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Amer. Malac. Bull. 32(2): 1–15 (2014) http://www.bioone.org/toc/malb/32/2

Description of four new species of Ledella Verrill and Bush, 1897 (Pelecypoda:
Nuculanidae) off the coast of Brazil, using a morphometric approach

Diniz Corrêa Paone Viegas, Natalia Pereira Benaim and Ricardo Silva Absalão

Universidade Federal do Rio de Janeiro (UFRJ), Instituto de Biologia, Departamento de Zoologia, Laboratório de Malacologia, Ilha do
Fundão, Rio de Janeiro, Brazil.CEP: 21941-901.
Correspondence, Diniz Corrêa Paone Viegas: diniz.viegas@gmail.com

Abstract: As a result of deep-water oil prospecting in the Campos Basin, Brazil, a rich diversity of protobranchiate pelecypods has been
revealed. In this study, we identified four new species of Ledella and conducted a morphometric study to corroborate their delimitation. The
resulting conchological-based statistic models are in 95% agreement with the traditional criteria for the right valves and 88.33% for the left
valves. Certain variables were particularly relevant in these analyses; these variables expressed the importance of the shape of the rostrum and
hinge plate for the right valves and the general shell shape and hinge plate for the left valves. The hinge plate variables are not typically used
as diagnostic characters of protobranchiate pelecypods and, when applied, they are not given much importance. However, our statistical
analyses stressed the importance of these characters. Therefore, we strongly suggest that future descriptions of protobranchiate pelecypods
place greater emphasis on these characters.

Key words: Biodiversity, deep sea, hinge plate, morphometry, Bivalvia.

As a consequence of oil prospecting in the Campos Basin Recently, Sharma et al. (2013) completed a major mo-
(20–23°S) by the oil company Petrobras, deep-water samples lecular analysis of the Protobranchia, showing that the genus
have been obtained, which have increased our knowledge of Ledella is not monophyletic. However, for the use of species
deep-sea mollusks from the southeast of Brazil over the last delimitation, we have analyzed all the species here described
10 years (e.g., Domaneschi and Lopes 1990, Absalão et al. as a unit.
2003, Caetano et al. 2006, Zelaya et al. 2006, Simone and Like the majority of protobranchiate Pelecypoda, Ledella
Cunha 2008, Oliveira and Absalão 2009, Passos and Birman has a conservative shell shape, subtle interspecific differences
2009, Figueira and Absalão 2010). and a lack of clear boundaries between genera. The species
Protobranchia is the most abundant group of Pelecypo- within this genus form groups or clusters with conchological
da in the deep-sea (400–5000 m), which constitutes the larg- similarity, as occurs in Yoldiella Verrill and Bush, 1897(La
est habitat on Earth, with at least 94% of the seabed lying Perna 2004, Benaim and Absalão 2011a). The species present-
below the permanent thermocline (Allen and Sanders 1996a). ed in this paper are no exception to this rule, and due to their
Recently, Benaim and Absalão (2011a, b, c) increased the similar shell outline, a morphometric analysis was required to
number of protobranchiate Pelecypoda known in Brazilian corroborate their delimitation into different species. There-
waters by approximately 27%. fore, this paper contributes to the knowledge of the genus
Ledella Verrill and Bush, 1897 is a genus that has a com- Ledella from Brazil by describing four previously unknown
plex taxonomic history, including changes of the type spe- species together with their morphometric data.
cies (Warén 1978, 1981, ICZN 1985, Allen and Hanna
1989). Information about this group in Brazilian waters is
currently lacking. The most recent Brazilian catalogue of MATERIAL AND METHODS
mollusks (Rios 2009) listed only Ledella solidula (Smith,
1885)and Ledella semen (Smith, 1885), with the latter given The samples used in the study were collected during
under the generic name of Nuculana Link, 1807. This does three projects. First, the “Live Resources of the Economic Ex-
not reflect the present knowledge of this genus in Brazil. clusive Zone” (Revizee) project, which took place during
Five species of Ledella are currently known to be present in 2001 (Table S1) and was sponsored by the Brazilian govern-
Brazilian waters: L.semen and L. solidula, as reported by ment as part of international demands to retain Brazilian au-
Smith (1885); Ledella ultima (Smith, 1885) and Ledella orixa thority over the 200 miles of territorial sea. This project extended
(Dall, 1927), as reported by Allen and Hanna (1989); across the Brazilian continental margin and the methodological
and Ledella acinula (Dall, 1889), as reported by Allen and details are given in Amaral and Rossi-Wongtschowski (2004)
Sanders (1996b). and Lavrado and Ignacio (2006). Second, the “Environmental
1
2 AMERICAN MALACOLOGICAL BULLETIN · ·
32 2 2014

Characterization of Campos Basin, RJ, Brazil” (Oceanprof)

project, which was conducted during the years 2002 and 2003
(Table S1). Finally, “Habitats Project – Campos Basin Envi-
ronmental Heterogeneity”, which was developed during 2008
and 2009 (Table S2). The two latter projects were sponsored
directly by Petrobras and were carried out using the research
vessel Astro–Garoupa equipped with a box corer. Details
about the associated methodological procedures can be found
in Lavrado and Brasil (2010). All biological material was sort-
ed under magnification and the best-preserved specimens of
each species were examined and photographed under a scan- Figure 1. Measurement scheme for the general shell shape. Total
ning electron microscope (ZEISS EVO 40) at the Gerência de length and height (L and H, respectively); dorsal and ventral height
Bioestratigrafia e Paleoecologia Aplicada (BPA) of the Petro- (DH and VH, respectively); length of the antero and postero-dorsal
margin (lam and lpm, respectively); and lengths of the four lines be-
bras Research Center (Centro de Pesquisas da Petrobras –
ginning near the shell center and radiating toward the four corners
CENPES). Live-collected specimens were very common in
of the rectangle delimiting the shell (R1, R2, R3, and R4).
the material obtained from the Habitats Project, which con-
stituted most of the examined material. All of the shells from
the Revizee and Oceanprof projects were empty. The speci-
mens studied have been deposited primarily in the mollusk stereomicroscope. The photographs were then analyzed to ob-
collection of the Instituto de Biologia, Universidade Federal tain the measurements, using the software Axio Vision release
do Rio de Janeiro (IBUFRJ), and secondarily at the Museu 4.6.3 (4.2007). To ensure the valves were parallel to the plane of
Nacional do Rio de Janeiro (MNRJ), the Museu Oceanográf- the camera lens, a platform made of playdough was used.
ico Eliézer de Carvalho Rios, Universidade Federal do Rio Measurements representing the general shell shape were
Grande (MOFURG), and the Muséum national d’Histoire taken as follows: total length (L); total height (H); dorsal
naturelle (MNHN), Paris. height (DH); ventral height (VH); length of the antero-dorsal
All of the Ledella samples studied here were first desig- margin (lam); length of the postero-dorsal margin (lpm); and
nated as morphospecies. We conducted an exploratory analy- lengths of the four lines beginning near the shell center and
sis based on the Population Aggregation Analysis (Davis and radiating toward the four corners of the rectangle delimiting
Nixon 1992) using1362 valves and taking into account the the shell (R1, R2, R3, and R4) (Fig.1). Similar approaches
“traditional” criteria used in bivalve taxonomy (Cox 1969, were used by Knudsen (1970), Rabarts and Whybrow (1979),
Bailey 2009, Killeen and Turner 2009), which are based on Warén (1989), Rhind and Allen (1992), Allen et al. (1995),
conchological characters. The identifications were made Bonfitto and Sabelli (1995), Fuiman et al. (1999), Benaim and
through comparisons with figures of the type species and de- Absalão (2011b), and Benaim et al. (2011).
scriptions available in the literature (Smith 1885, Laghi 1984, The measurements taken for the hinge plate were as fol-
Allen and Hanna 1989, Laghi and Palazzi 1989, La Perna lows: length of the anterior and posterior parts of the hinge
2004,Rios 2009,Allen and Sanders 1996b). plate (ahp and php, respectively); width of the anterior part of
Morphological orientation in pelecypods is usually re- the hinge plate and its teeth (wap and wat, respectively); and
lated to a series of axes (Bailey 2009). To determine dorsal width of the posterior part of the hinge plate and its teeth (wpp
and ventral height (DH and VH, respectively), we used the and wpt, respectively), as previously conducted by Benaim and
antero-posterior orientation based on an imaginary line pass- Absalão (2011b). Furthermore, ten additional measurements
ing under the adductor muscles scars, which divides the body were taken with the valves in dorsal view, replicating the meth-
into dorsal and ventral areas (Allen 1985). od of Benaim et al. (2011): width of the valve (W); total length
of the hinge plate (lhp); dorsal length of the anterior and pos-
Measurements terior parts of the hinge plate (dahp and dphp, respectively);
Considering the inequivalve condition of these species, height of the highest tooth of the anterior and posterior parts of
the valves were separated to obtain measurements of both left the hinge plate (hat and hpt, respectively); and lengths between
and right valves. Whenever possible, the valves measured the center of the highest tooth of the anterior and posterior
came from the same individual. This was not always feasible parts of the hinge plate to the umbo (alu and plu, respectively)
because the measurements of the hinge teeth require that and to the respective margins of the valve (alm and plm, re-
they are well-preserved. spectively) (Fig.2). These hinge plate measurements taken
The valves were photographed, together with a scale, using in the dorsal view have been used only once previously, by
a Canon Power Shot G10 camera attached to a Zeiss Stemi SV11 Benaim et al. (2011) on the genus Yoldiella.
 FOUR NEW SPECIES OF LEDELLA FROM BRAZIL 3

Figure 3. Measurement scheme for the shape of the rostrum. Height

of the rostrum (Hr); length of the rostrum (Lr); dorsal and ventral
Figure 2. Measurement scheme for the hinge plate and width of the heights of the rostrum (Hrd and Hrv, respectively); and profundity
valve. A, length of the anterior and posterior parts of the hinge plate of the rostrum’s sinuosity (Prs).
(ahp and php, respectively); width of anterior part of the hinge plate
and teeth (wap and wat, respectively); and width of the posterior
part of the hinge plate and teeth (wpp and wpt, respectively). B, analysis. To avoid this problem, in addition to redundancy
total length of the hinge plate (lhp); dorsal length of the anterior between the variables themselves, we performed a prelimi-
and posterior parts of the hinge plate (dahp and dphp, respectively);
nary correlation analysis among all variables and excluded
and lengths between the center of the highest tooth of the anterior
and posterior parts of the hinge plate to the umbo (alu and plu, re-
one of each pair of values with strong correlation (r ≥ 0.9).
spectively) and to the respective margins of the valve (alm and plm, The normality of variables was verified through normal prob-
respectively). C, width of the valve (W); and height of the highest ability plots and, when necessary, appropriate transforma-
tooth of the anterior and posterior parts of the hinge plate (hat and tions were implemented to normalize the data.
hpt, respectively). Discriminant function analysis was used to determine if
these morphospecies could be objectively identified through the
defined variables and, if so, which of the analyzed features were
An additional set of five measurements was implement- the most important for their discrimination. Forward stepwise
ed in this study: height of the rostrum (Hr), measured by a procedures were employed to select the most useful discriminat-
line that borders the first tooth of the posterior hinge plate ing variables. To perform these analyses, we standardized the
and goes straight from the dorsal to the ventral margins of the morphometric data as described by Romesburg (1984). All sta-
valve; length of the rostrum (Lr), measured perpendicularly tistical procedures were performed using STATISTICA software
from the Hr line to the edge of the rostrum; dorsal and ven- (version 7.0), StatSoft, Inc., Tulsa, Oklahoma.
tral heights of the rostrum (Hrd and Hrv, respectively), using The analyses were conducted for two separate subgroups:
the Lr line to divide the dorsal and ventral regions of the ros- the right valves and the left valves. Taking into account the
trum; and profundity of the rostrum’s sinuosity (Prs), which inequivalve condition of these species, a joint analysis would
is the greatest linear distance from the lower point of the sin- not have provided information about which group of valves
uosity to the imaginary line of continuity of the valve (Fig. 3). could determine the morphospecies more accurately nor
We also calculated 22 ratios: ahp/lam, alm/plm, alu/alm, about the features of each valve which are the most important
dahp/dphp, H/L, hat/dahp, hat/hpt, hat/W, hpt/dphp, hpt/W, for morphospecies discrimination.
Hr/Lr, Hr/H, Hrd/Hrv, lhp/L, Lr/L, Lr/plm, php/lpm, plu/
plm, W/L, W/lhp, wap/ahp and wpp/php. Taxonomic Discussion
Each morphospecies considered in this study was first
Morphometric analyses compared with the co-generic species previously reported in
One could argue that the above ratios will inevitably be Brazilian waters. They were then compared with the co-
strongly correlated with the variables that were utilized in the generic species that have similar shell shapes and, finally, with
calculation of these ratios, leading to redundancy in the the other morphospecies described in this work.
4 AMERICAN MALACOLOGICAL BULLETIN · ·
32 2 2014

Only Recent species were used to discuss the new species Table 1. Standardized coefficients for canonical variables for the
described herein. Thus, somewhat similar species such as Le- right valves analysis. The most relevant variables in each axis are
della peraffinis (Seguenza, 1877) (Miocene), whose neotype marked in dark gray; the second most representative variables in
was figured by Laghi and Palazzi (1989), Ledella pusio (Philippi, each axis are marked in light gray. The variables which made minor
contributions are marked in italics. Df, discriminant function.
1844) (Pliocene), which was reviewed and figured by Laghi
(1984), and Ledella seminulum (Seguenza, 1877) (Pleistocene-
Df1 Df2 Df3
Pliocene) and Ledella nicotrae (Seguenza, 1877) (Pliocene),
both figured by La Perna et al. (2004), were not considered in Prs 0.84685 0.57349 0.08940
the discussion. Besides the chronological distance between W 0.40454 0.14923 1.79748
the Recent and fossil species, all of these fossil species are ahp/lam -0.34177 0.51987 0.54561
from the Mediterranean Sea. lhp/L 0.75184 -0.27905 0.44486
Plu -0.48379 0.59059 0.82643
Alm 0.59928 -0.29880 2.74110
Hr/Lr -0.98651 -1.26410 -0.70248
RESULTS Ahp 0.20366 0.22406 -2.66803
H/L 0.36787 -0.48827 -0.05886
A traditional taxonomic descriptive approach suggested hat/dahp 0.62724 0.05129 0.82733
the existence of four morphospecies which provided the base Alu 0.26413 -0.56459 0.05034
for the morphometric procedures. Hat -1.01679 0.29528 -0.27112
Lpm 0.74758 -0.52086 -0.45604
Right Valves Hrd 0.34414 2.25031 0.70813
For this analysis, only nine variables were excluded due alm/plm -0.59847 0.09829 -2.33328
to strong correlation with the other variables. The excluded Hrd/Hrv -0.70831 -1.84689 -0.70744
Lr -1.73270 -1.34534 -2.74996
variables were: L, H, DH, lam, php, R2, R3, R4 and lhp. The
Hpt -1.16240 0.20600 -3.62005
remaining 44 variables were included in the analysis and 25 of
Wpt 0.44496 -0.28454 0.51637
them were retained in the final model, which was in 95% Wpp -0.36244 0.81991 -0.11739
agreement with the traditional criteria (Table 1). Wat -0.02904 0.70197 -0.09151
This rate of agreement was achieved using three discrim- wap/ahp -0.35423 0.75593 -4.37649
inant functions (df). The first one (df1) represents the role of Lr/plm 1.04844 -0.61189 2.15145
the shape of the Rostrum (Prs, Lr and Lr/plm) in the discrim- Wap 0.54872 -1.48020 4.63718
ination. Df2 represents the joint role of the Rostrum Height hpt/W 0.61434 -0.61345 2.37309
(Hrd e Hrd/Hrv) and Hinge Plate Width (wap and wpp) in Eigenval 3.70612 1.51830 1.05303
the discrimination. Df3 represents the role of the Hinge Plate Cum.Prop 0.59039 0.83225 1.00000
(hpt, wap/ahp, wap and hpt/W) in the discrimination. These
three roles were also perceived during the traditional and
more intuitive taxonomic discrimination of the individuals. tooth of the anterior part of the hinge plate to its respective
Three variables (ahp/lam, H/L and wpt) made a relative- margin (alm); df2 represents the role of the expansion of the
ly small contribution to the model, but we decided to retain antero-ventral region of the valve (R2) in the discrimina-
them because excluding them from the analysis generated a tion; df3 represents the role of the ratio between the size of
model with only 14 variables and 86.66% agreement with the the highest tooth of the posterior part of the hinge plate and
traditional criteria, which is 8.34% lower than the first result. the width of the valve (hpt/W) in the discrimination. In ad-
dition, in all three functions the ratio between the width of
Left Valves the valve and its length (W/L) was one of the two main vari-
For this analysis, only nine variables were excluded due ables: the ratio between the dorsal length of the hinge plate
to strong correlation with the others. These variables were: L, and the length of the valve (lhp/L) was one of the main vari-
H, DH, lam, php, R3, R4, lhp and hpt. The remaining 44 vari- ables in df1 and one of the two main variables in df2 and
ables were included in the analysis and 24 of these were re- df3, and the ratio between the width of the valve and the
tained in the final model, which showed 88.33% agreements dorsal length of the hinge plate (W/lhp) was one of the two
with the traditional criteria (Table 2), 6.67% less than the re- main variables in df2 and df3. The presence of these vari-
sult obtained for the right valves. ables in more than one discriminant function reinforces the
This rate of agreement was achieved using three dis- importance of the width of the valve, the length of the hinge
criminant functions. The first one (df1) represents the roles plate and the length of the valve to the discrimination of the
of the width of the valve (W) and of the distance of the highest morphospecies. The prominence of these variables in the
 FOUR NEW SPECIES OF LEDELLA FROM BRAZIL 5

Table 2. Standardized coefficients for canonical variables for the left

valves analysis. The most relevant variables in each axis are marked
in dark gray; the second most relevant variables in each axis are
marked in light gray. The variables that made minor contributions
are marked in italics. Df, discriminant function.

Df1 Df2 Df3

Hr/Lr 1.216731 0.33400 -0.42661
W -0.807000 -1.69353 0.66830
H/L -0.732078 0.54540 -1.49951
alm 1.332459 0.10327 -0.65290
php/lpm -0.274477 0.00366 0.09067
ahp/lam 0.060819 -0.33352 -0.18710
lhp/L -0.801864 -7.11341 -7.70510
Lpm -0.098203 1.17700 -0.46356
wpt -0.226754 0.42273 0.91259
ahp 0.661153 1.17329 1.74951
R1 -0.193882 -1.79420 0.91022
Prs -0.292743 -0.40283 0.10688
hat/hpt 0.286672 0.15125 -0.35170
W/L 1.921255 10.84032 13.32499
W/lhp -0.658799 -6.81776 -6.09872
alu -0.755089 1.14047 1.01010
R2 0.829580 1.d64855 -1.60351
VH -0.474442 -1.70245 2.36178
hpt/W -0.125905 0.60649 4.35272
hpt/dphp 0.102861 -0.25046 -3.73286
alu/alm 1.292262 -1.03185 -1.65989
dphp 0.125179 -0.81548 -4.69959
wpp -0.150277 -0.42347 -0.34195
wat 0.430124 0.29482 -0.07997
Eigenval 2.318400 1.96972 0.66221
Cum.Prop 0.468332 0.86623 1.00000

in which the shape of the rostrum was important to the dis-

crimination whilst the width and length of the valve were
only minor contributors.
Six variables (php/lpm, ahp/lam, Prs, hat/hpt, php and
dahp) made relatively small contributions to the model, but
we again decided to retain these, in accordance with the first
analysis. Removing these variables from the analysis generat-
ed a model with 18 variables and with 82.5% of the cases in
agreement with the traditional criteria, which is 5.83% lower
than the first result.

Hinge Plate
Figure 4. Canonical analysis scatterplots for the right valves, com- We also decided to test the discriminatory power using the
bining all three discriminant functions (df). hinge plate features alone, following Benaim et al. (2011). The
analyses were again performed using the right and left valves in
separate subgroups, and in both cases, nine variables were in-
three discriminant functions shows the importance of the cluded in the model. For the right and left valves respectively, the
hinge plate and the width and length of the valve in this models showed 66.66% and 64.16% agreement with the tradi-
analysis, in contrast to the results of the right valve analysis, tional criteria (Table 3).
6 AMERICAN MALACOLOGICAL BULLETIN · ·
32 2 2014

the thickness of the posterior part of the hinge plate; df3 rep-
resents the relative positions of the highest teeth from ante-
rior and posterior parts of the hinge plate. For the left valves,
df1 represents the relative positions of the highest teeth from
anterior and posterior parts of the hinge plate; df2 represents
the shape of the posterior part of the hinge plate; df3 repre-
sents the relative position of the highest tooth of the anterior
part of the hinge plate on the anterior part of the valve.
Only two variables were important in both hinge plate anal-
yses: alm/plm, which indicates the position of the highest teeth
of the anterior and posterior parts of the hinge plate in relation
to the margin, and wpp, which is the width of the posterior part
of the hinge plate. The first variable may be influenced by the
antero-dorsal rostrum present in some of the morphospecies;
however, this was not measured. The second variable had been
perceived as an important character previously, during the initial
discrimination of these morphospecies.
Considering the morphological differences previously
detected during the initial analysis of the material and the
high support provided by the morphometric approach for
the distinction of our initial morphospecies, we describe the
four species herein.

Systematics
Order NUCULANIDA Carter, Campbell and Campbell,
2000
Family NUCULANIDAE

Ledella spocki sp. nov.

(Figs. 4 and 5 as Ledella sp. 1 and Fig. 6 as Ledella spocki
sp. nov.)

Holotype
MNRJ 30480, Habitats 7 #F6 R2, 08-VII-2008. 22º19′11″S;
40º05′44″W; 403m.

Paratypes
IBUFRJ 19238, Habitats 7 #H6 R2, 07-VII-2008.
21º44′21″S; 40º05′18″W; 402 m.
MOFURG 51797, Habitats 8 #C6 R3, 31-I-2009.
22º59′00″S; 40º48′28″W; 376.6 m.
MNHN IM-2010-21015, Habitats 7 #B6 R2, 04-VII-
2008. 23º12′31″S; 40º58′31″W; 447.4m.
Material examined (See supplemental document at: http://
www.bioone.org/doi/suppl/10.4003/006.032.0201/suppl_
Figure 5. Canonical analysis scatterplots for the left valves, combin- file/Viegas_2014_Suppl_docs.pdf)
ing all three discriminant functions (df).
Etymology
These rates of success were achieved using three discrim- This species is named after the fictional character
inant functions (df) for each analysis. For the right valves “Spock” from the Star Trek TV and movie series. The overall
analysis, the first one (df1) represents the tridimensional shape of its valves resembles the shape of the pointed ear of
shape of the anterior part of the hinge plate; df2 represents the Vulcans, a humanoid extraterrestrial species depicted in
 FOUR NEW SPECIES OF LEDELLA FROM BRAZIL 7

Table 3. Standardized coefficients for canonical variables for the hinge plate analyses. The most relevant variables are marked in gray. Df,
discriminant function.

Right Valves Left Valves

df1 df2 df3 df1 df2 df3
Hat -0.717286 -0.292718 -0.74520 dphp -1.01472 0.921452 0.37551
alm/plm -0.080687 -0.894922 -1.46122 alm/plm -1.51834 0.427152 0.84545
wap/ahp 0.843338 0.364217 -0.98901 ahp -1.04440 0.109696 -0.40674
plu/plm -0.476898 0.052819 1.15000 alu/alm -0.61262 0.470459 0.95693
alu/alm 0.199852 -0.538310 -1.10171 plu 0.79939 -0.198373 -0.41296
wpp -0.020109 1.016999 -0.13674 wpp 1.58973 0.734929 0.80325
wpp/php -0.474712 -0.401955 -0.05389 wpt -0.36763 -0.690460 -1.17862
wpt 0.034626 -0.873887 0.55696 wpp/php -0.85653 -0.017427 0.02813
wat -0.443634 0.447644 -0.00058 hat/dahp -0.36750 0.072022 0.24557
Eigenval 0.848731 0.477082 0.11853 Eigenval 0.98125 0.626145 0.09994
Cum.Prop 0.587624 0.917935 1.00000 Cum.Prop 0.57473 0.941466 1.00000

this series. Because “Mr. Spock” is the most famous Vulcan in can be distinguished on the basis of the less opisthogyrous umbo-
the series, the species has been named after him. nes and the more profound rostral sinuosity of L. spocki sp. nov.
Ledella spocki sp. nov. shares a long and acute rostrum
Distribution and a profound rostral sinuosity with L. solidula, but L. spocki
Only known from the Campos Basin, off the coast of Rio sp. nov. can be distinguished from the latter on the basis of
de Janeiro, Brazil. having no concentric lines and a rostrum that is shorter and
less downward-pointing than that of L. solidula.
Diagnosis Ledella spocki sp. nov. differs from L. ultima in that the
Thin shell with marked anterior and posterior shoulders. former has no concentric lines and presents thinner teeth on
Acute rostrum. Profound rostral sinuosity. the hinge plate and a more acute rostrum than L. ultima.
Ledella spocki sp. nov. differs from L. acinula in that the for-
Description mer has no concentric lines and presents a more acute rostrum
Shell oblong–elliptical, inflated, thin, inequivalve, inequilat- and a less inflated posterior region of the shell than L. acinula.
eral, H/L about 0.72; umbones prominent, opisthogyrous and Ledella spocki sp. nov. differs from L. orixa in that the
inwardly directed. Antero-dorsal margin convex; anterior mar- former presents less opisthogyrous umbones, a larger poste-
gin showing a shoulder. Rounded ventral margin forming a sin- rior region in comparison to the anterior region and a smaller
uosity leading to an acute rostrate posterior end which points anterior part of the hinge plate when compared to the poste-
downward; postero-dorsal margin almost straight, with a shoul- rior part.
der near the end of the hinge plate. Smooth outer surface. Hinge Ledella spocki sp. nov. can be distinguished from L. acuminata
plate interrupted by a small and almost rounded resilifer. Poste- (Jeffreys, 1870) in that the former has no concentric lines, a
rior part of the hinge plate longer and thinner than anterior one shorter hinge plate and presents thinner hinge teeth in relation
in adult specimens, and almost equal in juveniles. Width of the to the hinge plate thickness, compared with L. acuminata.
anterior and posterior rows of teeth occupying about 60% and
58%, respectively, of the total width of the hinge plate of the right Ledella legionaria sp. nov.
valves and about 59% and 55% of the anterior and posterior (Figs. 4 and 5 as Ledella sp. 2 and Fig. 7 as Ledella legion-
hinge plates, respectively, of the left valve. Shell microsculpture: aria sp. nov.)
the prodissoconch presents a weak granulation and the hinge
teeth present pustules on the side that faces the umbones. Maxi- Holotype
mum shell length: 1.94 mm. Prodissoconch height: 70μm. MNRJ 30481, Habitats 7 #F6 R2, 08-VII-2008. 22º19′11″S;
Prodissoconch length: 73μm. (Table 4) 40º05′44″W; 403m.

Remarks Paratypes
Ledella spocki sp. nov. is similar to L.semen with respect to the IBUFRJ19300, Habitats 9 #H6 R2, 05-II-2009. 21º44′21″S;
general shell shape and in the acuteness of its rostrum, but they 40º04′53″W; 404 m.
8 AMERICAN MALACOLOGICAL BULLETIN · ·
32 2 2014

segmentata (segmented plates) armor. Addition-

ally, its shell is thicker than those of the other
species described herein, as though the animal
wears a heavier armor than the others.

Distribution
Only known from the Campos Basin, Rio
de Janeiro, Brazil.

Diagnosis
Shell thick with prominent anterior and
posterior shoulders. Rostrum acutely rounded.
Profound rostral sinuosity, more profound in
the left valve than in the right one.

Description
Shell oblong–elliptical, inflated, thick, in-
equivalve, inequilateral, H/L about 0.74; umbones
prominent, opisthogyrous and inwardly directed.
Antero-dorsal margin convex, acute shoulder de-
limiting antero-dorsal and anterior margins,
merging to a convex ventral margin. Ventral mar-
gin forming a sinuosity leading to a rostrate poste-
rior end which points slightly downward;
postero-dorsal margin almost straight, with a
marked shoulder near the end of the hinge plate.
Hinge plate interrupted by a deep, rounded resil-
ifer. Posterior part of the hinge plate longer and
thinner than anterior one in adult specimens, and
almost equal in juveniles. Width of the anterior
and posterior rows of teeth occupying about 61%
and 56%, respectively, of the total width of the
Figure 6. Ledella spocki sp. nov. (Holotype MNRJ 30480). Internal view: A, right valve; hinge plate on the right valves and about 61% and
B, left valve. External view: D, right valve; E, left valve. Dorsal view: F, right valve; G, 57% to the anterior and posterior parts of the
left valve. Details: C, resilifer of the right valve. Prodissoconch (IBUFRJ 19239): H, hinge plates, respectively, on the left valve. Shell
overview; I, detail of the tip of the prodissoconch showing some granulation. Hinge microsculpture: the prodissoconch presents a
teeth (IBUFRJ 19247): J, overview; K, detail of the tooth showing the tooth pustules. weak granulation and the hinge teeth present pus-
Scales A, B, D–G: 1mm. C, H: 0.1mm.I: 0.01 mm. J: 0.2 mm. K: 0.02 mm. tules on the side that faces the umbones. Maxi-
mum shell length: 2.31 mm. Prodissoconch
height: 77 μm, length: 86 μm. (Table 4)
MOFURG 51798, Habitats 7 #F6 R1, 08-VII-2008.
22º19′10″S; 40º05′42″W; 402.3m. Remarks
MNHN IM-2010-21016, Revizee V #41, 20-VII-2001. Ledella legionaria sp. nov. differs from L.semen in having
21º13′S; 40º13′W; 1000m. less opisthogyrous umbones, a more profound rostral sinuos-
Material examined (See supplemental document at: http:// ity and a less acute rostrum than L. semen.
www.bioone.org/doi/suppl/10.4003/006.032.0201/suppl_ Ledella legionaria sp. nov. differs from L. solidula in hav-
file/Viegas_2014_Suppl_docs.pdf) ing no concentric lines and in presenting a prominent shoul-
der in the antero-dorsal region and a rostrum that is shorter
Etymology and less acute than that of L. solidula.
This species is named after the Roman legionnaires. The Ledella legionaria sp. nov. differs from L. ultima in hav-
shape of its rostrum resembles the outline of a shoulder ar- ing no concentric lines and in presenting thinner teeth on the
mor, similar to those used by the legionaries in their lorica hinge plate and a more acute rostrum than L. ultima.
 FOUR NEW SPECIES OF LEDELLA FROM BRAZIL 9

Ledella legionaria sp. nov. differs from L. acinula in hav- Material examined (See supplemental document at: http://
ing no concentric lines and in presenting a more acute ros- www.bioone.org/doi/suppl/10.4003/006.032.0201/suppl_
trum and a less inflated posterior region of the shell than L. file/Viegas_2014_Suppl_docs.pdf)
acinula.
Ledella legionaria sp. nov. differs from L. orixa in pre- Etymology
senting less opisthogyrous umbones, a larger posterior region This species is similar in shape to the ear of an elf from
than the anterior region and a smaller anterior part of the Norse mythology, so we decided to name this species after
hinge plate in comparison to the posterior part. those elves.
Compared to Ledella sublevis Verrill and Bush, 1898, L.
legionaria sp. nov. has a similar shell shape, but a less smooth Distribution
postero-dorsal margin and a longer posterior part of the Only known from the Campos Basin, Rio de Janeiro,
hinge plate when compared to the anterior part, with thinner Brazil.
hinge teeth than L. sublevis.
Compared to Ledella jamesi Allen and Hanna, 1989, L. Diagnosis
legionaria sp. nov. has a similar shell shape, with a shoulder Thin shell with anterior and posterior shoulders usually
on the postero-dorsal margin, but L. legionaria sp. nov. has a absent; when present, shoulders not prominent. Acute ros-
longer rostrum and no concentric lines. trum. Profound rostral sinuosity.
Ledella lusitanensis Allen and Hanna, 1989, also has a
similar shell shape when compared to L. legionaria sp. nov., Description
but the latter has a shoulder in the postero-dorsal margin and Shell oblong–elliptical, inflated, thin, inequivalve, inequi-
a more inequilateral hinge plate, with the posterior part of the lateral, H/L about 0.72; umbones prominent, opisthogyrous
hinge plate longer and with thinner teeth than the anterior and inwardly directed. Antero-dorsal margin convex; anterior
one. margin rounded, smoothly merging to the slightly convex ven-
Compared to Ledella spocki sp. nov., L. legionaria sp. tral margin. Ventral margin forming a sinuosity leading to a
nov. is larger, thicker, more inflated and has a less profound rostrate posterior end which points downward; postero-dorsal
rostrum sinuosity on the right valve. Ledella legionaria sp. margin almost straight, sometimes weakly sloping downward
nov. also has a shorter rostrum and a more prominent pos- near the end of the hinge plate. Smooth surface. Hinge plate
terior shoulder. Ledella legionaria sp. nov. has longer poste- interrupted by a deep, rectangular resilifer. Posterior hinge
rior parts of the hinge plate on both valves, but longer plate longer and thinner than anterior one in adult specimens,
anterior parts of the hinge plate are present only on the right and almost equal in juveniles. Width of the anterior and poste-
valve. These species do not vary much from each other with rior rows of teeth occupying about 60% and 59%, respectively,
respect to the thickness of the hinge plate or that of the of the total width of the hinge plate of the right valves and
hinge teeth. Regarding the prodissoconch, they do not differ about 62% and 59% to the anterior and posterior hinge plates,
much in height and microsculpture, but the prodissoconch respectively, of the left valve. Shell microsculpture: the prodis-
of L. legionaria sp. nov. is longer in length than the prodis- soconch presents a weak granulation and the hinge teeth pres-
soconch of L.spocki sp. nov. ent pustules on the side that faces the umbones. Maximum
shell length: 1.86 mm. Prodissoconch height: 83 μm. Prodis-
Ledella elfica sp. nov. soconch length: 104 μm.(Table 4)
(Figs. 4 and 5 as Ledella sp. 3 and Fig. 8 as Ledella elfica
sp. nov.) Remarks
Compared to Ledella semen, L. elfica sp. nov. has a similar
Holotype shell shape, with a smooth outline and an acute rostrum, but
MNRJ 30482, Habitats 7 #H6 R2, 07-VII-2008.21º44′21″S; L. elfica sp. nov. has less opisthogyrous umbones and a more
40º05′18″W; 402m. profound rostral sinuosity.
Ledella elfica sp. nov. is similar to L. solidula in that both
Paratypes have an acute rostrum and a profound rostral sinuosity, but
IBUFRJ 19371, Habitats 7 #H6 R1, 06-VII-2008. they can be distinguished because L. elfica sp. nov. does not
21º42′02″S; 40º06′15′W; 404.7 m. have concentric lines and its rostrum is shorter and points
MOFURG 51799, Revizee Central V #52, 21-VII-2001. less downwards than that of L. solidula.
21º46′S; 40º05″W; 450 m. Ledella elfica sp. nov. differs from L. ultima in having no
MNHN IM-2010-21017, Habitats 7 #F6 R2, 08-VII- concentric lines and in presenting thinner teeth on the hinge
2008. 22º19′11″S; 40º05′44″W; 403m. plate and a more acute rostrum than L. ultima.
10 AMERICAN MALACOLOGICAL BULLETIN · ·
32 2 2014

no concentric lines, a shorter hinge plate and the

hinge teeth occupy less of the hinge plate’s width
in comparison to L. acuminata.
Ledella elfica sp. nov. is similar to L. spocki sp.
nov. in shape and thickness of the shell and in the
length of the rostrum, but they can be distinguished
because L. elfica sp. nov. is smaller in size (H, L and
W), the anterior part of its hinge plate is thinner
and longer on both valves and it has a less high and
less acute rostrum with a less profound rostral sin-
uosity than L. spocki sp. nov. Ledella elfica sp. nov.
also has a hinge plate (measured in dorsal view)
that is longer in the right valve and shorter on the
left one, compared to L. spocki sp. nov. Besides that,
the hinge teeth of L. elfica sp. nov. are longer, in
both valves, than those of L. spocki sp. nov. Regard-
ing the prodissoconch, they do not differ much in
height and microsculpture, but the prodissoconch
of L. elfica sp. nov. is larger in length than the
prodissoconch of L.spocki sp. nov.
Ledella elfica sp. nov. is similar to L. legionaria
sp. nov. with respect to the general shell shape,
but they can be distinguished because L. elfica sp.
nov. has a longer and more acute rostrum than
L. legionaria sp. nov. and its rostral sinuosity is
more profound on the right valve and less pro-
found on the left valve compared to L. legionaria
sp. nov. With regard to the hinge plates, the
hinge teeth of L. elfica sp. nov. occupy a larger
part of the hinge plate thickness on the posterior
part of the hinge plate than that of L. legionaria
sp. nov. Regarding the prodissoconch, they do
Figure 7. Ledella legionaria sp. nov. (Holotype MNRJ 30481). Internal view: A, right
not differ much in height and microsculpture,
valve; B, left valve. External view: D, right valve; E, left valve. Dorsal view: F, right
but the prodissoconch of L. elfica sp. nov. is larg-
valve; G, left valve. Details: C, resilifer of the right valve. Prodissoconch (IBUFRJ
19284): H, overview; I, detail of the tip of the prodissoconch showing some granula- er in length than the prodissoconch of L. legionaria
tion. Hinge teeth (IBUFRJ 14637): J, overview; K, detail of the tooth showing the sp. nov.
tooth pustules. Scales A, B, D–G: 1 mm. C, H, J: 0.1 mm. I: 0.02 mm. K: 0.05 mm.
Ledella manati sp. nov.
(Figs. 4 and 5 as Ledella sp. 4 and Fig. 9 as
Ledella manati sp. nov.)
Ledella elfica sp. nov. differs from L. acinula in having no
concentric lines and in presenting a more acute rostrum and Holotype
a less inflated posterior region of the shell than L. acinula. MNRJ 30483, Revizee V #504, 02-VII-2001. 14º28′S;
Ledella elfica sp. nov. differs from L. orixa in presenting 38º54′W; 278m.
less opisthogyrous umbones, a larger posterior region than
the anterior region and a smaller anterior part of the hinge Paratypes
plate when compared to the posterior part. IBUFRJ 19398, Habitats 7 #C6 R1, 04-VII-2008.
Ledella elfica sp. nov. is similar to L. acuminata in the 22º59′00″S; 40º48′25″W; 387.4 m.
general outline of the shell, with an acute rostrum, a pro- MOFURG 51800, Revizee Central V #1, 03-VII-2001.
found rostral sinuosity and the presence of a smooth shoul- 13º04′S; 38º21′W; 450 m.
der on the postero-dorsal margin, which is found in some MNHN IM-2010-21018, Habitats 7 #F6 R2, 08-VII-
specimens of L. elfica sp. nov. However, L. elfica sp. nov. has 2008. 22º19′11″S; 40º05′44″W; 403m.
 FOUR NEW SPECIES OF LEDELLA FROM BRAZIL 11

Diagnosis
Thick shell usually without anterior and
rostral shoulders, but, when present, not promi-
nent. Rostrum acutely rounded. Shallow rostral
sinuosity.

Description
Shell oblong–elliptical, inflated, thick, in-
equivalve, inequilateral, H/L about 0.72; umbo-
nes prominent, opisthogyrous and inwardly
directed. Antero-dorsal margin convex; anterior
margin with a shoulder, rounded ventral margin.
Ventral margin forming a shallow sinuosity lead-
ing to a rostrate posterior end; postero-dorsal
margins lightly convex, sometimes with a weak
shoulder near the end of the hinge plate. Smooth
surface. Hinge plate interrupted by a deep,
rounded resilifer. Posterior hinge plate longer
and thinner than anterior one in adult specimens,
and almost equal in juveniles. Width of the ante-
rior and posterior rows of teeth occupying about
59% and 58%, respectively, of the total width of
the hinge plate of the right valves and 61% and
57% to the anterior and posterior hinge plates,
respectively, of the left valve. Shell microsculp-
ture: the prodissoconch presents a weak granula-
tion and the hinge teeth present pustules on the
side that faces the umbones. Maximum shell
length 2.02 mm. Prodissoconch height: 90 μm.
Prodissoconch length: 118 μm. (Table 4)

Figure 8. Ledella elfica sp. nov. Internal view (Holotype MNRJ 30482): A, right valve; Remarks
B, left valve. External view (IBUFRJ 19364): D, right valve; E, left valve. Dorsal view Ledella manati sp. nov. differs from L. semen
(IBUFRJ 19363): F, right valve; G, left valve. Details: C, resilifer of the right valve. in having less opisthogyrous umbones and a less
Prodissoconch (IBUFRJ 19324): H, overview; I, detail of the tip of the prodisso- acute rostrum than L. semen.
conch showing some granulation. Hinge teeth (IBUFRJ 19385): J, overview; K, de-
Ledella manati sp. nov. differs from L. solidula
tail of the tooth showing the tooth pustules. Scales A, B, D–G: 1 mm. C, H: 0.1 mm.
I: 0.01 mm. J: 0.2 mm. K: 0.05 mm.
in having no concentric lines and a rostrum that is
shorter and less acute than that of L. solidula.
Ledella manati sp. nov. differs from L. ultima
in having no concentric lines and in presenting thinner teeth
Material examined (See supplemental document at: http:// on the hinge plate and a more acute rostrum than L. ultima.
www.bioone.org/doi/suppl/10.4003/006.032.0201/suppl_ Ledella manati sp. nov. differs from L. acinula in having
file/Viegas_2014_Suppl_docs.pdf) no concentric lines and in presenting a more acute rostrum
and a less inflated posterior region of the shell than L. acinula.
Etymology Ledella manati sp. nov. differs from L. orixa in presenting
Manati is a Caribbean word for the sea cow. Because sea less opisthogyrous umbones, a larger posterior region than
cows are large, but possess a smooth contour that is similar to the the anterior region and a smaller anterior part of the hinge
shells of this species, we decided to name it Ledella manati sp. nov. plate when compared to the posterior part.
Ledella manati sp. nov. is similar to L. sublevis in having
Distribution a shallow rostral sinuosity, a smooth surface and a thick shell,
Known from the Campos Basin, Rio de Janeiro, Brazil but L. manati sp. nov. can be distinguished by its less acute
and for Trindade Island, Espírito Santo, Brazil. rostrum and thinner hinge teeth.
12 AMERICAN MALACOLOGICAL BULLETIN · ·
32 2 2014

hinge plate with no thickness difference to that

of L. spocki sp. nov. The prodissoconch of L.
manati sp. nov. is larger than that of L. spocki sp.
nov. in both height and length, but there is no
difference regarding its microsculpture.
Ledella manati sp. nov. differs from L.
legionaria sp. nov. in having a longer and higher
rostrum, with a rostral sinuosity that is about
half the profundity as that of L. legionaria sp.
nov. on the left valve, but about 66% less pro-
found on the right valve. Ledella manati sp. nov.
also has a longer and thinner anterior part of the
hinge plate, but a shorter and thicker posterior
part when compared to L. legionaria sp. nov.
The hinge teeth occupy almost the same propor-
tion of the hinge plate thickness in both species.
Regarding the prodissoconch, they do not differ
much in height and microsculpture, but the
prodissoconch of L. manati sp. nov. is larger in
length than the prodissoconch of L. legionaria
sp. nov.
Ledella manati sp. nov. differs from L. elfica
sp. nov. inhaving a shorter and less acute ros-
trum, with a rostrum sinuosity with about half
the profundity as that of L. elfica sp. nov. Addi-
tionally, L. manati sp. nov. has a shorter and
thicker posterior part of the hinge plate, but an
anterior part with no difference in length or
thickness compared to L. elfica sp. nov. Regard-
ing the prodissoconch, they do not differ much
in height and microsculpture, but the prodisso-
conch of L. manati sp. nov. is larger in length
Figure 9. Ledella manati sp. nov. Internal view (Holotype MNRJ 30483): A, right than the prodissoconch of L. elfica sp. nov.
valve; B, left valve. External view (IBUFRJ 19435): D, right valve; E, left valve. Dorsal
view (IBUFRJ 14673): F, right valve; G, left valve. Details: C, resilifer of the right
valve. Prodissoconch (IBUFRJ 14637): H, overview; I, detail of the tip of the prodis- DISCUSSION
soconch showing some granulation. Hinge teeth: J, overview; K, detail of the tooth
showing the tooth pustules. Scales A, B, D–G: 1 mm. C, H: 0.1 mm. I: 0.02 mm. J:
One of the major problems associated with
0.2 mm. K: 0.01 mm.
the traditional taxonomical approach is its sub-
jectivity, which generates a relationship of de-
pendency between the expertise of the
Ledella manati sp. nov. is similar to Ledella verdiensis Allen taxonomist and the reliability of the taxonomical decisions.
and Hanna, 1989 in having a similar shell shape with a shal- In general, the taxonomist notes, somewhat intuitively, many
low rostral sinuosity, but it can be distinguished on the basis differences between the taxa he is working on. The main dif-
of having no strong concentric lines, a less high shell, a thick- ficulty lies in making these differences clear to the readers.
er hinge plate and a smoother postero-dorsal margin than The morphometric approach itself does not solve this prob-
L. verdiensis. lem, because the taxonomist cannot handle the distinct mea-
Ledella manati sp. nov. differs from L. spocki sp. nov. in surements taken from all over the shell to determine which
having a shorter and less acute rostrum, with a rostral sinuos- ones are relevant for discriminating among the taxa. A tech-
ity that is about half the profundity as that of L. spocki sp. nique to reduce this myriad of potentially important variables
nov., and a longer and thinner anterior part of the hinge plate to a manageable number of morphologically understandable
in relation to the shell size, but a shorter posterior part of the factors is fundamental for obtaining objectivity in taxonomic
 FOUR NEW SPECIES OF LEDELLA FROM BRAZIL 13

Table 4. Average values of the larval shell measurements and length of the adult shell. H(p) – Height of the prodissoconch I; SD, standard
deviation; L(p), Length of the prodissoconch I; n, number of measured valves; L (s), Length of the adult shell; Min, Minimum value; Max,
Maximum value.

H(p) SD L(p) SD n L (s) Min Max n

Ledella spocki sp. nov. 0,071 0,005 0,073 0,004 20 1,674 1,336 1,937 60
Ledella legionaria sp. nov. 0,078 0,006 0,086 0,004 20 1,891 1,367 2,318 60
Ledella elfica sp. nov. 0,083 0,006 0,104 0,004 20 1,632 1,291 1,869 60
Ledella manati sp. nov. 0,091 0,004 0,118 0,003 20 1,708 1,401 2,027 60

work. The discriminant analysis is not the only statistical variables. In other words, the variables that are associated
technique that is able to do this, but the method has a clear with the shell (valve) shape are essential for producing a sat-
and simple mathematical basis (Klecka 1982). In our case, it isfactorily high rate of agreement between traditional taxon-
reduced the original 44 variables to a few comprehensible omy and this morphometric-based approach.
factors. The larval shell of these species present some microscu-
Considering the right valves, the variables Prs, Lr and lpture. The prodissoconch I presents a weak granulation
Hrd/Hrv, which are all related to the rostrum, were found to (Figs. 6H, 6I, 7H, 7I, 8H, 8I,9H and 9I) that cannot be per-
be critical characters in discriminating among these species. ceived without the use of an electron microscope. The nepio-
Moreover, a complementary set of variables (wap, wap/ahp, conch cannot be clearly defined for these species, because
wpp, hpt, and hpt/W), which are all related to the hinge plate, either it is undifferentiated from the adult shell or it does not
was shown to be critical for discrimination. An additional set present a clear boundary with it.
of morphological variables providing minor contributions Regarding the prodissoconch, it is also important to notice
completed the analysis. The rate of agreement between tradi- that the prodissoconch I of these species are smaller than ex-
tional taxonomy and this morphometric-based approach was pected for protobranchs (Gustafson and Reid 1986) and, since
95%. the prodissoconch II is not visible, it probably means that the
Analysis of the left valves produced similar results. Due eggs are brooded or lecithotrophic (Zardus and Martel 2002).
to differences between right and left valves within the same Since no brooding was observed in the samples here analyzed,
specimen (inequivalve condition), a somewhat different set this suggests these species simply have rather small eggs.
of variables was selected to maximize the distinction among The SEM images also revealed the presence of microscu-
the four taxa. The most relevant variables were: W, lhp/L, lpture on the hinge teeth of all four species described herein
W/L, alm and R2. This set of variables carries information (Figs.6J, 6K, 7J, 7K, 8J, 8K, 9J and 9K). Malchus and Sartori
about the general shape of the valve, in contrast to the right (2012) describe similar teeth pustules in Nuculanidae, but the
valves analysis, which indicated different characters as being origin and function of these pustules are unknown (Malchus,
important. In the left valves analysis, the rate of agreement pers. comm.). They may generate more attrition between the
between traditional taxonomy and this morphometric-based teeth, so that they are harder to slide, turning the shell harder
approach was 88.33%. to open by predators, but a study about its function must be
The second set of analyses, which was based on hinge done prior to defining its actual function. This information is
plate characters alone, revealed the importance of these fea- not mentioned for the other Ledella species to which we com-
tures for distinguishing the four species: the rates of agree- pare our species, but we do not know if it is present or not,
ment between traditional taxonomy and this since the detection of this character is very specific and was
morphometric-based approach were 66.66% for the right not done for the other species yet.
valves and 64.16% for the left valves. This finding corrobo-
rates the importance of including the hinge plate structure in
morphometric analysis, as stated by Benaim et al. (2011) in CONCLUSIONS
their study of Yoldiella species. This outcome is not surprising
because hinge characters were important in the complete The description of these four species improves the un-
morphometric analyses performed earlier in this study. How- derstanding of the biodiversity of protobranchs of the Cam-
ever, despite being highly important, the hinge plate charac- pos Basis, contributes with some morphological aspects of
ters alone have a lower rate of agreement between the Ledella species and corroborates the importance of including
traditional taxonomical approach and the morphometric ap- specific hinge plate details in the characterization of proto-
proach than the analyses using the full set of morphometric branch species.
14 AMERICAN MALACOLOGICAL BULLETIN · ·
32 2 2014

ACKNOWLEDGMENTS Bonfitto, A. and B. Sabelli. 1995. Yoldiella seguenzae, a new species

of Nuculanidae (Bivalvia; Nuculoida) from the Mediterranean
We are deeply grateful to Dr. Paolo G. Albano for the Sea. Journal of Molluscan Studies 61: 21–27.
exchange of literature, to Dr. Nikolaus Malchus for the valu- Caetano, C. H. S., V. Scarabino, and R. S. Absalão. 2006. Scaphopoda
able contribution to improve this article, to Petrobras SA for (Mollusca) from the Brazilian continental shelf and upper
slope (13º to 21ºS) with description of two new species of the
the material used in this study and for the SEM images and to
genus Cadulus Philippi, 1844. Zootaxa (online)1267: 1–47.
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77–87.

Submitted: 12 June 2013; accepted: 26 December 2013;

final revisions received: 19 April 2014
 NUMBER 1 OF 1

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Amer. Malac. Bull. 32(2), Supplemental materials, 1–4 (2014)
http://www.bioone.org/doi/suppl/10.4003/006.032.0201/suppl_file/Viegas_2014_Suppl_docs.pdf

Description of four new species of Ledella Verrill and Bush, 1897 (Pelecypoda:
Nuculanidae) off the coast of Brazil, using a morphometric approach

Diniz Corrêa Paone Viegas, Natalia Pereira Benaim and Ricardo Silva Absalão

Universidade Federal do Rio de Janeiro (UFRJ), Instituto de Biologia, Departamento de Zoologia, Laboratório de Malacologia, Ilha do
Fundão, Rio de Janeiro, Brazil. CEP: 21941-901.
Correspondence, Diniz Corrêa Paone Viegas: diniz.viegas@gmail.com

Supplemental Materials 2009), 1 individual; IBUFRJ 19267 (Habitats 8 #C6 R2, 2009),
3 individuals; IBUFRJ 19268 (Habitats 9 #I6 R1, 2009) 2 indi-
Material examined (See Tables S1 and S2 for coordinates) viduals; IBUFRJ 19269 (Habitats 8 #D8 R3, 2009), 1 individ-
ual; IBUFRJ 19270 (Habitats 7 #H6 R3, 2008), 1 individual;
Ledella spocki sp. nov. IBUFRJ 19271 (Habitats 6 #I9 R1, 2008), 1 individual; IBU-
IBUFRJ 13696 (Revizee V #1 profound, 2001), 2 valves; FRJ 19272 (Habitats 7 #I6 R3, 2008), 1 individual; IBUFRJ
IBUFRJ 16835 (Oceanprof II #79 South, 2003), 6 valves; IBU- 19273 (Habitats 8 #B6 R2, 2009), 1 individual; IBUFRJ 19274
FRJ 18258 (Oceanprof I #44 North, 2002), 2 valves; IBUFRJ (Habitats 8 #A6 R3, 2009), 2 individuals; IBUFRJ 19275
19232 (Habitats 7 #F6 R2, 2008), 1 individual; IBUFRJ 19233 (Habitats 8 #C6 R1, 2009), 6 individuals; IBUFRJ 19276
(Habitats 7 #H6 R2, 2008), 2 individuals; IBUFRJ 19234 (Habitats 7 #A6 R3, 2008), 2 individuals; IBUFRJ 19277
(Habitats 8 #A6 R2, 2009), 3 individuals; IBUFRJ 19235 (Habitats 7 #D6 R3, 2008), 1 individual; IBUFRJ 19278 (Hab-
(Habitats 8 #D6 R3, 2009), 1 individual; IBUFRJ 19236 (Hab- itats 8 #A6 R1, 2009), 1 individual; IBUFRJ 19279 (Habitats 7
itats 7 #D6 R1), 1 individual; IBUFRJ 19237 (Habitats 7 #B6 #C6 R3, 2008), 1 individual.
R2, 2008), 2 individuals; IBUFRJ 19238 (Habitats 7 #H6 R2,
2008), 1 individual; IBUFRJ 19239 (Habitats 9 #H6 R1, 2009), Ledella legionaria sp. nov.
2 individuals; IBUFRJ 19240 (Habitats 7 #B6 R2, 2008), 1 in- IBUFRJ 13742 (Revizee V #41, 2001), 6 valves; IBUFRJ
dividual; IBUFRJ 19241 (Habitats 8 #F6 R1, 2009), 3 indi- 14527 (Revizee V #25A, 2001), 1 valve; IBUFRJ 17605
viduals; IBUFRJ 19242 (Habitats 8 #C6 R3, 2009), 5 (Oceanprof II #44 North, 2003), 2 valves; IBUFRJ 19280
individuals; IBUFRJ 19243 (Habitats 7 #B6 R2, 2008) 2 indi- (Habitats 7 #F6 R2, 2008), 10 individuals and 1 valve; IBUFRJ
viduals; IBUFRJ 19244 (Habitats 8 #C6 R2, 2009) 2 individu- 19281 (Habitats 7 #H6 R1, 2008), 2 individuals; IBUFRJ
als; IBUFRJ 19245 (Habitats 8 #B6 R3, 2009), 1 individual; 19282 (Habitats 7 #B6 R2, 2008), 10 individuals; IBUFRJ
IBUFRJ 19246 (Habitats 6 #F6 R3, 2008), 5 individuals; IBU- 19283 (Habitats 8 #C6 R3, 2009), 2 individuals; IBUFRJ
FRJ 19247 Habitats 7 C6 R1 8 individuals; IBUFRJ 19248 19284 (Revizee V #41, 2001), 16 valves; IBUFRJ 19285 (Habi-
(Habitats 7 #B6 R1, 2008), 9 individuals; IBUFRJ 19249 tats 7 #D6 R1, 2008), 5 individuals; IBUFRJ 19286 (Habitats
(Habitats 8 #B6 R3, 2009), 10 individuals; IBUFRJ 19250 7 #B6 R2, 2008), 7 individuals; IBUFRJ 19287 (Habitats 7 #B6
(Habitats 8 #A6 R1, 2009), 6 individuals; IBUFRJ 19251 R3, 2008), 10 individuals; IBUFRJ 19288 (Habitats 7 #A6 R1,
(Habitats 8 #D6 R1, 2009), 6 individuals; IBUFRJ 19252 2008), 2 individuals; IBUFRJ 19289 (Habitats 7 #F6 R1,
(Habitats 8 #B6 R2, 2009), 1 individual; IBUFRJ 19253 (Hab- 2008), 4 individuals; IBUFRJ 19290 (Habitats 7 #H6 R2,
itats 7 #H6 R2, 2008), 1 individual; IBUFRJ 19254 (Habitats 7 2008), 2 individuals; IBUFRJ 19291 (Habitats 7 #B6 R1,
#B6 R3, 2008), 1 individual; IBUFRJ 19255 (Habitats 8 #B6 2008), 15 individuals; IBUFRJ 19292 (Habitats 7 #A6 R2,
R2, 2009), 1 individual; IBUFRJ 19256 (Habitats 7 #C6 R3, 2008), 3 individuals; IBUFRJ 19293 (Habitats 7 #B6 R1,
2008), 1 individual; IBUFRJ 19257 (Habitats 8 #B6 R2, 2009), 2008), 2 individuals; IBUFRJ 19294 (Habitats 7 #H6 R2,
1 individual; IBUFRJ 19258 (Habitats 8 #B6 R2, 2009), 1 in- 2008), 6 individuals; IBUFRJ 19295 (Revizee V #52, 2001), 2
dividual; IBUFRJ 19259 (Habitats 8 #C6 R3, 2009), 1 indi- valves; IBUFRJ 19296 (Habitats 8 #C6 R1, 2009), 3 individu-
vidual; IBUFRJ 19260 (Habitats 8 #F6 R1, 2009), 1 individual; als; IBUFRJ 19297 (Habitats 7 #F6 R2, 2008), 1 individual;
IBUFRJ 19261 (Revizee V #41, 2001), 3 valves; IBUFRJ 19262 IBUFRJ 19298 (Habitats 7 #C6 R2, 2008), 16 individuals;
(Habitats 7 #A6 R1, 2008), 1 individual; IBUFRJ 19263 (Hab- IBUFRJ 19299 (Habitats 8 #C6 R2, 2009), 3 individuals; IBU-
itats 8 #B6 R3, 2009), 1 individual; IBUFRJ 19264 (Habitats 8 FRJ 19300 (Habitats 9 #H6 R2, 2009), 2 individuals; IBUFRJ
#D6 R2, 2009), 6 individuals; IBUFRJ 19265 (Habitats 7 #B6 19301 (Habitats 7 #C6 R3, 2008), 11 individuals; IBUFRJ
R3, 2008), 1 individual; IBUFRJ 19266 (Habitats 9 #F6 R3, 19302 (Habitats 8 #F6 R1, 2009), 4 individuals; IBUFRJ 19303
1
2 AMERICAN MALACOLOGICAL BULLETIN · ·
32 2 2014

(Habitats 7 #C6 R1, 2008), 17 individuals; IBUFRJ 19304 itats 7 #A6 R3, 2008), 7 individuals; IBUFRJ 19353 (Habitats
(Habitats 9 #H6 R1, 2009), 2 individuals; IBUFRJ 19305 7 #B6 R1, 2008), 1 individual; IBUFRJ 19354 (Habitats 7 #B6
(Habitats 7 #H6 R3, 2008), 1 individual; IBUFRJ 19306 (Hab- R1, 2008), 1 individual; IBUFRJ 19355 (Habitats 7 #B6 R2,
itats 7 #I6 R3, 2008), 3 individuals; IBUFRJ 19307 (Habitats 7 2008), 1 individual; IBUFRJ 19356 (Habitats 7 #B6 R2, 2008),
#B6 R2, 2008), 9 individuals; IBUFRJ 19308 (Habitats 7 #F6 1 individual; IBUFRJ 19357 (Habitats 7 #B6 R2, 2008), 1 in-
R3, 2008), 3 individuals; IBUFRJ 19309 (Habitats 7 #C6 R1, dividual; IBUFRJ 19358 (Habitats 7 #B6 R3, 2008), 5 indi-
2008), 1 individual; IBUFRJ 19310 (Habitats 7 #C6 R2, 2008), viduals; IBUFRJ 19359 (Habitats 7 #B6 R3, 2008), 3
1 individual; IBUFRJ 19311 (Habitats 7 #C6 R1, 2008), 2 in- individuals; IBUFRJ 19360 (Habitats 7 #C6 R1, 2008), 1 indi-
dividuals; IBUFRJ 19312 (Habitats 7 #D6 R3, 2008), 5 indi- vidual; IBUFRJ 19361 (Habitats 7 #C6 R2, 2008), 8 individu-
viduals; IBUFRJ 19313 (Habitats 9 #F6 R3, 2009), 1 als; IBUFRJ 19362 (Habitats 7 #C6 R2, 2008), 1 individual;
individual; IBUFRJ 19314 (Habitats 8 #D6 R1, 2009), 1 indi- IBUFRJ 19363 (Habitats 7 #C6 R3, 2008), 6 individuals; IBU-
vidual; IBUFRJ 19315 (Oceanprof I #44 North, 2002), 1 valve; FRJ 19364 (Habitats 7 #D6 R1, 2008), 3 individuals; IBUFRJ
IBUFRJ 19316 (Habitats 7 #C6 R3, 2008), 1 individual; IBU- 19365 (Habitats 7 #D6 R3, 2008), 5 individuals; IBUFRJ
FRJ 19317 (Habitats 7 #F6 R1, 2008), 1 individual; IBUFRJ 19366 (Habitats 7 #F6 R1, 2008), 6 individuals; IBUFRJ 19367
19318 (Habitats 7 #H6 R1, 2008), 1 individual; IBUFRJ 19319 (Habitats 7 #F6 R1, 2008), 1 individual; IBUFRJ 19368 (Hab-
(Habitats 8 #A6 R1, 2009), 3 individuals; IBUFRJ 19320 itats 7 #F6 R2, 2008), 2 individuals; IBUFRJ 19369 (Habitats
(Habitats 8 #A6 R3, 2009), 2 individuals; IBUFRJ 19321 7 #F6 R2, 2008), 1 individual; IBUFRJ 19370 (Habitats 7 #H6
(Habitats 8 #B6 R2, 2009), 12 individuals; IBUFRJ 19322 R1, 2008), 2 individuals; IBUFRJ 19371 (Habitats 7 #H6 R1,
(Habitats 8 #B6 R3, 2009), 15 individuals; IBUFRJ 19323 2008), 2 individuals; IBUFRJ 19372 (Habitats 7 #H6 R1,
(Habitats 8 #B6 R3, 2009), 2 individuals; IBUFRJ 19324 2008), 2 individuals; IBUFRJ 19373 (Habitats 7 #H6 R2,
(Habitats 8 #C6 R1, 2009), 16 individuals; IBUFRJ 19325 2008), 4 individuals; IBUFRJ 19374 (Habitats 7 #H6 R2,
(Habitats 8 #C6 R3, 2009), 8 individuals; IBUFRJ 19326 2008), 1 individual; IBUFRJ 19375 (Habitats 7 #H6 R3, 2008),
(Habitats 9 #F6 R3, 2009), 2 individuals; IBUFRJ 19327 1 individual; IBUFRJ 19376 (Habitats 7 #I6 R1, 2008), 1 indi-
(Habitats 7 #H6 R2, 2008), 4 individuals; IBUFRJ 19328 vidual; IBUFRJ 19377 (Habitats 7 #I6 R2, 2008), 1 individual;
(Habitats 7 #I6 R1, 2008), 2 individuals; IBUFRJ 19329 (Hab- IBUFRJ 19378 (Habitats 8 #A6 R1, 2009), 1 individual; IBU-
itats 7 #I6 R1, 2008), 3 individuals; IBUFRJ 19330 (Habitats 7 FRJ 19379 (Habitats 8 #B6 R3, 2009), 3 individuals; IBUFRJ
#I6 R2, 2008), 3 individuals; IBUFRJ 19331 (Habitats 7 #I6 19380 (Habitats 8 #C6 R1, 2009), 1 individual; IBUFRJ 19381
R2, 2008), 1 individual; IBUFRJ 19332 (Habitats 8 #A6 R2, (Habitats 8 #D6 R1, 2009), 1 individual; IBUFRJ 19382 (Hab-
2009), 2 individuals; IBUFRJ 19333 (Habitats 8 #A6 R3, itats 8 #F6 R1, 2009), 1 individual; IBUFRJ 19383 (Revizee V
2009), 1 individual; IBUFRJ 19334 (Habitats 8 #B6 R3, 2009), #25A, 2001), 4 valves; IBUFRJ 19384 (Revizee V #41, 2001), 4
3 individuals; IBUFRJ 19335 (Habitats 8 #C6 R1, 2009), 3 in- valves; IBUFRJ 19385 (Revizee V #52, 2001), 114 valves; IBU-
dividuals; IBUFRJ 19336 (Habitats 8 #C6 R2, 2009), 1 indi- FRJ 19386 (Revizee V #504, 2001), 6 valves; IBUFRJ 19387
vidual; IBUFRJ 19337 (Habitats 8 #F6 R1, 2009), 1 individual; (Revizee V #1 profound, 2001), 6 valves.
IBUFRJ 19338 (Habitats 8 #E6 R1, 2009), 1 individual; IBU-
FRJ 19339 (Habitats 6 #C6 R1, 2008), 1 individual; IBUFRJ Ledella manati sp. nov.
19340 (Habitats 6 #F6 R3, 2008), 3 individuals; IBUFRJ 19341 IBUFRJ 13378 (Revizee V #509, 2001), 2 valves; IBUFRJ
(Habitats 6 #Canac6 R1, 2008), 1 individual; IBUFRJ 19342 14070 (Revizee V #41C, 2001), 4 valves; IBUFRJ 14344 (Re-
(Habitats 9 #H6 R3, 2009), 2 individuals; IBUFRJ 19343 vizee V #41C, 2001), 4 valves; IBUFRJ 14569 (Revizee V #41C,
(Habitats 9 #I6 R3, 2009), 1 individual; IBUFRJ 19344 (Habi- 2001), 4 valves; IBUFRJ 14673 (Revizee V #41C, 2001), 6
tats 9 #Canac7 R1, 2009), 1 individual; IBUFRJ 19345 (Re- valves; IBUFRJ 15989 (Oceanprof II #69 South, 2003), 2
vizee V #25A, 2001), 2 valves; IBUFRJ 19346 (Oceanprof II valves; IBUFRJ 19388 (Oceanprof II #79 South, 2003), 6
#79 South, 2003), 1 valve. valves; IBUFRJ 19389 (Habitats 3 #C8 R3, 2008), 2 individu-
als; IBUFRJ 19390 (Habitats 6 #F6 R3, 2008), 1 individual;
Ledella elfica sp. nov. IBUFRJ 19391 (Habitats 7 #A6 R1, 2008), 1 individual; IBU-
IBUFRJ 16841 (Oceanprof II #79 South, 2003), 4 valves; FRJ 19392 (Habitats 7 #A6 R2, 2008), 3 individuals; IBUFRJ
IBUFRJ 17131 (Oceanprof I #44 North, 2002), 1 valve; IBU- 19393 (Habitats 7 #B6 R1, 2008), 5 individuals; IBUFRJ
FRJ 17715 (Oceanprof II #79 South, 2003), 5 valves; IBUFRJ 19394 (Habitats 7 #B6 R2, 2008), 2 individuals; IBUFRJ
19347 (Habitats 6 #H6 R1, 2008), 5 individuals; IBUFRJ 19395 (Habitats 7 #B6 R2, 2008), 1 individual; IBUFRJ 19396
19348 (Habitats 9 #H6 R2, 2009), 1 individual; IBUFRJ 19349 (Habitats 7 #B6 R3, 2008), 1 individual; IBUFRJ 19397 (Hab-
(Habitats 7 #A6 R1, 2008), 3 individuals; IBUFRJ 19350 itats 7 #C6 R1, 2008), 4 individuals; IBUFRJ 19398 (Habitats
(Habitats 7 #A6 R2, 2008), 4 individuals; IBUFRJ 19351 7 #C6 R1, 2008), 1 individual; IBUFRJ 19399 (Habitats 7 #C6
(Habitats 7 #A6 R2, 2008), 1 individual; IBUFRJ 19352 (Hab- R2, 2008), 3 individuals; IBUFRJ 19400 (Habitats 7 #C6 R2,
 FOUR NEW SPECIES OF LEDELLA FROM BRAZIL 3

2008), 1 individual; IBUFRJ 19401 (Habitats 7 #C6 R3, 2008),

3 individuals; IBUFRJ 19402 (Habitats 7 #D6 R1, 2008), 1 in-
dividual; IBUFRJ 19403 (Habitats 7 #D6 R1, 2008), 2 indi-
viduals; IBUFRJ 19404 (Habitats 7 #D6 R1, 2008), 1
individual; IBUFRJ 19405 (Habitats 7 #F6 R1, 2008), 3 indi-
viduals; IBUFRJ 19406 (Habitats 7 #F6 R1, 2008), 1 individu-
al; IBUFRJ 19407 (Habitats 7 #F6 R2, 2008), 5 individuals;
IBUFRJ 19408 (Habitats 7 #F6 R2, 2008), 3 individuals; IBU-
FRJ 19409 (Habitats 7 #H6 R1, 2008), 1 individual; IBUFRJ
19410 (Habitats 7 #H6 R2, 2008), 1 individual; IBUFRJ 19411
(Habitats 7 #H6 R2, 2008), 2 individuals; IBUFRJ 19412
(Habitats 7 #H6 R2, 2008), 1 individual; IBUFRJ 19413 (Hab-
itats 7 #I6 R1, 2008), 1 individual; IBUFRJ 19414 (Habitats 7
#I6 R1, 2008), 1 individual; IBUFRJ 19415 (Habitats 7 #I6 R2,
2008), 2 individuals; IBUFRJ 19416 (Habitats 7 #I6 R3, 2008),
1 individual; IBUFRJ 19417 (Habitats 8 #A6 R1, 2009), 3 in-
dividuals; IBUFRJ 19418 (Habitats 8 #A6 R2, 2009), 1 indi-
vidual; IBUFRJ 19419 (Habitats 8 #A6 R3, 2009), 1 individual;
IBUFRJ 19420 (Habitats 8 #B6 R2, 2009), 3 individuals; IBU-
FRJ 19421 (Habitats 8 #B6 R3, 2009), 1 individual; IBUFRJ
19422 (Habitats 8 #C6 R2, 2009), 1 individual; IBUFRJ 19423
(Habitats 8 #C6 R3, 2009), 5 individuals; IBUFRJ 19424
(Habitats 8 #C6 R3, 2009), 1 individual; IBUFRJ 19425 (Hab-
itats 8 #F6 R1, 2009), 1 individual; IBUFRJ 19426 (Habitats 8
#F6 R1, 2009), 1 individual; IBUFRJ 19427 (Habitats 8 #F6
R2, 2009), 1 individual; IBUFRJ 19428 (Habitats 9 #F6 R3,
2009), 1 individual; IBUFRJ 19429 (Habitats 9 #H6 R1, 2009),
2 individuals; IBUFRJ 19430 (Habitats 9 #H6 R2, 2009), 1
individual; IBUFRJ 19431 (Habitats 9 #Canac6 R2, 2009), 1
individual; IBUFRJ 19432 (Revizee V #1 profound, 2001), 4
valves; IBUFRJ 19433 (Revizee V #25A, 2001), 4 valves; IBU-
FRJ 19434 (Revizee V #52, 2001), 29 valves; IBUFRJ 19435
(Revizee V #504, 2001), 14 valves.

Table S1. Material examined from the Oceanprof and Revizee campaigns.

Campaign Station Date Latitude Longitude Depth (m)

Oceanprof I 44 10-XII-2002 22°10′43″S 39°54′46″W 754
Oceanprof II 44 01-VII-2003 22°10′43″S 39°54′45″W 749
Oceanprof II 69 18-VI-2003 22°31′11″S 40°15′12″W 743
Oceanprof II 79 01-VI-2003 22°20′22″S 40°01′24″W 755
Revizee V 1 03-VII-2001 13°04′S 38°21′W 450
Revizee V 25A 28-VI-2001 19°38′S 38°46′W 230
Revizee V 41 20-VII-2001 21°13′S 40°13′W 1000
Revizee V 41C 10-VII-2001 20°30′S 29°16′W 360
Revizee V 52 21-VII-2001 21°46′S 40°05′W 450
Revizee V 504 02-VII-2001 14°28′S 38°54′W 278
Revizee V 509 01-VII-2001 15°47′72″S 38°21′97″W 2076
4 AMERICAN MALACOLOGICAL BULLETIN · ·
32 2 2014

Table S2. Material examined from the Habitats campaigns.

Campaign Station Replica Date Latitude Longitude Depth (m)

Habitats 3 C8 R3 10-V-2008 23°01′32″S 40°45′20″W 975
Habitats 6 Canac6 R1 27-VI-2008 21°50′02″S 40°06′13″W 481
Habitats 6 I9 R1 29-VI-2008 21°11′06″S 40°03′06″W 1313
Habitats 7 A6 R1 03-VII-2008 23°37′55″S 41°19′40″W 390.2
Habitats 7 A6 R2 03-VII-2008 23°37′57″S 41°19′42″W 390.6
Habitats 7 A6 R3 03-VII-2008 23°37′57″S 41°19′42″W 388.9
Habitats 7 B6 R1 04-VII-2008 23°12′32″S 40°58′30″W 451
Habitats 7 B6 R2 04-VII-2008 23°12′31″S 40°58′31″W 447.4
Habitats 7 B6 R3 04-VII-2008 23°10′22″S 40°56′46″W 424.2
Habitats 7 C6 R1 04-VII-2008 22°59′00″S 40°48′25″W 387.4
Habitats 7 C6 R2 04-VII-2008 22°59′00″S 40°48′25″W 386.8
Habitats 7 C6 R3 04-VII-2008 22°59′01″S 40°48′24″W 399.7
Habitats 7 D6 R1 08-VII-2008 22°33′35″S 40°26′38″W 396.1
Habitats 7 D6 R3 11-VII-2008 22°33′33″S 40°26′40″W 393.4
Habitats 7 F6 R1 08-VII-2008 22°19′10″S 40°05′42″W 402.3
Habitats 7 F6 R2 08-VII-2008 22°19′11″S 40°05′44″W 403
Habitats 7 F6 R3 08-VII-2008 22°19′10″S 40°05′41″W 404.6
Habitats 7 H6 R1 06-VII-2008 21°42′02″S 40°06′15″W 404.7
Habitats 7 H6 R2 07-VII-2008 21°44′21″S 40°05′18″W 402
Habitats 7 H6 R3 07-VII-2008 21°44′21″S 40°05′18″W 401.6
Habitats 7 I6 R1 06-VII-2008 21°13′38″S 40°14′58″W 417.6
Habitats 7 I6 R2 06-VII-2008 21°13′38″S 40°14′58″W 412
Habitats 7 I6 R3 06-VII-2008 21°13′37″S 40°14′58″W 414.5
Habitats 8 A6 R1 01-II-2009 23°37′57″S 41°19′41″W 390.7
Habitats 8 A6 R2 01-II-2009 23°37′58″S 41°19′41″W 400.5
Habitats 8 A6 R3 01-II-2009 23°37′57″S 41°19′38″W 390.5
Habitats 8 B6 R2 01-II-2009 23°10′23″S 40°56′45″W 432
Habitats 8 B6 R3 01-II-2009 23°10′23″S 40°56′48″W 418
Habitats 8 C6 R1 31-I-2009 22°59′01″S 40°48′27″W 380.6
Habitats 8 C6 R2 31-I-2009 22°59′00″S 40°48′26″W 393.6
Habitats 8 C6 R3 31-I-2009 22°59′00″S 40°48′28″W 376.6
Habitats 8 D6 R1 31-I-2009 22°33′35″S 40°26′37″W 400
Habitats 8 D6 R2 31-I-2009 22°33′35″S 40°26′37″W 401
Habitats 8 D6 R3 31-I-2009 22°33′36″S 40°26′37″W 383
Habitats 8 E6 R1 31-I-2009 22°25′58″S 40°17′35″W 380
Habitats 8 F6 R1 30-I-2009 22°19′02″S 40°05′27″W 383.8
Habitats 8 F6 R2 30-I-2009 22°19′03″S 40°05′28″W 386
Habitats 8 D8 R3 22-I-2009 22°40′57″S 40°17′36″W 1013
Habitats 9 F6 R3 10-II-2009 22°19′10″S 40°05′42″W 400
Habitats 9 H6 R1 05-II-2009 21°44′21″S 40°04′59″W 405
Habitats 9 H6 R2 05-II-2009 21°44′21″S 40°04′53″W 404
Habitats 9 H6 R3 05-II-2009 21°44′21″S 40°04′59″W 402
Habitats 9 Canac6 R2 07-II-2009 21°50′02″S 40°05′55″W 780
Habitats 9 Canac7 R1 06-II-2009 21°47′26″S 40°01′55″W 474.8