Nuculidae (Bivalvia) from King George Island (South Shetland Islands) with the description of a new species of Ennucula. - Iberus 43(1): 83-92

Abstract

During the 31 st Soviet Antarctic Expedition in 1986/87, two species of nuculids have been found in Ardley Cove, a bay on King George Island, one of which is new to science and is described herein: Ennucula ardleyana n. sp. This species was found at two stations alive as single individuals in the intertidal zone and in a water depth of 100 meters. Nucula falk-landica Preston, 1912, the second species, is already known from this area. It was found at six stations with soft bottom and coarse sediment in water depths between 35 and 90 metres in Ardley Cove and Fildes Strait (between King George Island and Nelson Island).

Full text

Nuculidae (Bivalvia) from King George Island (South
Shetland Islands) with the description of a new species of
Ennucula
Nuculidae (Bivalvia) de la Isla Rey Jorge (Islas Shetland del Sur) con
la descripción de una nueva especie de Ennucula
Michael L. ZETTLER1& Andreas BICK2
urn:lsid:zoobank.org:pub:956D2D4E-ADE5-4829-BE50-264C58F3CFC4
Recibido el 30-IX-2024. Aceptado el 18-XI-2024
ABSTRACT
During the 31st Soviet Antarctic Expedition in 1986/87, two species of nuculids have been
found in Ardley Cove, a bay on King George Island, one of which is new to science and is
described herein: Ennucula ardleyana n. sp. This species was found at two stations alive as
single individuals in the intertidal zone and in a water depth of 100 meters. Nucula falk-
landica Preston, 1912, the second species, is already known from this area. It was found at
six stations with soft bottom and coarse sediment in water depths between 35 and 90
metres in Ardley Cove and Fildes Strait (between King George Island and Nelson Island).
RESUMEN
Durante la 31ª Expedición Antártica Soviética en 1986/87, se encontraron dos especies de
nucúlidos en Ardley Cove, una bahía de la isla Rey Jorge, una de las cuales se describe como
nueva para la ciencia: Ennucula ardleyana n. sp. Esta especie se encontró viva en dos esta-
ciones como individuos aislados en la zona intermareal y a una profundidad de 100 metros.
Nucula falklandica Preston, 1912, la segunda especie, ya se conoce en esta zona. Se encon-
tró en seis estaciones con fondo blando y sedimento grueso en profundidades de entre 35 y
90 metros en Ardley Cove y el estrecho de Fildes (entre la Isla Rey Jorge y la Isla Nelson).
KEY WORDS: Mollusca, Southern Ocean, Antarctica, King George Island.
PALABRAS CLAVE: Mollusca, Océano Austral, Antártida, Isla Rey Jorge.
INTRODUCTION
Species of Nuculidae Gray, 1824
(Bivalvia: Protobranchia) live in all
oceans of the world from intertidal to
abyssal zones, mostly in silty mud
(H
UBER
2010). Presently, about 170 valid
recent species in 11 genera are known
1Leibniz Institute for Baltic Sea Research (IOW), Warnemünde, Germany, michael.zettler@io-warnemuen-
de.de. Orcid ID: https://orcid.org/0000-0002-5437-5495
2Universität Rostock, Institut für Biowissenschaften, Allgemeine & Spezielle Zoologie, Universitätsplatz 2, D-
18055 Rostock, Germany; bick-andreas@t-online.de. Orcid ID: https://orcid.org/0000-0003-2356-6106
© Sociedad Española de Malacología Iberus, 43 (1): 83-92, 2025
83

(M
OLLUSCA
B
ASE EDS
. 2024). The genus
Nucula Lamarck, 1799 contains 86 recent
species (slightly more than 50 % of all
species of this genus) accepted as valid
(M
OLLUSCA
B
ASE EDS
. 2024). With 48
species, the genus Ennucula Iredale,
1931 also makes a significant contribu-
tion to the total number of species.
South of the Polar Front, only repre-
sentatives of the genera Nucula, Pronocula
and Ennucula are known in recent times
(W
HITTLE E T AL
. 2012). The record of
nuculid species in Antarctic waters is rel-
atively rare (P
OWELL
1965; D
ELL
1972;
L
INSE ETAL
. 2006). In particular, the scarcity
or absence of nuculid species in otherwise
quite extensive studies on the Antarctic
molluscs indicates their relative rarity
(N
ICOL
1966; E
GOROVA
1982, 1984; A
RNAUD
ET AL
. 1986; M
ÜHLENHARDT
-S
IEGEL
1989;
H
AIN
1990; A
BSHER
& F
EIJÓ
1998; N
ARCHI
ET AL
. 2002; A
LDEA
& T
RONCOSO
2008;
G
ORDILLO ET AL
. 2017; O
SORNO
-A
RANGO
& C
ANTERA
-K
INTZ
2021).
In contrast to the two abyssal species
Nucula austrobenthalis Dell, 1990 and
Pronucula notobenthalis (Thiele, 1912),
only Nucula falklandica Preston, 1912 is a
relatively common species, especially in
the Malvinas/Falkland Islands, South
Shetland Islands, South Orkney Islands
and Antarctic Peninsula (E
NGL
2012).
Other nuculid species do not reach the
Antarctic and are restricted to the more
distant peripheral regions such as the
Strait of Magellan, the Malvinas/Falk-
land Islands, South Georgia or the Ker-
guelen Islands, such as Nucula kerguelen-
sis Thiele, 1912, Nucula pisum G. B.
Sowerby I, 1833, Ennucula puelcha (A.
d’Orbigny, 1842), Ennucula grayi (A.
d’Orbigny, 1846), Ennucula eltanini Dell,
1990 or Ennucula georgiana Dell, 1964
(see T
HIELE
1912; D
ELL
1964, 1990; L
INSE
1999; V
ILLARROEL
& S
TUARDO
1998; F
OR
-
CELLI
2000; V
ALENTICH
-S
COTT ET AL
.
2020; Z
ELAYA
2005).
MATERIAL AND METHODS
The nuculids reported in this study
were collected by the second author in
the eastern part of the Fildes Peninsula
on King George Island, South Shetland
Islands, near the Russian Antarctic
station ‘Bellingshausen’ during the 31st
Soviet Antarctic expedition (1985 to
1987). All stations were located in
Maxwell Bay. Water entering this bay
originates from the Weddell Sea by east
wind drift or from the Bellingshausen
Sea by west wind drift, depending on
seasonal water circulation and prevail-
ing wind direction (G
ORDON
& N
OWLIN
1978). Water temperature and salinity
were not measured but are likely similar
those found by T
IAN ET AL
. (2015). Water
temperature in Ardley Cove was there
in the range 0.72-1.36°C (mean: 1.02°C)
and the salinity from 33.97 to 34.14.
The Fildes Strait, one of the sampling
areas, is characterised by an extremely
strong, tide-dependent current. The
bottom of this area consists mainly of
hard substrate, stones and rubble. The
sediment of Ardley Cove, the second
sampling area, consists of fine sand and
silt, and in the intertidal zone (the tidal
range is about 1 to 1.5 metres), third sam-
pling area, there are boulder fields and
littoral pools with rocky bottoms, some of
which are covered with brown algae. All
samples, except those from the intertidal
zone, were collected using a dredge with
an edge length of 0.3 m. The contents
were sieved through a sieve with a mesh
size of 0.5 mm. Samples from the inter-
tidal zone were taken during low tide
using a plastic tube corer (5 cm2). This
method is extensively described by B
ICK
& A
RLT
(2013). These samples were
washed through a sieve with a mesh size
of 0.2 mm. All samples were fixed in 4 %
borax-buffered formalin.
The molluscs from these samples
were hand-sorted under a low-magnifi-
cation binocular (Zeiss Discovery.V8)
and colour photographs were taken of
selected samples at IOW using a micro-
scope camera (Axiocam 105 color).
High-resolution imaging was done on
selected samples using the Scanning
Electron Microscope at IOW (FESEM
Zeiss Merlin VP compact with Gemini I
electron column; acceleration voltage 5-
15 KV; 5-axis stage with 3-70° tilt func-
tion; detectors: SE, VPSE, InLens SE,
Iberus, 43 (1), 2025
84

BSD; Magnification: 12-2.000 000). The
SEM samples were coated with Iridium
or Gold, by using a Cressington sputter
coater for high resolution imaging.
The holotype and paratypes of the
newly described species are retained in
the Senckenberg Museum, Frankfurt am
Main (SMF). The material of Nucula falk-
landica) is kept in the reference collection
at the Leibniz Institute for Baltic Sea
Research (IOW), Warnemünde,
Germany.
Z
ETTLER
& B
ICK
: Nuculidae from King George Island (South Shetland, Antarctica)
85
SYSTEMATICS
Remarks: Our material agrees well
with the description by D
ELL
(1990). The
maximum shell length is 3.4 mm. The
shell surface is sculptured with growth
folds and radial ribs. The prodissoconch
has a length of 360 µm. The perios-
tracum is yellowish-brownish and
sometimes appears covered with ferrug-
inous deposits. The hinge bears 7-8
anterior and 4-5 posterior teeth. The
chondrophore is strong, trapezoidal and
directed forwards.
D
ELL
(1964) used material that exists
in the British Museum (Natural History)
under the incorrect designation Nucula
pisum G.B. Sowerby I, 1833 from the
Malvinas/Falkland Islands, noting that
the type material cannot be found, the
description by P
RESTON
(1912) is vague
and inaccurate and the illustration of
the outer shell view is not good. A
syntype of the species is housed at
Museum of Natural History in Paris
(MNHN-IM-2000-38316). Thankfully,
photos were taken at our request, which
allowed us to clearly confirm our identi-
fication of N. falklandica.
E
NGL
(2012) illustrated Nucula
minuscula Pfeffer, 1886 sensu Melvill &
Standen, 1907 (NMS 1921.143.686)
sampled on the South Orkney Islands
and kept in the National Museum Scot-
land, and identified it as N. falklandica.
He also illustrated N. falklandica from
the 26th Soviet Antarctic Expedition
from King George Island, housed in the
Zoological Museum Hamburg.
R
AUSCHERT
(1991) published his find-
ings from the 26th (1980-1982) and 30th
(1984-1986) Soviet Antarctic Expedition
from Ardley Cove near King George
Island. The provenance of these records
is almost exactly the same as that of the
present study. At that time, he detected
the species at 12 stations in water depths
between 5 and 70 metres. In the present
study we observed this species between
35 and 70 m.
Class B
IVALVIA
Linnaeus, 1758
Family N
UCULIDAE
Gray, 1824
Genus Nucula Lamarck, 1799
Nucula falklandica Preston, 1912 (Figs. 1-10)
Nucula falklandica Preston, 1912: 637, pl. 21, Fig. 3; C
ARCELLES
& W
ILLIAMSON
1951: 322; P
OWELL
1960: 169; Powell 1965: 367; D
ELL
1964: 139, Figs. 1-17; D
ELL
1990: 5, Figs. 8-9; R
AUSCHERT
1991:
14; V
ILLARROEL
& S
TUARDO
1998: 127, Figs. 1, 27-28, 68-69, 102-103; E
NGL
2012: 37, pl. 1, fig. 2;
Z
ELAYA
2015: 254
Not: L
INSE
1997: 45 (as Nucula sp. with clear concentric-radial network structure); L
INSE
1999
(based on L
INSE
1997): 404; L
INSE
2002: 115, pl. 17, Figs. 9.1.2 (1-5)
Material examined: South Shetland Islands, King George Island • 7 individuals; 62.2332°S, 58.9985°W;
70 m; 30 Jan. 1986; Fildes Strait, Stn. 7, dredge. • 6 individuals; 62.2000°S, 58.9472°W; 50 m; 9 Jan.
1987; Ardley Cove, Stn. 27, dredge. • 6 individuals; 62.2032°S, 58.9410°W; 35 m; 13 Jan. 1987; Ardley
Cove, Stn. 30, dredge. • 4 individuals; 62.2032°S, 58.9395°W; 40 m; 7 Feb. 1987; Ardley Cove, Stn.
40, dredge. • 1 individual; 62.2032°S, 58.9397°W; 50 m; 1 Mar. 1987; Ardley Cove, Stn. 56, dredge.
• 10 individuals; 62.2041°S, 58.9295°W; 90 m; 7 Mar. 1987; Ardley Cove, Stn. 58, dredge.

The records by L
INSE
(1997, 1999 and
2002) for the Magellan region do not refer
to Nucula falklandica. The description pro-
vided and the images given in L
INSE
(2002) clearly show a different Nucula
species, probably Nucula pseudoexigua Vil-
larroel & Stuardo, 1998, although the dis-
tinction of that species from Nucula carlot-
tensis Dall, 1897 is still unclear (see
V
ALENTICH
-S
COTT ET AL
. 2020).
Iberus, 43 (1), 2025
86
(Right page) Figures 1-10. Nucula falklandica Preston, 1912, from King George Island, South
Shetland Islands. 1, 2: left valve, Fildes Strait, Stn. 7, depth 70 m (length 2.1 mm); 3: posterior
view of a complete shell, same locality (height 3.3 mm); 4: right valve, same locality (length 2.9
mm); 5: hinge of a right valve, same locality; 6, 7: specimen from Ardley Cove, Stn. 27, depth 50
m, views from right valve and dorsal side (length 3.1 mm); 8, 9: exterior of right valve and interior
of left valve, same locality (length 2.6 mm); 10: hinge of another left valve, same locality.
Diagnosis: Small species of ~1.8 mm,
with posterior umbo. Umbonal angula-
tion of ~120°. Ligament almost horizon-
tal. Hinge with 4 anterior and 3 poste-
rior teeth.
Description: Small elongated shell,
length up to 1.8 mm; height up to 1.3
mm (holotype length 1.5 mm, height 1.1
mm), equivalve, subequilateral,
rounded triangular outline, umbo poste-
riorly; surface whitish, transparent, with
light yellowish glossy periostracum.
Prodissoconch: Flattened, with
nearly circular outline, smooth; limit
with dissoconch clear; length between
184 and 190 µm and in height between
143 and 146 µm (Figs. 17, 18).
Dissoconch: Globose, subtruncated
in outline; pointed protruding umbo
posterior of midline inclined posteri-
orly; convex dorsal- and ventral
margins, prolonged anteriorly (Figs. 11,
12). Surface only sculptured by fine
irregular commarginal growth stages
and numerous fine growth lines. Lunula
and escutcheon inconspicuous (Figs. 15,
16). Internally smooth and glossy with
sharp margin (Figs. 13, 14); ventral
margin smooth. Anterior and posterior
adductor muscle scars not visible.
Pallial line without sinus. Posterior and
anterior parts of hinge plate with 3-4
strong teeth, rectangular in cross-
section, projecting vertically from hinge;
teeth near umbo very short. Resilifer
elongated-trapezoid (Fig. 15).
Ecology: As only 3 individuals were
found in 2 very different habitats, little
can be said about the ecology of this
species. One animal was found in inter-
tidal shallow water between stones. Two
animals were found in a water depth of
100 m with muddy sediment. The differ-
ent sediments and depths could be an
Genus Ennucula Iredale, 1931
Ennucula ardleyana n. sp. (Figs. 11-18)
urn:lsid:zoobank.org:act:D27A5879-F148-4891-8CFF-68581EA9F25D
Type material. Holotype: South Shetland Islands, King George Island, 1 live-collected specimen
(Figs. 11-12); 62.2025°S, 58.9187°W; 100 m; 28 Feb. 1986; Ardley Cove, Stn. 12, dredge; SMF 367275.
Paratypes: • 1 live-collected specimen (Figs. 13-16); same data as holotype; SMF 367276. • 1 live-
collected specimen (Figs. 17-18); 62.1969°S, 58.9357°W; Intertidal zone; 12 Feb. 1987; Ardley Cove,
Stn. 47, core; SMF 367277.
Type locality: South Shetland Islands, King George Island, 62.2025°S, 58.9187°W; 100 m; 28 Feb.
1986; Ardley Cove, Stn. 12, dredge.
Etymology: The name refers to the type locality in Ardley Cove, King George Island, South Shet-
land Islands.

Z
ETTLER
& B
ICK
: Nuculidae from King George Island (South Shetland, Antarctica)
87
Figuras 1-10. Nucula falklandicaPreston, 1912, de la Isla del Rey Jorge, Islas Shetland del Sur. 1, 2:
valva izquierda, estrecho de Fildes, Stn. 7, profundidad 70 m (longitud 2,1 mm); 3: vista posterior de
una concha completa, misma localidad (altura 3,3 mm); 4: valva derecha, misma localidad (longitud
2,9 mm); 5: charnela de una valva derecha, misma localidad; 6, 7: ejemplar de Ardley Cove, Stn. 27,
profundidad 50 m, vistas de la valva derecha y del lado dorsal (longitud 3,1 mm); 8, 9: exterior de la
valva derecha e interior de la valva izquierda, misma localidad (longitud 2,6 mm); 10: charnela de
otra valva izquierda, misma localidad.
12
3
45
678
910 1 mm
1 mm

Iberus, 43 (1), 2025
88
Figures 11-18. Ennucula ardleyana n. sp., King George Island, South Shetland Islands. 11, 12: holo-
type, Ardley Cove, Stn. 12, depth 100 m (length 1.5 mm); 13-16: paratype (length 1.8 mm), same
locality as holotype; 17, 18: paratype (length 1.7 mm), Ardley Cove, Lapidary Point, Stn. 47, inter-
tidal zone.
Figuras 11-18. Ennucula ardleyana n. sp., de King George Island, Shetland del Sur. 11, 12: holotipo, Ardley
Cove, Stn. 12, profundidad 100 m (longitud 1,5 mm); 13-16: paratipo (longitud 1,8 mm), misma locali-
dad que el holotipo; 17, 18: paratipo (longitud 1,7 mm), Ardley Cove, Lapidary Point, Stn. 47, zona inter-
mareal.
11
1 mm
12
13 14
15 16
17 18

indication that this species is much
more widespread, but could often be
overlooked due to its small size.
Differential diagnosis: We present a
comparison with the known species of
Ennucula from Sub-Antarctic and adja-
cent territories of South America: the
South Georgian Ennucula georgiana Dell,
1964 and species from the Strait of Mag-
ellan or Malvinas/Falkland Islands,
such as Ennucula puelcha (A. d’Orbigny,
1842), Ennucula grayi (A. d’Orbigny,
1846) and Ennucula eltanini Dell, 1990.
Ennucula georgiana is larger (9-15
mm in length), heavier, dorsally and
ventrally rounded, with posterior
margin subangled about in the middle.
It has 12 anterior and 7 posterior teeth.
A specimen 2.7 mm long pictured at the
Natural History Museum Rotterdam
(RMNH.MOL.463998) already has 7
anterior and 4 posterior teeth. The resil-
ifer is large, triangular and projecting
anteroventrally. However, it is not
certain that the identification is correct,
since the location is Weddell Sea and not
South Georgia. Ennucula georgiana is
known so far from South Georgia (D
ELL
1964; Z
ELAYA
2005).
Ennucula grayi, known from central
Chile, Patagonia (including Strait of
Magellan) and the Malvinas/Falkland
Islands (D
ELL
1964; L
INSE
1997; V
ILLAR
-
ROEL
& S
TUARDO
1998; F
ORCELLI
2000;
Z
ELAYA
2005; V
ALENTICH
-S
COTT ET AL
.
2020), is also distinctly larger (up to 20
mm), subovate, and is a very robust
species. It has 20 anterior and 7 poste-
rior teeth. The resilifer is large, conical
and projecting anteroventrally. Even
very small individuals (1.8 to 2 mm
long) already have 5 or 6 anterior and 3
to 4 posterior teeth. The most striking
difference between E. grayi and the E.
ardleyana n. sp. described here is the
shape of the resilifer (robust conical pro-
jecting from the hinge line into the
mantle cavity, versus elongated trape-
zoid lying on the hinge line) and the
outer shape of the shell, which already
differs from E. ardleyana n. sp. in this
juvenile size. At 211 to 224 µm, the
prodissoconch is larger than that of the
newly described species.
Ennucula puelcha occurs in Brazil,
Uruguay, Argentina, Malvinas/Falk-
land Islands, and the Strait of Magellan
(D
ELL
1964; V
ILLARROEL
& S
TUARDO
1998; F
ORCELLI
2000; P
ASSOS
& M
AGAL
-
HÃES
2011; P
ASSOS ET AL
. 2024).
According to S
IMONE
(2009) and V
ALEN
-
TICH
-S
COTT ET AL
. (2020), this species is
restricted to the Atlantic Ocean from
Brazil to northern Argentina, with
uncertain occurrence in southern
Argentina and Chile. It is also larger
(up to 13 mm); globular. It has 18 ante-
rior and 8 posterior teeth. The main
difference is its robust conical resilifer
and its relatively blunt umbo (S
IMONE
2009).
Ennucula eltanini is only known from
the Strait of Magellan and western
Tierra del Fuego from water depths
between 307 and 544 m (D
ELL
1990;
Z
ELAYA
2005). We examined material
from the Strait of Magellan from depths
between 253 and 462 m. It is a small
species reaching 4.1 mm in length and
3.2 mm in height. The umbos are promi-
nent and the resilifer is small, roundish
and upright. It has 8-10 anterior and 5-7
posterior teeth. Juveniles of E. eltanini
with lengths of 1.5 to 2 mm already had
6 to 8 anterior and 3 to 4 posterior teeth,
which distinguishes it from E. ardleyana
n. sp. With a diameter of 204 to 216 µm,
the prodissoconch was larger than that
of the newly described species.
All species mentioned above are the
only known species of the genus Ennu-
cula from the sub-Antarctic region near
the Antarctic Peninsula (Scotia Sea,
Patagonia (including Strait of Magellan)
and Malvinas/Falkland Islands) and are
more than 1000 km away from King
George Island and separated by the
Bellingshausen and Scotia Sea with
water depth deeper 3000 m. Ennucula
ardleyana n. sp. differs from all of them
by its smaller size, the subtruncated
shape and the much smaller number of
anterior and posterior teeth and the
elongated trapezoid resilifer lying on
the hinge line. Considering the small
size of this species, the relative robust-
ness of the anterior hinge plate is strik-
ing.
Z
ETTLER
& B
ICK
: Nuculidae from King George Island (South Shetland, Antarctica)
89

DISCUSSION with E. elongata, regarding the tooth
formula and the resilifer structure. The
shape of the shell, pointed posteriorly in
E. elongata and rounded in E. ardleyana
n. sp., is clearly different, although the
size is the same. In addition, radial stria-
tions are visible in E. elongata (see R
HIND
& A
LLEN
1992), but not in E. ardleyana n.
sp. As mentioned in the introduction, no
Ennucula species were found during the
numerous Antarctic expeditions, alt-
hough many of them also visited the
study area in Fildes Strait and Ardley
Cove (e.g. R
AUSCHERT
1991; E
NGL
2012).
It is very likely that E. ardleyana n. sp.
was overlooked in earlier studies due to
its small size.
ACKNOWLEDGEMENTS
We would like to thank Sascha
Plewe (Leibniz Institute for Baltic Sea
Research, Rostock) for his help in crea-
ting the SEM images. The Zoological
Museum in Hamburg provided us with
comparative material of Ennucula elta-
nini, E. grayi and E. puelcha, for which
we are grateful to the curator Prof. Dr.
Bernhard Hausdorf. We would like to
thank the curators Dr. Philippe Bouchet
and Pierre Lozouet from the Museum of
Natural History in Paris who, at our
request, took photos of a syntype of
Nucula falklandica.
Iberus, 43 (1), 2025
90
Two species of the Nuculidae were
found in waters near King George
Island in this study: Nucula falklandica
and Ennucula ardleyana n. sp.
The morphology of the studied spe-
cimens of N. falklandica is as reported by
D
ELL
(1990). Our material is 3.4 mm in
maximum length and it has up to 8
anterior and 5 posterior teeth compared
to 7-9 anterior and 5 posterior teeth
reported by D
ELL
(1990). The surface is
sculptured with some growth lines and
clearly visible stripes.
The present study has led to the dis-
covery of a previously undescribed
species and the first recent record of a
species of the genus Ennucula of South
Shetland Islands.
Of the 48 recent species of the genus
Ennucula known worldwide, only very
few are known with shell sizes smaller
than 3 mm (H
UBER
2010). The four sma-
llest are Ennucula interflucta (Marinco-
vich, 1973) from northern Chile with 1.5
mm, the Caribbean Ennucula elongata
(Rhind & Allen, 1992) with 2 mm, the
Atlantic Ennucula granulosa (A. E.
Verrill, 1884) with 2.5 mm and the North
American Ennucula similis (Rhind &
Allen, 1992) with 3 mm. Ennucula inter-
flucta lives in the intertidal zone, the
others in water depths below 500 m. The
greatest similarity can be recognised
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