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Culture and morphology

of some New Zealand
barnacles (Crustacea:
Cirripedia)
a b
M. F. Barker
a
Department of Zoology , University of
Auckland , Private Bag, Auckland, New
Zealand
b
Marine Research Laboratory , R. D. Leigh,
New Zealand
Published online: 30 Mar 2010.

To cite this article: M. F. Barker (1976) Culture and morphology of some New
Zealand barnacles (Crustacea: Cirripedia), New Zealand Journal of Marine and
Freshwater Research, 10:1, 139-158, DOI: 10.1080/00288330.1976.9515605

To link to this article: http://dx.doi.org/10.1080/00288330.1976.9515605

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 N.Z. Journal of Marine and Freshwater Research 10 (1) : 139-58. March 1976

CULTURE AND MORPHOLOGY OF SOME

NEW ZEALAND BARNACLES
(CRUSTACEA: CIRRIPEDIA)
M. F. BARKER

Department of Zoology, University of Auckland, Private Bag, Auckland,

New Zealand, and Marine Research Laboratory, R. D. Leigh,
New Zealand
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ABSTRACT

The larval stages of the acorn barnacles Chamaesipho columna, Chamaesipho

brunnea, Elminius plicatus, Elminius modestus, Balanus trigonus, and Tetraclita
purpurascens are described from larvae reared in the laboratory. A key and a
table are included for separating larval stages and species.
Skeletonema costatum appears to have a wide application as a food in the
rearing of larvae of the Balanidae.
The two Chamaesipho species show typical chthamalid characters of hispid
antennal setae and unilobed labrum. The similarity between the larvae of
Elminius plicatus and Tetraclita purpurascens supports the classification of E.
plicatus as a tetraclitid.

INTRODUCTION

Zooplanktonologists have often grouped larval stages of barnacles as

unidentified "barnacle nauplii" or "barnacle cyprids". Such action
masks the ecological information that could be obtained if specific
identification was possible: some species of barnacles, for example, are
solely intertidal as adults and may be confined to wave-exposed coasts
or harbours. Barnacles seem good candidates for indicators of the origin
and movements of plankton, so long as their larvae can be easily
identified.
The larval stages of 35 barnacle species have been described, mainly
from northern hemisphere waters. There are six free-swimming and feed-
ing naupliar stages, followed by a cyprid stage which settles on the
substrate before metamorphosis to the adult form. Most descriptions have
been based on planktonic material, and the main difficulty of this
approach has been in relating larvae and adult species. Also, a large
number of larvae must be obtained before the larval stages can be
separated by means of size-frequency distribution. A more reliable
method of identifying the larval stages is to rear the larvae from known
parents. A problem inherent in this method is that under culture con-
ditions such features as size and pigmentation may differ from those
found in "wild" larvae from the plankton.

Received 26 June 1975.

 140 N.Z. JOURNAL OF MARINE & FRESHWATER RESEARCH [MARCH

Since first attempts to raise larvae by Bate (1851), more recent

workers have successfully reared cirripede larvae through all the larval
stages (e.g., Tighe-Ford et al. 1970).
Larval stages of some New Zealand balanomorph barnacles have
previously been studied. Elminius modestus larvae were described by
Knight-Jones & Waugh (1949) and B. trigonus larvae by Sandison
(1954) from the plankton of European and South African waters
respectively. Foster (1967) gave details of nauplius I and II of Chamae-
sipho columna, C. brunnea, Tetraclita purpurascens, Elminius plicatus,
E. modestus, and Balanus vestitus and Joll (unpublished 1961) described
the first two nauplii of C. columna.
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In this study, larvae of Chamaesipho columna, C. brunnea, Elminius

plicatus, E. modestus, Balanus trigonus, and Tetraclita purpurascens are
described. Although all the larval stages of E. modestus and B. trigonus
have been previously described they are included for completeness and
because some differences from earlier descriptions have been found in
the New Zealand specimens examined.
The planktonic distribution and abundance of the larval stages of
species described here, will be discussed in a later paper.

MATERIALS AND METHODS

The culture technique for the mass rearing of larvae was similar to
that used by Tighe-Ford et al. (1970). Mature ovigerous lamellae were
removed from the mantle cavity of adult barnacles collected in the Leigh
area, and placed in a fingerbowl of seawater. Nauplii I that hatched
gathered on the side of the fingerbowl nearest a light source. They were
pipetted off into 5-litre pyrex beakers containing 3 litres of filtered sea-
water at a density of 2-3 larvae per millilitre.
At this stage algal food was added, and, to some cultures, 0.4 ml of
Crystamycin (Glaxo Laboratories) solution. This antibiotic solution
contains 300 mg of sodium penicillin G and 0.5 g of streptomycin
sulphate base in* 2 ml of distilled water. The beakers were covered with
plastic and kept at constant temperature (±0.5°c) in a water bath.
Light from neon strip lights and nearby windows entered the cultures.
A 5-cm-diameter strip of black polythene plastic was placed around the
outside of the culture vessel at the water level to prevent the photo-
positive larvae from becoming trapped in the meniscus. The water and
food in the culture vessels was replaced every 2 or 3 days, but on some
occasions, particularly when large numbers of larvae were present at
the beginning of an experiment, algal food was added more frequently.
A magnetic stirrer was used twice daily to resuspend settled algae.
The algal species used as food included uni-algal but not bacteria-free
cultures of the flagellates Isochrysis galbana and Dunaliella primolecta,
and of the diatoms Skeletonema oostatum and Phaeodactylum tri-
cornutum. These were all grown in standard Erdschreider culture medium
 1976] BARKER—BARNACLE LARVAL MORPHOLOGY 141

and pasteurised sea water (Wisely 1960) in 5-litre pyrex Erlenmeyer

flasks kept under constant illumination from neon strip lights. Fresh
cultures were prepared weekly. A 4-litre culture generally reached a
sufficient density for use as food after 7-8 days at 20°c. The density
of algal cells used in experiments was normally undetermined, food
being added until the culture was cloudy. However on a number of
occasions, the density of Skeletonema cultures was determined at
2-3 X 10 6 cells.mH. Between 250 and 500 ml of this algal culture was
added to 3 litres of water in the barnacle culture vessels giving a density
of approximately 2-3 X 105 cells.mh1.
Every 1 or 2 days a quantity of larvae were removed for examination.
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These were preserved in 5% formalin in seawater or 70% alcohol.

Formalin was found to be the better preservative as osmotic differences
between alcohol and seawater tended to distort the naupliar carapace,
particularly of more advanced stages.
Nauplii and cyprids were stained in 1% Gentian violet and cleared in
25% glycerol in 75% seawater (Jones & Crisp 1954) and microscopically
examined with phase-contrast lighting. Drawings were made with the
aid of a camera lucida and from photomicrographs. The following
measurements were made with an ocular micrometer: carapace width -
the greatest width of the body behind the fronto-lateral horns; carapace
length - in nauplius I, II, and III from the tip of the caudal spine to the
frontal margin of the carapace; in nauplius IV, V, and VI from the
frontal margin to the hind margin of the carapace, not including the
carapace spines (because flexure of the caudal spine and abdominal
process made total length measurements inaccurate).
Cyprid length was measured from the anterior to the posterior margin
of the bivalve carapace. Cyprid depth measurements were from dorsal
to ventral margins of the carapace measured at the deepest point.

TERMINOLOGY USED IN DESCRIBING CIRRIPEDE LARVAE

Cirripede nauplii (Fig. 1) can be distinguished from other crustacean

nauplii by the presence of paired fronto-lateral horns (f.l.h.). A broad
triangular carapace (car.) is continued posteriorly in the first three
naupiliar stages as a caudal spine (c.s.), but in nauplius IV, V, and VI,
the carapace possesses a definite posterior border and the caudal spine
articulates with the carapace. In some species a pair of spines project
from the posterior carapace border (p.c.s.). Ventral to the caudal spine
lies the abdominal process (a.p.), from the terminal end of which pro-
ject the furcal rami (f.r.). One or more pairs of lateral ventral spines
(l.v.s.) may also be present on the abdominal process.
All nauplii possess a central anterior nauplius eye (c.e.) and in the
last nauplius (VI) a pair of antero-lateral cyprid eyes develop, but may
not always be pigmented and obvious.
Just anterior to the nauplius eye a pair of fine frontal filaments (f.f.)
project beyond the anterior margin of the carapace. Ventrally and
medially a single-lobed or trilobed labrum (la.) projects posteriorly,
 142 N.Z. JOURNAL OF MARINE & FRESHWATER RESEARCH [MARCH
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Lateral view

Ventral view

FIG. 1—Terminology of barnacle nauplii as used in this paper:

f.f. = frontal filament ex. ^ exopodlte
f.l.h. = fronto-lateral horn. set. : = seta
c.e. = central eye setii. = setule
antu. = antennule p.c.s. — posterior carapace spine
3a. — Iabnim a.p. ~ abdominal process
ant. = antenna l.v.s. = lateral ventral spine
ma. = mandible f.r. = furcal ramus
car. — carapace c.s. = caudal spine
en. ~ endopodite

TABLE 1—Time taken (days) for the appearance of the larval stages in the most pro-
ductive cultures of the barnacle species named (* = Dunaliella added; f = Skele-
tonema added; - = no larvae detected; *** — time unknown)
WATER DAYS TO APPEARANCE OF
TEMPERATURE STAGE
SPECIES ALGAL FOOD (°c) I II III IV V
VI cy
Chamaesipho columna Isochrysis 20 1 1 4 4 7 12 17
Chamaesipho brunnea Isochrysis 20 1 1 4 4 10 12 -
Isochrysis 25 1 1 5 14 - - —
Skeletonema 20 1 1 5 - - — -
Skeletonema 25 1 1 - - - - -
Isochrysis
and others 20 1 1 4 7 11* 14f 33
Elminius plicatus Skeletonema 20 1 1 3 4 6 7 8
Isochrysis 20 1 1 6 - — - -
Dunaliella 20 1 1 3 6 8 - -
Elminius modestus Skeletonema 20 1 1 2 3 4 5 6
Skeletonema 25 1 1 4 4 8 8 -
Skeletonema 30 1 1 _ _ _ _ _
Balanus trigonus Isochrysis 20 1 1 6 9 13 19 -
Skeletonema 20 1 1 6 6 6 8 15
Tetraclita purpurascens Skeletonema 20 1 1 6 6 8 13
Phaeodactylum 20 1 1 •L 4 - — —
Dunaliella 20 1 1 3 6 — -
 1976] BARKER—BARNACLE LARVAL MORPHOLOGY 143

0-1 mm
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Mandible

FIG. 2- -Setation of the appendages of Tetraclita purpurascens IV to show the

derivation of the setation formula.

with the nauplius mouth at its base. Ventral and lateral views of a stage
IV nauplius larvae are shown in Fig. 1. All nauplius stages have paired
uniramous antennules (antu.), biramous antennae (ant.), and biramous
mandibles (ma.); the shape and arrangement of setae on the limbs of a
Stage IV nauplius is shown in Fig. 2. The three pairs of appendages are
used in swimming and in feeding, and the antennules are also sensory.
The arrangement and number of setae on these appendages and
whether the setae (set.) bear setules (setu.) or not, varies with the
naupliar stage, but is constant for a particular stage within a species and
shows very little variation between the same stages of different species.
The setation formula of Bassindale (1936) has been followed in this
present paper. For example the mandible shown in Fig. 2 has a setation
formula 0.1.4-0.4.3.4.3.G; bold numerals refer to terminal setae.

RESULTS OF LARVAL CULTURES

Cyprids of all six species of barnacles were reared. The results of the
most successful cultures for each species under the different conditions
tested are given in Table 1. In Chamaesipho columna and C, brunnea
larval mortality was high, and only one cyprid of each species was
 144 N.Z. JOURNAL OF MARINE &. FRESHWATER RESEARCH [MARCH

III

I
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IV

VI

cy

FIG. 3—Outline drawings of the larval stages of Chamaesipho columna, all drawn
to the same scale. Ventral (left) and lateral (right) aspects of each naupliar
stage are shown.
 1976] BARKER—BARNACLE LARVAL MORPHOLOGY 145

obtained; however, it was possible to obtain additional planktonic speci-

mens of these species for examination.
In the descriptions of larval stages below, comments on structure are
mainly confined to features of diagnostic importance. Measurements
given are of larvae reared in culture except for some cyprid larvae from
the plankton. Most of the wild larvae examined were within the size
ranges given.

Chamaesipho columna Fig. 3, Tables 2 & 3

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NAUPLII: Carapace nearly as broad as long in II-VI without posterior carapace

spines in III, IV, and V. Frontolateral horns directed ventrally and slightly
posteriorly, well developed in I. Abdominal process and caudal spine weakly
developed in I, the former about two-thirds the length of the latter in II-VI;
number of spines on abdominal process increases from 1 pair in II and III to 4
pairs in V. In VI the bulbous base of the abdominal process bears 6 pairs of
small distinct spines, the developing cyprid thoracic appendages. Labrum unilobed,
with two lateral teeth terminally. Setation of appendages shown in Table 2.
Antenna with hispid seta on basal endopod segment.
CYPRID: Blunt anterior and pointed posterior end. Dorsal margin rounded with
a distinct mid dorsal lump.

Chamaesipho brunnea Fig. 4, Tables 2 & 3

NAUPLII : Carapace as broad as long in II-VI (not I), with posterior carapace
spines in IV-VI. Frontolateral horns prominent and directed ventrally and slightly
anteriorly in II-VI, very long and posteriorly directed in I. Abdominal process
and caudal spine weakly developed in I, the former about two thirds the length
of the latter in II-VI. Spines on abdominal process identical to those in C.
columna. Labrum unilobed with two lateral teeth terminally. Setation of append-
ages shown in Table 2. Antenna with hispid seta on basal endopod segment.
CYPRID: Similar shape to C. columna, but not so rounded. Posterior end of
carapace may have prominent area of dark reddish-purple pigmentation.

Ekninius plicatus Fig. 5, Tables 2 & 3

NAUPLII : Carapace triangular, especially in I—III; length slightly less than twice
the width. Posterior carapace spines present in IV-VI. Fronto-lateral horns
prominent, projecting anteriorly in II-VI. Frontal filaments present in stages
II-VI. Abdominal process and caudal spine weakly developed in I, the former
about two-thirds the length of the latter in II-VI. Furcal rami diverge at an angle
greater than 40°. Labrum trilobed, with a fairly short central lobe. Setation of
appendages (Table 2) typically balanid with characteristic comb-like setae.
CYPRID: Pointed posterior and anterior ends. Dark purple pigmentation towards
posterior end, shown as a stippled area in Figure 5.

Elminius modestus Fig. 6, Tables 2 & 3

NAUPLII: Carapace triangular, length less than twice the width. Short posterior
carapace spines present in IV-VI. Short fronto-lateral horns directed anteriorly
in II-VI. Frontal filaments present in stages II-VI. Abdominal process and
caudal spine weakly developed in I, the former half tfae length of the latter in
If, two-thirds the length in IV-VI. Labrum trilobed, with median lobe extending
well beyond lateral lobes. Setation of appendages shown in Table 2.
CYPRID: Dorso-ventrally compressed carapace, and pointed anterior and posterior
ends, giving a fusiform shape.

Sig—10
 146 N.Z. JOURNAL OF M A R I N E & FRESHWATER RESEARCH [MARCH

111
II
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FIG. 4—Outline drawings of the larval stages of Chamaesipho brunnea, all

drawn to the same scale. Ventral (left) and lateral (right) aspects of each
naupliar stage are shown.

OPPOSITE
FIG. 5—Outline drawings of the larval stages of Elminius plicatus, all drawn to
the same scale. Ventral (left) and lateral (right) aspects of each naupliar
stage are shown.
Downloaded by [Nara Women's University] at 22:09 08 March 2015

1976]

11
ill
BARKER—BARNACLE LARVAL MORPHOLOGY
147
 148 N.Z. JOURNAL OF MARINE & FRESHWATER RESEARCH [MARCH

TABLE 2—Setation formulae of the appendages of the reared nauplius larval stages of
Chamaesipho columna (C. c), Chamaesipho brunnea (C. b.), Eliminius plicatus
(E. p.), Eliminius modestus (E. m.), Balanus trigonus (B. t.), and Tetraclita
purpurascens (T. p.) (formulae in parentheses are differences found by earlier
workers).

STAGE SPECIES ANTENNULE ANTENNA MANDIBLE

C.c. 0.4.2.1.1 0.1.4-0.3.2.2.2.G 0.1.3-0.3.2.2.2.G

(0.3.2.1.1) (0.1.4-0.2.1.2.2.G) (0.1.3-0.3.2.2.2. G)
C.b. 0.4.2.1.1 0.1.4-0.3.2.2.2.G 0.1.3-0.3.2.2.2.G
E.m. 0.4.2.1.1 0.1.4-0.3.2.2.2.G 0.1.3-0.3.2.2.2.G
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E.p. 0.4.2.1.1 0.1.4-0.3.2.2.2.G 0.1.3-0.3.2.2.2.G

B.t. 0.4.2.1.1 0.1.4-0.3.2.2.2.G 0.1.3-0.3.2.2.2.G
(-0.3.2.1.2.G)
T.p. 0.4.2.1.1 0.1.4-0.3.2.2.2.G 0.1.3-0.3.2.2.2.G

C.c. 0.4.2.1.1 0.2.5-0.3.2.2.3.G 0.1.4-0.3.2.3.2.G

(0.3.2.2.1) (0.2.4-0.3.1.2.2.G) (0.1.4-0.3.2.3.2.G)
C.b. 0.4.2.1.1 0.2.5-0.3.2.2.3. G 0.1.4-0.3.2.3.2.G
II E.m. 0.4.2.1.1 0.2.5-0.3.2.2.3.G 0.1.3-0,3.2.3.2.G
(0.1.6-0.3.2.2.3.G)
E.p. 0.4.2.1.1 0.2.5-0.3.2.2.3.G 0.1.4-0.3.2.3.2.G
B.t. 0.4.2.1.1 0.2.5-0.3.2.2.3.G 0.1.3-0.3.2.3.2.G
T.p. 0.4.2.1.1 0.2.5-0.3.2.2.3.G 0.1.4-0.3.2.3.2.G

C.c. 1.4.2.1.1 0.2.5-0.3.2.2.4.G 0.1.4-0.3.3.3.3.G

C.b. 1.4.2.1.1 0.2.5-0.3.2.2.4.G 0.1.4-0.3.3.3.3. G
E.m. 1.4.2.1.1 0.2.5^0.3.2.2.4.0 0.1.4-0.3.3.3.3.G
III (0.1.6-0.3.2.2.4.G)
E.p. 1.4.2.1.1 0.2.5-0.3.2.2.4.G 0.1.4-0.3.3.3.3.G
B.t. 1.4.2.1.1 0.2.5-0.3.2.2.4.G 0.1.4-0.3.3.3.3.G
T.p. 1.4.2.1.1 0.2.5-0.3.2.2.4.G 0.1.4-0.3.3.3.3.G

C.c. 1.1.4.2.1.1 0.2.7^0.5.3.3.4.G 0.1.4-0.5.3.4.3.G

C.b. 1.1.4.2.1.1 0.2.7-0.5.3.3.4.G 0.1.4-0.S.3.4.3.G
IV E.m. 1.1.4.2.1.1 0.2.7-0.5.3.2.4.G 0.1.4-0.4.3.3.3.G
E.p. 1.1.4.2.1.1 0.3.6-0.5.3.2.4.G 0.1.4-0.4.3.4.3.G
B.t. 1.1.4.2.1.1 0.3.6-0.5.3.2.4.G 0.1.4-0.4.3.4.3.G
T.p. 1.1.4.2.1.1 0.3.6-0.5.3.2.4.G 0.1.4-0.4.3.4.3.G

C.c. 1.1.1.4.2.1.1.1 0.2.9-0.5.3.4.4.G 0.1.5-0.5.4.4.3.G

C.b. 1.1.1.4.2.1.1.1 0.2.9-0.5.3.4.4.G 0.1.5-0.5.4.4.3.G
E.m. 1.1.1.4.2.1.1.1 0.3.8-0.5.3.2.4.G 0.1.5-0.4.4.4.3.G
V (2.1.4.2.1.1.1)
E.p. 1.1.1.4.2.1.1.1 0.4.7-0.5.3.2.4.G 0.1.5-0.4.4.4.3.G
B.t. 1.1.1.4.2.1.1.1 0.4.7-0.5.3.2.4.G 0.1.5-0.4.4.4.3.G
(-0.4.3.4.3.G)
T.p. 1.1.1.4.2.1.1.1 0.3.7-0.5.3.2.4.G 0.1.5-0.4.4.4.3.G

C.c. 1.1.1.4.2.1.2.1 0.3.9-0.5.4.4.4.G 0.1.5-0.6.4.4.3.G

C.b. 1.1.1.4.2.1.2.1 0.3.9-0.5.3.4.4.G 0.1.5-0.5.4.4.3.G
E.m. 1.1.1.4.2.1.2.1 0.4.8-0.5.3.2.4.G 0.1.5-0.4.4.4.3.G
VI (2.1.4.2.1.2.1)
E.p. 1.1.1.4.2.1.2.1 0.5.7-0.5.3.2.4.G 0.1.5-0.4.4.4.3.G
B.t. 1.1.1.4.2.1.2.1 0.4.7-0.5.3.2.4.G 0.1.5-0.4.4.4.3.G
T.p. 1.1.1.4.2.1.2.1 0.3.7-0.5.3.2.4.G 0.1.5-0.4.4.4.3.G
 1976] BARKER—BARNACLE LARVAL MORPHOLOGY 149

TABLE 3—Size-ranges of reared larval stages of Chamaesipho columna (C. c), Chamae-
sipho brunnea (C. b.), Elminius plicatus (E. p.), Balanus trigonus (B. t.) and
Tetraclita purpurascens (T. p.) (sizes in parentheses are those given by earlier
workers; * = particularly large T. purpurascens VI obtained in one culture).

STAGE SPECIES CARAPACE LENGTH (mm) CARAPACE WIDTH (mm)

C.c. 0.20-0.23 (0.21 ±0.03) 0.10(0.10±0.02)

C.b. 0.19-0.21 0.09-0.10
I E. m. 0.21-0.23 (0.24-0.26) 0.10-0.11 (0.11-0.14)
E.p. 0.30-0.32 0.12-0.14
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B. t. 0.21-0.23 (0.24) 0.10(0.14)

T.p. 0.29-0.31 0.13-0.14

C. c. 0.30-0.32 (O.35±O.05) 0.16-0.17 (0.17±0.03)

C.b. 0.35-0.37 0.18-0.19
II E. m. 0.36-0.38(0.36-0.43) 0.15-0.16(0.15-0.17)
E.p. 0.47-0.50 0.19-0.20
B. t. 0.36-0.38(0.36) 0.16-0.17(0.14)
T.p. 0.45-0.47 0.19-0.20

C.c. 0.37 0.21

C.b. 0.42-0.44 0.24-0.25
III E. m. 0.41-0.45(0.35-0.43) 0.18-0.20(0.18-0.20)
E.p. 0.56-0.59 0.23-0.24
B. t. 0.45-0.46(0.38) 0.22-0.23 (0.19)
T.p. 0.53-0.55 0.22-0.24

C.c. 0.27-0.29 0.24-0.27

C.b. 0.30-0.32 0.30-0.32
IV E. m. 0.28-0.29(0.29-0.35) 0.22-0.24(0.22-0.25)
E.p. 0.34-0.36 0.26-0.30
B. t. 0.33-0.36(0.30) 0.30-0.31 (0.26)
T.p. 0.33-0.35 0.24-0.26

C.c. 0.33-0.35 0,29-0.31

C. b. 0.40-0.41 0.40-0.41
V E. m. 0.34-0.37(0.34-0.44) 0.27-0.29(0.26-0.31)
E.p. 0.45-0.50 0.37-0.40
B. t. 0.46-0.49(0.38) 0.38-0.42(0.32)
T.p. 0.43 0.33-0.34

C.c. 0.37-0.40 0.35-0.36

C.b. 0.50-0.52 0.50-0.51
VI E. m. 0.44-0.48(0.42-0.55) 0.35-0.37 (0.33-0.41)
E.p. 0.55-0.56 0.45-0.47
B. t. 0.59-0.61 (0.57) 0.49-0.51 (0.43)
T.p. 0.50-0.52(0.67-0.69)* 0.40^0.43 (0.54-0.55)*

C.c. 0.44-0.48 DEPTH 0.21-0.26

C. b. 0.58-0.60 0.28-0.29
Cyprid E. m. 0.51-0.58(0.54-0.56) 0.23-0.26
E.p. 0.65-0.70 0.30-0.35
B. t. 0.63-0.66 0.30-0.33
T.p. 0.55-0.58 0.24-0.26
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150
N.Z. JOURNAL OF MARINE & FRESHWATER RESEARCH
[MARCH
 1976] BARKER—BARNACLE LARVAL MORPHOLOGY 151

Balaniis trigonus Fig. 7, Tables 2 & 3

NAUPLII: Carapace triangular in I—II, but rather "square" in outline in III-VI,
with a convex anterior margin. Well-developed posterior carapace spines in
IV-VT. Prominent fronto-lateral horns directed posteriorly in I, horizontally in
II, and anteriorly in III-VI. Frontal filaments present in II-VI. Abdominal process
and caudal spine weakly developed in I, the former increasing in length during
development from half the length of the caudal spine in II to nearly equal in VI.
Labrum trilobed, median lobe does not extend beyond lateral lobes. Setation of
appendages shown in Table 2.
CYPRID: Large, with rounded anterior end and dorsal margin.
Tetraclita purpurascens Fig. 8, Tables 2 & 3
NAUPLII: Carapace triangular. Short posterior carapace spines in IV-VI. Fronto-
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lateral horns prominent, directed posteriorly in I, anteriorly in II-VI. Frontal

filaments present in II-VI. Abdominal process and caudal spine weakly developed
in I, the former about two-thirds the length of the latter in. II-VI. Furcal rami
diverge at an angle of less than 30°. Labrum trilobed, median lobe does not
extend beyond lateral lobes. Setation of appendages shown in Table 2.
CYPRID: Shape similar to E. plicatus. Pointed anterior and slightly less pointed
posterior ends.

DISCUSSION
LARVAL CULTURE
The type and concentration of algal food appears to be a critical
factor in the rearing of cirripede larvae. Moyse (1960) and Moyse &
Knight-Jones (1967) found that they could rear cyprids of the boreal
species Balanus balanoides with diatoms as food but not with flagellates,
and that those of the tropical Chthamalus steliatus and oceanic Lepas
anatifera could be reared on flagellates but not on diatoms. They suggest
that this might reflect some general trends in the distribution of the
various phytoplankton groups because " . . . diatoms are particularly
well known for their contributions to the plankton and abyssal oozes of
high latitudes" . . . "while dinoflagellates and coccolithophores contribute
more to the plankton of warmer seas and summer regions" (Moyse &
Knight-Jones 1967, p. 605).
Their results are interesting when compared to the diet on which
Chamaesipho columna and C. brunnea cyprids were reared. C. columna
was reared to the cyprid stage on the flagellate Isochrysis galbana. C.
brunnea proceeded to nauplius VI on a diet of equal amounts of the
flagellates Isochrysis and Dunaliella primolecta. Metamorphosis to the
cyprid stage did not occur until the diatom Skeletonema was supplied to
the sixth stage nauplii; thus, diatoms may be important in the diet of
later stage larvae of this species. In cultures with Skeletonema as the
only food source, development did not proceed past nauplius III.
The best results in the present study were obtained with Skeletonema
costatum. Although some development of nauplius stages occurred with
the flagellates Isochrysis galbana and Dunaliella primolecta and the

OPPOSITE
Flo. 6—Outline drawings of the larval stages of Elminius modestus, all drawn to
the same scale. Ventral (left) and lateral (right) aspects of each naupliar
stage are shown.
 152 N.Z. JOURNAL OF M A R I N E & FRESHWATER RESEARCH [MARCH

III
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FIG. 7—Outline drawings of the larval stages of Balanus trigonus, all drawn to
the same scale. Ventral (left) and lateral (right) aspects of each naupliar
stage are shown.
 1976] B A R K E R — B A R N A C L E LARVAL MORPHOLOGY 153
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FIG. 8—Outline drawings of the larval stages of Tetraclita purpurascens, all

drawn to the same scale. Ventral (left) and lateral {right) aspects of
each naupliar stage are shown.
 154 N.Z. JOURNAL OF MARINE & FRESHWATER RESEARCH [MARCH

diatom Phaeodactylum tricornutum, development never proceeded to the

cyprid. Phaeodactylum, although ingested by the nauplii was apparently
not digested, as faecal pellets contained compacted but intact plant
cells. This may explain the low survival rate which Wisely (1960)
obtained with Elminius modestus. Other workers (Moyse 1960, 1961,
1963; Moyse & Knight-Jones 1967; Hudinaga & Kasahara 1941; and
Tighe-Ford et al. 1970) have all reported successful development of a
variety of barnacle species using S. costatum and it appears to have a
wide application as a food in the rearing of larvae of the Balanidae.

DESCRIPTION OF LARVAL STAGES

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The identification of cirripede larvae to species has long been a

problem to the planktonologist. The setation formula introduced by
Bassindale (1936) is useful in determining the different stages of a given
species, but has limited value in distinguishing between species. This is
largely because of the similarity in setation of many species (Costlow &
Bookhout 1958) and also because of variation within a naupliar stage
of a given species (Norris et al. 1951, Bassindale 1936). The present
study has shown that there may be differences in the naupliar setation
within a species described both from local specimens and those collected
from other areas. This has been found for Chamaesipho columna (Joll
unpublished 1961) from South Island waters, in Elminius modestus
(Knight-Jones & Waugh 1949) from European waters, and in Balanus
trigonus (Sandison 1954) from South Africa (Table 2).
In C. columna and B. trigonus differences are due either to variations
in the larvae examined or, more likely, to an oversight of setae by
previous authors. The different setation found for E. modestus is
probably due to a different interpretation of what constitutes a group
of setae, rather than to a structural difference in the larvae. The text
diagrams of Knight-Jones & Waugh (1949) certainly indicate identical
setation to New Zealand nauplii.
Although setation formulae do in some cases indicate differences, they
tend to be impractical when dealing with large numbers of specimens
because larvae have to be dismembered and examined under high
magnification. For the planktonologist the most convenient means of
identification would be based on more readily recognisable morphological
details, such as shape of carapace, degree of development of spines on
abdominal process, shape of labrum, angle of divergence of furcal rami,
and details of setation obvious under a binocular microscope. Size has
been found to be of limited value as a diagnostic character (Pyefineh
1948, Bassindale 1936, Sandison 1967). Crisp (1962) found great varia-
tion in size of nauplius larvae of B. balanoides from different latitudes
and suggests that size is related to temperature, with larger sizes in
cooler waters. The cyprid larvae, too, has a wide size-range. In the
present study, cyprids obtained from the plankton were often larger
than those obtained by culture methods. Even larvae of a particular
species from the same culture under controlled conditions showed
considerable variations in size.
 1976] BARKER—BARNACLE LARVAL MORPHOLOGY 155

Nevertheless, it is possible to recognise most nauplii of species en-

countered in this study. Stage I nauplii are difficult to distinguish, but
are rarely found in plankton samples. Characters used to separate larval
stages are given in Table 4, and those used to separate species in the
naupliar stages are given in the key provided:

KEY TO IDENTIFICATION OF NAUPLII OF SIX BARNACLE SPECIES

1. Unilobed labrum, no frontal filaments Chthamalidae 2.
Trilobed labrum, frontal filaments present Balanidae 3.
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2. Carapace rounded, no posterior carapace spines in

IV—VI Chamaesipho columna
Carapace square, posterior carapace spines present in
IV-VI Cbrunnea
3. Labrum with protuberant centre lobe Elminius modestus
Labrum with short centre lobe 4.
4. Carapace only slightly longer than wide Balanus trigonus
Carapace considerably longer than wide 5.
5. Angle between furcal rami 40° E, plicatus
Angle between furcal rami 30° Tetraclita purpurascens
The close similarity of cyprid larvae, however, has been noted by a
number of workers (Moyse 1961, Norris & Crisp 1954, Costlow & Book-
hout 1957, 1958; Crisp 1962, Pyefinch 1948) and their specific identifica-
tion in plankton samples remains a problem. In some descriptions, the
cyprids have been omitted, either because they could not be reared or
were not readily assignable on temporal and spatial grounds to a
particular species from plankton samples. Most descriptions have utilised
general shape, colour, size, and the position of the median and paired
eyes within the carapace. Size is an unreliable character and the position
of the eyes appears to vary according to the degree of extension of the
antennules. The number of setae and spines, and the pattern of setation
and setulation of the six pairs of thoracic limbs are all the same in some
species (Moyse 1961). Significant differences could not be found in the
present study, but the carapace shape and pigmentation seemed to be
reliable diagnostic features, although even these may vary on occasions
(Moyse 1961): when used in conjunction with relative size, a fairly
reliable identification can be made.
The morphology of stage III-VI nauplii of Chamaesipho spp. has not
been described before. Other chthamalids of which the larvae have been
described are Chthamalus stellatus (Bassindale 1936), C. dentatus and
Octomeris angulosa (Sandison 1954), and C. aestuarii (Sandison 1967).
Diagnostic features shown by all the chthamalid nauplii so far described
are the hispid seta of the antenna and the unilobed labrum. The posterior
carapace spines characteristic of nauplii IV-VI of balanid species are
only present in C. brunnea and O. angulosa. The nauplii of the two
species of Chamaesipho differ in the posterior carapace spines mentioned
above, the prominence, size, and direction of the fronto-lateral horns,
and in that there is one less seta on the endopodite of the mandible and
antenna of nauplius VI of C. brunnea.
 156 N.Z. JOURNAL OF MARINE & FRESHWATER RESEARCH [MARCH

TABLE 4—Features of diagnostic importance for larval stages of Chamaesipho

columna, C, brunnea, Elminius plicatm, E. modestus, Balanus trigonus and
Tetraclita purpurascens reared in the laboratory

Nauplius I
Setae of appendages without setules. Setation formula of antennule 0.4.2.1.1.
Reduced caudal spine and abdominal process.
Nauplius II
Some setae of appendages setulate. This also occurs in all subsequent stages.
Setation formula of antennule 1.4.2.1.1. 1 pair of lateral spines on ventral side of
abdominal process.
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Nauplius III
Setation fomulae of antennule 1.4.2.1.1. 1 pair of lateral spines on ventral side
of abdominal process. Posterior border to carapace and developing carapace
spines of stage IV may be visible through exoskeleton.
Nauplius IV
Setation formula of antennule 1.1.4.2.1.1. Posterior border to carapace which
may carry carapace spines. This is also present in all subsequent stages. 2 pairs
of lateral spines on ventral side of abdominal process.
Nauplius V
Setation formula of antennule 1.1.1.4.2.1.1.1. 3 pairs of lateral spines on ventral
side of abdominal process Additional small spines may also be present, but are
often difficult to distinguish under low magnification. Characteristics of nauplius
VI may make their appearance. Muscle blocks of future cyprid thoracic segments
are frequently discernible within the swollen abdominal process.
Nauplius VI
Setation formula of antennule 1.1.1.4.2.1.2.1. A row of 6 pairs of spines appear
on base of abdominal process. Paired anteriolateral eyespots may be present in
older larvae. Penultimate segment of antennule swollen in position of cyprid
attachment organ. Paired muscle blocks of cyprid "thorax" visible within
abdomen.
Cyprid
Distinct bivalved carapace, 1 pair of modified antennules and 6 pairs of
thoracic appendages.

Of the other species described here, usually classified in the Balanidae,

there is a striking similarity between the nauplii of E. plicatus and T.
purpurascens. Recent studies of adult balanomorphs have suggested that
the tetraclitids should deserve full familial rank, and that Elminius
plicatus is rightly classified as a tetraclitid (Foster, in press). The
similarities noted between larvae of E. plicatus and an undoubted
tetraclitid supports these adjustments.

ACKNOWLEDGMENTS

I wish to thank Dr W. Ballantine for facilities provided by the Leigh Marine

Laboratory, and Dr B. A. Foster for encouragement and helpful criticism of the
manuscript.
 1976] BARKER—BARNACLE LARVAL MORPHOLOGY 157

LITERATURE CITED

BASSINDALE, R. 1936: The developmental stages of three English barnacles,

Balanus balanoides, Chthamalus stellatus and Verruca stroemia. Pro-
ceedings of the Zoological Society of London 106: 51-14.
BATE, C. S. 1851: On the development of the Cirripedia. Annals and Magazine
of Natural History 8, ser. 2: 324-32.
COSTLOW, J. D. & BOOKHOUT, C. G. 1957: Larval development of Balanus
eburneus in the laboratory. Biological Bulletin, Woods Hole 112;
313-24.
1958. Larval development of Balanus amphitrite var. denticulata Broch
Downloaded by [Nara Women's University] at 22:09 08 March 2015

reared in the laboratory. Biological Bulletin, Woods Hole 114: 284-95.

CRISP, D. J. 1962: The planktonic stages of the Cirripedia Balanus balanoides
(L.) and Balanus balanus (L.) from north-temperate waters. Crust-
aceana 3: 207-21.
FOSTER, B. A. 1967: The early stages of some New Zealand shore barnacles.
Tane 13: 33-42.
FOSTER, B. A. in press: The Marine Fauna of New Zealand. Barnacles (Cirri-
pedea: Thoracica). N.Z. Oceanographic Institute Memoir 69.
HUDINAGA, M. & KASAHARA, H. 1941: Larval development of Balanus amphitrite
hawaiiensis. Zoological Magazine (Tokyo) 54: 108-18.
JOLL, J. unpublished 1961: Some aspects of development and life cycle of
Chamaesipho columna. M.Sc. thesis lodged in University of Canter-
bury Library, Christchurch, N.Z. 100 pp.
JONES, L. W. G. & CRISP, D. J. 1954: The larval stages of the barnacle Balanus
improvisus Darwin. Proceedings of the Zoological Society of London
123: 765-80.
KNIGHT-JONES, E. W. & WAUGH, G. D. 1949: On the larval development of
Elminius modestus Darwin. Journal of the Marine Biological Associa-
tion of the U.K. 28: 413-28.
MOYSE, J. 1960: Mass rearing of barnacle cyprids in the laboratory. Nature,
London 185: 120.
1961: The larval stages of Acasta spongites and Pyrogoma anglicum
(Cirripedia). Proceedings of the Zoological Society of London 137:
371-92.
1963: A comparison of the value of various flagellates and diatoms
as food for barnacle larvae. Extrait du Journal du Conseil Inter-
national Pour L'Exploration de la Mer 28 (2) : 175-87.
MOYSE, J. & KNIGHT-JONES, E. W. 1967: Biology of cirripede larvae. Proceedings
of Symposium on Crustacea, Marine Biological Association of India
2: 595-611.
NORRIS, E. & CRISP, D. J. 1954: The distribution and planktonic stages of the
cirripede Balanus perforatus Bruguiere. Proceedings of the Zoological
Society of London 123: 393-409.
NORRIS, E., JONES, L. W. G., LOVEGROVE, T. & CRISP, D. J. 1951: Variability in
larval stages of cirripedes. Nature London 167: 444-5.
PYEFINCH, K. A. 1948: Methods of identification of the larvae of B. balanoides
(L.), B. crenatus Bruguiere and Verruca stroemia (O. F. Muller).
Journal of the Marine Biological Association of the U.K. 27: 451-63.
 158 N.Z. JOURNAL OF MARINE & FRESHWATER RESEARCH [MARCH

SANDISON, E. E. 1954: The identification of the nauplii of some South African

barnacles with notes on their life histories. Transactions Royal Society
of South Africa 34: 69-101.
1967: The naupliar stages of Balanus pallidus stutsburi Darwin and
Chthamalus aestuarii Stubbings (Cirripedia: Thoracica). Crustaceana
13: 161-74.
TIGHE-FORDE, D. J., POWER, M. J. D. & VAILE, D. C. 1970: Laboratory rearing
of barnacle larvae for antifouling research. Heligolander Wissenschaft-
liche Meeresuntersuchungen 20: 393-405.
WISELY, B. 1960: Experiments on rearing the barnacle Elminius modestus to the
settled stage in the laboratory. Australian Journal Marine and Fresh-
water Research 11: 42-54.
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