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Antennulary Sensory Organs in Cyprids of Octolasmis
and Lepas (Crustacea: Thecostraca: Cirripedia:
Thoracica): A Scanning Electron Microscopic Study
Mikkel Blomsterberg,1 Jens T. Høeg,1* William B. Jeffries,2 and Niklas C. Lagersson1
1
Zoological Institute, University of Copenhagen, DK-2100, Copenhagen, Denmark
2
Dickinson College, Carlisle, Pennsylvania 17013, USA
ABSTRACT Cypris larvae of the pedunculate barnacles
Octolasmis angulata (Poecilasmatidae), Lepas australis,
L. pectinata, and Dosima fascicularis (Lepadidae) were
studied with scanning electron microscopy, focusing on
the sensory setae and the attachment disc on the anten-
nules. The antennules of O. angulata did not exhibit any
remarkable trait, but carry the same number of setae as
seen in most other thoracicans. The third segment is bell-
shaped and quite distinct from the second and its attach-
ment disc is surrounded by a skirt. We found several
potential synapomorphies in antennulary morphology be-
tween cyprids of the lepadid species but none of them were
shared with the cyprids of Octolasmis; the list of unique
lepadid characters includes: one additional, preaxial seta
on the second segment; multiple similar (up to eight)
postaxial setae (PS3) on the third segment, unlike all
other thoracicans, where there is only a single PS3; the
third segment consists almost entirely of the attachment
disc, which is distended and surrounded by two parallel
rows of radial setae; on the fourth segment the terminal
seta E is diminutive. We found no traits in cyprids of
Octolasmis that seem to be adaptations to their attach-
ment site within the branchial chamber of swimming
crabs and, in particular, no similarities with cyprids of
rhizocephalan barnacles, many of which also attach in the
gill chamber. The synapomorphies between cyprids of the
lepadid species may be adaptations to their life in the
neuston. J. Morphol. 260:141–153, 2004.
© 2004 Wiley-Liss, Inc.
KEY WORDS: Octolasmis; Lepas; cypris; phylogeny; an-
tennule; morphology; sensilla; SEM; settlement
Members of the genus Octolasmis (family Po-
ecilasmatidae) are pedunculate barnacles that live
mainly as epibionts on portunid crabs. Species of
Octolasmis must not only find a suitable host, but
must also complete their entire life-cycle between
two successive molts of the host crab. Many species
attach within the branchial chamber and in this
they resemble those parasitic barnacles (Rhizo-
cephala) that invade their host crabs through the
thin cuticle of the gill filaments. A close relationship
between Octolasmis and the Rhizocephala is not at
issue, but due to their specialized epibiosis they are
nevertheless very interesting analogs and a study of
© 2004 WILEY-LISS, INC.
JOURNAL OF MORPHOLOGY 260:141–153 (2004)
their biology may very well shed light on the evolu-
tion into parasitism.
In all Cirripedia it is the task of the cypris larva to
locate a suitable substratum and carry out the ac-
tual attachment (Lagersson and Høeg, 2002). The
cypris larvae of rhizocephalan barnacles and of some
thoracican species are now well studied by means of
scanning and transmission electron microscopy.
These studies have revealed that rhizocephalan cyp-
rids possess an armament of sensory organs that
differs from those seen in thoracican barnacles and
may well be related to their parasitic lifestyle (Nott
and Foster, 1969; Walker, 1985; Glenner et al., 1989;
Clare and Nott, 1994; Moyse et al., 1995; Høeg and
Rybakov, 1996). Within the Thoracica, some species
with an epibiotic life style such as commensals with
corals also possess rather specialized cyprids.
Within the last two decades many articles have
dealt with the ecology and life-cycle of species of
Octolasmis (Walker, 1974; Jeffries et al., 1985, 1989,
1992; Gannon, 1990; Gannon and Wheatly, 1992;
Voris et al., 1994; Jeffries and Voris, 1996), but none
have focused on the morphology of the cypris larvae.
It is therefore unknown to what extent cyprids of
Octolasmis reflect the specialized epibiotic lifestyle.
Any study of morphological adaptation must occur
within an evolutionary framework, but unfortu-
nately the phylogenetic position of Octolasmis (and
the family Poecilasmatidae) is not well established.
The Poecilasmatidae are habitually put “close” to
the Lepadidae, the latter being specialized barnacles
living in the neuston. The principal similarity be-
tween poecilasmatids and the lepadids is the pres-
Contract grant sponsor: the Danish Natural Science Research
Council SNF; Contract grant number: 9901769 (to JH).
*Correspondence to: Prof. Jens T. Høeg, Department of Zoomorphol-
ogy, Zoological Institute, University of Copenhagen, Univer-
sitetsparken 15, DK-2100, Copenhagen, Denmark.
E-mail: jthoeg@zi.ku.dk
DOI: 10.1002/jmor.10131
142 M. BLOMSTERBERG ET AL.
Fig. 1. Octolasmis angulata. Whole cyprid, lateral view. Both antennules are extended.
ence of five shell plates, but this is nothing more
than a very plesiomorphic trait within the Thoracica
(Glenner et al., 1995) that cannot by itself argue for
a close phylogenetic relationship. Very recent stud-
ies based on 18S rDNA sequences support a sister
group relationship between Octolasmis and Lepas
(Mizrahi et al., 1998; Perl-Treves et al., 2000; Harris
et al., 2000) but as yet such a relationship has not
been confirmed by morphological evidence.
Larval characters offer an independent set of
traits for phylogenetic analysis of the Cirripedia and
they are particularly useful because they can be
used for all cirripedes irrespective of whether they
have shell plates or not (such as the shell-less Rhi-
zocephala and Acrothoracica). The literature is re-
plete with detailed descriptions of the naupliar
stages of cirripedes, but until recently this informa-
tion has mostly been used for the identification of
such larvae from the plankton. Few studies have
placed the naupliar morphology in an evolutionary
perspective (Moyse, 1987; Korn, 1995) and until re-
cently none have used cladistic methods. This
changed dramatically when Newman and Ross
(2001) presented a comprehensive analysis of tho-
racican phylogeny based on a matrix of 30 taxa and
59 characters from the setation of the naupliar ap-
pendages, but they did not include Octolasmis.
Characters from the terminal larval stage, the
cyprid, offer another suite of characters. Unfortu-
nately, almost all studies of larval development in
Cirripedia have either omitted the cyprids or at best
illustrated them as uninformative outlines of the
body, and this is also true for studies on Octolasmis
(see Lang, 1976). A recent series of articles have
shown how SEM studies of cyprids provide a wealth
of characters that vary in a biologically and phylo-
genetically interesting manner (Walker, 1985;
Jensen et al., 1994a,b; Moyse et al., 1995; Glenner
and Høeg, 1995; Høeg and Rybakov, 1996; Rykabov
et al., 2002). We have used SEM to study cyprids of
Octolasmis angulata and to compare them with cyp-
rids from three species of the Lepadidae. We put
special emphasis on the sensory organs of the an-
tennules as these structures are used in substrate
choice of the settling larvae and present a large
number of characters for phylogenetic analysis (Nott
and Foster, 1969; Glenner et al., 1989; Clare and
Nott, 1994; Høeg and Rybakov, 1996; Kolbasov et
al., 1999; Høeg and Kolbasov, 2002). Our aim is not
to use single characters in phylogenetic analysis, but
rather to increase the morphological database on
larval (cyprid) cirripedes, both for use in future cla-
distic analyses and to understand how cypris sen-
sory organs correlate with very specific settlement
substrata as seen for species of Octolasmis.
MATERIALS AND METHODS
Cyprids of Octolasmis angulata (Aurivillius, 1894) were ob-
tained live from Phuket, Thailand, and fixed in formalin. Cyprids
of the lepadid species Lepas australis (Darwin, 1851), Lepas pec-
tinata Spengler, 1793, and Dosima (Lepas) fascicularis (Ellis and
Solander, 1786) were the same as used in the studies of Jensen et
al. (1994b) and Moyse et al. (1995). None of the larvae were
relaxed before fixation. Specimens for SEM were dehydrated
through an alcohol series, transferred from absolute alcohol to
acetone, critical point-dried in CO2, and studied in either a Jeol
JSM-840 or a Jeol FEG-GUN JSM-6335F, both situated at the
Zoological Museum, University of Copenhagen. All images were
saved digitally. Measurements were performed on the SEM mi-
crographs that presented the best angle — not necessarily the
images presented here. All measurements were approximate due
to the inaccuracy inherent in SEM micrographs.
RESULTS
The antennules consist of four segments in all the
examined species, but the first segment was always
 CYPRIS LARVAE OF OCTOLASMIS AND LEPAS 143

Fig. 2. Octolasmis angulata; antennule. A: Second and third segment, mediolateral view. One postaxial seta 2 (PS2) and one
postaxial seta 3 (PS3) are positioned in the same plane. PS2 and PS3 are similar in length. Both exhibit sparse setulation. PS3 is well
separated from the attachment disc. Note the wrinkled cuticle in the constriction immediately below the attachment disc, indicating
that this part of the cuticle is very soft or thin. Note also the row of spicules on the second segment (arrow). B: Attachment disc, oblique
distal view. A dense carpet of microcuticular projections (⫽ “villi”) covers the disc surface, also in close proximity to the protuberant
axial sensory organ (ASO). The margin of the disc is irregularly rounded. Bases of four subterminal setae (STS) are arranged in a row
on the fourth segment. C: PS2; lightly setulated, sited in a small depression. D: Close-up of A. The axial sense organ with a prominent
terminal pore stands among the carpet of microcuticular projections. E: Margin of attachment disc, seen from the inside of the disc.
A radial seta (RS), prominent, tubular-shaped, and with a terminal pore (arrow) is situated between the microcuticular projections and
the skirt. Note that the skirt lacks microcuticular projections. F: Margin of attachment disc, lateral view. The disc is bordered by a
distinct skirt of thin cuticle with radial reinforcement ribs (arrows). AD, attachment disc; ASO, axial sensory organ; KK, Knockando;
PS2, postaxial seta 2; PS3, postaxial seta 3; RS, radial seta; STS, subterminal setae.
144 M. BLOMSTERBERG ET AL.

Figure 3
 CYPRIS LARVAE OF OCTOLASMIS AND LEPAS
positioned within the carapace and therefore was
not visible with SEM. The gross morphology of the
antennules (Fig. 1) resembles that in other cir-
ripedes, so we focus primarily on differences in the
setation and the shape of the attachment disc.
Octolasmis angulata
The antennules consist of four segments both in
Octolasmis angulata and in the three lepadid spe-
cies described below, but as the first is hidden by the
carapace we only describe the morphology of the
second, third, and fourth segments. In O. angulata
the second segment is an elongated cylinder taper-
ing distally toward the articulation with the third
segment (Figs. 2A, 3A). The third segment is con-
spicuous, quite distinct from segment 2, and is bell-
shaped. Its attachment disc is orientated at an
oblique angle with respect to the long axis of seg-
ment 2 and faces ventrodistally. The same is true for
the attachment discs in cyprids of Lepas australis, L.
pectinata, and Dosima fascicularis. The fourth seg-
ment is attached proximolaterally on the third. Our
naming of antennulary setae follows the scheme
used by Glenner and Høeg (1995) and Moyse et al.
(1995).
Second segment. The second segment is orna-
mented with scattered groups of small cuticular
combs in which the teeth face distally (Fig. 2A). The
only seta is the postaxial seta 2 (PS2), which sits
ventrodistally but is still somewhat removed from
the articulation with the third segment. PS2 is ⬃50
␮m long, slightly curved, and carries small setules
(Fig. 2C). Here and below we use “setule” for such
small outgrowths on setae, although we could not
observe any basal articulation.
Third segment. On the third segment the only
seta outside the attachment disc is the single post-
axial seta 3 (PS3) (Fig. 2A). It sits proximoventrally
on the segment about midway between the articula-
tion to segment 2 and the rim of the attachment disc.
PS3 resembles PS2 in being ⬃50 ␮m long and hav-
ing small setules.
Fig. 3. Octolasmis angulata; antennule. A: Right antennule;
lateral view of segments 2– 4. B: Close-up of segment 4. Four
identical subterminal setae (STS) exhibit pores (arrows); note the
distinct terminal plateau, rimmed by the slightly raised cuticle
and carrying five terminal setae A–E. A and B are identical and
are attached close together. C is hidden. D is close to A,B and is
bag-shaped with a conspicuous ringed surface pattern. E is at-
tached slightly apart and with a basal socket. C: Close-up of the
spout-like tip of seta D. Arrow marks pore. D: Segment 4, lateral
view of apical end showing attachment of setae. Note the ob-
liquely cut off, small ledge carrying the row of four subterminal
setae and the distinct, inverted cone-shaped socket of terminal
seta D. The tiny seta C is obscured in this view. E: Base of
terminal setae showing the diminutive seta C. Seta B is obscured
by its twin. AD, attachment disc; STS, subterminal setae on
segment 4; A–E, terminal setae on segment 4.
145
Attachment disc. The third segment is conspic-
uously constricted below (proximally to) the attach-
ment disc and the cuticle of the constricted zone has
a wrinkled surface, indicating that it is thinner and
more flexible than the general integument of the
segment (Fig. 2A). The attachment disc (Fig. 2B) is
almost circular, ⬃30 ␮m in diameter, and is densely
carpeted with microcuticular projections (cuticular
“villi” sensu Moyse et al. [1995] and earlier authors).
The disc is surrounded by a continuous, low rim of
thin cuticle without microcuticular projections (Fig.
2E); i.e., a skirt sensu Moyse et al. (1995). Seen from
the outside, this skirt seems to have a rather regu-
larly spaced reinforcement of thicker cuticle (Fig.
2F).
The axial sense organ (seta) occupies the center of
the disc. It is a cylindrical, elongated structure, ⬃3
␮m in length, with a conspicuous terminal pore (Fig.
2D). It emerges directly from among the microcu-
ticular projections, which prevented us from observ-
ing its basal-most part. These microcuticuticular
projections also obscured most other putative setae
on the disc and all exit pores of the cement gland and
antennulary glands. In particular, we observed no
postaxial sense organ (seta).
A few radial setae were observed close to the pe-
rimeter. They are simple elongated setae projecting
1–2 ␮m beyond the rim of the skirt and each has a
distinct terminal pore (Fig. 2E).
Fourth segment. The fourth segment sits
proximolaterally on the third segment (Fig. 3A)
and has an inverted cone-shape, being rather nar-
row at the base but increasing in width toward the
apical end (Fig. 3B). It carries four subterminal
and five terminal setae. The subterminal setae are
sited as a transverse row on a distinct ledge very
close to the apical end. They are all similar, ⬃20
␮m long, tapering distally and terminating in a
pore. Orientation of the four setae differed be-
tween specimens. The subterminal setae lack set-
ules, but have a single distinct spine on their
convex side.
The apical end of segment 4 is blunt, forming a
circular plateau, encircled by a low rim (Fig. 3D).
The cuticle on the plateau is somewhat wrinkled,
indicating that it is thinner and more flexible than
in the rest of the segment (Fig. 3B). The five termi-
nal setae project distally from the plateau. They
comprise (Fig. 3B,D,E) two similar plumose setae A
and B, a very short and simple seta C, a conspicu-
ously ornamented seta D, and a long and simple seta
E. Setae A and B sit very close together, while seta
D is adjacent to these two. Seta C sits almost be-
tween D and the AB pair, while seta E sits slightly
but distinctly apart from the other setae. Setae A
and B are ⬃100 –120 ␮m long, with long setules
toward the distal end. Seta C is only 3 ␮m long, with
a terminal pore. Seta D has a complex shape. It sits
on a conspicuous, inverted cone-shaped socket with
a smooth cuticle. From this projects a bilaterally
146 M. BLOMSTERBERG ET AL.

Figure 4
 CYPRIS LARVAE OF OCTOLASMIS AND LEPAS
compressed, 30 – 40 ␮m long sac-like structure orna-
mented with a conspicuous pattern of closely spaced
and ring-shaped bars. The seta ends in a spout-like
structure (Fig. 3C), probably with a terminal pore.
Lepas australis
The carapace has a distinct mid-dorsal hinge.
There are five pairs of lattice organs of the pore-field
type with an anterior terminal pore in LO1-2 and a
posteriorly situated terminal pore in LO-3-5 (for de-
tails, see Jensen et al., 1994b)
Second antennulary segment. The postaxial
seta 2 (PS2) is situated ventrodistally on the second
segment very close to the articulation to the third
segment (Fig. 5C,D). It is comparable to the PS2 in
Octolasmis angulata, but carries more prominent
rows of setules and is encircled by an annulus ⬃20
␮m from its base. The second segment carries an-
other, much smaller seta. This preaxial seta (PrS) is
also sited very close to the articulation to the third
segment, but on the dorsal side slightly medial of the
midline (Fig. 4A).
Third segment. In both Lepas australis, L. pec-
tinata, and Dosima fascicularis, the third segment
consists of little more than the attachment disc
that forms a distended platform at the end of the
segment (Figs. 4A, 6C, 7A). At some angles of view
the disc therefore seems to sit almost directly at
the end of the extensive arthrodial cuticle connect-
ing segments 2 and 3; thus, segment 4 and a row
of several postaxial setae 3 emerge more or less
directly under the rim of the disc. In cyprids of L.
australis, the number of postaxial setae 3 (PS3)
varied from six to eight when they could be
counted (Fig. 4A–E). No cyprid had both anten-
nules extended, so we could not decide whether
the number can also differ between the left and
right antennule. The PS3 setae are longer (200 –
300 ␮m) than the PS2 setae and their setulation is
also more pronounced. As in Octolasmis angulata,
the PS3 setae are attached proximoventrally on
segment 3.
147
Attachment disc. The oval-shaped attachment
disc measures ⬃70 ⫻ 120 ␮m and is densely cov-
ered with microcuticular projections. It is circum-
scribed by two different and concentric rims sep-
arated by a distinct groove. The inner rim is the
swollen margin of the attachment disc and, unlike
a skirt, it therefore carries microcuticular projec-
tions (Fig. 4A–E). On the outward-facing side of
the rim these projections merge into a system of
wrinkles. The outer rim is smooth. The central
part of the attachment disc is somewhat depressed
and surrounded by the swollen inner so that it
resembles a deep bowl. Both the fourth segment
and the PS3 setae sit just outside the outer rim.
The axial sense organ sits at the center of the disc
(Fig. 4G), where it projects from a small area devoid
of microcuticular projections but with scattered
small pores. From this central area extend several
narrow and similarly naked lanes toward the perim-
eter of the disc (Fig. 4G). We observed a small post-
axial organ in one specimen. There are two separate
rows of radial setae around the perimeter of the
attachment disc (Fig. 4D,F), but we could not deter-
mine the exact number in either series. The inner
row sits on the swollen, inner rim, while the outer
row sits in the aforementioned groove between the
inner and outer rims. Both inner and outer radial
setae are simple and ⬃20 –30 ␮m long.
Fourth segment. Four similar subterminal se-
tae, ⬃80 ␮m long, sit in a row on the subterminal
ledge (Fig. 5A); they are slightly curved distally
and have terminal pores. The distal plateau car-
ries five terminal setae (Fig. 5A,B), which differ
somewhat from those in Octolasmis angulata. Se-
tae A and B are similar, 200 –250 ␮m long and
plumose except for the proximal part. Seta C is
only 5 ␮m long; it sits on a broad base but tapers
rapidly and terminates in a pointed tip. Seta D is
a broad, long, and flattened sac-like structure,
ornamented with a reticular structure of ridges,
all the way from base to tip except for the socket.
Seta E is only 15 ␮m long and inserts very close to,
or perhaps actually on, the socket of seta D.

Fig. 4. Lepas australis; antennulary segment 3. A: Oblique ventral view of the distal part of the antennule. Distal part of segment
2 with postaxial seta 2 (PS2) (partially obscured) and additional, more dorsally sited preaxial seta (PrS). Attachment disc on segment
3 is circumscribed by the raised, inner rim. Both the general disc surface and the inner rim are covered by a dense carpet of
microcuticular projections. A second, naked outer rim is separated by a distinct groove from the inner rim. In the center of disc is the
prominent, pointed axial sensory organ (ASO). Radial setae (RS) sits around the periphery of the disc. Six postaxial setae 3 (PS3) are
attached closely adjacent in a row just outside the outer rim, close to the articulation to segment 2. Segment 4 protrudes on the lateral
side of segment 3. B: As A, but the specimen carries seven PS3 setae. C: As A,B, but this specimen has eight PS3 setae. D: Detail of
A; close-up of the proximal part of segment 3 with six PS3 setae. Note how one outer radial seta (white arrow) sits in the groove
between the inner and outer rim of the attachment disc. Inner radial seta is almost obscured (black arrow). The inner rim is distended
and covered with microcuticular projections that most peripherally turn into wrinkles on the surface. Outer rim lacks projections.
E: Detail of B with seven PS3 setae. F: Rim of the attachment disc and a double row of radial setae. Inner and outer rim are separated
by a groove. The radial setae of the inner row are sited on the inner rim among microcuticular projections; the radial setae of the outer
row outer are sited in the groove between the inner and outer rim. G: The pointed axial sensory organ, lateral view. Its base is
surrounded by naked cuticle. The terminal pore is marked with an arrow. AD, attachment disc; ASO, axial sensory organ; PrS; preaxial
seta; PS2, postaxial seta 2; PS3; postaxial seta 3; RS, radial seta; STS, subterminal setae.
148 M. BLOMSTERBERG ET AL.

Fig. 5. Lepas australis; antennulary structures. A: Segment 4, oblique view. A row of four identical, subterminal setae (STS) are on the
ledge and, partly obscured on the terminal plateau, the five terminal setae A–E (in opposite direction). B: Same antennule as depicted in
A. Note seta B; diminutive seta C on a broad base; bag-shaped, ornamented seta D; and seta E. Seta E in L. australis is very short and appears
to insert on the base of D. C: Postaxial seta 2 (PS2), with an annulus (arrow). The seta is fairly short and sparsely setulated. D: Detail of
C showing the PS2 seta and the postaxial setae 3 (PS3). The PS2 seta appears to be setulated in rows, while the PS3 apparently have their
much heavier setulation arranged more uniformly. AD, attachment disc; PS2, postaxial setae 2; PS3, postaxial setae 3; STS, subterminal setae.

Fig. 6. Lepas pectinata. A: Whole cyprid. The hinge region exhibits a conspicuous crest. The carapace is sculptured except for an area
just outside the compound eye (arrow). The two frontal horns protrude anteroventrally. B: Close-up of A. The frontal horn has the same
sculpture as the carapace, but with an added cover of hair-like setae. A thin, naked sheath of cuticle surrounds the gland pore (arrow). C:
Distal part of the antennule; end-on view of attachment disc. The second segment carries one postaxial seta 2 (PS2) and one hook-shaped
preaxial seta (PrS). Note segment 3 with five postaxial setae 3 (PS3) sited just under the attachment disc. The attachment disc is circular
with a dense carpet of microcuticular projections. The inner and outer rims of the attachment disc are difficult to distinguish due to the angle
of view. D: Hook-shaped preaxial seta with distinct terminal pore (arrow) (detail from C). E: The axial sense organ is sited in a depression
and has a distinct terminal pore (arrow). It sits in an area relatively free of microcuticular projections. Arrowheads point to small pores. ASO,
axial sense organ; FH, frontal horn; PrS, preaxial seta; PS2, postaxial setae 2; PS3, postaxial setae 3; RS, radial setae.
CYPRIS LARVAE OF OCTOLASMIS AND LEPAS 149

Figure 6
150 M. BLOMSTERBERG ET AL.

Figure 7
 CYPRIS LARVAE OF OCTOLASMIS AND LEPAS
Lepas pectinata
Carapace. In lateral view the carapace has a
characteristic shape (Fig. 6A). It is very high ante-
riorly but tapers posteriorly toward the pointed end.
A conspicuous crest runs all along the dorsal midline
and curves anteroventrally; it splits in two at the
anterior mantle opening and continues on each side
almost to the level of the frontal horns. The hinge
line between the two carapace valves lies within the
crest, which is heavily sculptured by a row of irreg-
ular knobs. The remaining carapace is sculptured
with a reticular pattern of ridges or knobs, except
from the area outside the compound eye, where the
cuticle is smooth. The frontal horns are very conspic-
uous and sculptured more or less like the remaining
carapace cuticle. Near the tip (Fig. 6B) the horns
carry numerous, small, and simple setae. The ter-
minal opening is surrounded by a sheath of smooth,
thin, and flexible cuticle (for more details, see Elfi-
mov, 1995).
Second antennulary segment. As in Lepas aus-
tralis, the sturdy second segment carries both a
large postaxial seta 2 (PS2) and the much smaller
preaxial seta near the dorsal side (Fig. 6D). In L.
pectinata it is evident that the small seta has a
terminal pore (Fig. 6D).
Third segment. We could only count the number
of postaxial seta 3 (PS3) in two cyprids and only on
one antennule in each. One specimen carried only a
single PS3 seta, while the other had a row of five
identical PS3 setae (Fig. 6C). Whether single or mul-
tiple, the PS3 setae are ⬃100 ␮m long and are fur-
nished with numerous small, short setules. The sin-
gle PS2 seta occurs close to the PS3 setae and differs
in having fewer setules.
Attachment disc. The attachment disc is circu-
lar and the centrally placed axial organ (Fig. 6E) is
in an area of the disc almost devoid of microcuticular
projections. Adjacent to the organ is a distinct
groove, while small pores are scattered on the naked
surface. The axial sense organ has a broad base with
Fig. 7. Dosima fascicularis, antennule. A: Distal part of left
antennule, oblique view. Note one postaxial seta (PS2), one pre-
axial seta (PrS), and four postaxial setae 3 (PS3). Observe the full
view of attachment disc with a characteristic wavy rim with two
rows of peripheral radial setae. The distally placed setae of the
inner row are most prominent. B: Detail of right antennule of the
larva in A. Note five PS3 setae. C: Apical end of segment 4.
Observe five terminal setae (A–E). A and B are similar, both long
and plumose; D is long, bag-shaped, and ornamented; C and E are
diminutive. Four subterminal setae (STS) all similar being long,
setuled, distally hooked and with a terminal pore. D: Detail of C;
note plateau with terminal setae (A–E) surrounded by a high
cuticular rim. A and B are simple basally. C is a small, pointed
seta sitting on a very broad base. D exhibits a socket that is
clearly distinguished from the reticulated bag-shaped part. E is
small, very close to D and has a terminal pore. Observe subter-
minal setae (STS) on a ledge distinct from the distal plateau and
clearly setulated. ASO, axial sense organ; PrS, preaxial seta; PS2,
postaxial setae 2; PS3, postaxial setae 3; STS, subterminal setae.
151
a few microcuticular projections and tapers into a
narrow part terminating in a distinct pore. In our
specimens the axial organ was in a reclined position,
but this could be an artifact of preservation. We
observed no postaxial organ. A row of radial setae
that are situated at the margin of the disc among the
outermost microcuticular projections is easily distin-
guished. We did not observe an outer row of radial
setae with absolute certainty. The true margin of
the disc has the form of a swollen rim, as in Lepas
australis, so the whole attachment disc again has a
bowl-shaped appearance.
Fourth segment. Segment 4 carries four subter-
minal (80 ␮m) and five terminal setae. The terminal
seta D (⬃200 ␮m) has a reticulated surface orna-
mentation and setae C and E are both very small
(not illustrated). The sizes and general appearance
are very much like those described for Lepas aust-
ralis.
Dosima fascicularis
In this species we focused exclusively on the an-
tennules. The second segment resembles that de-
scribed for the two other lepadid species, including
the presence of a preaxial seta (Fig. 7A). In the third
segment the morphology of the attachment disc dif-
fers from what we observed in Lepas. It has a circu-
lar shape, but unlike cyprids of L. australis and L.
pectinata, the margin is not swollen but thin and
wavy. Moyse et al. (1995) described a marginal zone
devoid of microcuticular projections, but we found
that they continue to the very rim of the attachment
disc. There seem to be two rows of radial setae, as in
L. australis, although we only observed a few be-
longing to the outer row. The radial setae of the
inner row are by far the most conspicuous among the
species examined by us. They are sturdier than in
Lepas and Octolasmis and project far beyond the
rim, especially those situated at the distal end of the
disc.
We could only count the number of postaxial setae
(PS3) in one cyprid, where one antennule had five
PS3 setae (Fig. 7B). The other antennule had only
four, but due to the mounting of the larva we could
not determine whether a seta had been lost.
Fourth segment. The four subterminal setae are
very long (⬃80 ␮m), both in an absolute sense and
relative to the other setae and the whole segment
(Fig. 7C). Additionally, they carry minute setules
that continue as a small ridge down the side of the
seta. The five terminal setae resemble those in
Lepas, but the rim of the terminal plateau is more
raised in Dosima fascicularis. In addition, seta E
has a distinct terminal pore.
DISCUSSION
Antennules
The cypris antennules in the three species of
Lepas share some unique characteristics, while the
152 M. BLOMSTERBERG ET AL.
TABLE 1. Character states for antennules in cyprids of Octolasmis, Lepas, and Dosima
Octolasmis Lepas
angulata australis
PrS on segment 2 ⫺ ⫹
segment 3 distinct yes no
number of PS3 1 max. 8
rows of RS 1 2
periphery of AD skirt two rims
seta E diminutive no yes
Boldface character states are putative apomorphies for the Lepadidae. Based on present results and Moyse et al. (1995). AD,
attachment disc on third antennulary segment; PS3, postaxial seta on third antennulary segment; PrS, preaxial seta on second
antennulary segment; RS, radial setae on third antennulary segment; seta E, one of four terminal setae on fourth antennulary
segment.
antennule of Octolasmis angulata cyprids is more
similar to those previously described in other tho-
racicans. The following characters in antennulary
setation are found in all three species of lepadids,
but not in O. angulata nor in any other thoracican
species (Table 1): 1) An extra, small seta, the preax-
ial seta, distally on the second segment in addition
to the single PS2 seta. This seta has rarely if ever
been explicitly described before, but it may not be
unique to lepadids because Glenner and Høeg (1995)
observed a minute spine or seta in exactly the same
position in Balanus amphitrite; 2) multiple PS3 se-
tae on the third segment, in contrast to the single
one seen in all other thoracicans; 3) an extra, outer
row of radial setae around the attachment disc (al-
though we are not absolutely sure of their presence
in Lepas pectinata); 4) terminal seta E on the fourth
segment is very small, whereas it has a length com-
parable to setae A, B, and D in O. angulata and
other thoracicans. This is the first time a diminutive
seta E is described, but it should be noted that
Willemoes-Suhm (1875) observed only three termi-
nal setae in Dosima (then: Lepas) fascicularis:
“…two olfactory hairs between which there is a
broad paddle-shaped appendage…,” since the two
diminutive setae C and E must have been next to
impossible to identify with techniques available to
him. The accuracy of this observation testifies to the
remarkable skill of this author, who sadly died be-
fore he could produce more contributions to cirripede
larval biology; and 5) the four subterminal setae on
the fourth segment are particularly long in the lepa-
dids investigated here, although this character is
harder to compare across the Thoracica.
Characters 2– 4 listed above are putative apomor-
phies for the family Lepadidae (Table 1). Lepadid
cyprids also share the possession of a rather char-
acteristic and very bowl-shaped attachment disc
that almost dwarfs the remaining part of the third
segment. As in most other thoracicans, the exits of
the cement gland are probably located in the depres-
sion around the axial sense organ. The smaller pores
observed in Lepas australis and L. pectinata could
be the exits of unicellular antennulary glands (see
Moyse et al., 1995). The shape is much more conven-
tional in Octolasmis angulata, where the disc sits on
Lepas Dosima
pectinata fascicularis Other thoracica
⫹ ⫹ variable?
no no yes
max. 5 max. 5 1
(2) 2 1
two rims wavy often skirt or velum
yes yes no
a conspicuous bell-shaped segment. In O. angulata
the attachment disc is furthermore encircled by a
cuticular skirt, as is also the case in many other
cirripede cyprids (Glenner et al., 1989; Moyse et al.,
1995). In both species of Lepas the disc has no skirt
but is encircled by a conspicuously swollen rim. This
feature is not found in Dosima fascicularis. In this
species Moyse et al. (1995) argued that the attach-
ment disc is encircled by a “skirt,” but we find that
this is just the thin and flexible rim of the attach-
ment disc itself. A true skirt is devoid of microcu-
ticular projections as in O. angulata.
We conclude that no characters in cypris morphol-
ogy point to a close relationship between Octolasmis
and the Lepadidae (Table 1), but this in no way
disqualifies this phylogenetic hypothesis. A sister
group relationship between Octolasmis and Lepa-
dids is unequivocally supported by the molecular
dataset and this is not contradicted by any morpho-
logical characters. In addition, nauplii of Octolamis
and lepadids share the possession of very long fron-
tal horns and of similarly very long spines on the
hind body (Lang, 1976), and both these traits are
probably additional apomorphies for an Octolasmis-
Lepadidae clade or perhaps a Poecilasmatidae-
Lepadidae clade.
The absence of similarities in cypris antennulary
morphology between Octolasmis and the Lepadidae
is perhaps linked to the very different nature of the
substrata on which they settle. All lepadids attach to
objects in the very specialized neustonic zone,
whereas cyprids of O. angulata settle in the
branchial chamber of crabs. The most conspicuous
character in the antennules of lepadids is perhaps
the multiplication of the PS3 seta, but it is still mere
speculation how this and other traits may relate to
the settlement site. Interestingly, we found no obvi-
ous antennulary adaptation to the specialized set-
tlement site in Octolasmis. In particular, we found
no striking parallels to the antennules of rhizo-
cephalan barnacles, many of which also settle in gill
chambers (Glenner and Høeg, 2002). The general
shape of the third segment and the presence of a
true skirt around the attachment disc are similari-
ties, but rhizocephalan cyprids differ in having spe-
 CYPRIS LARVAE OF OCTOLASMIS AND LEPAS
cialized sac-shaped setae, which Glenner et al.
(1989) suggest are chemosensory aesthetascs.
ACKNOWLEDGMENTS
We are very grateful to Dr. Nikalaj Scharff, Zoo-
logical Museum University of Copenhagen, for hav-
ing spearheaded a successful application for a JEOL
FEG-GUN JSM-6335F Microscope. Anonymous ref-
erees improved the text in several respects. We
thank the entire ’Crustacea Copenhagen’ group of
colleagues for continued interest and support.
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