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THE
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HISTORY
MUSEUM

VOLUME 63 NUMBER2 28 NOVEMBER 1997

‘OY 1997

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© The Natural History Museum, 1997

Zoology Series
ISSN 0968-0470 Vol. 63, No. 2, pp. 93-147

The Natural History Museum
Cromwell Road
London SW7 5BD Issued 28 November 1997

Typeset by Ann Buchan (Typesetters), Middlesex
Printed in Great Britain by Henry Ling Ltd., at the Dorset Press, Dorchester, Dorset

Bull. nat. Hist. Mus. Lond. (Zool.) 63(2): 93-122

The lucinid bivalve genus Cardiolucina | wis
(Mollusca, Bivalvia, Lucinidae): systematics,

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Issued 28 November 1997

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anatomy and relationships | oe BESENTED

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JOHN D. TAYLOR. AND EMILY A. GLOVER Pe Ny r
Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK abe

CONTENTS
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Fossil species of Cardiolucina

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PNG WE GE CIMEMES cece saaees s tian <sch eta cb edsh <aaeazstsnansh etss vere nssedeveascesacooscedenseotnsesseontseestesatcerseesiyaceseisanaaustsvetecessseanastneseasneeseaseancasee 121
PRESTR OTIC CS be ceasece cee eet ee a a ae ey Sc Soe oe Sa ado cence ate sadcainins conus nares orurabsnevecapcnnaesveadsaacshsuaareraasussuhtneserensacvne 121
SYNOPSIS. The marine lucinid genus Cardiolucina (usually known as Bellucina) comprises 11 living species of small, heavily

ornamented, subspheroidal bivalves. The type species of the genus is the Miocene fossil C. agassizi (Michelotti). A new
definition and description of the genus is given. All the living species and the type species are redescribed and illustrated. Two
species, C. australopilula from Western Australia and C. siquijorensis from the Philippines, are described as new. Anatomical
details are given for two species which both contain chemosymbiotic bacteria in the ctenidia. A preliminary phylogenetic
analysis using shell characters and Parvilucina as an outgroup, indicates three clades of Cardiolucina. Nine species are found
in the Indo-W. Pacific and two in the tropical Atlantic. The greatest diversity occurs around Australia where seven species live.
The two Atlantic species are similar to the widespread Indo-Pacific species C. semperiana. The earliest-known species of

Cardiolucina is from the Eocene, but the fossil record is very poor.

INTRODUCTION

Since the discovery of endosymbiotic sulphur-oxidising bacteria
contained in the gills, the biology of bivalves in the family
Lucinidae has attracted much attention (Fisher & Hand, 1984;
Dando, Southward, Southward, Terwilliger & Terwilliger, 1985;
Dando, Southward & Southward, 1986: Southward, 1986; Reid &
Brand, 1986; Distel and Felbeck, 1987; Cary, Vetter & Felbeck,
1989; Fisher, 1990; Reid, 1990). The symbiosis has so far been
recorded in about 25 species of Lucinidae and some species have
been shown to be nutritionally dependent on the association (Le
Pennec, Beninger & Herry, 1995). The discovery and investigation
of the symbiosis has been the key to understanding the many
unusual features of lucinoid morphology and habitat distribution
(Allen, 1958; Reid & Brand, 1986, Reid, 1990). Despite the bur-
geoning biological interest in lucinid bivalves, it is unfortunate
that the systematics of family are not well understood, with many
taxonomic groups, particularly at the supraspecific level, being
rather loosely defined. Most recent comprehensive systematic
treatments of Lucinidae (Chavan, 1969; Bretsky, 1976) both pre-
date the discovery of chemoautotrophy in the family and,
moreover, are based entirely on shell characters As Hickman
(1994) points out, there has been no attempt to incorporate the

© The Natural History Museum, 1997

potential wealth of anatomical and biological characters into sys-
tematic studies. Furthermore, following closer attention to lucinid
biology and the investigation of new habitats it is now being
recognised that there are many undescribed species.

In this paper, we review the species of the lucinid genus
Cardiolucina (better known as Bellucina), which are largely tropi-
cal, small, subspheroidal bivalves found in the Indo-Pacific and
Atlantic Oceans. The impetus for this revision stems from the
discovery of two coexisting lucinids in the shallow waters around the
Abrolhos Islands, Western Australia (Glover & Taylor, 1997). Both
species appeared to be undescribed and one with unusual periostracal
extensions has been assigned to a new genus Rastafaria (Taylor &
Glover, 1997). The other species, which forms a part of this paper,
was evidently a Cardiolucina and seemed similar to C. semperiana
Issel, which is widespread in the Indo-West Pacific. However,
preliminary research indicated that there is much taxonomic confu-
sion within the genus, including the priority of the name Cardiolucina
over Bellucina. This led us to revise and redescribe the type species,
provide a new diagnosis of the genus, to review the Recent species
from the Indo-Pacific and Atlantic Oceans and to describe new
species from Western Australia and the Philippines. We have also
attempted to identify within the genus Cardiolucina those shell and
anatomical characters that might be used in a wider study of lucinid
systematics.

94
ABBREVIATIONS

Institutions: AM — The Australian Museum, Sydney; ANSP —
Academy of Natural Sciences, Philadelphia; BMNH — The Natural
History Museum, London; MNHN — Muséum National d’ Histoire
Naturelle, Paris; IRSNB — Royal Belgian Institute of Natural Sci-
ences; NM — Natal Museum; NMW — National Museum of Wales,
Cardiff; NSMT — National Science Museum, Tokyo; SAM — South
Australian Museum, Adelaide; USNM — National Museum of Natu-
ral History, Washington; UMT — University Museum, Tokyo; WAM
— Western Australian Museum, Perth; ZMA — Zoological Museum,
Amsterdam; ZMC — Zoological Museum, Copenhagen.

Shell measurements: H= height of shell from umbone to ventral
margin, L = length of shell from anterior to posterior, T= tumidity of
shell i.e. maximum convexity measured on a single valve.

SYSTEMATIC DESCRIPTIONS

Family LUCINIDAE Fleming, 1828
Genus CARDIOLUCINA Sacco, 1901

Cardiolucina Sacco, 1901: 89. Published June 30 (see Sacco, 1904;
Marshall, 1991).

Bellucina Dall, 1901: 806. Published August 22nd (see Keen, 1971:
983). Lucina eucosmia Dall, 1901: 806, original designation.

TYPE SPECIES.
nation).

Cardium agassizi Michelotti, 1839 (original desig-

NOMENCLATURE. The species considered in this paper have usu-
ally been assigned to the genus Bellucina Dall, 1901. However, the
Miocene C. agassizi Michelotti, the type species of the genus
Cardiolucina Sacco 1901, has all the characters of Recent Bellu-
cina species (see description and figures of C. agassizi below).
Chavan (1937) recognised this and synonymized the two genera
giving Bellucina priority. Although published in the same year,
Sacco’s work appeared in June 1901 (Sacco, 1904, Marshall, 1991)
and Dall’s in August (see Keen, 1971). Cardiolucina therefore has
priority over Bellucina.

Because the name Bellucina has been commonly used, the con-
cept of this genus has to be considered in some detail. Dall (1901, p.
806) erected Bellucina as a section of the subgenus Parvilucina
within the genus Phacoides. The genus Parvilucina was originally
diagnosed as ‘Shell small, plump, often inequilateral; sculpture
more or less reticulate but not muricate, teeth small, common but all
usually present’ and the Section Bellucina was rather skimpily
distinguished as ‘Dorsal areas and sculpture strong.’ However, Dall
(1901) clearly designated Lucina eucosmia (= L. pisum Reeve) as
the type species and the concept of Bellucina must be based upon
this species. Many authors (e.g. Fischer, 1871, Lamy, 1920; Chavan,
1969; Britton 1972; Bretsky, 1976; Oliver, 1992) have considered B.
eucosmia to be synonymous with Bellucina semperiana (Issel,
1869). Consequently, the characters of the latter species have been
taken to represent Bellucina. Unfortunately, Reeve (1850) mixed
two distinct species in the original description and illustrations of
Lucina pisum which is clear from the syntype material. Reeve’s
figure 66a illustrates a specimen from Port Essington, Northern
Territory, Australia, and figure 66b shows a shell of another species
from Singapore. In the original description of Lucina pisum Reeve
clearly referred to the deeply incised lunule and fenestrate ornament

J.D. TAYLOR AND E.A. GLOVER

which are features of the Port Essington specimens. ‘Shell globosely
cordate, solid, thickly latticed with concentric and radiating ribs, of
which the interstices are pitted, posterior side grooved, anterior
short, lunule small, deeply excavated; whitish.’ For this reason one of
the syntypes from Port Essington has been selected as the lectotype
of Lucina pisum Reeve (see below under description of Cardiolucina
eucosmia). Because the name L. pisum is preoccupied (L. pisum
Sowerby, 1836) then C. eucosmia (Dall, 1901) becomes the valid
replacement name. The other Reeve syntypes from Singapore we
consider to be Cardiolucina semperiana (see below).

DIAGNOSIS. Shells small but solid, usually less than 14mm in
height, subcircular (Height/Length ratio around 1.0), moderately
inflated to subspheroidal (Tumidity/Length ratio between 0.38 and
0.48). Umbones prosogyrate, more or less central. Anterior sulcus
either absent or shallow and narrow. Posterior sulcus always
present and either deeply-indented or shallow. Lunule varies in
size and depth between species, but where visible it is generally
heart shaped. It can be deeply incised and large as in C. eucosmia,
or very small and hidden under the beaks as in C. australopilula.
Posterior dorsal area distinct, either concave or convex, usually
with concentric ribbing. Ornament of radial ribs and concentric
lamellae, relative strength of either varies between species. Con-
centric lamellae often raised and fluted. Intersection with radial
ribs produces a variety of ornament from fenestrate to beaded.
Hinge plate thick, dentition of single anterior and posterior lateral
and two cardinal teethin each valve. In the right valve of some
species one of the cardinal teeth can be either reduced or absent.
Ligament external and varying in length; can extend from beaks to
position of posterior lateral tooth, or may be two thirds of this
distance. Internal ventral margin finely to coarsely crenulate, cor-
responding to position of radial ribs. Often small denticles around
antero- and postero-dorsal margins. Anterior adductor muscle scar
quadrate to elongate, ventral tip only slightly detached from pallial
line. Mature individuals of spheroidal species often have the ad-
ductor muscle scars located on an internal buttress. Posterior
adductor scar short and rounded.

Juvenile shells are thinner, less inflated, more anteriorly extended
with radial ribs often visible from interior of shell. Interior ventral
margin not thickened.

ANATOMY. Based on observations of C. australopilula and C.
semperiana.

Mantle largely unfused except at posterior exhalant aperture.
Mantle margin thick and muscular. Mantle ‘gills’ absent. No fused
inhalant aperture. Exhalant aperture a muscular tube with flared
distal end, which can presumably be protracted. Ctenidia consist of
inner demibranchs only, filaments elongate and thick. Outer lamella
of demibranchs attached to posterior mantle just ventral to the
exhalent aperture. Labial palps are small ridges at edge of lips. Foot
cylindrical and vermiform with a posterior heel. Body wall muscu-
lar, with large hemispherical visceral pouch containing oocytes.
Rectum curves around dorsal side of the posterior adductor muscle
and opens near the posterior aperture.

Many of the anatomical features, such as the thickened inner
demibranchs, reduced labial palps and vermiform foot are similar to
those of other lucinids, although some characters may be less widely
distributed. A distinctive and possibly functionally important feature
is the attachment of the ctenidia to the posterior mantle. It is
uncertain how widely this character is distributed amongst the
Lucinidae. It was first illustrated by Reid & Brand (1986) for
Parvilucina tenuisculpta, but not mentioned by Allen (1958) in his
general survey of lucinoids, or by Morton (1979) for Fimbria and not
present in Rastafaria (Taylor & Glover, 1997). Large hemispherical

ANATOMY AND SYSTEMATICS OF CARDIOLUCINA

visceral lobes or pouches are not present in most Lucinidae (Allen,
1958), but are found in Parvilucina (Reid & Brand, 1986), and
species of Thyasiridae have several pouches (Allen, 1958). Most
Lucinidae seem to possess both inhalant and exhalant posterior
apertures, produced by mantle fusion (Allen, 1958). In Cardiolucina
there is no inhalant aperture, a state found also in Parvilucina (Reid
& Brand, 1986).

Cardiolucina species share three anatomical characters with
Parvilucina tenuisculpta, namely; the attachment of the posterior
ctenidia to the mantle, the hemispherical visceral pouches and the
lack of inhalant apertures. These characters may indicate a possible
relationship, but the distribution of most anatomical characters
amongst the Lucinidae is not well known.

GEOLOGICAL RANGE. Eocene (Lutetian) to Recent.

COMPARISON WITH OTHER GENERA. There has been a tendency to
place small, inflated lucinids with both radial and concentric orna-
ment, rather arbitrarily into a number of genera such as Phacoides,
Parvilucina, Linga and Bellucina. These decisions have usually
been made without reference to the characters of the type species.
For instance, the status of Bellucina in major taxonomic revisions
has varied as follows:

Phacoides (Parvilucina) section Bellucina — Dall, 1901: 806
Phacoides (Bellucina) — Lamy, 1920: 211

Linga (Bellucina) — Chavan, 1937: 205; 1969: 496
Parvilucina (Bellucina) — Britton, 1972: 15

Lucina (Bellucina) — Bretsky, 1976: 272-3

Cardiolucina (as Bellucina) has often been considered as a sub-
genus of Parvilucina Dall, 1901, type species Lucina tenuisculpta
Carpenter, 1864, which ranges from Alaska to Baja California.
Hickman (1994) redescribed and illustrated P. tenuisculpta and two
other west American species. Although they are small, rounded and
fairly inflated, Parvilucina species are thinner shelled and less tumid
(T/L 0.29) than all Cardiolucina (T/L 0.38-48) species. Most
Cardiolucina have a pronounced posterior sulcus, and sometimes a
narrow anterior sulcus, compared to the weak, shallow, posterior
sulcus of Parvilucina. Parvilucina has both radial and concentric
ribs; the radial ribbing consisting of closely-spaced, low riblets,
which are generally weak, but most prominent at the anterior and
posterior of the shell. The internal ventral margin of Parvilucina is
finely denticulate, compared with the coarser crenulations of
Cardiolucina. The lunule in Parvilucina is narrowly lanceolate and
asymmetric with the right valve overlapping the left, compared with
the shorter, heart-shaped lunule of Cardiolucina agassizi. The hinge
plate of Parvilucina is also thinner and the lateral teeth distant from
the cardinals. Although a major reappraisal of the relationships of
these genera is necessary, evidence suggests that the Parvilucina
taxa centred around the type species, represent a clade distinct from
Cardiolucina.

Another genus which should be considered in relation to
Cardiolucina is Radiolucina Britton, 1972, (type species Phacoides
amiantus Dall, 1901) from the Caribbean. Dall placed this species
and the similar West American species L. cancellaris Philippi, 1846
in Bellucina. Later, Keen (1971, p.121) recognised that they were
only superficially similar to the type species of Bellucina and
placed them in the subgenus Pleurolucina Dall, 1901, remarking
that American authors had been slow to recognise Dall’s error.
Bretsky (1976) confusingly gave an extensive diagnosis of the
genus Bellucina based on P. amiantus rather than the type species
Bellucina eucosmia. Britton (1972) had previously compared P.
amiantus with Bellucina semperiana (Issel) which he considered
the type species of Bellucina and recognised that P amiantus

95

represents a distinct and separate lineage within the Lucinidae. For
this reason, he erected a new subgenus Radiolucina. The type
species, R. amianta has a more elongate shell than most Bellucina
species H/L 0.92 (data from Bretsky, 1976, p. 273) compared with
about 1.0 for Cardiolucina species, and the shell is less tumid T/L
0.33 compared with 0.43 for Cardiolucina. Furthermore, R. amianta
has 8-12 broad radial ribs crossed by threadlike concentric lamel-
lae, often with intermediate secondary radial ribs in the ventral two
thirds of the shell. We agree with Keen (1971) and Britton (1972)
that Radiolucina amiantua belongs to a distinct and separate clade
which includes several fossil species from the south western USA.
Therefore, Bretsky’s (1976, p. 272) diagnosis of Bellucina based
on P. amiantus should be rejected.

Some American palaeontologists (Woodring, 1925; Gardner,
1926) used the generic name Cardiolucina for some Miocene luci-
nids with high beaks and lacking radial ribs. These species are now
placed in the genus Cavilinga Chavan, 1937 (see Olsson &
Harbison, 1953 p. 85; Bretsky, 1976 p. 265).

Chavan (1937, 1969) classified Bellucina as a subgenus of Linga
de Gregorio 1885, for which the type species is Lucina columbella
Lamarck, 1818, a Miocene fossil from France. This is a large, very
thick-shelled species with concentric ribs only and a strong posterior
sulcus unlike Cardiolucina.

SPECIES DESCRIPTIONS

Cardiolucina agassizi (Michelotti, 1839)
Fig. 1

Cardium agassizi Michelotti, 1839: 17.

Lucina agassizi — Michelotti, 1847: 404 pl. 4 figs 4,5, 7.

Lucina irregularis Eichwald, 1853: 82-83, pl. 5 fig. 4.

Cardiolucina agassizi (Michelotti) — Sacco, 1901: 89-90, pl. 10 figs
37-39.

Phacoides (Cardiolucina) agassizi (Michelotti) - Cossmann &
Peyrot, 1911: 688-689, pl. 28. figs 83-86.

TYPE MATERIAL. ?Museo di Paleontologia, Rome.

TYPELOCALITY. Not givenby Michelotti, but Sacco (1901) records
several localities in the Miocene of Italy.

DESCRIPTION. Shell small, solid, height to 7.4 mm, subcircular,
mean H/L 1.03, moderately inflated, mean T/L 0.39, tumidity to a
maximum of 2.7 mm on a single valve. Shell inequilateral, ex-
tended anteriorly. Deep posterior sulcus with fine concentric
lamellae. Shallow anterior sulcus. Lunule heart-shaped, small and
shallow. Escutcheon lanceolate with concentric lamellae and a
single strong radial ridge. Exterior sculpture of more than 25 faint,
radial, rounded ribs with narrow interspaces. There are about 20
prominent, concentric lamellae which are often slightly recurved.
Lamellae are extremely variable in thickness, degree of projection,
recurvature and width of interspaces. Hinge plate thick. Left valve
with single posterior lateral tooth, two cardinal teeth and single
anterior lateral. Right valve with single posterior and anterior
laterals and single cardinal. Ligament extending two thirds of the
distance from umbone to posterior lateral tooth. Inner margin with
28-30 crenulations and small denticles along anterodorsal and
posterodorsal margin. Anterior adductor muscle scar elongate, not
buttressed, with pallial line attached near to the ventral tip. Poste-
rior muscle scar ovate. Pallial line indistinct. Shell white.

In the material examined from France we found the concentric
ornament to be extremely variable from separated, thin lamellae to
clusters of thicker lamellae producing a rugose ornament (see Fig.

96

J.D. TAYLOR AND E.A. GLOVER

ox
ts
ox
= *
a
~
7

Fig. 1 Cardiolucina agassizi (Michelotti, 1839), middle Miocene (Langhian), St Jean de Marsacq, Landes, France. (coll Lozouet and Maestrati, MNHN )
a, exterior of right valve; b, exterior of right valve; c, interior of left valve; d, interior of right valve. Scale bars a = 750um, b =700um, c = 1.5mm, d=

500um.

1d). Similar variability of the concentric sculpture is seen in the
Recent species C. crassilirata and C. semperiana.

DISTRIBUTION. As well as the records of Sacco (1901), C. agassizi
is also recorded from the Tortonian of the Aquitaine Basin (Coss-
mann & Peyrot, 1911) and from the middle Miocene Korytnica
Formation of Poland (Friedberg, 1934).

MATERIAL EXAMINED. FRANCE: Middle Miocene (Langhian),
St Jean de Marsacq, Landes, France; ca 100 shells, Lozouet &
Maestrati coll (MNHN).

Cardiolucina australopilula Taylor and Glover, new
species

Figs 2-6

TYPE MATERIAL. Holotype: WAM reg number 1 10—96: H. 13.6, L.
12.8, T. 6.6 mm. Paratypes: WAM 111-96 2 preserved specimens
station 35; AM C.202975 1 preserved specimen, station 23; BMNH
1996086, 1996087, 1996088, 1996089 4 preserved specimens, sta-
tions 1, 33, 36, 37; BMNH 1996090 one dry shell station 36. (Station
data in Glover & Taylor, 1997).

TYPE LOCALITY. 9 km north of Beacon Island, Wallabi Group,
Houtman Abrolhos Islands, Western Australia, 113° 46'E, 28° 23'S,
depth 39m.

SHELL DESCRIPTION. Shell solid, small, height to 13.8 mm,
subcircular in outline, H/L mean 1.03, inflated, orbicular, T/L mean
0.47, tumidity to a maximum of 7.5 mm on a single valve. Shell
slightly inequilateral, extended anteriorly. Very shallow posterior
sulcus. Umbones pronounced, prosogyrate. Lunule extremely small
and shallow. Anterodorsal area slightly convex, heart-shaped in
outline and with fine concentric ribbing. Escutcheon broadly lan-
ceolate, convex and finely ribbed with concentric and a few radial
ribs. Exterior sculpture of about 20 radial ribs, rounded in profile,
separated by flat interspaces. Anteriorly, interspaces are as wide as
the ribs, elsewhere interspaces are about two thirds of the width of
radial ribs. Radial ribs have very fine radial striations (Fig. 3d).
Radial ribs are crossed by 34—36 thin, low, concentric lamellae and
their intersection produces a finely fenestrate ornamentation. Hinge
plate thick, left valve with single posterior lateral tooth, two cardinal
teeth, the anterior of these is robust and hooked (Fig. 3c) and a single
anterior lateral. Right valve with single posterior lateral

ANATOMY AND SYSTEMATICS OF CARDIOLUCINA

97

Fig. 2 Cardiolucina australopilula Taylor and Glover new species, Beacon Island, Wallabi Group, Houtman Abrolhos Islands, Western Australia. a,
Holotype (WAM 110-96), exterior of right valve; b, Holotype, exterior of left valve; c, Paratype (BMNH 1996090), exterior of left valve; d, Paratype
(BMNH 1996090), interior of left valve; e. interior of right valve of juvenile shell, Eagle Bluff, Peron Point, Shark Bay; f, Holotype dorsal area. Scale

bars a—d = 2.0mm, e = 2.5mm, f = 2.0mm.

tooth, two cardinal teeth, the posterior larger than the anterior, and a
single anterolateral. Cardinal teeth sit on a curved, ventrally project-
ing buttress. Ligament elongate, extending from beak to posterior
lateral tooth. Inner margin of shell thickened with 21 crenulations.
Small sinus on posterolateral margin corresponding to the position
of the posterior sulcus. Anterior adductor muscle scar narrow,
elongate and sited on a buttress with ventral third detached from the

pallial line. Posterior muscle scar ovate. Pallial line continuous.
Shell colour white.

Juvenile shells (Fig 2e) less inflated, more anteriorly extended.
External radial ribs visible on interior of shell.

ANATOMY. The mantle margin is thick and fused only at the
posterior exhalant aperture (Fig. 4), and there is no fused inhalant

98

J.D. TAYLOR AND E.A. GLOVER

Fig.3 Cardiolucina australopilula new species. a, Holotype, hinge in the right valve; b. Holotype, hinge in the left valve; c, Paratype, left valve, detail
of cardinal teeth; d, outer shell surface showing radial striations on radial ribs. Scale bars a=2.0mm, b=2.5mm, c=1.0mm, d=500ym.

aperture. The exhalant aperture consists of a muscular tube with a
flared distal end, which is inverted in preserved specimens, but can
presumably be protracted. The ctenidia comprise inner demibranchs
only. These have large, thick filaments which are fused between the
ascending and descending lamellae. To the posterior, the outer
lamellae are attached to the mantle margin just ventral to the
exhalant aperture. The attachment area forms a thin, muscular
septum. The labial palps consist of small folds at the edge of the thin,
ridge-like lips. The foot is cylindrical and vermiform, tapering to a
blunt point and extending posteriorly with a rounded heel. The
visceral mass is covered by a muscular wall with a single, large,
hemispherical visceral lobe on each side. The rectum runs around
the dorsal side of the posterior adductor muscle and opens into the
mantle cavity close to the exhalant tube.

COMPARISON WITH OTHER SPECIES. This species is similar in size,
shape and sculpture to C. quadrata (Prashad, 1932) (Fig. 16) but
differs in the greater prominence of the radial ribs and thinner,
concentric lamellae. C. australopilula is also more tumid and spheri-
cal. This species has been referred to as Bellucina cf. semperiana by
Slack-Smith (1990), but semperiana 1s smaller, with different orna-
mentation and hinge teeth.

HABITAT. Fine, well-sorted, calcareous sand between 30—40m, in
association with other bivalves Amusium balloti, Mimachlamys
australis and Rastafaria thiophila (Glover & Taylor 1997).

DISTRIBUTION. Western Australia, Houtman Abrolhos Islands to

Broome (Fig. 6).
ETYMOLOGY.

MATERIAL EXAMINED. WESTERN AUSTRALIA: Broome (AM
C.309449); Roebuck Bay, Broome (BMNH 1892.1.29.121);
Montebello Islands, Bunsen Channel, N. end of Trimoville I. 20°
23.8'S; 115° 32.43'E 3m (WAM); Barrow Island 20° 53'S, 115° 20'E
(WAM); Exmouth Gulf (WAM 3461-3468); Mauds Landing, N.W.
Cape, 3m (WAM); Turtle Beach, off N.W. Cape (AM C.309456);
Carnarvon (AM C.309455); Dirk Hartog Island, 10-20m (WAM
3475.68); Bernier Islands 24° 53.3'S 113° 17.5'E. (WAM); Shark
Bay, Peron Peninsula (WAM 249.94); Shark Bay, 5 km E.S.E. Cape
Heirisson, 18m (WAM); Shark Bay, Freycinet Estuary (WAM 252—
94). Houtman Abrolhos Islands: N. of Wallabi Islands, 40m (BMNH
acc 2384); Easter Group, 50m (WAM N-2777); Zeewyk Channel,
40m (WAM N-2822); Little North Island, Easter Group, 40m
(WAM); Rat Island, 30m (WAM wet material); East of Good Friday
Bay, 28° 39.5'S — 113° 49.5'E, 40m (WAM).

australopilula meaning “Australian little ball’.

ANATOMY AND SYSTEMATICS OF CARDIOLUCINA

# 1
i hi ai

A

3mm

Fig. 4 Cardiolucina australopilula anatomy. Right mantle and ctenidium
removed. Abbreviations aa — anterior adductor muscle, cme — cut mantle
edge, exa — exhalant aperture, ext — exhalant tube, fet — fused ctenidium
to mantle, ft — foot, Ict — left ctenidium, lp — labial palps, me — mantle
edge, pa — posterior adductor muscle, ppr — posterior pedal retractor
muscle, r — rectum, vl — visceral lobe.

Fig.5 Semidiagrammatic transverse section through the posterior
exhalant area of C. australopilula showing the inverted exhalant tube
and the connection of the ctenidium to the mantle. Abbreviations: cm —
connection between mantle and ctenidium, ct — ctenidium, ea — exhalant
aperture, iet — inverted exhalant tube, Im — longitudinal muscle, me —
mantle edge, n — nerve.

99

Fig. 6 Geographical distribution of Cardiolucina australopilula new
species.

Cardiolucina civica (Yokoyama, 1927)
Figs 7-9

Cardium civica Yokoyama, 1927: 179. pl. 48, figs 3 & 4.

Dentilucina (Bellucina) hedleyi Prashad, 1932: 163-164, pl. 5, figs
17 & 18.

Bellucina civica (Yokoyama, 1927) — Kuroda, Habe & Oyama,
1971: 394, pl. 118 figs 17 & 18.

Lucina (Codakia) semperiana (Issel, 1869) — Melvill & Sykes,
1898: pl. 3, fig.1.

TYPE MATERIAL. C. civica syntypes, University Museum, Tokyo,
CM 24913-24914, see Makiyama (1959); C. hedleyi holotype, I
individual 2 valves (ZMA).

TYPE LOCALTIES. C. civica was described as a Pliocene fossil from
Nagaya and Kakuma in Kaga Province, Japan. C. hedleyi from the
Philippines, Sulu Archipelago, Siboga station 105 (6° 8'N, 121°
19'E), depth 275 m.

NOMENCLATURE. We have examined the holotype of C. hedleyi
from Indonesia, which is very similar to both Recent specimens of
C. civica from Japan and the fossils figured by Yokoyama (1927) and
consider them conspecific.

SHELL DESCRIPTION. Shell solid, small, height to a maximum of
10.6 mm; subcircular, mean H/L 1.02, inflated, mean T/L 0.40,
tumidity to a maximum of 4.5 mm on a single valve. Shell slightly
extended anteriorly. There is a shallow posterior sulcus which has
concentric ribs. Lunule small and shallow. Anterodorsal area weakly
defined. Escutcheon slightly convex and finely ribbed. Exterior
sculpture of numerous (more than 30) fine, slightly rounded, radial
ribs with narrow interspaces. Radial ribs are crossed by many (more
than 25) conspicuous concentric lamellae, which are rounded in
profile and slightly recurved. Left valve with a single posterior
lateral, two cardinal teeth and a single posterior lateral; right valve
with a single posterior lateral, two cardinal teeth, the posterior

J.D. TAYLOR AND E.A. GLOVER

Fig.7 a-d Cardiolucina civica (Yokoyama, 1927) southern Honshu, Japan BMNH 1878.10.16.191 and e-f Dentilucina (Bellucina) hedleyi Prashad,
1932 Holotype, Sulu Archipelago, Philippines. a, exterior of left valve; b, interior of left valve; c, hinge in left valve; d, outer surface of shell showing
concentric and radial ribs; e, Holotype C. hedleyi exterior of left valve; f, Holotype C. hedleyi interior of left valve. Scale bars a=2.5mm, b=1.2mm,
c=500um, d=500um, e=2.0mm, f=2.5mm.

ANATOMY AND SYSTEMATICS OF CARDIOLUCINA

101

Fig. 8 Cardiolucina civica from Andaman Islands. Large specimen illustrated by Melvill and Sykes (1897, fig.1) as Lucina (Codakia) semperiana Issel
(BMNH 1898.4.30.7) a, Right valve exterior; b, Left valve interior. Scale bar a—b = 1.0mm.

cardinal large and the anterior very small. There is a single anterior
lateral tooth. Ligament short, extending from the beak to about half
way to posterior lateral tooth. Inner margin of shell thickened and
with 30 crenulations. Small denticles on anterodorsal margin. Ante-
rior adductor muscle scar elongate and not buttressed, pallial line
attached near ventral tip. Posterior adductor scar small and rounded.
Pallial line continuous. Shell grey white.

Juveniles less inflated and more extended anteriorly. Posterior

sulcus more pronounced and anterodorsal shell margin markedly
convex. Shell thinner with radial riblets visible from interior of shell.

A large specimen (BMNH 1898.4.30.7) from the Andaman
Islands, figured by Melvill and Sykes (1898 pl.3, fig.1) as Lucina
(Codakia) semperiana, has many of the characters of C. civica,
although larger than specimens from other localities. Other speci-
mens in the BMNH from Port Blair, Andamans are similarly
large.

Fig.9 Geographical distribution of Cardiolucina civica (Yokoyama).

102

J.D. TAYLOR AND E.A. GLOVER

Fig. 10 Cardiolucina crassilirata (Tate, 1886), Western Port, Victoria, Australia. BMNH 1963200. a, exterior of left valve; b, interior of left valve. Scale

bars a = 2.0mm, b = 2.5mm.

COMPARISON WITH OTHER SPECIES. C. civica is most similar to C.
crassilirata, but it has a shorter ligament and coarser marginal
crenulations.

HABITAT. Fine sandy and shelly bottoms mainly from shallow
water to 200m, with a few dead shells recorded up to 600 m.

DISTRIBUTION. Andaman Islands, North Western Australia, South
East Asia, Japan, to Fiji (Fig. 9).

MATERIAL EXAMINED. ANDAMAN ISLANDS: 11°37'N,
92°56’E, 357m (ANSP 292239); Port Blair, BMNH 1953.1.30.199—
201; BMNH 1898.4.30.7. AUSTRALIA: Western Australia, N.W.
of Port Hedland 19°13'S, 116°6'E, 271m & 19°24'S, 115°52'E,
238m (AM C.309458 & C.309453); N.W. Beagle Island, 29°43'S,
114°17'E, 274—283m (AM C.309465); N.W. Roebuck Bay, Broome
17°34'S 120°22'E, 188m (AM C.309457). PHILIPPINES: Luzon,
off Matocot Pt W., 370m (USNM 295697); Tayabas Bay, off Tay-
abas Lt, 200m (USNM 297745); Ragay Gulf, off North Burias,
185m (USNM 246286); off Balanja Pt, S.E. Mindoro, 280m (USNM
295581); Adyagan I. E. Masbate,190m (USNM 298134); off Pt
Dumurug, Masbate, 273m (USNM 298227); Linapacan Strait, 90m
(USNM 301994); off Pt Lauis, E. Cebu, 280m (USNM 300493); off
Gigantangan I., N.W. Leyte, 210m (USNM 298317); off Dammi I.,
Sulu Archipelago, 446m (USNM 299036); off Simaluc Island, Tawi
Tawi Is, 600m (USNM 299195); off Jolo, Jolo I., 283m (USNM
294618) & 37m (USNM 294584); 14°O1'N 120°18'E, 192m
(MNHN). CHINA: off Pratas Islands., China Sea, 272m, (USNM
296724). JAPAN: off southern Honshu 34°13'N, 136°13'E, 90—
100m (BMNH 1878.10.16.191); Kumano-Nada, off Kii Peninsula,
93m (NSMT 70534); off Jyogashima I., Miura Peninsula (NSMT
70535); off Goto Islands, 168m (NSMT 56297); off Tsushima
Islands, 112m (NSMT 56298); Usibuka, Anakusa Island (NSMT
70533); Sagami Bay (USNM 708071). IRIAN JAYA: Aoeri I.,
Geelvink Bay, 140 (ANSP 275808, 280332); Japen I., Geelvink
Bay, 30-40m (ANSP 277817, 276638, 279725); Schouten I., 18m
(ANSP 280204). NEW CALEDONIA: 19°07'S 163°22'E, 110-
200m (MNHN); 22°02'S 165°57'E 135-150m (MNHN); Grande

Récif Sud, 22°42'S 157°22'E, 35m (MNHN). FIJI: Viti Levu Bay 0—
5m near shore (USNM 878321).

Cardiolucina crassilirata (Tate, 1887)
Figs 10-11

Lucina crassilirata Tate, 1887a: 6—7, pl. 4 fig. 2.

Codokia crassilirata (Tate) — Cotton & Godfrey, 1938: 206 fig. 218.
Loripes crassilirata (Tate) — Pritchard & Gatliff, 1903: 139.
Bellucina crassilirata (Tate) - Macpherson & Gabriel, 1962: 327.

Fig. 11

Geographical distribution of Cardiolucina crassilirata (Tate).

ANATOMY AND SYSTEMATICS OF CARDIOLUCINA

Linga (Bellucina) crassilirata (Yate, 1887) — Lamprell & White-
head, 1992: pl. 21 figs 128a & b.

HOLOTYPE. South Australian Museum No. D.12957. L = 8.5 mm,
H = 8.3 mm, T = 7.0 mm.

TYPE LOCALITY. Streaky Bay, near Ceduna, South Australia.

SHELL DESCRIPTION. Shell small, solid, height to 8.4 mm, outline
circular, H/L mean 0.98; shell moderately inflated with T/L mean
0.41, tumidity of a single valve to a maximum of 3.4 mm. Shell
slightly inequilateral, extended anteriorly. Shallow sulcus poster-
iorly, with concentric lamellae. Lunule small and shallow. Escutcheon
slightly convex, with fine concentric ribs. Exterior sculpture of more
than 30 fine concentric lamellae, clustered in groups of 4—6 with
broader interspaces between the clusters. Radial sculpture of many
fine ribs visible in the interspaces of the concentric ribs. Hinge plate
narrow; left valve with single posterior lateral, two cardinal teeth and
single anterior lateral. Right valve with a single posterior and
anterior laterals and two cardinal teeth, posterior cardinal larger than
anterior cardinal tooth. Ligament long, extending from beaks to
posterior lateral tooth. Inner margin with around 60 fine crenulations.
Anterior adductor muscle scar, elongate, not buttressed, with ventral
quarter detached from pallial line. Posterior adductor muscle scar
ovate and not buttressed. Pallial line discontinuous. Shell greyish
white.

COMPARISON WITH OTHER SPECIES. See C. civica.

HABITAT. Shallow subtidal to 70m.
DISTRIBUTION. South Australia, Victoria and Tasmania (Fig. 11).

MATERIAL EXAMINED. AUSTRALIA: Victoria, BMNH
1906.9.19.70-2; Western Port (BMNH 1963200); BMNH
1911.12.19.11-12; Western Port (WAM); Western Port (AM
C.303440); Western Port, 15—18m (AM C.309469); Port Fairy,
beach (ANSP 135488); New South Wales, Between Eagle and
Crawfish Rock (AM); South Australia, Spencers Gulf (BMNH
1893.12.12.352-54); Hardwicke Bay (AM C.13463); Neptune L.,
70m (AM); South Australia (AM C.6677); Tasmania, Geography
Strait, 27m (AM C.38180).

Cardiolucina eucosmia (Dall, 1901)
Figs 12-13

Lucina pisum Reeve, 1850 (in part): pl. 11 fig. 66a (non Lucina
pisum Sowerby, 1836: 341).

Phacoides (Bellucina) eucosmia Dall, 1901: 806 (replacement name
for L. pisum Reeve).

Phacoides eucosmia Dall. — Hedley, 1909: 426, pl. 37 fig 16.

TYPE MATERIAL. Lectotype here selected BMNH 1963194/1 (Figs
12a—d). Paralectotypes BMNH 1963194/2-3; 1845.825.210-212;
1963552 (this is C. semperiana see p. 107)

TYPE LOCALITY. Restricted to Port Essington, Northern Territory,
Australia (modified from Reeve, 1850).

NOMENCLATURE. Many authors, for example Fischer (1871), Lamy
(1920), Chavan (1969), Britton (1972), Bretsky (1976) and Oliver
(1992) have considered Lucina semperiana Issel, 1869 a senior
synonym of P. eucosmia. However, Lucina pisum Reeve, for which
Dall gave the replacement name Phacoides eucosmia, is based on a
syntype series composed of two distinct species. For the reasons
given above in the generic section, we consider the specimens from
Port Essington to conform most closely with Reeve’s description

103

and concept of Lucina pisum and one of these has been selected as a
lectotype. The other syntypes from Singapore, now become
paralectotypes and we consider these to be the eastern Indian Ocean
form of C. semperiana (Fig. 21 a & b).

SHELL DESCRIPTION. Shell solid, small, height to 7.8 mm,
subcircular in outline, slightly extended in an anterolateral direction,
H/L mean 1.05; inflated, T/L mean 0.49, tumidity to a maximum of
3.8 mm on a single valve. Posteriorly, there is deep sulcus extending
from hinge to ventral margin. The sulcus and the posterior portion of
the shell has fine concentric ribbing with no radial ribs. The lunule
is ‘U-shaped and deeply incised to the ventral edge of the hinge
plate. Anterodorsal area is slightly convex, ovate in outline with fine
concentric sculpture. Posterior dorsal area broadly lanceolate in
outline, convex, with fine concentric ribbing. Exterior sculpture of
about 12 broad, radial ribs with rounded profile, somewhat flattened
on upper surface. Ribs narrower towards umbos. Interspaces between
ribs deep, narrower than ribs. Radial ribs crossed by 13-14 thin,
low, concentric lamellae which project slightly above radial ribs.
The intersection of concentric and radial sculpture gives a distinctive
fenestrate ornament. Hinge plate thick. Left valve has a single,
strong lateral tooth posteriorly, two thin, cardinal teeth, the posterior
tooth much larger than the anterior. Cardinal teeth are denticulate
(see Fig. 12d). A single anterior lateral lies close to perimeter of the
lunule. Right valve has a single posterior lateral, a large single
cardinal tooth and a single anterior lateral. Both valves have small
denticles around the dorsal margin of valve. Ligament elongate,
extending from beak to posterior lateral tooth. Inner shell margin
thickened, with 12 coarse crenulations. Small sinus on posterior
lateral margin corresponding to position of sulcus. Anterior adduc-
tor muscle scar elongate, sitting on a shallow buttress with the only
the ventral tip detached from the pallial line. Posterior adductor scar
ovate on a shallow buttress. Pallial line is thin and continuous. Shell
colour creamy white.

Juvenile shells (Figs 12 e & f) are less inflated and more extended
anteriorly. The lunule is widely opened in younger stages. Denticles
are absent on the dorsal inner margins. Radial ribs begin narrow and
broaden ventrally between each concentric rib. Radial ribs are
visible on interior of shells.

COMPARISON WITH OTHER SPECIES. Although C. pisiformis, C.
siquijorensis and C. australopilula have a similar fenestrate orna-
ment, C. eucosmia can be easily distinguished from all other species
by the deeply incised lunule.

HABITAT. Low intertidal to 100 m in fine sand and mud. Hedley
(1909) reports dredging the species in the Gulf of Carpentaria from
about 10 m in soft black mud, where it was abundant.

DISTRIBUTION. Northern Australia (Fig. 13).

MATERIAL EXAMINED. AUSTRALIA: Northern Territory and
Queensland, BMNH 1963490, 1963236, 1910.9.28.77-8; W. of
Cape York, 8m (BMNH 1887.2.9.2776-7); Gulf of Carpentaria
(NMW 1955.158); Sweers I. (AM C.75249, 78223); Forsyth I. (AM
C.14892); Horsey River mouth, 9m (AM C.15152), Karumba (AM
C.14963); Mapoon (AM C.14281); off Bountiful Islands (AM);
Gulf of Carpentaria (AM C.100498); mid-Gulf of Carpentaria, 66m
(AM); off Albany River (AM); Karumba Point (AM 1909); Albany
Passage, 8-25m (AM C. 36172); Port Essington (BMNH,; off Point
Charles, Darwin, 27-37 m (AM C.309468); Van Diemen’s Inlet
(NMW 1955.158); Van Diemen’s Inlet (AM C.15273); 100 miles N.
of Croker Island, Arafura Sea, 124m (AM C.309454): 150 km N. of
Coburg Peninsula, 108m (AM); Dinah Beach, Darwin (AM); Cro-
codile Research Station, Manningrida, Arnhem Land (AM);

J.D. TAYLOR AND E.A. GLOVER

Fig. 12 Cardiolucina eucosmia (Dall, 1901) Port Essington, Northern Territory, Australia. Figured specimens a—d are the lectotype of Lucina pisum
Reeve, BMNH 1963194/1. & Figure e-f C. eucosmia, Arafura Sea, Northern Territory, Australia, AM C.309454. a, exterior of right valve; b, exterior of
left valve; c, interior of left valve; d, hinge and lunule in left valve; e, juvenile, interior of left valve; f, juvenile, exterior of left valve. Scale bars a—c =
1.0mm, d = 500um, e = 650um, f = 1.0mm.

ANATOMY AND SYSTEMATICS OF CARDIOLUCINA

Fig. 13 Geographical distribution of Cardiolucina eucosmia (Dall).

Western Australia, Prince Frederick Harbour (WAM); Cape Leveque
(AM); Port Hedland (AM); Direction Island, off Onslow (AM
C.69334).

Cardiolucina pisiformis (Thiele, 1930)
Figs 14-15

Phacoides (Parvilucina) pisiformis Thiele, 1930: 592.

Phacoides (Parvilucina) pisiformis Thiele — Ponder, 1978: 439, pl. 2
figs 3 & 9.

Bellucina pisiformis (Thiele) — Slack-Smith, 1990: 135.

HOLOTYPE. Museum fiir Naturkunde, Berlin. Reg. no. 67726. L=
3 mm, H= 2.9 mm (details from Ponder, 1978).

TYPE LOCALITY. 5 km N.W. of Denham, Shark Bay, Western

Australia, 3 m.

SHELL DESCRIPTION. Shell very small, toa maximum height of 4.4
mm, outline circular, H/L mean 0.97; inflated, T/L mean 0.42,
tumidity of single valve to 2.0 mm. Inequilateral, slightly extended
anteriorly. Shallow posterior sulcus with concentric lamellae. Lunule
heart-shaped, slightly excavated, smooth. Anterodorsal area indis-
tinct. Posterodorsal area with concentric lamellae. Exterior sculpture
of 13-17 low, flattened, radial ribs with broad interspaces. Ribs
broader and more rounded towards anterior of shell. About 20 thin,
concentric lamellae project slightly above radial ribs. Sculpture
distinctly fenestrate in appearance. Hinge plate strong. Left valve
with single posterior lateral, two cardinal teeth, the posterior of these
large and robust, the anterior small. Single anterior lateral tooth.
Right valve with a single, strong, posterior lateral tooth, two cardi-
nals, the anterior large and the posterior very small and a single
anterior lateral. Ligament extends from beak to posterior lateral
tooth. Inner margin with about 13 coarse crenulations, which are
sometimes bifurcate with occasional small denticles in the
interspaces. Small, rounded denticles around anterior and posterior
dorsal margins. Anterior adductor muscle scar elongate, slightly
buttressed with only the ventral tip detached from the pallial line.
Posterior adductor muscle scar rounded. Pallial line continuous.
Shell white.

Juvenile shells (Fig.14 e) are more extended both anteriorly and
posteriorly, with a prominent convex anterodorsal area.

105

COMPARISON WITH OTHER SPECIES. This is a small distinctive
species with an exterior ornament similar to C. eucosmia and C.
siquijorensis. It is smaller, more circular in shape and lacks the
deeply-excavated lunule of C. eucosmia. C. siquijorensis has fewer
and broader radial ribs, a shallower lunule and a different dentition.

HABITAT. Shallow water to 240 m.

DISTRIBUTION. Western Australia (Fig. 15).

MATERIAL EXAMINED. AUSTRALIA: Western Australia, W. of
Rottnest Island, 110m (WAM 262-94); Point Peron Peninsula,
Shark Bay (WAM 257-94, 258-94, 259-94, 260-94,); Carnarvon
(WAM); N. of Homestead Point, Dirk Hartog Island (WAM 250-
94); Ningaloo Reef, N.W. Cape (WAM); Mangrove Bay beach,
N.W. Cape (WAM); Yardie Creek to Tantibiddi, N.W. Cape (WAM
wet material); N.W. of Port Hedland, 112m (AM C.309462):; N.W.
Shelf, W. of Roebuck Bay, 238m (AM C.309464); Broome (ANSP
233737); Northern Territory, N. of Cobourg Peninsula, 55m
(AM); 200km N. of Melville Island, 215m (AM).

Cardiolucina quadrata (Prashad, 1932)
Figs 16-17

Dentilucina (Bellucina) hedleyi var. quadrata Prashad, 1932: 164,
pl. 5 figs 19 & 20.

TYPE MATERIAL. Holotype, one individual two valves, L=10.5
mm, H=10.5 mm. Also 4 paratype valves. (ZMA).

TYPE LOCALITY. Siboga Expedition station 212 (5° 54.5'S, 120°
19.2'E), west of Salayar, Sulawesi, Indonesia, 462 m.

SHELL DESCRIPTION. Shell solid, small, height to 12.9 mm,
subcircular, H/L mean 0.97, moderately inflated, mean T/L 0.38,
tumidity to a maximum of 5.3 mm on a single valve. Shell
inequilateral, extended anteriorly. Shallow posterior sulcus with
concentric lamellae; umbones prominent. Lunule extremely small
and shallow. Anterodorsal area heart-shaped, slightly convex with
fine concentric lamellae. Escutcheon lanceolate, with fine concen-
tric lamellae and a few radial ribs. Exterior sculpture of about 25
fine, closely spaced, radial ribs which are rounded in profile. Radial
ribs crossed by 25-30 low, broad, concentric lamellae, the intersec-
tion giving a finely beaded appearance. Hinge plate thick. Left valve
with an single posterior lateral tooth, two cardinal teeth, the anterior
of which is hooked and sits on a rounded buttress, and a single
anterior lateral tooth. Right valve with single posterior lateral, two
cardinal teeth with the posterior larger and a single anterior lateral
tooth. Ligament elongate extending from beak to two thirds of
distance to the posterior lateral tooth. Inner margin with about 25
crenulations, some of those to the posterior bifurcating. Shallow
sinus on posterior margin. Anterior adductor muscle scar elongate,
not buttressed, slightly detached at the ventral tip from the pallial
line. Posterior adductor muscle scar ovate. Pallial line continuous.
Shell colour white.

COMPARISON WITH OTHER SPECIES. Although this species name
was first introduced by Prashad (1932) as a variety of Dentilucina
(Bellucina) hedleyi = C. civica) it is clearly distinct. In many
characters, such as hinge teeth, it resembles C. australopilula, but
differs in shape and external sculpture, the latter having a fenes-
trate ornament compared with the finely beaded sculpture of C.
quadrata.

HABITAT. A deeper water species from muds at 230-1280 metres.

J.D. TAYLOR AND E.A. GLOVER

Fig. 14 a—d, f Cardiolucina pisiformis (Thiele, 1930) Bandicoot Bay, Barrow Island, Western Australia, WAM 268-94 and Fig. e C. pisiformis, Peron
Peninsula, Shark Bay, Western Australia WAM 258-94. a, exterior of right valve; b, exterior of left valve; c, interior of left valve; d, interior of right
valve; e, juvenile, exterior of shell; f, exterior of shell surface showing posterior sulcus. Scale bars a—d = 1.0mm, e = 500um, f = 300um.

ANATOMY AND SYSTEMATICS OF CARDIOLUCINA

Fig. 15 Geographical distribution of Cardiolucina pisiformis (Thiele).

DISTRIBUTION. Indonesia and Philippines (Fig. 17).

MATERIAL EXAMINED. INDONESIA: Kai Islands, 05°23'S
132°37'E, 436-413m & other stations in same area between 250-—
450m (MNHN, Bouchet and von Cosel manuscript).
KALIMANTAN: Sibuko Bay, 762 & 589m (USMN 302240).
PHILIPPINES: 11°58'N 122°02'E, 486-55 1 m (MNHN) (see Cosel
& Bouchet, ms.); off Baliscasag I.,S. Bohol, 812m (USNM 298742);
Tligan Bay, N. Mindanao, 500m (USNM 298616) & 505m (USNM
298660): off Claveria, Burias, 390m (USNM 2800546): off N. Ticao
1.,420m (USNM 298025); off Ponson I., Camotes Islands, 650m
(USNM 298468); off Apo I., S. Negros, 460m (USNM 298759);
Sogod Bay, Leyte,1280m (USNM 298560).

Cardiolucina rugosa (Hedley, 1909)
Figs 18-19

Lucina (Myrtea) seminula Gould, 1862: 174 (non Lucina seminulum
Deshayes, 1858: 673).

Lucina (Codakia) seminula (Gould) — Smith, 1885: 180-181, pl. 13,
figs 5 & Sa.

Phacoides rugosus Hedley, 1909: 427.

Lucina seminula Gould — Johnson, 1964: 148, pl. 28 fig. 3.

TYPE MATERIAL. Lectotype selected by Johnson (1964, pl 28 fig.
3) (USNM 553). Hedley (1909) introduced the replacement name P.
rugosus because L. seminula is preoccupied. The specimen from the
type locality illustrated in Fig. 18 was given by Gould to Cuming.

TYPE LOCALITY. Hong Kong Harbour, 9-18 m.

SHELL DESCRIPTION. Shell solid, small, height to a maximum of
3.1mm, subcircular in outline, H/L mean 0.92; moderately inflated,
T/L mean 0.38, tumidity to a maximum of 1.2 mm ona single valve.
Shell inequilateral and extended towards anterior. Pronounced pos-
terior sulcus with concentric lamellae. Anteriorly, there is a narrow
sulcus. Lunule small, heart-shaped. Anterodorsal area shallowly
concave with fine concentric lamellae. Escutcheon lanceolate and
slightly convex. Exterior sculpture of about 20 thick, slightly
recurved, concentric lamellae. Interspaces between concentric la-

107

mellae are variable in width. Concentric lamellae often extended
into short flanges at margin of posterior sulcus. Radial ribs indis-
tinct, flattened and closely spaced. Hinge plate thick. Left valve with
single posterior lateral, two cardinal teeth, the anterior of which is
thicker and triangular and a single anterior lateral. Right valve with
a single posterior lateral tooth, two cardinal teeth, the posterior large
and the anterior a thin plate, with a single anterior lateral tooth.
Ligament very short, extending from beak to less than half way to
posterior lateral tooth. Inner margin with 20-21 sharp crenulations.
Small denticles on posterior and anterior dorsal margin. Shallow
sinus on posterior margin corresponding to the position of the
posterior sulcus. Anterior adductor muscle scar elongate, not but-
tressed, detached from the pallial line only at the ventral tip. Posterior
adductor muscle scar ovate. Pallial line continuous. Shell colour
creamy white.

COMPARISON WITH OTHER SPECIES. Cardiolucina rugosa is closely
similar to C. semperiana, but it is smaller, less tumid with more
prominent, recurved concentric lamellae and greater anterior exten-
sion.

HABITAT. 10—250 m witha few dead shells from 540 m, sublittoral
silts and muds.

DISTRIBUTION.

MATERIAL EXAMINED. HONG KONG: North Lantau (BMNH;
BMNH 1963233; NMW acc. 55.158; AM C.33806; 9-18m (USNM
611842, 2 paratypes); (USNM 24252); (USNM 714821). CHINA:
off Pratas Islands, 268m (USNM 296955). KALIMANTAN: off
Silungan I., 539m (USNM 291027). PHILIPPINES: (USNM
21109); Pescador I. Tanon Straits, 412m (USNM 293876): Bohol
(BMNH); off Sombrero I., Balayan, 216m (USNM 312342); S.E. of
Bantayan I., 60m (USNM 293115); Destacado I., 218m (USNM
298090); off N. Cebu, 125m(USNM 293266); Pujada R., E.
Mindanao, 253m (USNM 294765); off Tochanhi Pt, Tawi Tawi
Islands, 90m (USNM 292997) & 35m (USNM 283372); 11°43'N,
122°34'E, 93-99m (MNHN). AUSTRALIA: Queensland, Cape
York (BMNH 1887.2.9.2772-5); Hope Islands, 10-18m (BMNH
1910.9.28.68-71 and BMNH 1963259); Palm Island (NMW acc.
55.158); Gulf of Carpentaria 16°58'S 140°53'E, 9m (AM C.015275);
Albany Passage, Torres Strait, 9-30m (AM C.36171); Palm Island
(AM C.1049); Low Isles, 27m (AM); Cape Sidmouth (AM C.
2594); Endeavour Reef, 40m (AM C.44668); Hervey Bay (AM);
Barney Point, Port Curtis (AM C.21805); Facing Island (AM C.4428);
Quoin Island, Port Curtis (AM); Northern Territory, Darwin (AM).
PAPUA NEW GUINEA: 0.5 mile off Fairfax Harbour, Port Mores-
by, 15—18m (AM C.309466).

Queensland, Australia to Hong Kong (Fig. 19).

Cardiolucina semperiana (Issel, 1869)
Figs 21-24

Lucina pisum Reeve 1850 (part), pl. 11, fig. 66b (non L. pisum
Sowerby 1836).

Lucina semperiana Issel, 1869: 82-83.

Phacoides (Bellucina) semperiana (Issel) — Lamy 1920: 211-213.

Dentilucina (Bellucina) macassari Prashad 1932: 163 pl. 5 figs 13-
16.

Lucina semperiana (Issel) — Bouchet & Danrigal 1982: 16, fig. 19.

Bellucina semperiana (Issel) — Oliver 1992: 99, pl. 19 fig. 7.

TYPE MATERIAL. ‘The species concept of C. semperiana was based
on the figure in Savigny (1817, pl. 8, fig 12) and one specimen from
Suez in the Issel Collection (now lost); the single shell of this species
in the Savigny collection (MNHN) illustrated by Bouchet & Danrigal

J.D. TAYLOR AND E.A. GLOVER

Fig. 16 Cardiolucina quadrata (Prashad, 1932). a—d, holotype of Dentilucina (Bellucina) var. quadrata Prashad, 1932, Salayar, Sulawesi, Indonesia and
e-f, C. quadrata, Kai Islands, Indonesia, BMNH. a, exterior of right valve; b, exterior of left valve; c, interior of left valve; d, interior of right valve; e,
exterior shell surface; f, hinge showing hooked anterior cardinal tooth in the left valve. Scale bars a-e = 2.0mm, f = 1.0mm.

ANATOMY AND SYSTEMATICS OF CARDIOLUCINA

Fig. 17 Geographical distribution of Cardiolucina quadrata (Prashad).

(1982, fig. 19) is here designated as lectotype. The holotype of
Dentilucina (Bellucina) macassari Prashad is located at ZMA, reg.
number 3.32.090 (Fig. 21 c & d).

TYPELOCALITY. C. semperiana from Suez, Red Sea. C. macassari
from Makassar, South Sulawesi, Indonesia (Siboga Expediton sta-
tion 71).

SHELL DESCRIPTION. Shell solid, small, height to a maximum of
7.7 mm, sub-circular in outline H/L mean 0.98; inflated, T/L mean
0.46, tumidity of single valve to 4.6 mm. Inequilateral, extended

109

Fig. 19 Geographical distribution of Cardiolucina rugosa (Hedley).

anteriorly. Deep sulcus posteriorly with prominent concentric la-
mellae. Posterodorsal margin of the sulcus marked by a ridge with
prominent short lamellae. Lunule heart-shaped, shallow. Anteriorly,
there is a narrow sulcus. Escutcheon lanceolate with fine concentric
sculpture. Exterior sculpture of 20—30 low, rounded radial ribs with
narrow interspaces. These are crossed by 15—24 projecting, concen-
tric lamellae which are often irregularly spaced. In some specimens,
concentric lamellae are thickened with steep, ventral edges, giving a
stepped appearance to the shell. Hinge plate thick. Left valve with
single posterior and anterior lateral teeth and two cardinal teeth, of

Fig. 18 Cardiolucina rugosa (Hedley, 1909), Hong Kong Harbour, BMNH 1963233. Specimen of Lucina seminula Gould 1862 from type locality, given
by Gould to Cuming. a, exterior of left valve; b, interior of right valve. Scale bar a—b = 1.0mm.

J.D. TAYLOR AND E.A. GLOVER

Fig. 20 a—b Cardiolucina semperiana (Issel, 1869), Gulf of Suez, Red Sea BMNH 1870.1.19.4, c-f C. semperiana, Tuléar, Madagascar, BMNH. a,
exterior of right valve; b, interior of left valve; c, dorsal view articulated shell; d, exterior of right valve; e, interior of left valve; f, hinge showing details
of the dentition with denticulate posterior cardinal tooth. Scale bars a—b = 1.0mm, c = 750um, d—-e = 1.0mm, f = 500um.

ANATOMY AND SYSTEMATICS OF CARDIOLUCINA

111

Fig. 21

Cardiolucina semperiana (Issel, 1869). Figure a—b Paralectotype of Lucina pisum Reeve, 1850 BMNH 1963552, Singapore and c—d Holotype of

Dentilucina (Bellucina) macassari Prashad, 1932, Makassar, Indonesia, ZMA. a, exterior of right valve; b, interior of left valve; c, exterior of right valve;
d, interior of left valve with attempted drill hole on the posterior part of the umbo. Scale bars a—d = 1.0mm.

which the anterior is the smaller. Right valve with single posterior
and anterior laterals and two cardinals, the posterior being the larger.
Cardinal teeth have small denticles. Ligament extends two thirds of
the distance from the beak to the posterior lateral tooth. Inner margin
with 20-21 crenulations and there are also small, rounded denticles
along anterodorsal and posterodorsal margins. Both anterior and
posterior shell margins have shallow sinuses corresponding to the
sulci. Anterior adductor muscle scar elongate, buttressed in the
more tumid specimens, detached from the pallial line only at the
ventral tip. Small pedal retractor muscle scar at dorsal tip of anterior
adductor scar. Posterior adductor muscle scar ovate. Pallial line
continuous. Shell creamy white.

ANATOMY. (Fig. 22 ) Posterior inhalant aperture absent, exhalant
aperture a small inverted tube. Ctenidia large and thick, inner
demibranchs only, with a thin, shallow, ventral food groove. Gill
filaments long and broad, with large central space crossed by
transverse muscle strands (Fig. 22 c). Bacteriocyte zone packed with
ovoid bacteria about 2.0—2.5 um in length (Fig. 22 d). Labial palps
consist of small papilla at edge of lips. Foot cylindrical, vermiform,

with a much-corrugated outer surface, a pointed tip and a small
rounded heel to the posterior. Ctenidia attached to posterior mantle
by a thin muscular sheet (Fig. 22 f). Body wall with a large irregular
visceral pouch in the antero-ventral position.

GEOGRAPHICAL VARIATION. This species is extremely variable
both within populations and across its range from the northern end of
the Red Sea to the Philippines. In Southeast Asia semperiana are
generally smaller, but relatively more tumid, as shown in Fig. 23,
which plots ratio of length over tumidity against mean height of
specimens from Red Sea, Madagascar, India, South East Asia. The
holotype of macassari is the most tumid of all the specimens
measured, but is probably an abnormal individual which has sur-
vived attempted drilling at the posterior dorsal margin. C. semperiana
from Southeast Asia also tend to have fewer, but heavier, concentric
lamellae (Figs. 21).

HABITAT. Live material collected from shallow water (up to 50 m),
a few dead shells from up to 350 m, mud and sand.

COMPARISON WITH OTHER SPECIES. C. semperiana is similar to C.
rugosa but is larger, more tumid and has more prominent radial ribs.

112 J.D. TAYLOR AND E.A. GLOVER

Fig. 22 Cardiolucina semperiana (Issel, 1869), Sinai Coast, Gulf of Suez, Egypt. NUW 1982.068. a, interior of right valve showing thick inner
demibranch of the ctenidium and cylindrical, vermiform foot; b, right valve showing pronounced visceral pouch and the foot with strongly corrugated
surface and small, rounded heel; c, section showing broad gill filaments and transverse muscle strands crossing the spaces within the filaments.

d, bacteriocyte zone with ovoid bacteria; e, labial palps (Ip) and mouth (m); f, ctenidia attached to the posterior mantle by a thin muscular sheet.
Abbreviations: c=ctenidium, f=foot, v=visceral pouch, h=heel, lp=labial palps, m=mantle, a=attachment of ctenidium to mantle. Scale bars a = 1.0mm, b
= 500um, c = 40um, d = Sum, e = 150um, f = 250um.

ANATOMY AND SYSTEMATICS OF CARDIOLUCINA 113

8.0
7.5
7.0
6.5
e SEAsia
= 6.0 oO Arabia
@ 5.5 4 Madagascar
5.0 © India
Vv macassari
45
4.0
3.5
3.0

co, of 40 42 44 AG .48 £50 .52 54 56
Tumidity/length

Fig. 23. Geographical shape variation of Cardiolucina semperiana. The specimen of C. macassari is the holotype.

Fig. 24 Geographical distribution of Cardiolucina semperiana (Issel).

114

J.D. TAYLOR AND E.A. GLOVER

Fig. 25

Cardiolucina siquijorensis Taylor and Glover new species, Siquijor, Philippines. a, Paratype (USNM 812176), exterior of right valve; b, Holotype

(USNM 812176), exterior of left valve; c, Paratype, interior of left valve; d, Holotype, detail of lunule; e, Paratype, hinge of left valve showing posterior
lateral tooth with pronounced rim on the socket. Scale bars a—c = 500um, d = 150m, e = 250um.

DISTRIBUTION. Northern Red Sea, southern Africa to Philippines

(Fig. 24).

MATERIAL EXAMINED. EGYPT: Gulf of Suez (BMNH
1870.1.19.4); Suez (BMNH 1963492): Nuweiba, Sinai (AM 1954).
YEMEN: Aden (BMNH 1902.12.30.754-6 & 1963235); Aden
(AM C.33727); (USNM 608854). OMAN: Qurm, Muscat (NMW
Z.1993.061.0434); Masirah (NMW Z.1993.061.0427 & 0433).
ARABIAN GULF: (USNM 636963); Bahrain (NMW
1993.061.043); Linyah (NMW acc. 55.158); Qatar (BMNH acc.

2258); Abu Dhabi (BMNH). ZANZIBAR: Nyange I., 20m (ANSP
251213); 1.5 miles S.W. of Ras Shangani, 18—20m (ANSP 213493);
1 mile N. Bawi I., 22m (ANSP 250912). KENYA: Mombasa
(BMNH acc. 2223), (NMW 1955.158). TANZANIA: Msimbasi
Bay, Dar es Salaam 5m (AMC.309448). MOZAMBIQUE: Bazaruto
I., 8-10 m (NM K5317 & G4458). SOUTH AFRICA: Durban Bay,
(NM E834 & A2357); Shelly Beach, Izotsha, Natal (NM9898).
MADAGASCAR: N.W. Madagascar, 30m (AM C.70660); Tuléar
(MNHN); 40 miles W.S.W. Tuléar, 33m (USNM 761189); Nossi Bé
(USNM 633927); Nossi Bé, shallow water to 35m, (ANSP 261127,

ANATOMY AND SYSTEMATICS OF CARDIOLUCINA

259587, 259028, 260541, 261338, 259901, 261873). MAURI-
TIUS: 1.5m N.W. Black River, 15m (ANSP 273431). PAKISTAN:
Karachi (BMNH acc. 1831). INDIA: 80 miles S.W. Bombay 17°
54'N 72 °27’E, (USNM 716676); Madras (BMNH 1953.1.30.190-—
4); Madras (BMNH 89.2.17.23-6); Tondi, Madras (NMW acc
55.158); Manapad (BMNH 1953.1.30.26); Mandapam, Gulf of
Mannar, 3m (ANSP 302433); Tuticorin (BMNH 1953.1.30.188—
89); S. of Krusadai I., Gulf of Mannar, 18m (ANSP 301196);
Andaman Islands (AM C.040397, C.040597); Andaman Islands
(NMW 55.158). SRI LANKA: Colombo (AM); Trincomallee, 29m
(AM); E., S., and N. coasts (AM C.309467); BMNH 1963491; Palk
Strait, 22m (AM); 9°34.5'N 80°39'E (AM C.309461); northern tip of
Eluvativu I., (ANSP 211085). BURMA: 57 miles N.W. Tavoy L.,
Anadamans Sea, 39m (ANSP 293254). THAILAND: Koh
Kahdat(AM C.031287); Phuket (USNM 661156); south end Light-
house I., Phuket, 5m (ANSP 286500); 25 miles N.N.W. Phuket, 42m
(ANSP 291734). MALAYSIA: Mersing Island, E. Malaya, 3.5m
(AM). SINGAPORE: BMNH acc. 2172; Singapore (AM C.3437);
NMW acc 55.158. INDONESIA: Kalimantan (AM C.309470).
PHILIPPINES: Corregidor, Bay of Manila, intertidal (BMNH;
Manila Bay, 18m (ANSP 281443); Manila Bay, 18m (ANSP 328655);
off Antonia I., E. Panay, 40m (USNM 293697); N. of Marinduque,
353m (USNM 295197); Lataau I., 33m (USNM 236058); Malampaya
Sound, Palawan, 25—35m (USNM 297395); off Tinkata I., Tawi
Tawi Islands, 30m (USNM 257677); VIETNAM: Vung Tau, Chilins
(ANSP 330797). AUSTRALIA: Western Australia, Shark Bay
(AM C.69245); Carnarvon, 7.6m (AM); Carnarvon (AM C.69270);
210 miles N. of Broome, 230m (AM); 110 miles N. of Melville
Island, Arafura Sea (AM); 72 miles N.W. of Dampier Island,110m
(AM 2405).

Cardiolucina siquijorensis Taylor and Glover, new species
Fig. 25

TYPE MATERIAL. Holotype, articulated specimen (Fig. 25b) USNM
812176. H=3.4 mm, L=3.5 mm. Paratypes, USNM 812177, 812169.

TYPELOCALITY. Siquijor, Solong-on, Siquijor I., Visayan Islands,
Philippines (09° 13'10"N 123° 27'30"E) depth 2-3 m.

SHELL DESCRIPTION. Shell very small, toa maximum height of 3.5
mm, outline circular H/L 0. 97, inflated, T/L 0.39, tumidity of single
valve to 1.3 mm. Shallow posterior sulcus with concentric lamellae.
Anterior sulcus absent. Lunule shallow, heart-shaped. Anterodorsal
area indistinct, posterodorsal area with concentric lamellae. Exte-
rior sculpture, fenestrate, of 1 1-12 broad, straight-sided, radial ribs.
Interspaces wide, but narrower than the ribs. Ribs are slightly
broader towards the anterior. About 14—16 thin, concentric lamellae,
project slightly above radial ribs. Hinge plate robust. Left valve with
single posterior lateral tooth, the dorsal edge of the socket extended
into a sharp rim (Fig. 22 e); two cardinal teeth, triangular in shape
and more or less equisized; single anterior lateral tooth. Right valve
with single anterior lateral tooth, a single central cardinal tooth and
a robust posterior lateral tooth. Ligament extends from beak to the
dorsal end of the posterior lateral tooth (Fig. 22 e). Inner margin with
12 coarse crenulations, and fine beading extending to the hinge
margins. Anterior adductor muscle scar elongate, not buttressed,
with only ventral tip detached from pallial line. Posterior adductor
muscle scar rounded. Pallial line continuous. Shell creamy white.

COMPARISON WITH OTHER SPECIES. This species is similar in size
and ornament to C. pisiformis but is distinguished by fewer and
broader radial ribs, the shallower lunule, the two equal-sized cardi-
nal teeth in the left valve and the single cardinal in the right valve.

115
HABITAT AND DISTRIBUTION. As for type locality.

MATERIAL EXAMINED. PHILIPPINES: Siquijor I., collected with
type material (USNM 812177, 812176, 812169).

Cardiolucina sp.
Fig. 26

In addition to the species described, we have four lots from the
Philippines consisting of rather badly preserved, disarticulated shells
and two rather better preserved juvenile shells. These are distinct in
a number of shell characters from the other Cardiolucina species.
Because the adult specimens are badly preserved and because
juveniles often differ considerably from adults, we are reluctant to
describe a new species based on this material.

The adult shells have maximum height of 10.8mm, with tumidity
of a single valve to 2.7mm and H/L 0.98, T/L 0.5. The posterior
sulcus is broad and shallow, the anterior sulcus absent. About 17-18
prominently scalloped, concentric lamellae, with more than 20 fine,
rounded, radial ribs, which do not cross the concentric lamellae. The
interspaces between the ribs are narrow, but relatively deep and the
intersection of the concentric and radial ornament produces a deeply-
pitted appearance. The lunule is heart shaped and relatively deep. In
the left valve, there is a single posterior lateral, two triangular
cardinal teeth, and a single posterior lateral. The inner margin of the
shell has about 25 crenulations.

This species is similar to C. semperiana but can be distinguished
by the lack of the anterior sulcus, the shallower posterior sulcus, the
pitted appearance of the shell surface and the more rounded
anterodorsal area.

MATERIAL EXAMINED. PHILIPPINES: off Sibugay I., E. of
Masbate, 198 m (USNM 292666); Daram Channnel, W. Samar, 48 m
(USNM 292830); off Matocot Point, W. Luzon, 400 m (USNM
295706); off Corregidor Light, 73 m (USNM 294200).

ATLANTIC OCEAN SPECIES

Cardiolucina lamothei (Dautzenberg, 1913)
Figs 27 a & b, 28

Lucina lamothei Dautzenberg, 1913: 100-101, pl. 3 figs 50-54.

Phacoides (Parvilucina) congoensis Thiele & Jaeckel, 1931: 220—
221, pl. 4 fig. 91.

Phacoides (Bellucina) lamothei (Dautzenberg) — Lamy 1920 : 215.

TYPE MATERIAL. Figured specimens of L. lamothei—two syntypes
from Tamara, Archipel de Los, Guinée, marked TYPE in
Dautzenberg’s handwriting (MNHN). Numerous other specimens
from localities mentioned by Dautzenberg (1913) are housed in the
Royal Belgian Institute of Natural Sciences (IRSNB inventory I.G.
10591). Our figured specimen is from a sample sent by Dautzenberg
to Tomlin (NMW 1955.158).

TYPE LOCALITY. West Africa, Tamara, Archipel de Los, Guinée.

SHELL DESCRIPTION. Shell solid, very small, height to maximum
of 4.0 mm. Subquadrate, inequilateral, slightly extended anteriorly.
Not inflated. Anterior sulcus shallow leading to marginal sinus.
Umbones are conspicuous, prosogyrate. Lunule very small, under-
neath beaks. Anterodorsal area indistinct and slightly convex.
Escutcheon with concentric lamellae. Exterior sculpture of 15-18
prominent, narrow concentric lamellae, which are slightly reflexed.
About 40 fine, radial ribs in the interspaces between concentric

116

J.D. TAYLOR AND E.A. GLOVER

Fig. 26
bars a = 400um, b = 500um, c=1mm.

lamellae. Hinge plate thick. Left valve with single, hooked anterior
lateral tooth, two cardinals and single posterior lateral tooth. Right
valve with similar. Indentations on both anterior and posterior
margins which correspond to the position of sulci. Inner margin with
about 40 crenulations. Anterior adductor muscle scar elongate, not
buttressed. Pallial line attached to lower third of anterior adductor
muscle scar. Posterior adductor scar reniform in outline. Pallial line
continuous. Shell colour grey-white.

COMPARISON WITH OTHER SPECIES. ‘This species is similar to C.

. rehderi but has a greater number of fine radial ribs.
HABITAT. Fine muddy sand from 4—200 m.
DISTRIBUTION. Senegal to Northern Angola (Fig. 28).

MATERIAL EXAMINED. AFRICA: Guinea, Los Islands (NMW
1955.158).

Cardiolucina rehderi (Britton, 1972)

Fig.27c &d
Parvilucina (Bellucina) rehderi Britton, 1972: 7-9, fig. 3.
Lucina (Parvilucina) rehderi (Britton) — Rios, 1994: 252.

TYPE MATERIAL. Holotype, USNM 208255, L = 4.2 mm, H = 4.2
mm, T = 4.1 mm (paired valves), also 9 paratypes.

a—-b, Cardiolucina sp. Corregidor, Philippines (USNM 294200); c, Cardiolucina sp Sibugay Island, Masbate, Philippines (USNM 29266). Scale

TYPE LOCALITY. Brazil, no further details.

SHELL DESCRIPTION. Shell small, height to 4.2 mm. Subcircular;
moderately inflated, maximum tumidity on a single valve to 1.9 mm.
Inequilateral, slightly extended anteriorly. Broad posterior sulcus.
Posterior dorsal area slightly convex, with distinct elevation of
concentric lamellae. Shallow anterior sulcus, anterior dorsal area
indistinct. Lunule shallow, heart-shaped. Exterior sculpture of about
30 indistinct radial ribs most clearly visible in the interspaces
between concentric lamellae, ribs separated by shallow, narrow
interspaces. Concentric lamellae,14—16, prominent, thickened la-
mellae often appear scalloped at position of intersection with radial
ribs. Hinge plate thickened. Left valve with single anterior lateral
tooth; two cardinal teeth and single posterior lateral. Right valve
with similar. Ligament short, extending about half distance from
beak to posterior lateral tooth. Inner margin with about 20 fine
crenulations. Anterior adductor muscle scar longer than wide but
relatively short with pallial line joining near ventral tip. Posterior
adductor muscle scar rounded. Shell colour white. A more extensive
description of this species is given by Britton (1972).

COMPARISON WITH OTHER SPECIES.
to C. lamothei.

This species is closely similar

HABITAT. Unknown.

ANATOMY AND SYSTEMATICS OF CARDIOLUCINA

7 eRe **
" . ee &. % he ae

117

Fig. 27 Cardiolucina lamothei (Dautzenberg, 1913) and C. rehderi (Britton, 1972). a, C. lamothei Iles de Los, nr Conakry, Guinea (NMW 1955.158) Left
valve. P35.082; b, left valve interior; c, C. rehderi Britton, left valve, Paratype NMNH,; d, left valve interior. Scale bar a—d = 1.0mm.

Fig. 28 Geographical distribution of Cardiolucina lamothei
(Dautzenberg). Data from Dautzenberg (1913).

DISTRIBUTION. Known only from type material reported on by
Britton (1972).

FOSSIL SPECIES OF CARDIOLUCINA

Apart from the description of the type species C. agassizi, we have
not attempted a comprehensive review of the fossil record. How-
ever, the species listed below have the characters of Cardiolucina.
Other fossil species that we do not consider to belong the genus are
listed separately.

Phacoides (Parvilucina) ligatus Cossmann and Pissarro,
1904. (pages 17-18, pl. 7, figs 1-4).

Described from the Eocene (Lutetian) of Hauteville, Cotentin Pe-
ninsula, France. This small species has a thick shell and prominent
concentric lamellae with fine radial ribbing similar to that of
Cardiolucina agassizi. Chavan (1937: 207) first suggested that this
species should be classified as a Bellucina. On the basis of the
published figures, we consider that this is probably the earliest
known Cardiolucina.

Lucina nuciformis Tate, 1887b: 144 (figured in Tate, 1886,
p. 158, pl. 12 figs.10a & b).

118

This Miocene species from Blanche Point, Aldinga Bay, South
Australia, is very similar to the Recent Cardiolucina crassilirata
(Tate).

Linga (Bellucina) gonzalesi Shuto,1971 (pp. 30-31; Plate
1 figs 8,11,13—17; text fig. 6)

Described from the Lower Pliocene Cabatuan Formation of Panay
Island, Philippines. This species is well described and illustrated. It
has about eight prominent, radial ribs with a fenestrate ornament like
the Recent Cardiolucina eucosmia, C. pisiformis andC. siquijorensis.
The intersections of the radial ribs and concentric lamellae are
‘strongly granulated’, a feature found in none of the three Recent
species.

Lucina polli Icke & Martin, 1907. (p. 250, plate 18 figs 40,
40a)

Described from the Pliocene of Nias Island, Indonesia. From the
illustrations, this is a definite Cardiolucina species with fine radial
ribbing and prominent concentric lamellae, similar in form to C.
rugosa.

Cardiolucina agassizi var. regularior Sacco, 1904 (p. 90,
pl. 20 figs. 40-42) and var. constricta Sacco,1904 (p. 90 pl.
20 fig. 43).

Without a morphological analysis of C. agassizi populations it is
difficult to evaluate these names, but they would seem to be forms of
a variable species.

SPECIES EXCLUDED FROM CARDIOLUCINA.
Listed under original generic assignments as published.

Phacoides (Bellucina) actinus Dall, 1903: 1385, pl. 52 fig.
3. Also Woodring (1925: 126-127; pl. 17 figs 5-8).

Described from the middle Miocene Bowden Formation of Jamaica,
this species has 16—22 radial ribs and fine concentric lamellae.
However, the lunule is lanceolate and the hinge different from
Cardiolucina species. We consider that it may belong to the same
clade as Radiolucina amianta (Dall) (see below).

Phacoides (Bellucina) amiantus Dall, 1901: 826-7, pl. 39
fig.10.

Although Dall included this Caribbean species in Bellucina and
Bretsky (1976) used it as the basis of her diagnosis of the genus, it
differs in many features from the type species of Cardiolucina and
Bellucina. Britton (1972) discussed this at some length and pro-
posed the new generic name Radiolucina with P. amiantus as the
type species.

Phacoides (Bellucina) tuomeyi Dall, 1903: 1385, pl. 52
fig. 1.

Described from the late Miocene Duplin Marl of North Carolina,
this species is similar to the Recent Radiolucina amianta (Dall).

Phacoides (Bellucina) waccamawensis Dall,1903: 1385,
pl. 52 fig. 2.

Described from the Pliocene Waccamaw Formation of South Caro-
lina, this species is closely similar to the Recent Radiolucina ami-

J.D. TAYLOR AND E.A. GLOVER

anta (Dall) and was tentatively synonymised by Bretsky (1976).

Phacoides peritaphros Barnard, 1964: 476-477 fig. 22.
Syntypes: BMNH 1964264

Although Barnard regarded this southern African species as allied to
Cardiolucina lamothei (Dautzenberg), it lacks radial riblets, has a
shallow, open lunule and is more anteriorly extended. This species is
similar in many features to Cavilinga trisulcata (Conrad, 1841) (see
Olsson and Harbison 1953, pl.7 fig. 4), the type species of the genus
Cavilinga Chavan, 1937.

Lucina cancellaris Philippi, 1846: 21.

This is a West American twin species to the Caribbean Radiolucina
amianta. Keen (1971, p. 121) points out that ‘Dall unfortunately
assigned this species and its Caribbean twin L. amianta (Dall, 1901),
to the subgenus Bellucina, the type of which is an Indian Ocean form
only superficially similar, and American authors have been slow to
recognise the error.’

Lucina pulchella Lynge, 1909: 173, pl 3 figs. 13-15.

Lynge compared this species with L. seminula Gould, L. pisum
Reeve and L. semperiana Issel. We examined the type material
(Zoological Museum, Copenhagen) which has a different ribbing
pattern from any Cardiolucina and lacks posterior and anterior sulci.
This species may be an Epicodakia.

Lucina valida Smith, 1904: p.40 pl.3 fig.19.

This southern African species is very similar to and a probable senior
synonym of Phacoides peritaphros Barnard. Syntypes: BMNH
1903.12.19.1270-4.

Cardiolucina striatula var. taurotrigona Sacco, 1901: 90,
pl. 20 fig. 44-47.

Cardiolucina striatula var. ovatuloides Sacco, 1901 90 pl.
20 fig. 48.

Cardiolucina taurocrenulata Sacco, 1901: 90 pl. 20 fig. 50.

C. taurocrenulata var. aviculina Sacco, 1901: 90 pl. 20
figs 51 & 52.

Cardiolucina oligobliqua Sacco, 1901: 91 pl. 20 fig. 53.

All these taxa were described from the Miocene of Italy. They have
concentric ornament only and differ in shape from the type species.
In our opinion these taxa should all be excluded from Cardiolucina.

RELATIONSHIPS AMONGST THE SPECIES

In an attempt to establish relationships amongst the Cardiolucina
species a preliminary phylogenetic analysis using parsimony was
performed using a set of 12 shell characters. We used PAUP version
3.0, with further interpretation of characters with McClade version
3.04. Although it would have been desirable to include anatomical
characters these were available for only two species. We selected as
an outgroup the northeastern Pacific Parvilucina tenuisculpta (Car-
penter, 1864), the type species of the genus Parvilucina. Since the

ANATOMY AND SYSTEMATICS OF CARDIOLUCINA

S
o -~ &
a jana tae anh
[sth eof rian meh oe aa
TS Oe AGO! Ee IO er hS
5)
a ee ek oe
Ss ts gut amy ie Cr cement item gee las
10
9
8
7
4

pisiformis

119
outgroup
©
3 = ~ =
2 a = = &
2 Ks) * g 5
ae) 0 be = A)
5 3 3 g 2 3
S = = 2 g oO
7) As} iy 1s) 1s) ~
6 3
2
1

Fig. 29 Strict consensus tree of Cardiolucina species derived from two equally parsimonious trees. Outgroup is Parvilucina tenuisculpta. Character state
changes at numbered internal branches. Branch 1: char. 1, state 2>1; char. 3 state 3>0; char. 6 state 2>1. Branch 2: char. 3 state 0>2; char. 8 state 0>1;
char. 11 state O>1. Branch 3: char. 11 state 1>2. Branch 4: char. 7 state 1>0. Branch 5: char. 4 state 0>1; char.5S state O>1; char. 6 state 1>0; char. 12 state
2>0. Branch 6: char. 9 state 0>1. Branch 7: char. 1 state 1>0; char. 2 state 0>2. Branch 8: char. 12 state 2>1. Branch 9: char. 2 state 2>1. Branch 10:

char.10 state O>1.

Table 1. Characters and character state codings used in the phylogenetic analysis.

1. Posterior sulcus. This is an indentation in the posterior shell seen on
the outer surface extending from the umbo to the shell margin. It is present
in all Cardiolucina species, deeply incised in some (Fig. 20d), shallow in
others (Fig. 2a) and weak and poorly-defined in the outgroup.

States: 0 - deep prominent;1 - shallow; 2 - weak

2. Anterior sulcus. This is a similar indentation at the anterior end of the
shell. This is absent in some species, but where present (Fig. 20) never as
prominent as the posterior sulcus.

States: 0 - absent; | - narrow; 2 - broad

3. Lunule. Many species have a heart-shaped, shallow lunule (Fig. 20c) ,
but in C. eucosmia it is deeply excavated. In other species the lunule is
very small and tucked under beak (Fig.2) and in the outgroup it is
lanceolate.

States: 0 - shallow, heart-shaped; | - deeply excavated; 2 - tiny under
umbo; 3 - thin lanceolate

4. Radial ribs - number
States: 0 - less than 20; 1 - more than 20

5. Radial rib - shape

Some species have prominent ribs with straight sides and separated by
broad interspaces (Figs. 2,3). In others, the ribs are closer together with
low rounded profiles and narrow interspaces (Figs. 7,10).

States: 0 - rounded, narrow interspaces; | - straight-sided broad interspaces

6. Radial rib intersection with concentrics.

In some species, ribs appear to cross concentric lamellae without
interruption (Fig. 2d) whilst in others the ribs terminate at the intersection
with major concentric lamellae (Fig. 7d). In the outgroup the radial ribs
are extremely weak.

States: 0 - strong, cross concentrics ; | - weak, terminate at concentrics; 2 -
very weak

7. Concentric lamellae - number
Concentric lamellae are a prominent feature of many Cardiolucina species.
States: 0 - less than 20; 1 - more than 20

8. Shape of anterior cardinal tooth in left valve

The majority of species have a triangular wedge shaped tooth (Fig. 18b)
but in some species the anterior cardinal is a hook-shaped projection (Fig.
3109):

States: 0 - wedge; 1 - hooked

9. Posterior cardinal of left valve

Most species have a relatively thin tooth (Fig. 7c ), but in C. pisiformis ,
for example, it is much thicker (Fig. 14c ).

States: 0 - thin; 1 - thick

10. Denticles on cardinal teeth

This refers to small denticles seen at high magnification along the edges of
the posterior cardinal (Fig. 12d).

States: 0- absent; 1 - present

11. Hinge buttress

The hinge may be projected ventrally below the cardinal teeth to form a
rounded buttress (Fig 3c).

States: 0 - absent; | - slight; 2 - prominent

12. Margin crenulations - number

These are seen on the internal ventral margin of shell. These were counted
around the shell margin, fine beading not included.

States: 0 - coarse (n = 10-20); 1 - medium (n = 21-30); 2 - fine (31 + ).

120

Table 2 Matrix of Cardiolucina species and the outgroup Parvilucina
tenuisculpta and their character states. Character coding as in Table 1.

1

N
&

OQ 7 9 10 Ii

—
N

agassizi 0
australopilula l
civica 1
crassilirata 1
eucosmia ]
pisiformis 1
quadrata 1
rugosa 0
semperiana 0
siquijorensis 1
lamothei 0
rehderi 0
P.tenuisculpta 2

SnnowrrHrooracoco
WBOSCSCCONOHNNNO |] w
Be ROR RF RF OOH OF
SeoOOHMOoOCOoOKRHOOHO|U
NeEH Oe Ee HooHrHor
SOOCOCOOFOOHKHH OO]
SeooOrooorcecoce
SOSONuRH HK OOHKoCoOCSO
eecooocoonooeene
NHENOHKPHK RB OCONNHE=

Soo oom oot 1S StS

original designation by Dall (1901), Cardiolucina (as Bellucina) has
been widely listed as a subgenus of Parvilucina and the shells have
some features in common (descriptions in Hickman 1994). More-
over, there are some similar anatomical features, P tenuisculpta
having a visceral lobe resembling that found in C. semperiana and
C. australopilula, as well as the connection of the posterior ctenidia
to the mantle and the lack of a fused inhalant aperture (Reid and
Brand, 1986). The characters and their states are listed in Table 1.

The distribution of the various character states amongst the
Cardiolucina species and the outgroup is shown in Table 2.
Phylogenetic analysis using this matrix produced 2 equally parsimo-
nious trees of 26 steps with a consistency index of 0.731 and a
homoplasy index of 0.269. A strict consensus tree is shown in Fig.
29. The differences in the 2 trees resulted from the grouping of
C. civica and C. crassilirata as a single clade or as a trichotomy with
the C. australopilula /C. quadrata clade.

The analysis suggests that there are three distinct clades of
Cardiolucina species (Fig. 29). Clade 1 contains C. australopilula,

Table 3 Summary of shell characters in Cardiolucina species

J.D. TAYLOR AND E.A. GLOVER

crassilirata, civica and quadrata. Apomorphies which define this
clade at Branch 2 are; the tiny lunule, the hooked-shaped, anterior
cardinal tooth and the hinge buttress. Clade 2, separated at Branch 4
by a single apomorphy of having less than 20 concentric lamellae,
contains two major branches; that comprising the type species C.
agassizi and the species around C. semperiana. (Branch 7) and
another branch (Branch 5) containing C. eucosmia, pisiformis and
siquijorensis. This latter clade is defined by four apomorphies;
having less than 20 radial ribs, which are straight-sided and cross the
concentric lamellae and also possess coarse marginal crenulations.
The C. agassizi — lamothei clade is defined at Branch 7 by the
prominent posterior sulcus and a broad anterior sulcus.

This hypothesis of relationships should be regarded as tentative
because it is based on shell characters alone. Nevertheless, it is open
to falsification by additional or alternative character sets.

GEOGRAPHICAL DISTRIBUTION

Two species, Cardiolucina lamothei and C. rehderi are found on
opposite sides of the Atlantic, whilst the other nine are distributed
largely within the tropical Indo-West Pacific Province. One of these,
C. semperiana, has a wide range from the Gulf of Suez to Philip-
pines whilst the rest have more restricted distributions within the
central Indo-Pacific. Four species are endemic to Australia; C.
eucosmia, C. pisiformis and C. australopilula to the western and
northern coasts and C. crassilirata to the temperate southern coast.
Cardiolucina civica occurs from Japan through Indonesia to the
Andaman Islands in the west and Fiji in the east. C. quadrata is
restricted to deeper water around the Philippines and Indonesia.
Apart from C. rehderi, which is known from only one vague locality,
the species with the most limited range is C. siquijorenis from
Siquijor Island in the Philippines. All the species are largely tropical
in distribution, except for C. crassilirata from southern Australia.

Lunule
Sulcus Marginal small, under _heart-
Species H L T Radial ribs Concentric lamellae post. ant. crenulae beak shaped
agassizi 74 71 2.7 ¢.25 faint, closely c.20 width of interspaces deep narrow 28 shallow
spaced extremely variable
australopilula 13.8 12.7 7.5 c.20 interspacesalmost c.34-36 low, narrow shallow none 21 a
as wide as ribs
civica 10.6 104 45 >30 fine, closely >25 conspicuous shallow none >60 =
spaced
crassilirata Hi) tt3} >40 fine, closely >30 fine, clustered shallow none 31 as
spaced in groups of 4-6
eucosmia 7.8 7.7 3.8 c.12 broad, with c.13-14 low v.deep narrow 12 v. deep
wide interspaces
pisiformis 44 44 2 13-17 low, widely c.20 narrow shallow none 13 deep
spaced
quadrata 12.9 13.6 5.3 ¢.25 fine,closely 25-30 low, broad shallow none 25 *
spaced
rugosa 331 3.2 1.2 >25 faint, closely c.20 prominent deep narrow 20 shallow
spaced
semperiana Well 7.5 2.7 20-30 low, closely 15-24 interspaces variable deep narrow, 20-21 shallow
spaced in width ribbed
siquijorensis 3.5 3).5) 13 12 broad 14-16 thin shallow none 12 shallow
lamothei 4 310) ae >40 fine, closely 15-18 prominent deep broad 40 s
spaced
rehderi 42 42 1.9 >30 fine, faint 14-16 prominent deep broad 20 3

closely spaced

H=height, max mm, L=length, max mm, T=tumidity, max mm.

ANATOMY AND SYSTEMATICS OF CARDIOLUCINA

The phylogenetic analysis shows that Cardiolucina lamothei and
C. rehderi are closer to the C. semperiana clade of species than to the
C. australopilula or C. eucosmia clades. This suggests that the clade
containing C. lamothei-rehderi and C. semperiana had a widespread
Tethyean distribution prior to the (late Miocene) closure of the
connection between the Indo-Pacific and Atlantic Oceans. This is
confirmed by the occurrence of C. agassizi in the Middle Miocene of
south west Europe. The C. australopilula clade is restricted in
distribution to the central Indo-Pacific. The Miocene species C.
nuciformis Tate from South Australia is similar to the Recent C.
crassilirata from the same area suggesting a long occupancy.

CONCLUSIONS

In this revision of the systematics of Cardiolucinawe have redefined
the genus and its type species and recognised eleven living species
from the tropical Atlantic and Indo-W. Pacific provinces. The previ-
ous lumping of many taxa into C. semperiana underestimated
species diversity. Table 3 provides a summary of the characters of the
recognised species as an aid to identification. Anatomical characters
are known for only two species, C. australopilula andC. semperiana.
Comparison with other genera is difficult because of the lack of
information, but Parvilucina has some similar characters, such as
the lack of mantle fusion at the posterior inhalant area, the connec-
tion of the ctenidia to the posterior mantle and the large visceral lobe
(Reid and Brand, 1986). Further work is needed to establish the
distribution of these characters in other lucinid taxa.

Although the earliest Cardiolucina would seem to be the Eocene
species C. ligata (Cossmann & Pissarro), the fossil record, apart
from C. agassizi which is common in the middle Miocene of Europe,
is rather scanty. The only species we can confirm are Cardiolucina
nuciformis (Tate) from the Miocene and, from the Pliocene, C.
gonzalesi (Shuto) and C. polli (Icke & Martin). This paucity of
records is unsurprising because, even in Recent faunas, these small
lucinids are neglected.

The problems encountered during this systematic revision reflect
the current state of systematics within the family Lucinidae. Many
genera are defined only by a small number of shell characters, type
species of genera are often inadequately described and, moreover,
their characters are often ignored by systematists. Species diversity
is often greater than realised because taxa have been shoe-horned
into a small number of ‘well-known’ names. Relationships between
genera are generally unknown and rigorous phylogenetic analyses
are lacking. Bretsky (1970, 1976) produced a phenetic/phylogenetic
analysis of lucinid genera, with possible phylogenies for her various
lineages. Unfortunately, there are problems with these phylogenies
because she lumped together distinct genera and confused type
species, for example, using for Bellucina characters of Radiolucina
amianta instead of the type species B. eucosmia. Moreover, there is
no anatomical or biological information available for the vast major-
ity of species, especially those from the Indo-Pacific province.
Recent studies on the chemosymbiosis of lucinids have focused on
a small number of species from the Caribbean and temperate north
Atlantic and Pacific (reviewed in Fisher, 1990, Le Pennecet al.,1995),
but biological information from these studies has yet to be integrated
into lucinid systematics. Systematic and phylogenetic studies should
be providing the framework to test hypotheses concerning the
evolutionary history and radiation of the Lucinidae, but, at present,
these are inadequate.

121

ACKNOWLEDGEMENTS. Ian Loch, Jerry Harasewych, Tom Schiotte,
Graham Oliver & Alison Trew, Philippe Bouchet & Rudo von Cosel, Dick
Kilburn, Robert Moolenbeek, Fred Wells & Shirley Slack-Smith, Hiroshi
Saito and John Slapcinsky generously provided access to material and loan of
specimens in their care. Thierry Backeljau kindly sent information about the
Dautzenberg collection. Philippe Bouchet & Rudo von Cosel provided
access to unpublished information and commented on the manuscript. We
thank Graham Oliver for discussion and nomenclatural advice. Additionally
we are grateful to David Cooper for the thin sections of C. australopilula,
Harry Taylor for fine macrophotography, Nick Hayes for printing of SEM
photographs and Alex Ball for translating some descriptions.

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Issued 28 November 1997

A new species of water mouse, of the genus
Chibchanomys (Rodentia, Muridae,
Sigmodontinae) from Ecuador

PAULINA D. JENKINS ~

Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD, U.K.

ADRIAN A. BARNETT

School of Life Sciences, Roehampton Institute, West Hill, London SW15 5SN, U.K.

SyNopsIs. A new species of the rodent genus Chibchanomys is described from Ecuador on the basis of external and cranial
morphology. A phylogenetic analysis is used to assess the relationship of the new species to other ichthyomyine taxa.

INTRODUCTION

The New World rodents currently placed in the large and complex
subfamily Sigmodontinae (sensu Carleton & Musser, 1984) include
amorphologically and ecologically distinctive group of semi-aquatic
South American genera (Voss, 1988), for which it is convenient here
to use the name ichthyomyines. In a monograph of ichthyomyine
rodents, Voss (1988) included five genera: Ichthyomys Thomas,
1893 (four species), Anotomys Thomas, 1906a (monotypic), Rheomys
Thomas, 1906b (four species), Neusticomys Anthony, 1921 (four
species) and a new genus, Chibchanomys Voss, 1988 for an enig-
matic species of uncertain generic affinity. Chibchanomys trichotis
(Thomas, 1897) was originally placed in Jchthyomys but was subse-
quently assigned to Rheomys (see Tate, 1932; Cabrera, 1961), then
to Anotomys (see Handley, 1976). Subsequent to the revision by
Voss, an additional ichthyomyine species, Neusticomys mussoi Ochoa
& Soriano, 1991, has been described.

During the course of several zoological surveys of Las Cajas
Plateau, Ecuador from 1981 to 1987, five specimens of an undescribed
species of ichthyomyine rodent were captured. Observations were
made on two of these animals, which subsequently escaped, while
three specimens were donated to The Natural History Museum.
Another specimen was filmed for the 1992 BBC National Geo-
graphic wildlife film ‘Avenue of the Volcanoes’ (Jim and Theresa
Clare, personal communication). On the basis of external and
craniodental characters the study specimens agree most closely with
the generic diagnosis of Chibchanomys but are also sufficiently
similar in some features to Neusticomys to warrant a phylogenetic
analysis.

MATERIALS AND METHODS

The description of the new species is based on conventional morpho-
logical characteristics and the terminology used follows Voss (1988).
Specimens were measured using dial calipers, with all measure-
ments provided in millimetres. The skeletal elements remaining in
the skins were observed by means of X-rays.

A parsimony analysis (PAUP Version 3.0) was carried out to
determine the position of the new species relative to other
ichthyomyines. Details of the eighteen characters used, listed
below, are given more fully in Voss (1988 pages 440-442); the

© The Natural History Museum, 1997

hypothesised primitive state, using the criteria defined by Voss, is
scored as ‘0’.

. Pelage: glossy (0); dull (1).

. Ventral pelage countershaded: absent (0); present (1).

. Tail: unicolored (0); bicolored (1).

. Philtrum: present (0); absent (1).

. Pinnae: large, visible above fur (0); small, concealed in fur (1).

. Superciliary vibrissae: present (0); absent (1).

. Plantar pads of manus: hypothenar pad separate, not fused with
third interdigital pad (0); hypothenar and third interdigital pads
fused (1); hypothenar and thenar pads fused respectively with
adjacent third and first interdigital pads (2).

. Fringing hairs on pes: weakly developed (0); well developed (1).
9. Lower third molar: entoconid-hypoconid cusp pair distinct (0):

m3 peglike, entoconid-hypoconid cusp pair absent or reduced to
a small conule (1).

10. Nasal bones: long, produced anteriorly beyond premaxillae (0);
short, truncated behind premaxillae (1).

11. Supraorbital foramina: on the lateral surface of the frontals,
within orbital fossae (0); on the dorsal surface of the frontals
between the orbital fossae (1).

12. Carotid arterial supply (see Voss, 1988 page 296): pattern 1 (0);
pattern 2 (1); pattern 3 (2).

13. Orbicular apophysis of maleus: present (0); absent (1).

14. Metatarsal configuration: WI>IV >>> V>1(0);IV>I1> 1H,
V>I1(1).

15. Omohyoid muscle: present (0); absent (1).

16. Gastric glandular epithelium: present (0); restricted (1).

17. Gall bladder: present (0); absent (1).

18. Bacular cartilage: tridigitate, medial digit lacking a calcified

centre (0); single digit (1); tridigitate, medial digit grossly

swollen with calcified core (2).

ADM fLWND —

oo

The character states for the new species were assessed by PJ but the
character state assessments for the other taxa were taken directly
from Voss (1988 Table 45, page 441). Characters of the visceral and
reproductive systems (characters 15—18 above) were unobservable
in the new taxon because of the lack of whole bodies. In an initial
analysis, all character states were unordered; in a second analysis,
multistate characters were ordered (as by Voss): 0 ~ 1 — 2 for
characters 7, 12 but also for character 18. For character 18, the order
recommended by Voss (1988) was 0 — 1; 0 — 2, a sequence not
readily handled by the analysis and affecting only one generic group

124

(Rheomys); Voss (personal communication) recommended an alter-
native ordering of this character (1 > 0 — 2) so in a third analysis,
characters 7 and 12 were ordered as above and character 18 by this
alternative.

ABBREVIATIONS USED IN THE TEXT

BMNH - The Natural History Museum [formerly British Museum
(Natural History)]

M1, M2, M3 respectively first, second and third upper molars
ml,m2,m3 ___ respectively first, second and third lower molars

ABBREVIATIONS USED FOR TAXA IN THE
PAUP ANALYSIS:

Ale Anotomys leander Thomas, 1906a

Ctr Chibchanomys trichotis (Thomas, 1897)
Cor Chibchanomys undescribed species

Thy Ichthyomys hydrobates (Winge, 1891)
Ipi Ichthyomys pittieri (Handley & Mondolfi, 1963)
Itw Ichthyomys tweedii Anthony, 1921

Nmo Neusticomys monticolus Anthony, 1921
Nve Neusticomys venezuelae (Anthony, 1929)
Rme Rheomys mexicanus Goodwin, 1959

Rha Rheomys raptor Goldman, 1912

Rtt Rheomys thomasi Dickey, 1928

Run Rheomys underwoodi Thomas, 1906b
RESULTS

Chibchanomys orcesi, sp. nov.

HOLOTYPE. BMNH 82.816, adult male, skin and skull; collectors’
number 148; collected 22 August 1981 by members of the Oxford
Expedition to Las Cajas from Lake Luspa, Las Cajas, Provincia
Azuay, Ecuador, 02°50'S 79°30'W, altitude 3700m.

PARATYPES. BMNH 82.815, adult male, skin and skull; collectors’
number 146, other details as for the holotype. BMNH 84.349, adult
male, skin and skull; collectors’ number 78; collected 7 August 1983
by members of the Combined Universities Expedition to Ecuador
1983, from Lake Llaviucu, Zorracucho Valley, Las Cajas, Provincia
Azuay, Ecuador, 02°51'S 79°O1'W, altitude 3100m.

DIAGNOSIS

An ichthyomyine species belonging to the genus Chibchanomys in
the following combination of features. Dorsal pelage dull; small
pinnae concealed in pelage of head; tail longer than head and body;
manus with five separate plantar pads; hindfoot broad with well
developed fringing hairs; supraorbital foramina open laterally within
orbits; carotid circulation pattern 1.

Differing from Chibchanomys trichotis in the following charac-
ters. Rhinarium light brown; philtrum present; nasals medium in
length, barely projecting anterior to premaxillae; orbicular apophy-
sis of maleus present; upper incisors slightly inclined medially; M3
and m3 reduced in size; anteromedian flexid absent or barely
indicated on anteroconid of m1; metatarsals I] >IV > IL>V>L.

DESCRIPTION
Tail subequal to or slightly longer than head and body (see Table 1

P.D. JENKINS AND A. BARNETT

Table 1 External and cranial measurements of Chibchanomys trichotis
and C. orcesi. Dimensions given as mean, plus or minus standard
deviation, followed by range, with sample size in parentheses.

C. trichotis _C. trichotis_ C. trichotis C. orcesi
Venezuela Colombia Peru Ecuador
Head and body 113.5 +5.50 105 + 1.63
length 105-120 (4) 125 (1) 102 (1) 103-107 (3)
Tail length 126.8 + 7.36 113.3 +6.18
115-133 (4) 131(1) 123 (1) 108-122 (3)
Hindfoot length 31.8 + 1.09 22 +2.16
30-33 (4) 30) 3312)) “3iha) 19-24 (3)
Ear length Uz WS) WLS) Sey
6-10 (4) 8 (1) 6 (1) 9.5-14 (3)
Weight (in grams) - - - S72 2-83
3541 (3)
Ratio of tail 1.12 + 0.02 1.08 + 0.06
length to head 1.10-1.15 (4) 1.05 (1) 1.21 (1) 1.02—1.16(3)
and body length
Ratio of tail 4.9+0.12
length to condylo- 4.7—5.0 (4) Sel) 5.2 (1) 4.3, 4.7 (2)
incisive length
Ratio of hindfoot 0.28 + 0.00 0.21 + 0.02
length to head 0.28-0.29 (4) 0.26 (1) 0.30 (1) 0.18—0.23 (3)
and body length
Condyloincisive 25.7 +1.01
length 24.3-26.9 (4) 25.9 (1) 23.8 (1) 23.0, 24.6 (2)
Diastema Length 6.3 + 0.54 - 33/5029
5.7-7.0 (3) 5.7 (1) 5.0-5.7 (3)
Length of upper 4.4+0.08 4.2 + 0.05
molars 43-45 (3) 4.4(1) 4.2 (1) 4.14.2 (3)
Incisive foramina 5.0 + 0.27 4.5 + 0.09
length 4.6-5.3(4) 5.1 (1) 4.5 (1) 4.44.6 (3)
Breadth of 1.2+0.11 1.4+ 0.05
incisor tips 1.0-1.3 (4) cl.2,1.3(2) 1.0(1) 1.4-1.5 (3)
Breadth of incisive 1.9+0.15 2.1 + 0.09
foramina 1.7-2.1 (4) 2.4 (1) 1.9 (1) 2.0-2.2 (3)
Breadth of 3.1+0.29 2.2 + 0.08
palatal bridge 2.7-3.4 (3) - PHO) ((l)) 2.12.3 (3)
Nasal length 9.0 + 0.41 9.3 + 0.29
8.5-9.5 (3) 8.1 (1) 8.5 (1) 8.9-9.6 (3)
Nasal breadth 3.0 + 0.05 3:2 O05
2.9-3.0(4) 3.0(1) 2.8 (1) 3.1-3.2 (3)
Interorbital breadth 4.7 + 0.13 4.6 + 0.08
45-4.8(4) 4.9(1) 4.3 (1) 4.54.7 (3)
Zygomatic breadth 13.4 + 0.66
12.3-14.1 (4) c13.9 (1) 11.7 (1) c12.8 (1)
Braincase breadth 13.4+0.27
13.0-13.7 (4) 13.8 (1) 12.4 (1) 1.9, 12.0 (2)
Ratio of inter- 0.35 + 0.02
orbital breadth to 0.33-0.37 (4) 0.36 (1) 0.35 (1) 0.38, 0.39 (2)
braincase breadth
Breadth of 1.1+0.05 1.1+0
zygomatic plate 1.0-1.1 (4) 1.0,1.2(2) 1.0(1) 1.1 (3)
Breadth of first 1.6+0.05 1.3+0.05
upper molar 1.5-1.6 (3) 1.5 (1) 1.4 (1) 1.3-1.4 (3)
Height of upper 4.6 + 0.33 4.3-4.5 (3)
incisor 4.1-5.0(4) 5.3 (1) 3.9 (1) 44+ 0.08
Depth of upper 1.3+0.12 1.2-1.3 (3)
incisor 11-144) 12) 1.0 (1) 1.3 +0.05
Breadth across 74+0.18
occipital condyles 7.2—7.6 (4) 7.6 (1) TeSialy) 6.6, 6.7 (2)

A NEW SPECIES OF CHIBCHANOMYS

Fig. 1 Live specimen of Chibchanomys orcesi.

Fig. 2 Skull of Chibchanomys orcesi (BMNH 1982.816) from left to right in dorsal, ventral and lateral view.

126

for measurements). Pelage soft, dense and woolly, dark brownish
grey dorsally, light grey ventrally; tail greyish brown, densely
haired, grey brown and brown hairs predominate proximally, with a
proportional increase of buff and cream hairs distally, extending
beyond tip in a short pencil. Distal portion of muzzle light grey in
young adults, cream in older individuals (age based on degree of
dental wear); rhinartum light brown in dry specimens; philtrum
present; conspicuous silvery-grey mystacial vibrissae present. Pin-
nae small, concealed by pelage; region of more-or-less conspicuous
light grey hairs ventro-lateral to pinnae. Manus with three interdigital
and two carpal pads. Well developed fringe of stiff hairs on margin
of metatarsus and digits of pes; claw of fifth digit extends beyond
first interphalangeal joint of fourth digit; claw of first digit reaches
midway along first phalange of second digit. See Fig. 1 for external
features visible in a photograph of a live specimen.

Skull (see Fig. 2) with moderately long nasals, overlapping nasal
orifice to conceal incisors in dorsal view but barely projecting
beyond premaxillae; rostrum short and narrow, naso-lacrymal cap-
sules evident in dorsal view; interorbital region moderately narrow
relative to braincase breadth (0.38, 0.39 (n = 2); frontals slightly
inflated, braincase moderately broad and long; posterior border of
incisive foramina between anterior roots of M1s, palatal foramina lie
between posterior roots of M1s; bullae slightly inflated; orbicular
apophysis of maleus present. Carotid circulation pattern 1, based on
osteological features (see Voss, 1988: 298).

Upper incisors moderately narrow, anterior enamel surface pale
buff, slightly inclined medially. No anteroloph on M1; small
posteroloph on M2; M3 small, protocone and paracone evident in
unworn dentition, posterior conule absent. Anteroconid of m1 sim-
ple or with slight indication of anteromedian flexid; no anterolophid
on m2; small posterolophids on ml and m2, small mesolophids
present or absent; m3 small, with small posterior basally positioned
conulid.

Metatarsal proportions: third metatarsal slightly longer than fourth,
fourth longer than second; all three far longer than first and fifth;
fifth longer than first. Configuration: II] 21V > Il > V >I.

ETYMOLOGY

This species is named in honour of Professor Gustavo Orcés, a
pioneer of Ecuadorian mammalogy. He was of great help to AB with
fieldwork organisation in Ecuador, and his kindness and knowledge
were a source of inspiration.

DISTRIBUTION AND ECOLOGY

Known only from Las Cajas Plateau, Ecuador, where specimens
have been recorded from three localities: Lake Luspa, Lake Llaviucu,
Zorracucho Valley and Lake Torreadora. All specimens were trapped
in close proximity to fast-flowing streams at altitudes ranging from
3100m to 4000m, in high-altitude moorland vegetation (paramo)
(see Barnett, 1992). For more precise details of the habitat at each
site and notes on diet see Barnett (1997).

COMPARISON WITH C. TRICHOTIS

The new species is similar in external appearance to C. trichotis,
except that the pelage is paler and slightly harsher, and the rhinarium
is light brown in dry specimens of C. orcesi, black in C. trichotis. A
philtrum is present in C. orcesi but absent in C. trichotis.
Chibchanomys orcesiis smaller in external size and averages smaller
in cranial size than all known specimens of C. trichotis, with the
exception of the single specimen from Peru (see below for com-
ments on the status of this specimen). Both species of Chibchanomys
are similar in external proportions, except that the hindfoot is
proportionately shorter in C. orcesi (see Table 1). The metatarsal

P.D. JENKINS AND A. BARNETT

configuration differs in the two species: IV > III > Il=V>Iin C.
trichotis; W121V > IL >> V > Lin C. orcesi. The two species differ in
the following cranial features: while the nasals of both species are of
comparable length, those of C. orcesi are slightly broader and barely
project anterior to the premaxillae, unlike those of C. trichotis,
which project anteriorly and conceal the incisors and nasal orifice in
dorsal view. In lateral view, the globose braincase of C. trichotis
rises abruptly in the frontal region, unlike the narrower and less
inflated braincase of C. orcesi; the braincase is slightly broader and
the breadth across occipital condyles is greater in C. trichotis (see
Table 1). The orbicular apophysis of the maleus is present in C.
orcesi but absent in C. trichotis. The upper incisors of C. orcesi are
less delicate and slightly broader than those of C. trichotis (see Table
1), the anterior enamel surface of C. trichotis is cream coloured and
not medially inclined unlike C. orcesi. The third upper molar is
smaller relative to M1 and M2, and m3 is smaller relative to m1 and
m2 in C. orcesi than in C. trichotis. The anteromedian flexid is
absent or barely indicated on the anteroconid of m1 in C. orcesi but
present in C. trichotis, dividing the anteroconid into small but
distinct lingual and labial conulids. The posterior conulid of m3 is
positioned more basally in C. orcesi than in C. trichotis (when
present).

According to Voss (1988) the young specimen that he identified as
C. trichotis from Peru differs from the northern specimens of C.
trichotis in several features: the braincase is much less inflated, the
occipital condyles are slightly broader, the bullae are somewhat
smaller and an indistinct philtrum is indicated; features that resem-
ble those of the new species. Voss mentioned that these differences
might indicate that southern populations of Chibchanomys are
phenotypically distinctive from their northern counterparts. Unfor-
tunately it has not proved possible to examine the Peruvian specimen,
although information on it was kindly provided by Mark Hafner
(personal communication). It is possible that the Ecuadorian and
Peruvian specimens are conspecific but additional material and
more extensive comparisons are required to elucidate the status of
the latter specimen.

COMPARISON WITH OTHER ICHTHYOMINE GENERA

Chibchanomys is readily distinguished from Anotomys, Ichthyomys
and Rheomys (see Voss, 1988). Chibchanomys and Neusticomys
differ from other ichthyomyines in showing carotid arterial circula-
tion pattern | and in the distribution of the glandular epithelium
around the stomach. Chibchanomys differs from Neusticomys in
having small pinnae concealed in the pelage (pinnae obvious in
Neusticomys); ventral countershading present (absent in
Neusticomys); tail longer than head and body (tail shorter than head
and body in Neusticomys); the hindfoot is broader with longer digits
and the fringing hairs are well developed (narrower with shorter
digits and less developed fringing hairs in Neusticomys). The new
species does however share several features with Neusticomys which
are not exhibited by C. trichotis, such as the similar metatarsal
configuration and presence of a philtrum, while the orbicular apo-
physis of the malleus is also present in some species of Neusticomys.

RESULTS OF THE PHYLOGENETIC ANALYSIS

There is evidence in support of the ichthyomyines as a monophyletic
group of the subfamily Sigmodontinae (sensu Carleton & Musser,
1984) (see Voss, 1988). In contrast, evidence in support of the
monophyly of the Sigmodontinae is lacking and a tribal level
classification of this subfamily, while convenient in many respects,
is unsatisfactory from a phylogenetic point of view, making difficult
the choice of satisfactory outgroups for phylogenetic analyses (see
Voss, 1988: 436-438, 1991: 33-37; Carleton & Musser, 1989: 53—
55; Voss & Carleton, 1993: 21-22). The necessity of making such a

A NEW SPECIES OF CHIBCHANOMYS

Table 2. Matrix showing character state distributions among 12
ichthyomyine species (for details see text and Voss, 1988). The character
state assessments for all taxa other than C. orcesi were taken directly
from Voss (1988 Table 45, page 441). Character states for the new
species were assessed by PJ except for those of the visceral and
reproductive systems (characters 15—18) which were unobservable in
the new taxon, so scored as ‘?’.

Characters
eee a ee Ome, (o 00) eNO) eats 4 tL 6, 17) 8

Taxa
Ale
Ctr
Cor
Thy
Ipi
Itw
Nmo
Nve
Rme
Rha
Rtt
Run

MSM S) (SS) (SSS Siete
St oo ee ee
Se) SSS) (Sa) La aay (=)
eoOoorc”joq°”ccocdmh6cOlULcOlmrR hE
eS Sa CS) ne
a a)
NrreNocCcC oOo So co -
oon ee ee
SiO SS 1S (Si oS ic
Oe OS OO On = OOS
NNNNOCONNN COCK
(SS = SS SV) (Se) fe)
—— eS eS OOKe Se HS Oe eS
SESS) SW SiS ays) (SS) a
— RO OOO RR SE IO
Oo Oo © © 6 Oi = 3S
NNONNOCCOCOCyN O-

choice has been avoided in the current study, since it is aimed at
determining the level of affinity of the new taxon to other
ichthyomyines, rather than seeking to add any new dimension to the
phylogenetic status of the ichthyomyines as a group. Instead an
hypothetical outgroup was constructed in which all character states
were assessed as primitive, which was used to root the trees.
Using branch and bound algorithms, a search was made of the
character data summarised in Table 2. In the analysis in which all
character states were unordered, the length of the shortest tree was
equal to 32 character state transformations and six trees were
retained. In each of the other analyses (with the multistate characters
7 and 12 ordered, and character 18 varyingly ordered) the length of
the shortest tree was equal to 33 character state transformations but

127

only three trees were retained. The variation in treatment of charac-
ter 18, was not considered to be particularly important in this study,
since character state 2 is exhibited only by taxa of the genus
Rheomys. In both of the latter analyses C. orcesi and C. trichotis are
non-monophyletic in all three trees and also in the semistrict consen-
sus of these trees (see Fig. 3). The only evidence of a monophyletic
generic grouping shown in the semistrict consensus tree is for
Ichthyomys, and this tree is similar in most respects to the most
parsimonius hypothesis of ichthyomyine relationships shown by
Voss (1988: Fig.88).

DISCUSSION

There is obvious conflict in that the results of the phylogenetic
analysis do not support the generic classification currently in use.
The morphological data is sufficiently persuasive to conclude that,
on the available material, the new taxon is correctly attributed to the
genus Chibchanomys as currently construed.

ACKNOWLEDGEMENTS. We are grateful to Jim and Theresa Clare for
unpublished information on Chibchanomys and to the late Professor Gustavo
Orcés for support and encouragement during fieldwork. We are indebted to
Robert Voss (American Museum of Natural History) who initially recognised
the uniqueness of the new species and who provided much helpful advice
when commenting on drafts of the manuscript. Particular gratitude is owed to
Darrell Siebert (Natural History Museum) for guidance with the phylogenetic
analysis, and for constructive comments and criticism of the manuscript.
Mark Hafner (Museum of Zoology of Louisiana State University) kindly
provided photographs and measurements of the specimen of C. trichotis from
Peru. As always the staff of the Natural History Museum freely provided
support; in particular we thank Richard Sabin (Mammal Group) and Deryck
Jones (Electronics) for the X-rays, Phil Hurst (Photographic Unit) for the
photographs, while Clive Moncrieff (Biometrics) patiently assisted with the
PAUP analysis.

Fig. 3 Semistrict consensus tree showing hypothetical phylogenetic relationship of Chibchanomys orcesi to other taxa of ichthyomyine rodents.

Consistency Index 0.636, Retention Index 0.786.

128

REFERENCES

Anthony, H.E. 1921. Preliminary report on Ecuadorean mammals No.1. American
Museum Novitates (20): 1-6.

1929. Two genera of rodents from South America. American Museum Novitates
(383): 1-6

Barnett, A. 1992. Notes on the ecology of Cryptotis montivaga Anthony, 1921 (Insect-
ivora, Soricidae), a high-altitude shrew from Ecuador. Mammalia 56 (4): 587-592.

1997. The ecology and natural history of a fishing mouse Chibchanomys spec.
nov. (Ichthyomyini: Muridae) from the Andes of southern Ecuador. Zeitschrift fiir
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BBC-National Geographic wildlife film ‘Avenue of the Volcanoes’

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Bull. nat. Hist. Mus. Lond. (Zool.) 63(2): 129-136

A new species in the asterinid genus Patiriella
(Echinodermata, Asteroidea) from Dhofar,
southern Oman: a temperate taxon in a
tropical locality

POD ~~ > Fr | a
ANDREW C. CAMPBELL _ 0 aaageilh ipscsiie deine
School of Biological Sciences, Queen Mary and Westfield College, University of London, Mile End Road,
London E]1 4NS, UK.

FRANCIS W.E. ROWE
Goldbrook Boarding Kennels and Cattery, Nuttery Vale, Cross Street, Hoxne, Suffolk, IP21 5BB, U.K.

CONTENTS

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SYNOPSIS. The status of the hitherto temperate-water asterinid genus Patiriella is briefly discussed, including in its diagnosis
details of actinal plate and ventral/lateral plate arrangement. A new species, P. paradoxa, is described from shallow water on the
Dhofar coast of Oman. The distribution of Patiriella on the Dhofar coast is discussed in relation to the peculiar oceanographic

Issued 28 November 1997

conditions and vicariant events in the region.

INTRODUCTION

Although one of us (ACC) made extensive collections along the
coasts of the Sultanate of Oman between 1983 and 1990 (Campbell
and Morrison, 1988 and Marsh and Campbell, 1991), the echinoderm
fauna of southern Arabia remains incompletely known. However,
those echinoderms which have been recorded are tropical species or
endemic species with tropical affinities (Clark and Rowe, 1971;
Price, 1982; Campbell and Morrison, 1988; Marsh and Campbell,
1991). The discovery (by FWER) of three specimens of a new
species of the predominently temperate-water, asterinid starfish
genus Patiriella Verril (1913) among the Omani shallow water (less
than 10m) collections from two sites in Dhofar therefore poses a
paradox.

Here we briefly discuss the status of the genus Patiriella and
describe fully the new species. This work attempts to explain the
presence of an otherwise temperate-water echinoderm genus in this
tropical location, given the peculiar local oceanic conditions which
include upwelling (see summary by Campbell and Morrison, 1988),
and past vicariant events.

The type specimens have been deposited in the Natural History
Museum, London U.K.

© The Natural History Museum, 1997

SYSTEMATIC DESCRIPTION

PATIRIELLA Verrill

Patiriella Verrill, 1913: 483; 1914: 263; Fisher, 1919: 410; H.L.
Clark, 1946: 134; Dartnall, 1971: 39; A.M. Clark and Courtman-
Stock, 1976: 80; A.M. Clark, 1983: 365; A.M. Clark and Downey,
1992: 192.

TYPE SPECIES. Asterina (Asteriscus) regularis Verrill, 1870
(1867); by original designation.

DIAGNOSIS
After A.M. Clark in A.M. Clark and Downey (1992), amended.

A genus of Asterinidae with five to about eleven short rays (R up
to c. 60mm); pentagonal to stellate in outline; aborally arched;
primary abactinal plates in two ‘fields’, a slightly irregular radial
(usually the midradial and first dorsal-lateral series on either side)
‘field’ and a regular lateral ‘field’ on either side; mid-radial abactinal
plates with proximal edge trilobed or simply crescentic; the proxi-
mal concave sides of abactinal plates subtend spaces with usually
one to several papular pores, separated by one to several small
secondary plates; abactinal armament comprising relatively few
(<40 per plate) very coarse, almost granuliform, multipillared

130

spinelets; actinal plates distinctly aligned in oblique series between
the second or third, and subsequent adambulacral plates and the
inferomarginal plates and delimiting a membranous, proximal, tri-
angular area which is usually filled by several plates; ventral-lateral
angle of rays supported internally by abactinal plates which meet the
actinal plates by virtue of the oblique alignment of both; towards the
ray base as the ventral-lateral angle becomes less acute with ray
depth, totally internalised plates, spanning between the abactinal
and actinal plates, can be found; actinal armament coarse, short
spines, mostly single, not more than two per plate; furrow spines
usually two (sometimes one) per plate; subambulacral spines one
(rarely two); suboral spines one to four or none; no pedicellariae.
Species included: Patiriella brevispina H.L. Clark, 1938;
Asterias calcar Lamarck, 1816; Asteriscus calcarata Perrier, 1869;
Asteriscus chilensis Lutken, 1859; Asterina dyscrita H.L. Clark,
1923; Asterias exigua Lamarck, 1816; Asterina fimbriata Perrier,
1875; Asterina gunni Gray, 1840; Patiriella inornata Livingstone,
1933; Asterina oliveri Benham, 1911; Patiriella parvivipara
Keough and Dartnall, 1978; Patiriella pseudoexigua Dartnall, 1971
(with subspecies pacifica (Hayashi, 1977, as Asterina); Asterina
(Asteriscus) regularis Verrill, 1870 (1867); Patiriella vivipara

A.C. CAMPBELL AND F.W.E. ROWE

Dartnall, 1969b; Patiriella paradoxa sp. nov.

Other species Patiriella nigra H.L. Clark, 1938 and Patiriella
obscura Dartnall, 1971 are considered to be conspecific with P
oliveri (Benham) and P. pseudoexigua Dartnall respectively by
Rowe (in Rowe and Gates, 1995); Patiriella tangribensis Domantay
and Acosta, 1970, is inadequately described and cannot confidently
be assigned to this genus.

REMARKS

The genus Patiriella Verrill, 1913, has had a rather chequered
history. It has been considered a valid taxon by most recent authors
(e.g. Fisher, 1919; H.L. Clark, 1928; 1938; 1946; Livingstone, 1933;
Madsen, 1956; Dartnall, 1971; A.M. Clark and Rowe, 1971; A.M.
Clark and Courtman-Stock, 1976; A.M. Clark, 1983; A.M. Clark
and Downey, 1992) or a synonym of Asterina Nardo, 1834 (e.g. H.L.
Clark, 1916; 1923; Hayashi, 1940; 1977; Mortensen, 1933 (as a
subgenus of Asterina)). The history and current status of Patiriella
has been most recently discussed by A.M. Clark (1983; 1992 (in
Clark and Downey)) who commented (1992:178) that “The very
coarse and abbreviated, almost granuliform, armament of the upper
side may warrant a supra-specific distinction from A. gibbosa of P.

Fig. 1 a, Scanning electron micrograph of the actinial surface of Asterina gibbosa from Plymouth U.K. Scale bar = 5mm . G = gonopore; b, Scanning
electron micrograph of the actinial surface of ‘Asterina’ cepheus from Wadi Haart, Sadh, Dhofar, Southern Oman. Scale bar = 5mm

A NEW SPECIES OF PATIRIELLA

regularis together with P gunni (Gray) and P. calcar (Lamarck)
from Australia, also P exigua (Lamarck) which extends from the
Indo-West Pacific into the S.E. Atlantic’. She thought reassessment
of the rank of Patiriella was best left to ‘one of several Australasian
specialists to determine . . .’, retaining the species exigua as a mem-
ber of the genus Patiriella. A new review of the family Asterinidae,
including a reappraisal of the status of Patiriella is being undertaken
by F.W.E.R. elsewhere. However, whilst in the present paper we
acknowledge that a very close relationship exists between the genera
Asterina, Patiriella and Patiria (Gray, 1840) (Patiria is considered a
synonym of Asterina by Hayashi (1940) and A.M. Clark (1983 (in
Clark and Downey, 1992)); we accept their separate generic status,
until the matter is more clearly resolved, such a resolution being
outside the scope of this paper '. In taking this stance, we have
amended A.M. Clark’s in Clark and Downey, 1992) diagnosis of
Patiriella to include a description of the alignment of actinal plates
(included by Verrill (1913) in his diagnosis of the genus) and the
internal alignment of actinal/abactinal plates at the ventral/lateral
angle (described by Verrill (1913) as a feature of the family
Asterinidae). As a taxonomic character, actinal plate alignment has
been largely ignored. The exception was Fisher (1917; 1919) who
thought this a useful character when distinguishing his new genus
Paranepanthia (type-species Nepanthia platydisca Fisher, 1913) in
which the plates are aligned obliquely across the actinal surface,
from Asterina in which he saw the actinal plates forming chevrons
across each interradial area, the plates being aligned only parallel to
the furrows along each ray. Fisher (1919) was, however, comparing
Paranepanthia platydisca with the species ‘Asterina’ cepheus (Miiller
& Troschel), and ‘A’. coronata (von Martens) and not the type
species A.gibbosa. In accepting Asterina in this sense, Fisher was
clearly not familiar with the fact that in the type-species, A. gibbosa,
the actinal plates, as in Patiriella and Paranepanthia, are clearly
aligned obliquely between furrow and margin Fig la & b (there is no
question, however, that Paranepanthia is a valid genus (F.W.E.R.).
A.M. Clark (1971, in A.M. Clark and Rowe) noted Fisher’s work
when identifying two specimens of an undetermined species of what
she considered a Paranepanthia from Zanzibar. She considered
(1971:71) the importance of actinal plate arrangement, as a func-
tional and taxonomic character, required investigation. However,
A.M. Clark (1983) did not expand further on this matter in her more
recent revision of the family Asterinidae. Although a recent survey
of the family by one of us (RW.E.R.) does show actinal plate
arrangement to be a useful character in distinguishing some of the
genera, actinal plate arrangement undoubtedly has a functional role
which is probably micro-habitat related. We include description of
the internal structure of the ventral/lateral angle in our diagnosis
following A.M. Clark (1983) who concluded that differences in
arrangement of these internal plates were important in indicating
taxonomic affinities, a conclusion supported herein, though on the
basis also, that it may relate to ray shape within the family.

Patiriella paradoxa sp. nov.

DIAGNOSIS. Stellate species of Patiriella with five rays; abactinal
plates of aboral, radial ‘field’ tri-quadrilobed, forming a delicate
reticulum; papular areas each with three to six papulae and one to
two secondary plates; no suboral spines; proximal actinal plates
each with a single spine.

' Species attributed to Asterina s.s. from outside the Atlantic region are NOT conge-
neric with the type-species, A. gibbosa (Pennant), and require reallocation to other
existing or new asterinid genera according to Rowe (in Rowe and Gates, 1995); the
suggested recognition of Asterinides Verrill (1913) (type-species A. folium (Liitken)) as
a valid subgenus of Asterina, by A.M. Clark (1983; 1992 (in Clark and Downey)) is
supported, but at generic level, by Rowe (in Rowe and Gates, 1995).

131

HOLOTYPE. BMNH (dry) BMNH 1997.1016

TYPE LOCALITY. 0.5 km southeast of Wadi Haart and about 4km
north east of Sadh village, Sultanate of Oman (17° 04'N, 55° 06'B),
intertidal, just below level of the conspicuous barnacle Tetraclita
squamosa rufotincta, collected together with ‘Asterina’ cepheus
(Miiller & Troschel, 1842), A.C. Campbell 4 May 1987 (210050201)
(fig.2).

MATERIAL. In addition to the holotype, two paratypes (dry).
Paratype 1 BMNH 1997.1017, collected at the same locality with the
holotype. Paratype 2 BMNH 1997.1018, Raaha, 2.5km west of Wadi
Ayn, Sultanate of Oman (16° 58'N, 54° 50'E), c. 8m depth, on rocks
amongst corals, coll. A.C. Campbell, 5 Dec., 1986 (210050202)
(fig.2).

EtYMOLOGY. The species name (Lat. paradoxum) refers to the
unexpected occurrence of this predominently temperate-water ge-
nus along an otherwise tropical coastline.

DESCRIPTION OF HOLOTYPE. (Figs 3 a & b; 4a & b). Specimen
stellate in outline, R=17.8mm, r=9.0mm, R/r=1.98; br=10.3mm
(across base of ray between first superomarginal on each side), R/
br=1.73. Orally flat, aborally arched. Rays more or less elongate
triangular, tapering from a relatively wide base to a rounded tip.
Centre of disc delimited by a complete ring of prominent, spinelet-
bearing plates, outside of which a second, more or less complete ring
of less prominent plates evident. The prominent, ovate madreporite
occurs in interradius CD.

Beyond the central disc, the imbricating primary abactinal plates
form two ‘fields’ along the rays. Radial ‘field’ comprising an
irregular, zig-zag series of mid-radial (carinal) plates and the first
dorsal-lateral row on each side. Shape of this ‘field’ elongate-leaf-
shaped along the ray, tapering proximally and distally and widest
at about ’R. Denuded plates deeply notched, mostly quadrilobed
(X-shaped), but, at least between ¥2—4R, some mid-radial plates
tri-lobed (Y-shaped), the plates together forming a relatively deli-
cate reticulum. Papular areas, between the plates, are relatively
large, up to 0.8mm diameter, each subtending 3-6 papulae
between which | or 2 minute, spinelet-bearing secondary plates
usually occur. Primary plates with crystal bodies, except on their
proximal, crescentic ridge which carries 5—7 granuliform spinelets
in a single series; spinelets range from c. 0.30mm long x 0.12mm
wide (straight sided) to about 0.37mm long x 0.14mm wide (these
larger spinelets becoming club-shaped (0.18mm wide) towards
their tip). Lateral ‘field’ comprising about 10 regularly arranged
rows of plates at the base of the rays, closely imbricate with a deep
proximal notch subtending 1-3 papulae occurring in the proximal
half of the first 5 rows of plates, the papulae not extending to the
superomarginal line. Lateral ‘field’ plates with crystal bodies and
from 2-7 spinelets.

Ventral-lateral margin sharply delimited, however neither
inferomarginals nor superomarginals significantly larger than im-
mediately adjacent actinal or abactinal plates respectively. Infero-
marginal plates aligned in the same plane as the actinal surface,
slightly protrude laterally and bear a group of 2—3 minute spinelets.
Superomarginal plates aligned vertically, correspond with the
inferomarginals below them and bear 1—2 spinelets.

About 10 rows of actinal plates counted; the first two extending to
the tip of the ray; plates are aligned both parallel to the furrow but
also distinctly obliquely across the actinal surface between the
adambulacral plates and inferomarginal plates. Oblique alignment
accentuated both by the actinal spination and narrow, shallow

A.C. CAMPBELL AND F.W.E. ROWE

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A NEW SPECIES OF PATIRIELLA

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Fig.3 a, Photograph of Patiriella paradoxa, Holotype, from Wadi Haart, Sadh, Dhofar, Southern Oman. Abactinial view. Scale in mm; b, Photograph of
Patiriella paradoxa, Holotype, from Wadi Haart, Sadh, Dhofar, Southern Oman. Actinial view. Scale in mm.

furrows occurring between the oblique lines of plates. First com-
plete oblique line of plates arises opposite the second adambulacral
plate in each furrow, creating a triangular area bounded by the first
adambulacral plates, distal edge of the oral plates and first oblique
series in each actinal intermediate area. This proximal area occupied
by 2 (interradii BC, DE and EA), 3 (interradius AB) or 4 (interradius
CD) plates. A small circular patch of non-calcified skin (up to
0.6mm diameter) occurring adjacent to the distal edge of the oral
plates in interradii AB, CD and EA. Actinal plates each bear a single,
tapering spinelet (up to 0.64mm long x 0.30mm at base) on the
proximal 80% of the actinal surface, the remaining distal plates with
2 small spinelets.

There are 28-29 pairs of adambulacral plates to each furrow.
These are twice as wide as long. First 12—14 plates bear 2 tapering
furrow spines, proximalmost more slender and shorter than distalmost
spine (on adambulacral 3: distalmost spine measures 1.0mm long x
0.35mm wide, at base tapering 0.20mm wide near tip; proximalmost
spine measures 0.78mm long x 0.23mm wide at base, tapering to
0.14mm wide near tip). Proximalmost spine becomes rapidly smaller
and peg-like towards the 12th—14th plate, beyond which the
adambulacral plates each bear a single furrow spine. A single,
stouter, subambulacral spine (on adambulacral 3: 0.87mm long x
0.29mm wide, more or less cylindrical) occurs on each adambulacral
plate. The oral plates each with 4 oral (furrow) spines of which the
apicalmost is longest. No suboral spines.

Gonopores not occurring on the oral surface and cannot be
distinguished aborally.

CoLour. Dried holotype is a uniform, pale ‘museum’ buff colour.
Colour in life is not recorded.

PARATYPE 1. BMNH 1997.1017 has R=19.0mm, r=7.9mm.

R/r=2.4; br=9.4mm, R/br=c.2. Generally very similar in appearance
to the holotype, differing only in minor detail. Two furrow spines
occurring only on the 1st-4th adambulacral plates, thereafter the
plates with a single furrow spine. Small patch of non-calcified skin
occurs adjacent to the distal edge of the oral plates in each interradius,
this proximal actinal triangle being filled by 1 (interradii CD, DE) or
2 (interradii EA, AB, BC) plates.

PARATYPE 2. BMNH 1997.1018 has R=19.0mm, r=10.00mm,
R/r=1.9; br=11.2 mm, R/br=1.7. The dry specimen is contorted. It
differs from the holotype and paratype | in the following features.
Rays are slightly broader at their bases relative to their lengths.
Aborally, centre of disc not delimited by a prominent ring or rings of
plates. Radial ‘field’ of abactinal plates is compact, plates closely
imbricating with only the ridge evident and bearing 5-7 spinelets.
Papular areas contain 1-4 papulae and occasionally a single second-
ary plate. Actinal spinulation coarser, otherwise the plate arrangement
closely similar to the other type specimens. Actinal proximal trian-
gle filled by 1 or 2 plates, no patches of non-calcified skin evident.
Some 36 pairs of adambulacral plates along each furrow; each
bearing a single furrow and single subambulacral spine, with the
exception of the first plate with a second, smaller proximal furrow
spine. Furrow and subambulacral spines flattened along their length
with a squared-off tip. Several furrow spines spaced along length of
the furrow, with bifid tip. Oral plates and spines are similar to those
of both the holotype and paratype 1. The paratype 2 is uniformly
light grey in colour.

HABITAT. The holotype and paratype | were collected on 4.5.87,
0.5km. south east of Wadi Haart and about 4 km. north east of Sadh
village. These were collected, together with ‘Asterina’ cepheus
intertidally on a gently shelving exposed rocky shore with tide pools
just below the level of the conspicous barnacle Tetraclita squamosa

A.C. CAMPBELL AND F.W.E. ROWE

Fig. 4 a, Scanning electron micrograph of the abactinial surface of Patiriella paradoxa from Wadi Haart, Sadh, Dhofar, Southern Oman. Scale bar: 5mm;
b, Scanning electron micrograph of the actinial surface of Patiriella paradoxa from Wadi Haart, Sadh, Dhofar, Southern Oman. Scale bar: 5mm.

rufotincta. During the monsoon period, July-September, the middle
shore is richly cloaked with the green alga Ulva sp. and brown algae
develop on the lower shore. These growths are burnt off by Decem-
ber, and then the intertidal remains almost alga free until the next
south west monsoon. It is worth noting that perennial beds of the
kelp Ecklonia radiata have been recorded off Sadh head (Barratt et
al., 1986) approximately 5 km. to the south west of this site.
Ecklonia may actually occur much closer as fragments were found
washed up on the beach.

Paratype 2 was collected on 5.12.86 at Raaha 2.5 km. west of
Wadi Ayn at 8 m depth on rocks amongst corals. This site faced south
south east and comprised a sandy cove bordered to the east by a ridge
of metamorphic rock sloping down to sand at 10 m. The rock ridge
was well covered with many scleractinian colonies, especially
Acropora sp. During the south west monsoon period this ‘coral
garden’ became completely overgrown with the brown alga
Sargassopsis zanardini. At the time of the collection the S. zanardini
growths had broken up and dispersed. No Ecklonia radiata was seen
growing in the immediate vicinity.

DISTRIBUTION. Known only from the type localities on the coast

of Dhofar, southern Oman.

REMARKS. Based on arrangement and shape of skeletal plates and
their armament, paradoxa is without doubt congeneric with Patiriella
regularis, the type species of Patiriella. It differs from that species,
as with the majority of its congeners in the form of the radial ‘field’
of abactinal plates, the high number of papulae per papular area and
absence of suboral spines. The absence of suboral spines is shared
with four species from the southern Australian coast. Of these P.
parvivipara is a small, precociously viviparous, pentagonal, cush-
ion-shaped star; P. brevispina and gunni have six, non-projecting
rays, and are more or less cushion-shaped and hexagonal in outline.
Additionally, these species differ from P. paradoxa in the increased
frequency of two spines on each of the proximal actinal intermediate
plates, and gunni possesses two subambulacral spines per
adambulacral plate. P paradoxa may appear to be most closely
related to P. calcar, differing most obviously in having a more
delicate abactinal plating and armament and having 5 instead of 7—
11 short rays. Clearly the geographical isolation of paradoxa from
its congeners has resulted in the evolution of a combination of
characters which isolates it within the genus.

A NEW SPECIES OF PATIRIELLA

ZOOGEOGRAPHICAL CONSIDERATIONS.

The genus Patiriella, as accepted herein, is distributed predominently
in temperate seas. No fewer than 10 of the 15 species included occur
in Australian waters (Rowe & Gates, 1995). Six of the species
(brevispina, calcar, gunni, inornata, parvivipara and vivipara) are
endemic to Southern Australia. Two species range across the Tasman
Sea between southeastern Australia and New Zealand (regularis) or
Lord Howe Island and Kermadec Islands (oliveri). One species
(pseudoexigua) is essentially tropical, ranging from northeastern
Australia north to Japan. The somewhat ubiquitous P. exigua ranges
from southern Australia, westward across the southern Indian Ocean
to St. Helena Island off southwestern South Africa in the southern
Atlantic. Of the other species, dyscrita is endemic to South Africa;
calcarata is endemic to Juan Fernandez Islands; chilensis occurs
between Chile and Peru and, fimbriata is distributed from southern
Chile to the Falkland-Magellan area and southern Argentina (A.M.
Clark, 1993). With the exception of fimbriata which has a known
depth range from intertidal to c. 300m, species of Patiriella are
essentially intertidal, occurring at most to about 30m depth (A.M.
Clark, 1993).

The occurrence of a species of Patiriella isolated on the Dhofar
coast of Oman, in the tropical, northwestern Indian Ocean, is
difficult to explain; whether as the result of distributive, accidental or
vicariant events.

To seek explanation by a distributive means requires knowledge
of reproductive strategies within the genus. Unfortunately, we have
been unable to determine reproductive strategy in PR paradoxa
through examination of the gonads, since the three specimens
collected to date were preserved and dried. Since gonopores do not
occur on the oral surface (see p. 133), and it is clearly not viviparous,
then we assume aboral gonopores occur in Pparadoxa and the
strategy involves either planktotrophic or lecithotrophic larvae.
However, life histories of at least seven species of Patiriella occur-
ring in Australia, including P. regularis which also occurs in New
Zealand have been determined (Byrne, 1991; 1992; Byrne and
Barker, 1991). Of these, regularis exhibits an indirect/planktotrophic
developmental pattern with feeding bipinnaria and brachiolaria
larvae; gunni, calcar, pseudoexigua exhibit a direct/lecithotrophic
developmental pattern with planktonic non-feeding brachiolaria
larvae; exigua exhibits direct/lecithotrophic developmental pattern
with a benthic non-feeding brachiolaria larva, vivipara and
parvivipara exhibit a direct/viviparous pattern of development as
intra-ovarian brooders, without larvae. The occurrence on the the
Dhofar coast of Oman of Patiriella originating via larval distribution
from either southern Africa or the Australasian region, even though
the nearest congeners are P. dyscrita (S. Africa) and P. pseudoexigua
(recorded as P. exigua by Koehler, 1910), from the northeastern
Indian Ocean, is difficult to envisage. The present-day water cur-
rents of the Indian Ocean must have been established for at least the
last 15—10my, following separation of the Indian Ocean (see Adams,
1981; van Andel, 1981). The isolation of P paradoxa, on the Dhofar
coast suggests a genetic isolation which is not receiving input from
other parts of the generic range.

Anexplanation of the origin of the temperate-water genus Patiriella
on the tropical Omani coast in association with a secondary agent is,
however, not at first sight, unreasonable to propose. For instance,
Dartnall (1969a) considered the New Zealand species P. regularis
had been introduced into Tasmanian waters amongst oyster spat. The
distribution of P. exigua from southern African to St. Helena Island,
in the south Atlantic, has been attributed, by Mortensen (1933) to
probable transport on the holdfasts of the kelp Ecklonia, which

135

though not established itself on St. Helena was washed ashore there.
A.M. Clark (1992, in A.M. Clark and Downey) considered this a
reasonable supposition since exigua is not recorded from Ascension
Island, to the north of St. Helena where the current is from the east.
The occurrence of Ecklonia radiata, otherwise known only from
around the coasts of South Africa, Australia and New Zealand,
which has been recruited to the southeastern coast of Arabia via the
deep Antartic current from the south (Sheppard, 1992, in Sheppard,
Price and Roberts) might have provided a ‘raft’ for Patiriella to reach
the Dhofar coast, originating either from South Africa or the
Patiriella-species-rich southern coast of Australia. However, such
‘rafting’ or ‘accidental’ introductions tend only to extend species
range, as it has done with Ecklonia. To seek a solution here for
Patiriella requires invocation of the subsequent evolution of the
species P. paradoxa and extinction of its ancestor on the coast of
Dhofar. Once again, the genetic and geographic isolation of P.
paradoxa would not appear to support such an arguement of its
origin on the Dhofar coast, even from its geographically nearest
congeners (see above).

The lack of fossil history of the family Asterinidae does not
preclude ancient history. It may be more productive, therefore to
seek a linkage between the distribution of Patiriella (particularly
considering its very close relationship with Asterina s.s., from the
Atlantic, and Patiria, from the North Pacific) and vicarient events in
the Indian Ocean. The oceanographic and geological configurations
of the area appears to have been relatively stable for at least the last
15-10 my (Powell et al., 1981; Adams 1981).

Coincident with this time the complete opening of Drake’s Pas-
sage (25-15 mya) established the circum-Antarctic current and a
sharp drop occurred in surface and bottom sea-water temperatures
(van Andel, 1981). It can be concurred that the present-day areas of
upwelling, including those within the Indian Ocean were also
established at that time. This is of interest for two reasons. Firstly, a
major area of upwelling is known off the coast of Dhofar (fig. 2).
This produces turbulent, nutrient-rich surface waters and almost
temperate conditions (minimum recorded water temperature of
15.9°C near Sadh; Savidge, et al., 1986) which prevail in the coastal
region of Dhofar for at least 4 months of the year (June—mid
September) (see Currie et al., 1973; Campbell and Morrison, 1988;
Miller and Morris, 1988 for details) and this is coincident with the
known distribution of P paradoxa. Secondly, the general temperate-
water distribution of species of Patiriella (as included herein),
Asterina s.s. (including only species gibbosa, phylactica, stellifera
and possibly pancera; according to FWER, unpublished) and Patiria
(including species miniata, pectinifera and possibly minor; accord-
ing to FWER, unpublished), which appear (with the exceptions of
Patiriella pseudoexigua and Asterina stellifera along the tropical
west coast of Africa, part of its range) not to extend into locations
much, if at all, above the 20°C isotherm (distribution data taken from
A.M. Clark, 1993). The implication of the present-day distribution
pattern of these three genera is that their ancestor was more wide-
spread in cooler parts of the Tethyan system (possibly during the
later part of the Oligocene epoch (c. 40-25 mya) (see van Andel,
1981)) and before its closure, following which event the structural
differences recognised in separating the three genera would appear
to have evolved in the fairly discrete geographical isolation which
occurred between them. Although the distribution of its congener
pseudoexigua across the tropics from northeastern Australia to
southern Japan is somewhat confounding, the isolated, endemic
occurrence of P. paradoxa on the southern Arabian coast might be
explained in terms of its being a relic. This is all the more likely due
to the influence of local seasonal upwelling, providing more temper-
ate conditions, suitable for the genus, at least for part of the year, the

136

species clearly having evolved to survive year-round conditions on
the coast in this tropical part of the world. Upwelling along the west
coast of Africa may well account for the extension of Asterina
stellifera into more tropical regions in the western Atlantic part of its
distribution. It will be of interest now to investigate whether other
shallow temperate-water echinoderms (other than the globally ubiq-
uitous ophiuroid, Amphipholis squamata) (see Price, 1982) or other
invertebrate taxa (see Barratt et al, 1984) might be found to occur in
the Dhofar region, which might support this thesis, particularly since
no other temperate-water distributed taxa appear to have been
recorded from the area.

ACKNOWLEDGEMENTS. A.C.C gratefully acknowledges the diving, field
and laboratory support given by Brian Carnell, Paul Fletcher, Mike Morrison,
Robert Whitcombe, and the Biology Department of Sultan Qaboos Univer-
sity. The field work was financed with a grant from Sultan Qaboos University,
Muscat, Sultanate of Oman.

We also wish to thank two unnamed referees for their valuable and
constructive criticisms of the manuscript. Any remaining errors are the sole
responsibility of the authors.

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Bull. nat. Hist. Mus. Lond. (Zool.) 63(2):137—147

Issued 28 November 1997

Morphological observations on Oncaea
mediterranea (Claus, 1863) (Copepoda,
Poecilostomatoida) with a comparison of Red
Sea and eastern Mediterranean populations

R. BOTTGER-SCHNACK,,

Institut fiir Meereskunde am der Universitat Kiel, Diisternbrooker Weg 20, D-24105 Kiel, Germany

R. HUYS

Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK.

SYNOPSIS. The taxonomic status of the allegedly cosmopolitan Oncaea mediterranea (Claus, 1863) is reviewed. Comparison
of Claus’ fragmentary original description and the subsequent redescription by Giesbrecht in 1892 revealed significant
differences which might lead to taxonomic confusion. Most authors have adopted Giesbrecht’s identification, but it is clear that
several, as yet unnamed, species have been recorded in the literature under the wrong name O. mediterranea. The species is
redescribed in detail on the basis of material from the Red Sea and eastern Mediterranean. The occurrence of two size morphs in
the eastern Mediterranean populations is briefly discussed. O. mediterranea sensu Bourne (1889) is regarded as species

inquirenda in the genus.

INTRODUCTION

Oncaea mediterranea (Claus, 1863) is one of the most widely
distributed oncaeids (Copepoda: Poecilostomatoida) and according
to the literature assumes a cosmopolitan distribution. It has been
recorded in epi- and mesopelagic layers of all three main oceans
between approximately 60° N and 60° S (Malt, 1983a: Fig. 14;
Heron and Bradford-Grieve, 1995). In some areas such as the
offshore waters of the Lebanon (Malt ef al., 1989) it represents the
most common oncaeid.

O. mediterranea is a well studied species. The naupliar develop-
ment has been documented by Hanaoka (1952), and Malt (1983c)
discussed the polymorphism and pore signature pattern of the
species. Paffenhofer (1993) obtained quantitative information on
reproduction rates and longevity of adult females. It was found that
early copepodids of O. mediterranea ingested about 100% of their
body weight daily when feeding at relatively high food levels.
Average reproduction rates of field-collected females ranged from
5.3 to 13.3 nauplii day~'. According to Paffenhofer (1993) neither
nauplii, nor copepodids or adults of O. mediterranea create a
feeding current, and because of their limited swimming perform-
ance the encounter with food has to be created either by motile food
particles or by the copepod jumping repeatedly to locate a food-rich
environment. O. mediterranea has been observed on discarded
appendicularian houses (e.g. Alldredge, 1972) which probably serve
as a major constituent of detritus and a food source for copepods in
the epi- and mesopelagic zones (Ohtsuka et al., 1996).

Like in many other so-called cosmopolitan planktonic species the
taxonomy of O. mediterranea is potentially confusing. Most identi-
fications of this species are based on the redescription of Giesbrecht
(1892) from the Gulf of Naples, which was excellent by contempo-
rary standards, and do not consider the type description of Claus
(1863, as Antaria mediterranea) from Messina. Comparison of
Claus’ and Giesbrecht’s text and illustrations casts certain doubts, as
to whether both descriptions are based on the same species. As part

© The Natural History Museum, 1997

of an ongoing study on Red Sea oncaeids the taxonomic history of O.
mediterranea is reviewed and the species is redescribed herein on
the basis of material from the eastern Mediterranean and various
regions in the Red Sea.

METHODS

Oncaeids were collected using a multiple opening-closing net with
a mesh size of 0.05 mm during cruise 5/5 of R/V Meteor in the
Southern and Central Red Sea and in the Eastern Mediterranean Sea.
A station list and sampling data are given in Table 1. The plankton
was initially fixed in 4% formaldehyde-seawater solution buffered
with hexamethylene tetramine and transferred after ca 2 years into a
preservation fluid of 5% propylene glycol, 0.5% propylene
phenoxetol, and 94.5% filtered seawater (Steedman, 1976). Speci-
mens were dissected in lactic acid, and mounted on slides in
lactophenol. All figures have been prepared using a camera lucida
on a Leitz Dialux differential interference contrast microscope.
Total body length and the ratio of prosome to urosome (excluding
caudal rami) were calculated as the sum of the middorsal lengths

Table 1 Station data of sampling with 0.05 mm mesh multiple opening
closing net during R/V Meteor Cruise 5.

Stn. Haul Date Time Latitude Longitude Depthof Total
No. No. (1987) (°N) (°E) Haul(m) water
depth (m)

Red Sea

663 17/4 20.07 Day 22°58.4' 37°19.4' 50-100 1200
703 39/5 03.08 Day 15°34.8' 41°54.9' 0-50 970

39/4 50-100
Eastern Mediterranean
44 31/5 24.01 Day 34°07.1' 31°54.7' 0-50 2530

138

of individual somites measured in lateral view. In the case of
telescoping somites these lengths are measured from the anterior to
the posterior margin. This approach differs from that traditionally
used in oncaeid taxonomy, where the telescoping of somites is not
considered in length measurements. Thus, sizes of the species in this
paper are not directly comparable to those of previous descriptions
(e.g. Heron, 1977; Heron and Bradford-Grieve, 1995) or earlier
studies in the Red Sea (Bottger-Schnack et al., 1989). Length data
given by the latter authors are only about 70% of the length pre-
sented in this paper, due to the excessive telescoping of somites in
the sorting medium.

Descriptive terminology for body and appendages follows that of
Huys and Boxshall (1991). Abbreviations used in the text are: ae =
aesthetasc; CR = caudal rami.

SYSTEMATICS

Order POECILOSTOMATOIDA Thorell, 1859
Family ONCAEIDAE Giesbrecht, 1892
Genus ONCAEA Philippi, 1843

Oncaea mediterranea (Claus, 1863)
(Figs 1-5)

SYNONYMS. Antaria mediterranea Claus, 1863; Oncaea medi-
terranea (Claus, 1863); Oncda mediterranea (Claus, 1863).

ORIGINAL DESCRIPTION. Claus (1863): 159-160, Tafel XXX, Fig.
IRGC), (ED)

OTHER DESCRIPTIONS. Giesbrecht (1892) [as Oncda mediterr-
anea]; Heron (1977) and Heron & Bradford-Grieve (1995) [as
Oncaea mediterranea].

TYPE LOCALITY. ‘Tyrrhenian Sea; Messina.

MATERIAL EXAMINED. (see Table I for locality data)

(1) The Natural History Museum, London: Southern Red Sea: Stn
703; R/V Meteor leg 5/5; collected with MSN 0.05 mm net
(Haul 39/4); depth 50-100 m: 102 Qand 10¢'C'in alcohol
(reg. nos 1996.1095—1114); leg. R. Bottger-Schnack;

(2) Dr Bottger-Schnack (personal collection):

(a) Southern Red Sea: Stn 703; R/V Meteor leg 5/5; collected
with MSN 0.05 mm net (Haul 39/4); depth 50-100 m; leg.
R. Béttger-Schnack: several 2 2 and oO’ Cin alcohol;

(b) Central-Northern Red Sea: Stn 663; R/V Meteor leg 5/5;
collected with MSN 0.05 mm net (Haul 17/4); depth 50—
100 m; leg. R. Bottger-Schnack: several 2 9 and O’C'in
alcohol;

(c) Eastern Mediterranean: Stn 44; R/V Meteor leg 5/1; col-
lected with MSN 0.05 mm net (Haul 31/5); depth 50-100
m; leg R. Béttger-Schnack: (i) small form: 1Qand 1
O'dissected on slides, 1 Qin alcohol; (ii) broad form: 2
2 QPdissected on slides, 1 Cin alcohol.

ADULT FEMALE
Body length (measured in lateral aspect; from anterior margin of
rostral area to posterior margin of caudal rami, calculated as sum of
individual somites): 1400 um

Exoskeleton well chitinized. Prosome 2.2 times length of urosome,
excluding caudal rami, 1.9 times urosome length including caudal

R. BOTTGER-SCHNACK AND R. HUYS

rami. Leg 2-bearing somite without conspicuous dorso-posterior
projection (Fig. 1B). Leg 3-bearing somite with conspicuous raised
pore protruding laterally (Fig. 1a). Other integumental pores on
prosome as indicated in Fig. 1A—B. Pleural areas of leg 4-bearing
somite with constricted posterolateral corners.

Proportional lengths (%) of urosomites are 9.7 : 60.7 : 8.2: 8.8:
12.6. Proportional lengths (%) of urosomites and caudal rami are 8.1
SOWA E Ochs Mes) = Os) 2 i>).

Genital double-somite nearly twice as long as maximum width
(measured in dorsal aspect) and twice as long as postgenital somites
combined; largest width measured at anterior third, posterior part
tapering gradually. Paired genital apertures located at about halfway
the distance from anterior margin of genital double-somite; arma-
ture represented by | spine and 1 diminutive spinule (Fig. 1H). Pore
pattern on dorsal surface as indicated in Fig. 1C.

Anal somite 1.3 times wider than long; about 2/3 the length of
caudal rami (Fig. 1C). Secretory pore discernible on either side of
anal opening. Anterior margin of anal opening (vestigial anal oper-
culum) with transverse row of minute denticles. Posterior margin of
somite finely serrate ventrally and laterally (Fig. 1D).

Caudal ramus (Fig. 1F) about 3.0 times as long as wide. Armature
consisting of 6 elements: antero- and posterolateral setae long,
spiniform and unipinnate along medial margin; outer terminal seta
long and plumose; inner terminal seta longest and plumose; terminal
accessory seta more than 2/3 the length of outer terminal seta and
more than twice the length of caudal ramus; dorsal seta about half
the length of terminal accessory seta, plumose and bi-articulate at
base. Inner margin of somite with fringe of long, setules. Dorsal
anterior surface (Fig. 1F) with secretory pore near insertion of seta
II. Dorsal surface covered with numerous small scales (Fig. 1F).

Antennule 6-segmented (Fig. 1E), relative lengths (%) of seg-
ments measured along posterior non-setiferous margin 6.2 : 20.6 :
46.4 : 10.3 : 6.2 : 10.3. Armature formula: 1-[3], 2-[8], 3-[5], 4-
[3+ae], 5-[2+ae], 6-[6+(1+ae)].

Antenna 3-segmented, distinctly reflexed (Fig. 2A). Coxobasis
with row of long, fine spinules or setules along outer and inner
margins and with few additional spinules on proximal and distal part
of outer (exopodal) margin; with bipinnate seta at inner distal corner.
Endopod segments unequal in length; proximal endopod segment
subtriangular forming outer lobate outgrowth bearing spinular patch,
with row of denticles along posterior inner margin. Distal endopod
segment slightly shorter than proximal exopod segment, with nar-
row cylindrical base articulating; with two patches of short spinules
along outer margin; lateral armature consisting of 1 unipinnate
spiniform seta and 3 curved setae, one of them sparsely pinnate, all
armature elements similar in length; distal armature consisting of 5
long unipinnate setae and 2 short naked setae; none of armature
elements spiniform or geniculate.

Labrum (Fig. 2B—C) distinctly bilobed. Distal margin of each
lobe with 4 strong dentiform processes medially, short row of long
fine setules latero-distally and row of minute spinules laterally.
Lobes separated by semicircular vertex covered anteriorly by over-
lapping rows of hyaline petaloid flaps. Posterior part of medial
incision with four rounded integumental thickenings. Anterior sur-
face (Fig. 2B) with short row of spinules either side of median
swelling, raised row of small strong denticles latero-posteriorly;
median swelling with large secretory pore posteriorly. Posterior
surface (Fig. 2C) with group of 3 secretory pores located on proxi-
mal part of each lobe and an additional one basally.

Mandible (Fig. 2D) with minute spinules on surface of coxa;
gnathobase with 5 elements: 1 at subdistal ventral corner, 2 along
distal margin and 2 along subdistal dorsal margin; ventral element
shorter than ventral blade, with long, fine setules along dorsal

MORPHOLOGY OF ONCAEA MEDITERRANEA 139

sa

SSS

me
SS

Ki
SSS

SSS

“tly
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SK

Fig. 1 Oncaea mediterranea (Claus, 1863), 9 (Red Sea). A, habitus, dorsal [a: lateral raised pore, enlarged]; B, same, lateral (appendages omitted), arrow
indicating position of lateral raised pore; C, urosome, dorsal; D, urosome, lateral; E, antennule; F, caudal ramus, dorsal; G, leg 5, dorsal; H, leg 6.

140

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R. BOTTGER-SCHNACK AND R. HUYS

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Key pt hhh bbb

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Ky

Fig. 2 Oncaea mediterranea (Claus, 1863), 9 (Red Sea). A, antenna; B, labrum, anterior; C, same, posterior; D, mandible, showing individual elements;
E, maxillule; F, maxilla, [f1, distal seta; £2, proximal seta]; G, maxilliped.

141

MORPHOLOGY OF ONCAEA MEDITERRANEA

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leg 2, anterior [b, terminal part of endopod];

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Fig.3 Oncaea mediterranea (Claus, 1863), 9 (Red Sea). A, leg 1, anterior [a, terminal part of endopod]; B

C, leg 3, posterior [c, terminal part of endopod]; D, leg 4, anterior [d, terminal part of endopod].

142

margin; ventral blade strong and spiniform, with row of setules on
posterior surface; dorsal blade strong and broad, with four dentiform
processes around distal margin; dorsal elements setiform, the shorter
hyaline, flat and densely setose, the longer multiipinnate.

Maxillule (Fig. 2E) indistinctly bilobed, with numerous spinules
on anterior and posterior surfaces. Inner lobe subcylindrical, with 3
elements: outermost one spiniform, swollen at base, fringed with
coarse spinules, others setiform and bipinnate; innermost one lo-
cated along concave inner margin at some distance from other
elements. Outer lobe with 4 elements; outermost element spiniform,
curved and unipinnate, longer than the following; other elements
with sparse spinules, innermost one shortest.

Maxilla (Fig. 2F) 2-segmented, comprising syncoxa and allobasis.
Syncoxa unarmed, surface ornamented with 2 spinular rows and 1
large secretory pore. Allobasis produced distally into slightly curved
claw bearing 2 rows of very strong spinules along medial margin;
outer margin with strong seta almost extending to tip of allobasal
claw, ornamented with few strong spinules distally and a thin
hyaline lamella bilaterally, tip of seta with tubular extension (Fig.
2f1); inner margin with slender pinnate seta and strong basally
swollen spine with double row of very strong spinules along the
medial margin and single row of shorter spinules along outer margin
(Fig. 2f2).

Maxilliped (Fig. 2G) 4-segmented, comprising syncoxa, basis
and 2-segmented endopod. Syncoxa unarmed, without surface orna-
mentation. Basis robust, inner margin with 2 spiniform bipinnate
elements nearly equal in length; fringe of long pinnules between
proximal seta and articulation with endopod; two rows of long
setules on anterior surface as illustrated in Fig. 2G. Proximal endopod
segment unarmed. Distal endopod segment drawn out into long
curved claw, with pinnules along proximal half of concave margin;
accessory armature consisting of minute, naked seta on outer prox1-
mal margin and unipectinate spine fused basally to inner proximal
corner of claw.

Swimming legs 1-4 biramous (Fig. 3A—D), with 3-segmented
rami. Spine and setal formulae as follows:

Coxa Basis Endopod Exopod
Leg 1 0-0 1-I 0-1; 0-1; 0,L5 0; I-1; Il,1,4
Leg 2 0-0 1-0 0-1; 0-2; LIL3 1-0; I-1; I,1,5
Leg 3 0-0 1-0 0-1; 0-2; LI1,2 0; I-1; IL,1,5
Leg 4 0-0 1-0 0-1; 0-2; LII,1 0; I-1; IL,15

Intercoxal sclerites well developed, without ornamentation. Coxae
and bases of legs 1-3 with surface ornamentation as shown in Fig.
3A-C. Bases with plumose (legs 1, 4), short naked (leg 2) or very
long naked, outer seta (leg 3); with anterior secretory pore near outer
proximal corner; inner portion slightly produced adaxially in legs 2—
4 (Fig. 3B—D). Inner basal seta on leg 1 long, spiniform and naked.
Respective legs without distinct length differences between rami
with endopod slightly longer than exopod. Bases of spines on
exopodal and endopodal segments surrounded by small spinules
which are most obvious around terminal endopod spines of legs 2
and 4 (Fig. 3b,d). Surface ornamentation of all segments with small
scales, similar to those on caudal ramus (Fig. 1F).

Exopods. Outer margin of exopod segments with well developed
serrated hyaline lamella; inner margin of proximal exopod segments
with long setules. Secretory pore present on posterior surface of
distal segments. Hyaline lamellae on outer exopodal spines well
developed. Terminal spine of leg 1 equal in length to (leg 1) or
shorter than (legs 2-4) distal exopod segment.

R. BOTTGER-SCHNACK AND R. HUYS

Endopods. Outer margin of endopod segments with fringe of long
setules. Inner seta of proximal endopod segment slightly swollen.
Distal endopod segments with single secretory pore on posterior
surface; distal margin of legs 2 and 3 produced into conical projec-
tion ornamented with minute spinules anteriorly (Fig. 3B,C,b,c).
Outer subdistal spine equal in length to distal spine in legs 2-4.
Outer margin of distal segment of leg 1 terminating in a long process
obscuring insertion of distalmost inner seta (Fig. 3a). Inner setae of
distal endopod segments of legs 2-4 with spinule comb along
proximal inner margin; this comb less obvious in leg 1; also present
on distal inner seta of middle endopod segment in leg 4.

Leg 5 (Fig. 1G) comprising small plumose seta arising from
lateral surface of somite, and small free segment representing exopod.
Exopod slightly longer than wide, bearing stout curved seta apically
and smaller slender seta inserted subdistally; surface covered with
minute denticles

Leg 6 (Fig. 1C, H) represented by operculum closing off each
genital aperture; armed with a spine and a short spinule.

Egg-sacs paired, oval-shaped; each sac containing approximately
40 eggs (diameter 50-65 pm).

ADULT MALE

Body length (measured in lateral aspect; from anterior margin of
rostral area to posterior margin of caudal rami, calculated as sum of
individual somites): 960 um. Sexual dimorphism in antennule,
antenna, maxilliped, legs 5—6, caudal ramus and in genital segmen-
tation.

Proportional lengths (%) of urosomites (excluding caudal rami)
9.4 : 70.0.: 3.4 : 3.8 : 3.0: 10.5 ; proportional lengths (%) of
urosomites (caudal rami included) 8.4 : 62.1 :3.0:3.3:2.7:8.1:
11.6. Leg 5-bearing somite with transverse row of denticles dorsally
(Fig. 4D). Caudal rami 1.8 times longer than wide, markedly shorther
than in female. Caudal setae with proportional lengths as in female.
Dorsal surface of genital somite covered with scale-like structures in
anterior half, arranged in a specific pattern (not figured). Surface of
genital flaps and ventral surface of anal segment ornamented with
several rows of small spinules (Fig. 4E). Dorsal surface of caudal
ramus covered with minute scales as in female.

Antennule (Fig. 4B) 4-segmented; distal segment corresponding
to fused segments 4—6 of female; relative lengths (%) of segments
measured along posterior non-setiferous margin 14.5 : 19.7: 42.1:
23.7. Armature formula: 1-[3], 2-[8], 3-[4], 4-[11+2ae+(1+ae)].

Antenna as in female, except for lateral armature on distal endopod
segment (Fig. 4G), with third spiniform element being much stouter
than in female and shorter than the 2 adjacent setae.

Maxilliped (Fig. 4C) 3-segmented, comprising syncoxa, basis
and 1-segmented endopod. Syncoxa without surface ornamentation,
unarmed. Basis robust, particularly inflated in proximal half form-
ing bulbous swelling; anterior surface with 2—3 transverse spinular
rows in addition to row of short flat spinules along inner margin
(Fig.4C); posterior surface with 3 rows of short spatulated spinules
of graduated length along palmar margin (Fig. 4C); with 2 small
naked setae inserted within the longitudinal cleft, the proximal one
being slightly longer than the distal one. Endopod drawn out into
long curved claw, concave margin unornamented; accessory arma-
ture consisting of short, unipectinate spine basally fused to inner
proximal corner of claw; claw with minute hyaline apex (Fig. 4c).

Legs 1-4 with armature as in female; variable number of spinules
on conical projection of endopods of legs 2 and 3.

Leg 5 (Fig. 4D, F) exopod not delimited from somite, general
shape and armature as in female; small plumose seta arising from
lateral surface of somite as in female.

Leg 6 (Fig. 4E) represented by posterolateral flap closing off

MORPHOLOGY OF ONCAEA MEDITERRANEA 143

a
Se—_
=.
C4 ej
C9
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Pa

SSS

SLOSS.

SSS

Fig.4 Oncaea mediterranea (Claus, 1863), (Red Sea). A, habitus, dorsal; B, antennule; C, maxilliped, anterior [c, tip of claw, showing hyaline lamella];
D, urosome, dorsal; E, urosome, ventral; F, same, lateral (spermatophores immature); G, antenna, distal endopod segment.

144

genital aperture on either side; covered by pattern of denticles as
shown in Fig. 4E; posterolateral corners protruding laterally so that
they are discernible in dorsal aspect (Fig. 4D).

Spermatophore oval, of variable size according to state of matu-
rity (Fig. 4F); swelling of spermatophore during development not
affecting shape or relative size of genital somite.

TAXONOMIC HISTORY

Claus (1863:159) presented a concise original description of Antaria
mediterranea with some illustrations of both sexes. Later Claus
(1866) identified two size groups upon re-examination of original
material from Messina: the large one [1.3 mm excluding CR setae]
being the one that was mentioned in the original description; the
small one [0.8—0.9 mm] which he also recorded from Nice; no
overlap was found between both size morphs from Messina.

Lilljeborg (1875) recorded a single specimen from Mosselbay
(Spitsbergen), and Car (1884) found it in plankton samples taken off
Trieste in the Adriatic Sea. Both authors, however, gave no descrip-
tions or figures and their records therefore remain unconfirmed.

Bourne (1889) [as Oncaea mediterranea] found 2 females in
surface waters off Plymouth. Giesbrecht (1892) regarded this record
doubtful and pointed out the discrepancy between the text and
figures with regard to the CR/anal somite length ratio. Careful
comparison of this character is hampered by the fact that Claus
(1863) only figured the female in lateral aspect and by the possibility
that the urosome is considerably telescoped in the specimen illustrated
by Bourne. Giesbrecht’s statement that Bourne’s and Claus’ speci-
mens differ fundamentally in maxilliped structure is equivocal since
he compared the former with what he had identified as O.
mediterranea rather than with the original type material. On the
basis of Brady’s (1883) illustrations [Challenger Expedition], Bourne
(1889) also tentatively regarded Onc@a obtusa (Dana) as a possible
synonym of O. mediterranea, however, as Giesbrecht (1892) and
subsequent authors suspected, the latter could as well be conspecific
with O. venusta Philippi.

Giesbrecht (1892) reviewed the earlier literature on Antaria and
Oncaea, and summarized the synonymies of the respective species
known at that time. Dana’s (1849) species Antaria obtusa and A.
crassimana were included under the synonymy of both O. venusta
and O. mediterranea, reflecting the author’s undecisiveness on this
matter. Giesbrecht redescribed O.-mediterranea on the basis of
material from Naples and distinguished two colour varieties flava
and rubra.

Comparison of Giesbrecht’s illustrations with Claus’ original
description, however, raises some doubts as to the conspecificity of
the Messina and Naples specimens. The major obstacle in this
comparison lies in the form, position and size of the setae on the
maxillipedal basis which is usually considered as an important
discriminant in oncaeid systematics. The issue is even more compli-
cated by the discrepancy between text and figures in Claus’ original
description. Claus (1863) stated that there are two ornated setae on
the palmar margin of the basis, yet in his figure (Tafel XXX, Fig. 6)
only one naked seta is illustrated. From the position of this seta,
being located halfway the inner margin, it is conceivable that Claus
has overlooked the proximal seta. This hints at the possibility that
Claus’ O. mediterranea is related to the ‘englishi species-group’
which includes O. ornata Giesbrecht, 1891, O. shmelevi Gordejeva,
1972, O. englishi Heron, 1977 and O. alboranica Shmeleva, 1979.
In these species the distal element is long, slender and minutely
pinnate, whereas the proximal one is spiniform and because of its

R. BOTTGER-SCHNACK AND R. HUYS

small size easily overlooked or misinterpreted as a spinule. In
Giesbrecht’s O. mediterranea, however, both elements are (1) of
about the same size and only half the length of the proximal seta in
the Messina material, and (2) positioned differently, i.e. the proxi-
mal one at 1/3 distance from the syncoxa-basis joint, the distal one
at 2/3 distance. A second possibility is that the long palmar seta in
Claus’ original illustration is in fact a maxillary element superim-
posed on the maxilliped since Claus believed that both appendages
represented the rami of a single limb, i.e. the maxilliped.

Other differences are found in the female leg 5 which is longer in
the original description and the shape of the genital double-somite in
lateral aspect which does not have the pronounced swelling anterio-
ventrally as shown in Giesbrecht’s illustration (Taf. 4, Fig. 16). The
male of Claus’ O. mediterranea shows an exceptionally long leg 5
exopod (his Taf. XXX Fig. 7) which might or might not be free. This
character has thus far been found in only a small number of Oncaea
species belonging to the notopus group, such as O. damkaeri Heron,
1977 and O. parila Heron, 1977 (Heron, 1977; Heron et al., 1984)
which also display a very long leg 5 exopod in the females. Species
of thenotopus group have a setation pattern on the female maxilliped
which is significantly different from that displayed in the englishi
group which raises the suspicion that Claus (1863) might well have
based males and females on different species.

As a result of this comparison it is clear that Claus’ original text
and drawings contain several internal inconsistencies and lack the
detail that is necessary to allow unequivocal identification. The
setation of the maxilliped is a potentially critical character in this
process as confirmation or refutal of Giesbrecht’s identification
depends on whether more weight is given to the text statement or to
the illustration. Given the fact that Claus’ figures of the other
cephalic appendages are similarly poor (setation elements are miss-
ing from every limb) it is preferred here to give more weight to the
text as this will lead to nomenclatural stability. Pending the rediscov-
ery of Claus’ types (which are in all probability lost) this admittedly
subjective decision is the best course of action. In view of the grossly
fragmentary original description in which the sexes were based on
two different — but unidentifiable — species and in the absence of
formal holotype designation the taxonomic problem is in our opin-
ion unsolvable. Moreover, it is considered highly unlikely that
collection of topotype material from Messina would be informative
as 130 years have lapsed since Claus’ discovery of the species in an
open pelagic environment that might have been subjected to major
changes since, such as the opening of the Suez Canal in 1869.

O. mediterranea (Claus, 1863) sensu Bourne (1889) is clearly
different from the Mediterranean material and is regarded here as
species inquirenda in the genus.

Other records of O. mediterranea.

O. mediterranea has been recorded from a wide range of localities
such as the Antarctic (Heron, 1977) and the Red Sea (Bottger-
Schnack, 1988). Many of its records, however, remain unconfirmed
such as the Red Sea records of Cleve (1900, 1903) and Thompson
and Scott (1903) [compiled by Halim (1969)]. Since most authors
have followed Giesbrecht’s identification and ignored Claus’ origi-
nal description it is likely that at least one, as yet unnamed, species
became established in the literature under the wrong name O.
mediterranea. For example, re-examination of material collected
during the Terra Nova and Challenger expeditions (deposited in The
Natural History Museum) proved to belong to at least two distinct
species differening in several aspects from O. mediterranea. Scott
(1894) recorded this species (‘1 or 2 females’) from the Gulf of
Guinea, but re-examination of his illustrations leave little doubt that

MORPHOLOGY OF ONCAEA MEDITERRANEA

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145

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Fig.5 Oncaea mediterranea (Claus, 1863), 2 , broad form (eastern Mediterranean). A, habitus, dorsal; B, same, lateral; C, caudal ramus.

he was dealing with a species of the englishi group. Supporting
evidence for this is found in the morphology of the antenna,
maxilliped and leg 4. Dakin and Colefax (1940) recorded the species
from New South Wales from a depth of 0-200 m were it was ‘rare’
but it is doubtful whether they were dealing with O. mediterranea
since material collected from the Great Barrier Reef and identified
with Claus’ species proved upon re-examination to belong to a
closely related undescribed species. Razouls (1974) described O.
mediterranea from the region of Banyuls-sur-mer (Golfe du Lion)
and summarized previous records from the Mediterranean Sea. His

drawing (Fig. 4G) of the male shows a terminal conical projection on
the endopod of swimming leg 4 which disproves his identification as
this is a character exclusively found in representatives of the conifera-
similis complex. Ferrari (1973, 1975) recorded the species from the
Gulf of Mexico and the northern Caribbean Sea and distinguished
three size variants, the status of which will be discussed below under
‘form variants’ of O. mediterranea. Boxshall (1977a) recorded O.
mediterranea from the Northeastern Atlantic near the Cape Verde
Islands and in another report (Boxshall, 1977b) gave detailed infor-
mation on their vertical distribution and diurnal vertical migration.

146

Many more records of O. mediterranea from different locali-
ties in the world ocean are known [see Malt (1983a) for a
review], but are not considered here, because they did not
include figures or a description that positively identified the
species.

FORM VARIANTS

In the eastern Mediterranean, O. mediterranea exhibits two
forms in both sexes: a smaller and more slender form and a
larger, more robust form. These forms differ only in the general
appearance and in the length : width ratio of the caudal ramus in
the females. The difference in general habitus between the two is
caused by the greater width of the leg 2-bearing and leg 3-
bearing somites in the larger form (Fig. 5A). In the females, the
length to width ratio of the caudal rami is larger in the broad form
(4:1) (Fig. 5C) as compared to the small form (3:1, cf. Fig. 1F),
whereas in the males no corresponding difference could be
found. No other morphological differences were discernible
between the two morphs. The small form of O. mediterranea
corresponds to the small variant described by Malt (1983a),
while the large form seems to be similar to her ‘typical’
mediterranea form. Only one size morph of O. mediterranea
was found in the Red Sea, which can be identified as the small
form found in the eastern Mediterranean and on the basis of the
length to width ratio of caudal rami seems to resemble the small
form of O. mediterranea recognized by Malt (1983b). A small
colourless form of O. mediterranea (in addition to the larger
‘orange-golden’ form) was found south of New Zealand by
Farran (1929) but no further description was provided.

Ferrari (1975) reported the existence of three distinct size
groups in O. mediterranea of the Gulf of Mexico. The two larger
forms differed only in size, but had a similar length to width ratio
of the caudal ramus of 4:1. On the basis of their morphometry
they might be assigned to the typical form of O. mediterranea
(sensu Malt). The smaller size group had a length to width ratio
of the caudal ramus of 3.0—3.5:1 and might thus be similar to the
small form sensu Malt and to that found in the Red Sea. The
smaller morph was initially regarded as a potential new species
(Ferrari, 1973) since it differed both in the proportional lengths
of body somites as well as in total body size. Subsequently,
Ferrari (1974), quoting O. conifera Giesbrecht, 1891 as an
exemplar of gross relative size variation in oncaeids, considered
it as merely another size group of O. mediterranea since not
structural differences could be detected in the appendages. The
recent outstanding work on the conifera-complex by Heron and
Bradford-Grieve (1995) has amply demonstrated that much of
the ‘variation’ in O. conifera can be explained by the fact that
morphologically similar species can co-exist and that this mor-
phological similarity can even cause anomalies in the mate
guarding configurations of certain species. It is conceivable that
many of the form variants represent genuine species which can
no longer be discriminated using traditional o-taxonomical
methods. For example, Malt (1983c) mapped the pore signature
pattern of the two female forms of O. mediterranea from the
North Atlantic but failed to reveal any significant difference
between them. It seems therefore that the question whether the
large morph found in the Eastern Mediterranean represents a
sibling species of O. mediterranea can perhaps only be resolved
by breeding experiments or alternative methods using molecular
data such as enzyme electrophoresis, immunological distance
methods or ribosomal RNA sequencing.

R. BOTTGER-SCHNACK AND R. HUYS

ECOLOGICAL NOTES

Geographical distribution.

O. mediterranea is distributed throughout the Red Sea (see review in
Halim (1969); Bottger-Schnack, 1990b, 1995). It was also found in
small mesh net samples from the northernmost part of the Gulf of
Aqaba (Bottger-Schnack, unpubl.).

Vertical distribution and vertical migration.

During summer and autumn, when a strong seasonal thermocline is
developed, the core of the O. mediterranea population during the
day is situated in the zone below the thermocline, at 50-150 m
(Bottger-Schnack, 1990a, unpubl.). Parts of the population migrate
into the upper 50 m during the night, with males showing a greater
proportion of migrating individuals than females. For females, a
bimodal vertical distribution can be found during these seasons, with
the lower part of the population dwelling in the 250-400 m layer, in
the core of the oxygen minimum zone. In the northern Red Sea the
mesopelagic populations of females are found at greater depths than
in the central area, corresponding to the regional differences in the
depth of minimal oxygen concentrations (Bottger-Schnack, 1990b).
These deep dwelling populations are not found during winter
(Bottger-Schnack, 1990b).

Seasonal variation in abundance (central Red Sea).

No consistent seasonal variation in abundance was found for O.
mediterranea in the central Red Sea, indicating that the populations
are not substantially recruited by those from the south (Bottger-
Schnack, 1995).

ACKNOWLEDGEMENTS. We wish to thank Prof. J. Lenz, Prof. D. Schnack
and Dr H. Weikert for support and help. Dr G.A. Boxshall (NHM) and Dr ED.
Ferrari (Smithsonian Institution) reviewed earlier drafts of the manuscript.
This study was supported by a Deutsche Forschungsgemeinschaft grant Le
232/18-1.

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MORPHOLOGY OF ONCAEA MEDITERRANEA

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CONTENTS

93 The lucinid bivalve genus Cardiolucina (Mollusca: Bivalvia: Lucinidae); systematics,
anatomy and relationships
John D. Taylor and Emily A. Glover

123 Anew species of water mouse, of the genus Chibchanomys (Rodentia: Muridae:
Sigmodontinae) from Ecuador
Paulina D. Jenkins and Adrian A. Barnett

129 Anew species in the asterinid genus Patiriella (Echinodermata: Asteroidea) from Dhofar,
southern Oman: a temperate taxon in a tropical locality
Andrew C. Campbell and Frasncis W.E. Rowe

137 Morphological observations on Oncaea mediterranea (Claus, 1863) (Copepoda:
Poecilostomatoida) with a comparison of Red Sea and eastern Mediterranean populations
R. Bottger-Schnack and R. Huys

Bulletin of The Natural History Museum :

ZOOLOGY SERIES
Vol. 63, No. 2, November 1997