See discussions, stats, and author profiles for this publication at: https://www.researchgate.

net/publication/266384674

The genera Limatula and Limea (Mollusca, Pelecypoda, Limidae) from deep
waters off Brazil

Article  in  Zootaxa · November 2008

DOI: 10.11646/zootaxa.1940.1.5

CITATIONS READS

12 444

2 authors:

Cleo Oliveira Ricardo Silva Absalão

Federal University of Rio de Janeiro Federal University of Rio de Janeiro
18 PUBLICATIONS   102 CITATIONS    96 PUBLICATIONS   769 CITATIONS   

SEE PROFILE SEE PROFILE

Some of the authors of this publication are also working on these related projects:

Morfoanatomia de Gastrópodes (Mollusca) Terrestres em Floresta Ombrófila Densa Montana do Parque Nacional da Serra dos Órgãos, Rio de Janeiro View project

All content following this page was uploaded by Cleo Oliveira on 11 November 2014.

The user has requested enhancement of the downloaded file.

 Zootaxa 1940: 48–58 (2008) ISSN 1175-5326 (print edition)
www.mapress.com / zootaxa/
Copyright © 2008 · Magnolia Press
ZOOTAXA
ISSN 1175-5334 (online edition)

The genera Limatula and Limea (Mollusca, Pelecypoda, Limidae)

from deep waters off Brazil

CLÉO DILNEI DE CASTRO OLIVEIRA1 & RICARDO SILVA ABSALÃO1,2

1
Departamento de Zoologia, Instituto de Biologia, Centro de Ciências da Saúde, Universidade Federal do Rio de Janeiro, Ilha do
Fundão, 21941-590 Rio de Janeiro, Rio de Janeiro, Brasil
2
Departamento de Zoologia, Universidade do Estado do Rio de Janeiro, Avenida São Francisco Xavier 524, Maracanã, 20550-900
Rio de Janeiro, Rio de Janeiro, Brasil

Abstract

Five species of the genus Limatula and one of Limea were found in a total of 83 samples taken from deep waters (700–
1950 m) off Brazil. Limatula laminifera (Smith, 1885) and Limea lirata Allen, 2004 were not previously reported from
Brazil. Limatula confusa (Smith, 1885) and L. louiseae Clarke, 1974, although already reported for Brazilian waters,
have their distribution extended southward. Limatula domaneschii n.sp. is described from the Campos Basin (˜22° S). It
is diagnosed by concentric lamellae strengthening towards the ventral margin of the shell, and absence of axial ornamen-
tation; an internal median groove is not present. A fifth Limatula species is present, and we strongly suspect that it is also
new to science, but a formal description is delayed until additional specimens are collected.

Key words: Limidae, Limatula, Limea, deep-water, geographic distribution, Southern Atlantic, Brazil, Bivalvia

Introduction

Members of the family Limidae are present in all the world seas and at most depths. This group usually com-
prises pelecypods with conchological variations of the same basic body plan. These shell characters are
important for taxonomy, and their distinguishing features usually include size; shell outline and ornamenta-
tion; internal ridge and/or fold; and, when present, the relative number of axial ribs. Ribs number has been
widely used as a species character in several limid genera (Mikkelsen & Bieler 2003). In 1968, Stuardo car-
ried out an extensive revision including conchological analysis, anatomy, distribution, revision of types, and
introduction of more than 80 new taxa (as manuscript names) to the family Limidae. Unfortunately, only a
small part of this study was published (Stuardo 1982). Nowadays, a considerable number of species have been
studied, with molecular analyses, detailed morphology, reproductive cycles, life history, respiration and filtra-
tion aspects, speciation and dispersal evidence (Gilmour 1974; Morton 1979, 2000; Lodeiros & Himmelman
1999; Page & Linse 2002; Allen 2004; Jarnegren & Altin 2006; Jarnegren et al. 2007). Stuardo (1968) distin-
guished, based on an extensive list of characters, two large groups of limid genera: Limaria-Limatula-Limea
and Ctenoides-Lima-Ancesta-Divarilima. For the Atlantic basin, Limatula and Limea were studied by Allen
(2004) concerning conchological and anatomical aspects.
In spite of the undoubted importance of conchological traits for taxonomy, not uncommonly definitions
based only on shell characters fail to establish clear boundaries among different taxa, even at the genus level.
In 1978, Fleming produced an extensive review and documented the conservative nature of the shell morphol-
ogy of Limatula, which makes the shell characters difficult to use for phylogeny reconstruction. The phyloge-
netic position, even at the family level, of the limids within the pteriomorph pelecypods is still under

48 Accepted by R. Bieler: 30 Oct. 2008; published: 24 Nov. 2008

discussion (Waller 1998; Steiner & Hammer 2000; Giribet & Wheeler 2002; Mikkelsen & Bieler 2003).
According to Allen (2004, p. 2593), the limid subgroups may overlap one another, because “it is such a
conservative group with such a continuum of form [...] that these subdivisions should be regarded with great
caution.” We agree with this opinion, and for this reason we prefer to retain the species cited here in their
usual genera.
The last Brazilian catalog of molluscs (Rios 1994) reported only three species in Limatula: Limatula con-
fusa (Smith, 1885); Limatula hendersoni Olsson & McGinty, 1958; and Limatula pygmaea (Philippi, 1845);
and one species in Limea: Limea browniana Dall, 1886. Among them, only L. confusa inhabits the deep
waters and will be considered at this time.
The goal of the present study was to revise the Brazilian deep water Limidae. The study forms part of a
major project on Brazilian deep water molluscs, especially from the Campos Basin in the state of Rio de Jan-
eiro, the main oil production region in Brazil.

Material and methods

The molluscs were taken from northern and southern regions off Brazil (700–1950 meters depth). Most of the
sampling stations were in the Bacia de Campos (Campos Basin) and were visited as part of the program
“Environmental Characterization of Campos Basin, RJ, Brazil”. Samples were obtained with a 0.25 m2 box
core, by the Research Vessel "Astro-Garoupa” belonging to Petrobras S.A. (a public Brazilian oil company).
A total of 82 samples was taken in this region, plus an additional sample from the continental slope off Amapá
State in northern Brazil. Each sample was washed in running seawater through a mesh of 300 μm, and the res-
idue placed in 70% ethanol. In the laboratory, this residue was sorted under magnification and the limids
picked out. Only empty shells were obtained. The generic characterization was adapted from Abbott (1974),
Rios (1994), and Allen (2004). Types were not examined when the original illustration (and/or description)
was good enough to identify the taxon. Since only one species was obtained from Amapá (north Brazil) any
kind of comparison between the two sites is precluded. All material is deposited in the Molluscan Collection
of the Departamento de Zoologia, Instituto de Biologia of the Universidade Federal do Rio de Janeiro (IBU-
FRJ).

Results

Limidae Rafinesque, 1815

Limatula Wood, 1839

Type species: Pecten subauriculatus Montagu, 1808

Description: Shell small, white or glassy, usually fragile, somewhat elongated and oblique, inflated, often
with axial ornamentation, beaks prominent, small auricles usually present, internal median groove may be
present, hinge without teeth.

Limatula louiseae Clarke, 1974

Figures 1–5

Limatula louiseae Clarke, 1974: p.17, fig. 9; Allen, 2004: p. 2601, fig. 10–12, 45a.

LIMATULA AND LIMEA FROM BRAZILIAN DEEP Zootaxa 1940 © 2008 Magnolia Press · 49
Description: Shell fragile, white or translucent, small (height mean = 1.61mm ± 0.38 s.d.; range 0.91–
2.23mm; n=50), height/length ratio 1.53, equivalve, equilateral, very inflated, elongated, ornamented exter-
nally with unevenly spaced concentric lines that are less apparent in the dorsal region, and many concentric
microscopic striae between these lines. Microscopic diagonal striae, which vary in number, on anterior and

FIGURES 1–5. Limatula louiseae Clarke, 1974. 1—External view, IBUFRJ 17809; 2—Internal view, IBUFRJ 17811);
3—Detail of sculpture, IBUFRJ 17809; 4—Hinge, IBUFRJ 17811; 5—Dorsal view, IBUFRJ 17810. FIGURES 6–9.
Limatula confusa (Smith, 1885). 6—External view, IBUFRJ 17804; 7—External view, IBUFRJ 17807; 8—Internal view,
IBUFRJ 17807; 9—Hinge, IBUFRJ 17807. FIGURES 10–12. Limatula domaneschii n.sp. 10—External view, IBUFRJ
17806 (Holotype); 11—Internal view, IBUFRJ 17801 (Paratype 1); 12—Hinge, IBUFRJ 17801(Paratype 1). Scale bars:
1–5: 250µm; 6–9: 500µm; 10–12: 100µm.

50 · Zootaxa 1940 © 2008 Magnolia Press OLIVEIRA & ABSALAO

posterior parts of shell. Internal median axial fold absent, auricles small and usually short and rounded, subau-
ricular sinus absent, ventral margin strongly rounded or angulated. Prodissoconch smooth and somewhat
detached (mean 109.00μm ± 7.38 s.d.; range 100–120 μm; n=50), interdissoconch not distinguishable.
Umbones prominent, hinge plate somewhat broad, ligamental pit triangular.
Geographic range: At Mid-Atlantic Ridge, close to equator and west of Newfoundland, off Surinam,
North America Basin. Depths ranging between 1500 and 4793 meters. Adapted from Clarke (1974) and Allen
(2004). Campos Basin (this study) represents the southernmost and the shallowest (1330 m) record of this spe-
cies, and the first record in the southeastern Atlantic Ocean.
Material examined: IBUFRJ 17809, off Bacia de Campos (21°58'36" S, 39°46'30" W, 1700 m),
10.VIII.01, [1 valve]; IBUFRJ 17810, off Bacia de Campos (22°37'02" S, 39°56'21" W, 1950 m), 23.XI.02, [1
spec.]; IBUFRJ 17811, off Bacia de Campos (21°58'36" S, 39°46'30" W, 1700 m), 10.VIII.01, [1 valve]; IBU-
FRJ 17824, off Bacia de Campos (22°38'53" S, 40°04'16" W, 1342 m), 14.VI.03, [1 valve]; IBUFRJ 17825,
off Bacia de Campos (22°03'28" S, 39°45'07" W, 1730 m), 08.V.02, [7 valves]; IBUFRJ 17826, off Bacia de
Campos (22°05'12" S, 39°42'41" W, 1930 m), 08.V.02, [4 valves]; IBUFRJ 17827, off Bacia de Campos
(22°09'11" S, 39°44'51" W, 1930 m), 08.V.02, [17 valves]; IBUFRJ 17828, off Bacia de Campos (22°04'53" S,
39°49'04" W, 1330 m), 09.V.02, [1 valve]; IBUFRJ 17829, off Bacia de Campos (22°05'45" S, 39°45'55" W,
1730 m), 09.V.02, [1 spec. + 4 valves]; IBUFRJ 17830, off Bacia de Campos (21°58'36" S, 39°46'30" W, 1700
m), 08.X.01, [3 valves]; IBUFRJ 17831, off Bacia de Campos (22°06'53" S, 39°44'14" W, 1930 m), 08.V.02,
[17 valves]; IBUFRJ 17832, off Bacia de Campos (22°08'24" S, 39°46'24" W, 1730 m), 09.V.02, [3 valves];
IBUFRJ 17833, off Bacia de Campos (22°41'04" S, 40°02'30" W, 1650 m), 23.XI.02, [1 spec.]; IBUFRJ
17834, off Bacia de Campos (21°52'44" S, 39°40'46" W, 1950 m), 11.XII.02, [8 valves]; IBUFRJ 17835, off
Bacia de Campos (21°57'16" S, 39°47'44" W, 1650 m), 14.XII.02, [20 valves]; IBUFRJ 17836, off Bacia de
Campos (21°57'27" S, 39°40'34" W, 1942 m), 27.VI.03, [23 valves]; IBUFRJ 17837, off Bacia de Campos
(22°30'22"S, 39°56'54" W, 1353 m), 21.VI.03, [2 valves]; IBUFRJ 17838, off Bacia de Campos (22°31'37" S,
39°55'14" W, 1630 m), 16.VI.03, [3 valves]; IBUFRJ 17839, off Bacia de Campos (21°52'41" S, 39°46'17" W,
1688 m), 26.VI.03, [12 valves]; IBUFRJ 17840, off Bacia de Campos (22°11'16" S, 39°43'45" W, 1968 m),
22.VI.03, [27 valves]; IBUFRJ 17841, off Bacia de Campos (22°28'46" S, 39°53'28" W, 1621 m), 17.VI.03, [7
valves]; IBUFRJ 17842, off Bacia de Campos (22°04'45" S, 39°41'58" W, 1910 m), 27.VI.03, [28 valves];
IBUFRJ 17843, off Bacia de Campos (22°36'12" S, 39°58'23" W, 1670 m), 13.VI.03, [2 valves]; IBUFRJ
17844, off Bacia de Campos (22°04'34" S, 39°52'05" W, 1030 m), 30.VI.03, [1 valve]; IBUFRJ 17845, off
Bacia de Campos (22°41'32" S, 40°00'47" W, 1906 m), 12.VI.03, [44 valves]; IBUFRJ 17846, off Bacia de
Campos (22°33'08" S, 39°54'21" W, 1934 m), 15.VI.03, [26 valves]; IBUFRJ 17847, off Bacia de Campos
(21°52'43" S, 39°40'42" W, 1941 m), 26.VI.03, [40 valves]; IBUFRJ 17848, off Bacia de Campos (22°41'11"
S, 40°02'20" W, 1623 m), 13.VI.03, [7 valves]; IBUFRJ 17849, off Bacia de Campos (22°30'35" S, 39°51'45"
W, 1970 m), 16.VI.03, [12 valves]; IBUFRJ 17850, off Bacia de Campos (22°04'45" S, 39°46'32" W, 1643
m), 27.VI.03, [3 valves]; IBUFRJ 17851, off Bacia de Campos (22°37'03" S, 39°56'20" W, 1945 m),
13.VI.03, [31 valves]; IBUFRJ 17852, off Bacia de Campos (22°36'12" S, 39°58'23" W, 1670 m), 13.VI.03, [4
valves]; IBUFRJ 17853, off Bacia de Campos (22°11'04" S, 39°47'05" W, 1654 m), 22.VI.03, [3 valves].
Remarks: The presence of diagonal striae on the anterior and posterior parts of the shell is the most
important diagnostic character of this species, and distinguishes it from all other species in the Atlantic Ocean.
According to Allen (2004), although rare, L. louiseae is widely distributed in the North Atlantic and south of
the equator. This taxon was abundant in our samples, with a total of 368 valves collected. The number of diag-
onal striae on the anterior and posterior parts of the shell is quite variable, and depends on the size of the shell;
often, small shells tend to have less conspicuous diagonal striae.

LIMATULA AND LIMEA FROM BRAZILIAN DEEP Zootaxa 1940 © 2008 Magnolia Press · 51
Limatula confusa (Smith, 1885)
Figures 6–9

Lima confusa Smith, 1885: p. 292, pl. XXIV, fig. 6, 6a.

Limatula confusa: Stuardo, 1968: pl. 16, fig. 88; Abbott, 1974: p. 454.

Description: Shell fragile, white, small (height mean = 3.28mm ± 2.06 s.d.; range 1.10–6.30mm; n=14),
height/length ratio 1.41, equivalve, slightly unequilateral, very inflated, outline suboval, thin axial ribs cover
entire shell, but axial ribs not evenly spaced. The axials are closer to each other on the dorsal side, and slowly
increase in the distance between them towards the ventral side, very fine concentric striae present, forming
small nodules where they cross the axial ribs. Internal axial ridge absent. Auricles small, with few ornaments,
subauricular sinus present, ventral region rounded or angled. Prodissoconch smooth (mean 177.50μm ± 16.69
s.d.; range 160–200 μm; n=14), interdissoconch hardly distinguishable in some specimens (mean 805.00μm
± 33.38 s.d.; range 760–860 μm; n=14). Umbones prominent, hinge plate large, ligamental pit broad, triangu-
lar.
Geographic range: North Carolina to the West Indies, off Azores, Northeast Brasil, off Pernambuco
State. Depths ranging between 60 to 2650 meters. Adapted from Abbott (1974), Rios (1994) and Allen (2004).
Campos Basin (this study), is the southernmost record in the Atlantic Ocean.
Material examined: IBUFRJ 17802, off Bacia de Campos (21°57'27" S, 39°40'34" W, 1942 m),
27.VI.03, [1 valve]; IBUFRJ 17803, off Bacia de Campos (21°52'41" S, 39°46'17" W, 1688 m), 26.VI.03, [1
valve]; IBUFRJ 17804, off Bacia de Campos (22°11'17" S, 39°43'45" W, 1950 m), 25.XI.02, [1 valve]; IBU-
FRJ 17807, off Bacia de Campos (21°58'36" S, 39°46'30" W, 1700 m), 10.VIII.01, [1 valve]; IBUFRJ 17812,
off Bacia de Campos (22°31'37" S, 39°55'14" W, 1630 m), 16.VI.03, [1 valve]; IBUFRJ 17813, off Bacia de
Campos (22° 24'32" S, 39°57'28" W, 1050 m), 20.XI.02, [1 valve]; IBUFRJ 17814, off Bacia de Campos
(22°36'12" S, 39°58'23" W, 1670 m), 13.VI.03, [1 valve]; IBUFRJ 17815, off Bacia de Campos (22°31'28" S,
40°03'50" W, 1043 m), 18.VI.03, [1 valve]; IBUFRJ 17816, off Bacia de Campos (21°57'27" S, 39°40'34" W,
1942 m), 27.VI.03, [1 valve]; IBUFRJ 17817, off Bacia de Campos (22°04'45" S, 39°46'32" W, 1643 m),
27.VI.03, [1 valve]; IBUFRJ 17818, off Bacia de Campos (22°11'04" S, 39°47'05" W, 1650 m), 25.XI.02, [1
valve]; IBUFRJ 17819, off Bacia de Campos (22°10'54" S, 39°52'19" W, 1050 m), 10.XII.02, [1 valve]; IBU-
FRJ 17820, off Bacia de Campos (22°05'45" S, 39°45'55" W, 1730 m), 09.V.02, [1 valve]; IBUFRJ 17821, off
Bacia de Campos (22°05'04" S, 39°50'02" W, 1230 m), 09.V.02, [1 valve]; IBUFRJ 17822, off Bacia de Cam-
pos (22°39'34" S, 40°08'22" W, 1200 m), 15.V.02, [1 valve].
Remarks: Apparently this species has not been collected since its description (from the Challenger Expe-
dition material), and a part of the type series was destroyed (Allen 2004, p. 2644). However, the description
and especially the type illustration in plate 24, figs. 6-6a (Smith 1885 p. 292), are good enough to allow us to
identify it. In our description we include details that were not mentioned in the original description because of
the limitations in optics of the period.
Limatula margaretae Allen, 2004 is the most similar taxon to L. confusa in the Atlantic basin: both spe-
cies share the same outline and type of ornamentation, but can be distinguished by the presence of axial ribs
over all the shell including the auricles in L. confusa (fig. 6–7), while the auricles are devoid of axial ribs and
the central and postero-central axial ribs are barely visible in L. margaretae. Besides that, the umbones at L.
margaretae are more prominent than in L. confusa.

Limatula domaneschii n.sp.

Figures 10–12

Description: Shell fragile, white, small (height mean = 1.07mm ± 0.15 s.d.; range 0.90–1.20mm; n=3),
height/length ratio 1.43, slightly inequilateral, moderately inflated, outline suboval, ornamented with rela-

52 · Zootaxa 1940 © 2008 Magnolia Press OLIVEIRA & ABSALAO

tively equidistant thin concentric ridges that become more apparent toward the ventral margin. Dorso-central
region without ornamentation. Secondary faint ridges may be present between the primary ridges. The con-
centric ridges extend toward the auricles, which become furrowed. With no external or internal axial orna-
mentation. Auricles small, symmetrical, and slightly inclined in relation to the antero-posterior axis, dorsal
edge raising upwards slightly, subauricular sinus absent, posterior margin slightly more curved than the ante-
rior margin. Ventral margin evenly curved. Prodissoconch smooth and rounded in shape, with conspicuous
edge (mean 170.00μm ± 5.00 s.d.; range 165–175 μm; n=3). Interdissoconch not distinguishable. Umbones
prominent. Hinge plate large, without teeth. Ligamental pit rounded.
Geographic range: Restricted to Campos Basin, Rio de Janeiro state, Brazil.
Material examined: Holotype: IBUFRJ 17806, off Bacia de Campos (22°24'30" S, 39°57'29" W, 1044
m), 20.VI.03, [2 valves]; paratype 1-IBUFRJ 17801, off Bacia de Campos (22°24'30" S, 39°57'29" W, 1044
m), 20.VI.03, [2 valves].
Etymology: This species is named in honor of Dr. Osmar Domaneschi, a Brazilian malacologist, for his
contribution to the knowledge of functional anatomy of the pelecypods and his contribution to the SBMa
(Sociedade Brasileira de Malacologia, Brazilian Malacological Society) for which he served more than 10
years as vice president.
Remarks: Because of the similarities among the genera of Limidae, we chose to compare our proposed
new taxon with all the species that are characterized mainly by the presence of concentric ornamentation,
regardless of their present generic allocation.
Limea lirata Allen, 2004 is more oblique and somewhat more rounded than L. domaneschii n.sp. More-
over, L. lirata is ornamented with numerous concentric lines, which are closer to each other and cover the
entire shell; the umbones are more slender and comparatively more projected beyond the hinge than the
umbones of L. domaneschii n.sp. The main difference is the very fine axial ridges in L. lirata, which are
absent in L. domaneschii n.sp.
Limatula domaneschii n.sp. can be distinguished from L. laminifera (Smith, 1885) by the presence, in the
latter, of an elongated, triangular ligamental pit, ornamentation over the entire shell, the presence of an auric-
ular sinus, a slender umbo, and a short hinge plate; whereas L. domaneschii n.sp. has a rounded ligamental pit,
the dorso-central region without ornamentation, no auricular sinus, and a comparatively large umbo and hinge
plate. Also, L. laminifera bears, internally, two thickened axial ridges at the middle of the shell, and axially
oriented slender striae, but both characters are absent in L. domaneschii n.sp. Additionally, L. domaneschii
n.sp. has the dorsal edge raising upwards slightly, which does not occur in any other species known from the
Brazilian coast and the surrounding area.
Limatula louiseae and L. domaneschii n.sp. share the concentric ornamentation, but the former species
shows the diagnostic microscopic diagonal striae that are absent in L. domaneschii n.sp.. Besides that, L. lou-
iseae is slender (height/length ratio 1.53), its prodissoconch is small (109μm) and the ligamental pit is triangu-
lar, whereas L. domaneschii n.sp, has a height/length ratio of 1.43, a prodissoconch of 170 μm and a rounded
ligamental pit.
A comparison with the 87 Limatula presumed species in Stuardo (1968) showed no close similarity with
L. domaneschii n.sp.; most of them have dominant axial ornamentation while L. domaneschii n.sp has exclu-
sively concentric ornamentation.

Limatula sp.
Figures 13–15

Description: Shell very fragile, white, small (height mean = 1.10mm ± 0.14 s.d.; range 1.00–1.20mm; n=2),
height/length ratio 1.47, slightly inequilateral, little inflated, outline suboval, dorso-central region smooth or

LIMATULA AND LIMEA FROM BRAZILIAN DEEP Zootaxa 1940 © 2008 Magnolia Press · 53
with irregularly spaced concentric cordlets, region near the edge of the shell ornamented with strong axial ribs
and concentric striae that cross the axial ribs, generating rounded nodules. The axial ribs are relatively equi-
distant and more visible on the ventral portion of the shell. Internally there are two slender axial ridges at the
middle of the shell. Auricles small, symmetrical and slightly inclined in relation to the antero-posterior axis,
forming an angle with the anterior and posterior margins. Subauricular sinus absent, anterior and posterior
margins approximately symmetrical. Ventral margin somewhat rounded. Prodissoconch smooth, with con-
spicuous edge (mean 122.50μm ± 7.50 s.d.; range 115–130 μm; n=2). Interdissoconch not distinguishable.
Umbones moderately prominent. Hinge plate large, without teeth. Ligamental pit broad, triangular.
Geographic range: Restricted to Campos Basin, Rio de Janeiro state, Brazil.
Material examined: IBUFRJ 17800, off Bacia de Campos (22°41'32" S, 40°00'47" W, 1906 m),
12.VI.03, [2 valves].

FIGURES 13–15. Limatula sp. 13—External view, IBUFRJ 17800; 14—Internal view, IBUFRJ 17800; 15—Hinge,
IBUFRJ 17800. Fig. 16. Limatula laminifera (Smith, 1885), External view, IBUFRJ 17808. Fig. 17–19: Limea lirata
Allen, 2004. 17—External view, IBUFRJ 17805; 18—Internal view, IBUFRJ 17805; 19—Hinge, IBUFRJ 17805. Scale
bars: all 250µm.

Remarks: The main diagnostic characters are: (a) the axial ribs, which are evenly spaced but restricted to
the mid-ventral edge of the valve (a view from inside is instructive, because the edge of the valve is crenulated
only in this region and smooth in the remaining part); (b) there is an internal axial fold at the middle of the
shell, barely visible externally; (c) the presence of a very large and smooth larval shell, not observed in any

54 · Zootaxa 1940 © 2008 Magnolia Press OLIVEIRA & ABSALAO

other species; and (d) there are very fine concentric striae that form small nodules (never spines nor scales)
where they cross the axial ribs. We strongly suspect that this may be an unnamed species, but decided to wait
for additional material to describe it.

Limatula laminifera (Smith, 1885)

Figure 16

Lima (Limatula) laminifera Smith, 1885: p. 293, pl. XXIV, fig. 7, 7a.
Limatula laminifera: Stuardo, 1968: p. 226. pl. 14, fig. 65; Abbott, 1974: p. 454; Allen, 2004: p. 2616. fig. 29–32, 46b.

Description: Shell fragile, white, small (height mean = 2.15mm ± 0.17 s.d.; range 2.04–2.40mm; n=4),
height/length ratio 1.37, equivalve, slightly inequilateral, moderately inflated. Ornamented with very fine,
irregularly spaced concentric striae and slender concentric lamellae, which become more foliaceous toward
the ventral margin. Internally with two thickened axial ridges, which correspond to two internal grooves in the
opposite valve. Auricles relatively small, subauricular sinus hardly visible. Dorsal margin straight and little
inclined from antero-posterior axis. Prodissoconch barely visible (mean 95.00μm ± 5.77 s.d.; range 90–
100μm; n=4), interdissoconch not distinguishable. Umbones prominent and triangular. Hinge plate narrow,
edentulous, with an elongated triangular ligamental pit.
Geographic range: Western North Atlantic, Florida to the West Indies, off the Azores, off Fernandina,
off Sombrero Island, off Culebra Island. Depths ranging between 137 to 1590 meters. Adapted from Smith
(1885), Abbott (1974), Allen (2004). Campos Basin (this study) represents the first record of this species for
the Brazilian coast and for the southeastern Atlantic Ocean.
Material examined: IBUFRJ 17808, off Bacia da Foz do Amazonas (03º25 'N, 48º03' W, 700 m),
14.XII.05, [1 valve]; IBUFRJ 17823, off Bacia da Foz do Amazonas (03º25 'N, 48º03' W, 700 m), 14.XII.05,
[4 valves].
Remarks: Both the original description of L. laminifera (Smith 1885) and a later redescription (Allen
2004) pointed out the near absence of axial sculpture, except for very faint lirae that may occur on the central
portion of the valves. In fact, the axial sculpture is barely visible in our specimens.

Limea Bronn, 1831

Type species: Ostrea strigilata Brocchi, 1814

Description: Shell small, white, usually fragile, ovate or suborbicular, slightly inflated, concentric striae or
even lamellae, axial ornamentation present, beaks prominent, auricles small, margins crenate, hinge with
series of short teeth on either side of resilifer.

Limea lirata Allen, 2004

Figures 17–19

Limea lirata Allen, 2004: p. 2622, fig. 36, 37, 46c.

Description: Shell very fragile, translucent, small (height mean = 1.15mm ± 0.07 s.d.; range 1.10–1.20mm;
n=2), height/length ratio 1.28, slightly inequilateral, little inflated, oval outline, ornamented with numerous
and quite slender concentric lines that are positioned close to each other and cover the entire shell, though the

LIMATULA AND LIMEA FROM BRAZILIAN DEEP Zootaxa 1940 © 2008 Magnolia Press · 55
lines are somewhat less evident toward the dorsal region. Axial ornamentation absent, fine internal axial ridge
and groove on both valves, and not perceptible on the external side. Auricles small, subauricular sinus absent,
postero and antero-dorsal margin curved in the hinge region, ventral margin strongly rounded. Prodissoconch
smooth (mean 153.00μm ± 3.00 s.d.; range 150–156 μm; n=2), interdissoconch not distinguishable. Umbones
prominent, hinge plate slender, ligamental pit rounded.
Geographic range: Restricted to North American Basin, at 2095 meters. Adapted from Allen (2004).
Campos Basin (this study) represents the southernmost and the shallowest (1044m) record of this species,
with the first record for Brazilian coast and the southeastern Atlantic Ocean.
Material examined: IBUFRJ 17805, off Bacia de Campos (21°52'43" S, 39°40'42" W, 1941 m),
26.VI.03, [2 valves].
Remarks: Although this is a rare species in our samples, with only three valves, the characteristic con-
centric ornamentation with the very conspicuous fine internal axial ridge and groove on both valves makes
this a distinct species.
Previously restricted to the North American Basin by Allen (2004) who collected 171 specimens from
only one station; since then, this species had not been cited. Our record, the first for the Brazilian and South
American coast, expands the area of occurrence to the South Atlantic.
The characterization of the genus Limea includes some characters, such as axial ornamentation and mar-
ginal crenulation, which are absent in L. lirata. But, since, as mentioned earlier, the definition of the limid
subgroups may be obscure and they probably do not form natural groups and even overlap one another, we
prefer not relocate this species now, avoiding the increase of unwarranted taxonomic mistakes.

TABLE 1. Diagnostic characters of Brazilian deep water Limatula and Limea. The symbol “_” means absence.

L. louiseae L. confusa L. domaneschii Limatula sp. L. laminifera Limea lirata

Mean shell height 1.61 ± 0.38 3.28 ± 2.06 1.07 ± 0.15 1.10 ± 0.14 2.15 ± 0.17 1.15 ± 0.07
(x mm ± s.d.)
Height/Length ratio 1.53 1.41 1.43 1.47 1.37 1.28
Prodissoconch 109.00 ± 7.38 177.50 ± 16.69 170.00 ± 5.00 122.50 ± 7.50 95.00 ± 5.77 153.00 ± 3.00
(x μm ± s.d.)
Interdissoconch _ 805.00 ± 33.38 _ _ _ _
(x μm ± s.d.)
Mean convexity 1.05 ± 0.29 2.44 ± 1.52 0.53 ± 0.12 0.30 ± 0.14 1.13 ± 0.05 0.60 ± 0.00
(x mm ± s.d.)
Ligamental pit Triangular broad, triangular Rounded broad, triangular elongated, trian- Rounded
gular
Depth range 1330–4793m 60–2650m 1044m 1906m 137–1590m 1941–2095m
Concentric ornamen- evenly spaced fine striae evenly spaced irregularly concentric lamel- numerous
tation spaced cordlets lae slender lines
Axial ornamentation _ Dominant _ strong at the ven- _ _
tral margin
Subauricular sinus _ Present _ _ hardly visible _
Microscopic diago- Present _ _ _ _ _
nal striae

Discussion

Allen (2004) reported the variability of shell characters but stressed the importance of shell ornamentation to
distinguish species. We agree with him, but unfortunately even the axial ribs are sometimes very variable and
a secure identification based only on empty shells may be a difficult task. For example, L. margaretae is said

56 · Zootaxa 1940 © 2008 Magnolia Press OLIVEIRA & ABSALAO

to have a central area devoid of axial ribs, but the SEM photo (Allen, 2004, 2630p., fig 45b) shows incipient
axial ribs, suggesting to us that a gradient of expression of this character could be present.
In an ideal situation, anatomy should be associated with conchological analysis, but this is hardly possible
because most specimens in this group are obtained only as separated valves or empty shells. Despite the limi-
tations of shell characters alone in appropriately describing species, it is noteworthy that, among Atlantic
Limatula species, only L. laminifera, L. lirata and L. louiseae share the concentric dominating pattern of shell
sculpture of Limatula domaneschii n.sp. and they possibly form a monophyletic group.
Among the six species here reported, one of Limea and five of Limatula, L. laminifera and L. lirata were
not previously reported from Brazil and one new species, Limatula domaneschii, is here described. This
shows the scarcity of faunal surveys in deep waters and the efforts that should be made at basic research
before broad biogeographical analyses can be performed.
In Table 1 we summarize the main diagnostic characters of the deep water species discussed herein.

Acknowledgements

We are grateful to Dr. John Allen for a very useful exchange of ideas, bibliography support and the type pho-
tos of Limatula margaretae, to Dr. Franklin Noel dos Santos who kindly gave us the specimens of Limatula
laminifera, to CENPES (Centro de Pesquisas da Petrobras (Brazilian Petroleum Co.)) for the SEM photo-
graphs and, finally, to Dra. Paula Mikkelsen and one anonymous reviewer for their critique and suggestions
that improved this manuscript.
This study was partially supported by CNPq (Conselho Nacional de Desenvolvimento Científico e Tec-
nológico) from Brazilian government and FAPERJ (Fundação de Amparo a Pesquisa do Estado do Rio de Jan-
eiro) from State of Rio de Janeiro government.

References

Abbott, R.T. (1974) American Seashells. 2nd.Van Nostrand Reinhold Co., New York, 411 pp.
Allen, J.A. (2004) The Recent species of the genera Limatula and Limea (Bivalvia, Limacea) present in the Atlantic, with
particular reference to those in deep water. Journal of Natural History, 38, 2591–2653.
Clarke, A.H. Jr. (1974) Molluscs from Baffin Bay and the northern Atlantic Ocean. Publications in Biological Oceanog-
raphy, National Museums of Canada. 7, 1–23.
Fleming, C.A. (1978) The bivalve mollusc genus: Limatula: a list of described species and a review of living and fossil
species in the southwest Pacific. Journal of the Royal Society of New Zealand. 8, 17–91.
Gilmour, T.H.L. (1974) The structure, ciliation, and function of the lips of some bivalve molluscs. Canadian Journal of
Zoology. 52, 335–343.
Giribet, G. & Wheeler, W. (2002) On bivalve phylogeny: a high-level analysis of the Bivalvia (Mollusca) based on com-
bined morphology and DNA sequence data Invertebrate Biology. 121(4), 271–324.
Jarnegren, J. & Altin, D. (2006) Filtration and respiration of the deep living bivalve Acesta excavata (J.C. Fabricius,
1779) (Bivalvia; Limidae). Journal of Experimental Marine Biology and Ecology. 334(1), 122–129.
Jarnegren, J.; Rapp, H.T. & Young, C.M. (2007) Similar reproductive cycles and life history traits in congeneric limid
bivalves with different modes of nutrition. Marine Ecology-An Evolutionary Perspective. 28(1), 183–192.
Lodeiros, C. & Himmelman, J. (1999) Reproductive cycle of the rock scallop Lima scabra (Pterioida: Limidae) and its
association with environmental conditions. Revista de Biología Tropical. 47(3), 411–418.
Mikkelsen, P.M., & Bieler, R. (2003) Systematic revision of the western Atlantic file clams, Lima and Ctenoides
(Bivalvia: Limoida: Limidae). Invertebrate Systematics. 17(5), 667–710.
Morton, B.S. (1979) A comparison of lip structure and function correlated with others aspects of the functional morphol-
ogy of Lima lima, Limaria (Platilimaria) fragilis, and Limaria (Platilimaria) hongkongensis sp. nov. (Bivalvia:
Limacea). Canadian Journal of Zoology. 57(4), 728–742.
Morton, B.S. (2000) The pallial eyes of Ctenoides floridanus (Bivalvia: Limoidea). The Journal of Molluscan Studies.
66(4), 449–455.

LIMATULA AND LIMEA FROM BRAZILIAN DEEP Zootaxa 1940 © 2008 Magnolia Press · 57
 Page, T.J. & Linse, K. (2002) More evidence of speciation and dispersal across the Antarctic Polar Front through molec-
ular systematics of Southern Ocean Limatula (Bivalvia: Limidae). Polar Biology. 25(11), 818–826.
Rios, E.C. (1994) Seashells of Brazil, 2nd. Fundação Universidade Federal do Rio Grande, Rio Grande, 368 pp.
Smith, E.A. (1885) Report on the Lamellibranchiata collected by H.M.S. Challenger during the years 1873–76. Chal-
lenger Reports, Zoology. 13, 1–341, 1–25 pls.
Steiner, G. & Hammer, S. (2000) Molecular phylogeny of the Bivalvia inferred from 18S rDNA sequences with particu-
lar reference to the Pteriomorphia. In: Harper, E.M.; Taylor, J.D. & Crame, J.A. (Eds.), The evolutionary biology of
the Bivalvia. The Geological Society of London, London, pp. 11–29.
Stuardo, J. (1968) On the phylogeny, taxonomy and distribution of the Limidae (Mollusca: Bivalvia). PhD thesis, Har-
vard University, Cambridge, MA, USA, 327 pp.
Stuardo, J. (1982) A new species of Ctenoides from the central Atlantic (Bivalvia: Limidae). Boletín de la Sociedad de
Biología de Concepción. Chile, 53, 145–149.
Waller, T.R. (1998) Origin of the molluscan clas Bivalvia and a phylogeny of major groups. In: Johnston, P.A. & Hag-
gart, J.W. (Eds), Bivalves: an Eon of Evolution; Paleobiological Studies Honoring Norman D. Newell. University of
Calgary Press, Calgary, 1–45 pp..

58 · Zootaxa 1940 © 2008 Magnolia Press OLIVEIRA & ABSALAO

View publication stats