
brasiliensis n. sp. sculpture: in the former it is composed of strongly marked,
When compared to Spheniopsis frankbernardi and S. regular concentric ridges, that are weak in the latter. Apart
sculpturata, differences are obvious from the external view, as from this, S. triquetra has a more solid shell and in the poste-
in Spheniopsis brasiliensis n. sp. the commarginal sculpture is rior slope there is a second radial ridge running from the
weak, while in the former species it is stronger [although with umbones to the posterior end, as stressed by Mikkelsen and
a variable degree of development in S. frankbernardi, as Bieler (2008); the Brazilian species is also fragile and only one
pointed out by Coan (1990) and Coan and Valentich Scott marked radial ridge is present at the edges of the escutcheon.
(2012)]. The shell of S. frankbernardi is also thick and the Genus Grippina Dall, 1912. Type-species: Grippina cali-
right hinge and its teeth are stronger, quite different from fornica Dall, 1912, by original designation.

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 216 AMERICAN MALACOLOGICAL BULLETIN · ·
33 2 2016

fine to coarse ribs, or these are absent.

Minute pits and calcareous periostra-
cal granules have being also recorded
(Marshall 2002). The right valve hinge
bears two short teeth and the left valve
is edentulous. The pallial sinus is short
and broad.

Grippina coronata new species

(Figs. 3, 4)
Type-material
Holotype: ZUEC BIV 6167, sta-
tion HAB11 E2 R2, 22°6′55.733″S,
40°38′58.263″W, Brazil - Campos
Basin, off Rio de Janeiro, “Habitats
Project” coll, 26/ii/2009, 56 m [1 shell
with disarticulated valves]. Dimensions:
Length: 1.67 mm; Height: 1.32 mm;
Width: 0.76 mm.
Paratypes: All collected in
Campos Basin, off Rio de Janeiro and
Espírito Santo States, Brazil, by the
“Habitats Project” station HAB13 I1
R2, 21°11′0.839″S, 40°28′27.312″W,
26 m [1 shell with disarticulated valves,
ZUEV-BIV 6168]; HAB13 H2 R3,
21°44′18.916″S, 40°17′14.472″W, 49 m
[1 specimen with soft parts, 6169];
HAB11 E2 R2, 22°6′ 55.733″S,
40°38′58.263″W, 53 m [3 pairs, 6170];
HAB11 F1 R3, 21°57′16.168″S,
Figure 2. Spheniopsis brasiliensis n. sp. (Inner view of shell). A–B, photomicrography of holo- 40°37′59.616″W, 26 m [2 pairs + 1
type (ZUEC BIV 6173), right and left valves, respectively. C–J, SEM views of paratypes; C–H, specimen with soft parts, 6171];
Intra-specific variation in outline; C, E, G, anteriorly broader specimen (ZUEC BIV 6187); D, HAB 13 Foz-30 R1, 21°34′12.752″S,
F, H, posteriorly pointed specimen (ZUEC BIV 6188). C, D, right valve; E, F, detailed view of 40°25′31.508″W, 29 m [1 pair + 1 spec-
the right hinge, exhibiting the anterior tooth continuous with a subdorsal lamella with anterior imen with soft parts, 6172]; HAB16 H1
and posterior grooves; note that in F the posterior tooth is more laminar in shape than in E. G, R1, 21°43′22.269″S, 40°31′52.609″W,
H, left valve. I, J, ventral and dorsal views (respectively) of internal ligament; the fibrous part 24 m [1 pair, MNRJ 26279]; HAB13 I1
can be distinguished from the calcified part (= lithodesma) (ZUEC BIV 6189, 6187). Abbre- R2, 21°11′0.839″S, 40°28′27.312″W,
viations: a, anterior tooth; fp, fibrous part; g, grooves; l, ligament; lt, lithodesma; p, posterior
26 m [2 pairs + 1 specimen with soft
tooth; sl, subdorsal lamella. Scale bars. A–B = 1000 μm; C, D, G, H = 300 μm; E, F = 100 μm;
I = 30 μm; J = 10 μm. (Color shown in electronic version only).
parts, MNRJ 26280]; HAB17 I2 R2,
21°11′1.370″S, 40°28′29.103″W, 26 m
Remarks: As noted by Coan (1990), the shell of Grippina [1 specimen with soft parts, MZSP
can be described as similar to Spheniopsis, the main difference 117138]; HAB13 Foz-21 R1, 22°6′21.254″S, 40°43′39.365″W,
being exhibited in the posterior end, that is not produced as 47 m [1 shell with disarticulated valves, MZSP 117139].
in the latter genus; instead, in the former, the shell is blunt Mounted on stubs: Stub A. (inds. 1–4) HAB11 D2 R1,
posteriorly. Additionally, at the external dorso-posterior sur- 22°12′52.897″S, 40°51′12.067″W, 52 m [3 pairs + 1 shell with
face (also called the “posterior slope” by Coan 1990) two low disarticulated valves, ZUEC-BIV 6178]; Stub B. (ind. 1)
radial ridges are generally present, one more dorsally bound- HAB17 Foz-21 R3 22°6′20.149″S, 40°43′41.039″W, 47 m
ing the escutcheon, and another just below this and com- [1 shell with disarticulated valves, 6179]; (ind. 2) HAB17
monly weaker. The commarginal sculpture is composed of Foz-10 R1, 21°55′26.077″S, 40°49′11.412″W, 21 m [1 shell with

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 TWO NEW SPECIES OF SPHENIOPSIDAE (BIVALVIA) FROM OFF BRAZIL 217

truncated posteriorly. Umbones promi-

nent, displaced posteriorly at a posi-
tion about 1/3 of the total shell length.
Lunule deep, wide, well demarcated;
escutcheon deep, wider, longer, bounded
by a radial rib in each valve; second rib
nearly absent. Prodissoconch large,
pronounced, with a rounded contour
and two central elevations. Dissoconch
with growth lines and randomly dis-
tributed, shallow micro-pits. Right
hinge plate short and narrow, with two
similar, peg-like, diverging teeth: an
anterior one continuous with a sub-
dorsal lamella, and a posterior one
somewhat closer and more robust
than the anterior in specimens with
higher shells. Left valve edentulous;
anterior and posterior margins slightly
thickened near the umbones. Ligament
deeply located on an oblique resilifer
placed below the umbones; its fibrous
part forms the antero-ventral con-
vex portion, and the calcified part
Figure 3. Grippina coronata n. sp. (Outer view of shell). A–B, photomicrography of holotype (lithodesma) forms the dorso-posterior
(ZUEC BIV 6167), left and right valve, respectively. C–H, SEM views of paratypes; C, dorsal concave portion. Anterior adductor
view (ZUEC BIV 6180); D, outer surface, right valve (ZUEC BIV 6183); E, F, prodissoconch, muscle scar elongated; posterior adduc-
dorsal and lateral view, respectively and dissoconch (ZUEC BIV 6180, 6181); G, detail of the
tor muscle scar oval, smaller than the
edge between the pro- and dissoconch, showing micro-pits restricted to the surface of the latter
(ZUEC BIV 6180); H, profile view of a broken valve showing micro-pits not crossing the shell
anterior. Pallial line and siphonal sinus
(ZUEC BIV 6182). Abbreviations: di, dissoconch; es, escutcheon; lu, lunule; mp, micro-pits; almost invisible.
pr, prodissoconch . Scale bars. A–B = 1000 μm; C–D = 100 μm, E = 30 μm; F, G = 20 μm;
H = 10 μm. (Color shown in electronic version only). Description
Shell small (up to 1.76 mm in
length), compressed (average dimen-
disarticulated valves, 6180]; Stub C. (ind. 1) HAB13 Foz-21 sions of ten largest specimens Length = 1.65 ± 0.2 mm, Height =
R1, 22°6′21.254″S, 40°43′39.365″W, 47 m [internal tissues, 1.3 ± 0.4 mm, Width = 0.74 ± 0.2 mm), ovate-trigonal to
6181]; Stub D. (ind. 1) HAB13 Foz-24 R1, 21°50′20.315″S, 40° subtrigonal, truncated posteriorly; slightly inequivalve, with
31′ 38.915″W, 27 m [1 pair, 6182]. the left valve dorsal margin fitting just below the right valve
Non-type material: (See supplemental document at: doi: dorsal margin (Figs. 3A–D). Umbones prominent, ortho-
10.4003/006.033.0207.s1). gyrate, displaced posteriorly at a position about 1/3 of the total
shell length. Valves fragile, whitish, translucent, not gaping,
Type-locality strongly inequilateral. Anterior dorsal margin straight, short;
Campos Basin, off Rio de Janeiro (22°6′55.733″S, anterior margin broad, uniformly rounded, smoothly conflu-
40°38′58.263″W), Brazil. ent with the ventral margin. Posterior dorsal margin straight,
Geographic distribution long, steep; posterior margin almost straight, oblique. Ventral
Hitherto, known only from the Rio de Janeiro and Espírito margin rounded anteriorly and nearly straight posteriorly. An
Santo States, Brazil. Bathymetric range (based on fifteen sam- intraspecific variation was observed in the contour of the
ples containing specimens with soft tissue): 21–53 meters. valves: some specimens are higher and more convex ventrally
(Figs. 4C–D), whereas others are longer, more pointed posteri-
Diagnosis orly and with an almost straight ventral margin (Figs. 4F–G).
Shell thin and delicate, small (maximum length of Lunule smooth, deep, wide, well demarcated; escutcheon
1.76 mm); valves with ovate-trigonal to subtrigonal outline, slightly smooth, deep, wider, longer, well defined in each valve

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 218 AMERICAN MALACOLOGICAL BULLETIN · ·
33 2 2016

shells (Figs. 4A, C, F, E, H). Deep

grooves are present between the ante-
rior and posterior dorsal margins and
their corresponding teeth in the right
valve (Figs. 4E, H). Left valve edentu-
lous, its anterior and posterior margins
being slightly thickened proximally
near the umbones (Figs. 4B, D, G);
when the valves are united, these mar-
gins fit into the right valve hinge sub-
marginal grooves. Ligament deeply
located on sunken oblique resilifers,
placed below the umbones (Fig. 4H);
fibrous part forming its antero-ventral
convex portion; trigonal-shaped,
calcified part (lithodesma) forming
its dorso-posterior concave portion.
Anterior adductor muscle scar elliptical,
elongated; posterior adductor muscle
scar oval, smaller than the anterior.
Retractor muscles scars not visible. Pallial
line and siphonal sinus almost invisible.
Etymology
From the Latin coronam, crown.
This name refers to the prodissoconch
Figure 4. Grippina coronata n. sp. (Inner view of shell). A–B, photomicrography of holotype that this Brazilian species bears; with
(ZUEC BIV 6167), right and left valves, respectively. C–H, SEM views of paratypes, showing its two characteristic central elevations
intra-specific variation; C, D, E, higher specimen (ZUEC BIV 6178); F, G, H, longer, posteri- it resembles a crown.
orly pointed specimen (ZUEC BIV 6179). C, F, right valve; D, G, left valve. E, H, detailed view
of the right hinge, showing the anterior tooth continuous with a subdorsal lamella with ante- Conchological remarks
rior and posterior grooves and posterior tooth; in H the ligament was maintained in place. Ab-
Grippina coronata n. sp. is distinct
breviations: a, anterior tooth; g, grooves; l, ligament; p, posterior tooth; sl, subdorsal lamella.
from G. californica, the latter having
Scale bars. A–B = 1000 μm; C, D, F, G = 200 μm; E, H = 100 μm. (Color shown in electronic
version only). thick valves with a subquadrate to
subtrigonal outline and an external
sculpture composed by commarginal
by the presence a radial rib running from the umbones up to ribs (Dall 1912, Coan 1990, Coan et al. 2000, Coan and
the posterior pointed end (Fig. 3C); second rib nearly absent. Valentich Scott 2012), different from the thin, ovate-trigonal
Prodissoconch I well preserved in young and adult individuals, to subtrigonal and externally smooth valves of the for-
large (254 ± 14μm, n = 7), pronounced, with a rounded con- mer. The hinge dentition is also different, much more
tour and two central elevations; limits of prodissoconch II not developed in G. californica than in Grippina coronata
visible (Figs. 3E–G). Periostracum thin, transparent, colorless. n. sp.
Apart from growth lines, at the external surface of the valves When compared with the New Zealand species, Grippina
there are randomly distributed, shallow micro-pits (absent in coronata n. sp. can be considered distinct from Grippina rex
the prodissoconch), which do not cross the shell (Figs. 3F–H). Marshall, 2002 and Grippina spirata Marshall, 2002, which
Inner margins smooth. Right hinge plate short and narrow, have thicker valves with an outline extended anteriorly, and
with two similar, peg-like, diverging teeth separated by the so appearing much more elongated. The shell of Grippina
resilifer; anterior tooth continuous with a sub-dorsal lamella, pumila Marshall, 2002 was characterized as moderately thick,
narrow in the proximal, umbonal portion, widening as it with umbones positioned at the mid-shell length position in
approaches the distal portion, near the dorsal edge of the ante- specimens less than ~1.6 mm long (Marshall 2002). This
rior adductor muscle scar; posterior tooth somewhat closer measurement is almost the maximum shell length of Grippina
and more robust than the anterior in individuals with higher coronata n. sp., in which the umbones are displaced posteriorly

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 TWO NEW SPECIES OF SPHENIOPSIDAE (BIVALVIA) FROM OFF BRAZIL
at a position about 1/3 of the total shell length. Grippina
aupouria grows up to 4.15 mm long, being only smaller than
G. rex (5.10 mm); both are obviously much larger than
Grippina coronata n. sp. In Grippina acherontis Marshall,
2002 (up to 2.45 mm long) calcareous periostracal granules
were observed at the external surface of the valves. In Grippina
globosa Marshall, 2002 (up to 1.43 mm long) there are pits in
the early dissoconch. These are important differences that
can be pointed out as distinctive from the Brazilian species, in
which micro-pits are distributed all over the dissoconch outer
surface; here, granules were not observed. Grippina punctata
Marshall, 2002, also a small species (up to 2.15 mm long), was
considered the thickest of all these New Zealand species,
and, thus, obviously different from the fragile Grippina cor-
onata n. sp. The right valve hinge dentition is also much
more developed in all New Zealand species than in Grippina
coronata n. sp.
CONCLUSIONS
Assessment of spheniopsid specimens with internal tis-
sues are scarce. Studies on this family are traditionally based
only on shell characters, which have been used for both taxo-
nomic and phylogenetic conclusions. Species distinctions are
currently made by observations of differences in the outlines
and external sculptures of the valves, as for example by Coan
(1990). The hinge structure of the right valve can also be
important, as shown here, for distinguishing Spheniopsis
brasiliensis n. sp. and Grippina coronata n. sp.
The structure of the internal ligament has been pivotal
in discussions about the phylogenetic affinities of the
Spheniopsidae. In all species of this family, it is attached to
sunken resilifers and there is a calcified supporting portion (=
lithodesma), characters which were considered by Marshall
(2002) for rejection of the former hypothesis of a relationship
with the Corbulidae, and for placing them among the
Anomalodesmata (an opinion also followed by Mikkelsen
and Bieler 2008). The exact position of spheniopsids in this
order, however, remains uncertain, and dependent on ana-
tomical and/or molecular data. Marshall (2002) suggested
affinities with the Cuspidariidae (Cuspidarioidea), based on
shell shape and external sculpture. In both Brazilian new spe-
cies, micro-pits were observed in the shell surface, their struc-
ture being similar to those recorded for other spheniopsids
(Marshall 2002) and cuspidariids (e.g., Oliveira and Absalão
2009, Absalão and Oliveira 2011). The presence of these pits
is probably another character to be considered as evidence of
relatedness among these families.
This work represents the first record of spheniopsids
species occurring in the southwestern Atlantic. Both Brazilian
species have small, fragile, shells, this being the most probable
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219
cause for them never being recorded before. Additionally,
they are presumably rare animals for only a few empty shells
and whole specimens were obtained. A similar case of descrip-
tion of rare, small, bivalve species from Brazil was made by
Passos and Machado (2014), this resulting from the great col-
lection effort made by the Habitats Project. Certainly, there
are more spheniopsids to be discovered, as noted by Redfern
(2001) and Mikkelsen and Bieler (2008) from the Atlantic,
and Marshall (2002) from the Pacific. Apart from new yet
unknown species, there are examples of both Spheniopsis and
Grippina species, which were originally described and attrib-
uted to other bivalve genera so that other members of this
enigmatic family may be discovered in the near future.
ACKNOWLEDGMENTS
This work was carried out with logistic and financial sup-
port provided by CENPES/Petrobras, under the “Habitats
Project”; we specially thank Dr. A. P. Falcão, who kindly invited
us to participate as taxonomists in this project. Financial sup-
port was provided by a scholarship from CAPES awarded to F.
M. Machado. Thanks are due also to A. C. S. Sprogis and S. M.
F. Ferraz (Laboratory of Microscopy – IB/UNICAMP), who
provided assistance in the techniques of microscopy; to E. V.
Coan and P. Valentich Scott (Santa Barbara Museum of
Natural History) for discussions about the identity of the spe-
cies and for having furnished important bibliographic mate-
rial; to R. S. Absalão and R. M. A. Figueira (UFRJ) who kindly
shared their opinions about the material from Campos Basin
in the beginning of this work; and to E. Strong (USNM) who
sent the pictures of the holotype of G. triquetra.
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Submitted: 15 September 2014; accepted: 14 December 2014;

final revisions received: 24 February 2015

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