J OURNAL OF C RUSTACEAN B IOLOGY, 34(5), 663-670, 2014
THE RISE AND FALL OF PYRGOPSELLA YOUNGI – REDISCOVERY OF
A LOST SPECIES
Yair Achituv ∗ and Noa Simon-Blecher
The Mina and Everard Goodman Faculty of Life Sciences, Bar-Ilan University, Ramat-Gan 5290002, Israel
ABSTRACT
The coral-inhabiting barnacle Pyrgopsella annandalei was collected in 1888 off the reefs of the Andaman Islands in the Indian Ocean,
diagnosed in 1906, and described in full in 1907. Since then, this barnacle has not been recorded. In 2006, several specimens of Pyrgopsella
were found embedded in the hermatypic coral Symphyllia radians. Based on morphological differences between this material and the
drawings and written description of P. annandalei, the specimens from Symphyllia were assigned to a new species, P. youngi. The discovery
of a single individual of Pyrgopsella in the collection of the Natural History Museum, London, labeled “cotype,” and its comparison to the
recent material from Symphyllia, revealed that the differences between P. annandalei and P. youngi represent no more than intraspecific
morphological variation. This conclusion is supported by a comparison of the DNA sequences of the CO1 and 12S rRNA genes from
specimens representing both morphological varieties. It is concluded that P. youngi is a junior synonym of P. annandalei, and the latter
name should be used in its place.
K EY W ORDS: junior synonym, Pyrgopsella annandalei, P. youngi, rediscovery
DOI: 10.1163/1937240X-00002257
I NTRODUCTION
Pyrgopsella annandalei (Gruvel, 1906) is an example of a
taxon that was recorded once and then not found or recorded
again for a century or even longer. The description of P.
annandalei was based on an unspecified number, probably
one, specimens of pyrgomatid barnacle dredged from the
vicinity of the Andaman Islands and subsequently stored in
the collections of the Indian Museum in Calcutta (Gruvel,
1906, 1907). Pyrgopsis proved to be a junior homonym and
Zullo (1967) proposed the replacement name Pyrgopsella
for it. In 1971, Rosell found a barnacle embedded in a
sponge (Rosell, 1973) described it as the new species Pyrgopsella stellula Rosell, 1973 and duplicated in Rosell, 1975
while suggesting that Pyrgopsella annandalei as well was
a sponge-inhabiting barnacle. A century after the description of P. annandalei, Achituv and Simon-Blecher (2006)
found several specimens of Pyrgopsella on the hermatypic
coral Symphyllia radians Milne-Edwards and Haime, 1849.
They thus confirmed that this genus is comprised of coralinhabiting barnacles, not sponge barnacles, and described
their find as the new species Pyrgopsella youngi Achituv
and Simon-Blecher, 2006. The sponge barnacle P. stelulla
was transferred to a new genus Pyrgospongia. Since then,
Pyrgopsella has been found again on another colony of the
same coral species.
Pyrgopsella youngi was separated from P. annandalei by
Achituv and Simon-Blecher (2006) on the basis of morphological differences between the material from Symphyllia radians and the description and drawings of Gruvel (1907)
(Fig. 1). The first difference concerns the inward-projecting
∗ Corresponding
internal tooth on the spur of the tergum, which was described
as truncate in P. annandalei whereas that found in P. youngi
is thin and pointed. The second difference relates to the distal
end of the penis: in P. annandalei it was described as pearlike and ornamented with short spines, while in P. youngi this
part of the penis is elongated, the pear-shaped tip is missing,
and it terminates as a cone carrying a few setae.
The material on which the description of the species is
based was sent to Gruvel in August 1905 (Gruvel, 1906),
by Nelson Annandale, who serviced in the Indian Museum
in Calcutta during the years 1904 to 1924. One specimen,
that described herein, was sent by Annandale to London
in 1907, and we can assume that the third one was left in
Calcutta. Nilsson-Cantell (1938) did not include Pyrgopsis
in his report on the cirripedes found in the collection of
the Indian Museum in Calcutta, and we assume that any
material of Pyrgopsis annandalei that might have been kept
there was lost [one of the present authors (Y.A.) did not
get any response to his letters of enquiry to the museum
in Calcutta]. The name on the label found inside the jar is
of the collector, James Wood-Mason, who worked in the
Indian Museum from 1863 to 1893. In 1887, he became the
superintendent there, and in 1888 he sailed on the Indian
Marine Survey steamship HMS Investigator, collecting,
among other samples, crustaceans (Alcock, 1902). It appears
that Pyrgopsella annandalei was collected by him during
that survey.
According to Gruvel (1907), three ‘echantillons’ were
dredged by the Indian Marine Survey steamship HMS Investigator at one locality at the reefs of the Andamans Islands
(11°49 30 N, 92°55 55 E) from approximately 90 m depth.
author; e-mail: [email protected]
© The Crustacean Society, 2014. Published by Brill NV, Leiden
DOI:10.1163/1937240X-00002257
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JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 34, NO. 5, 2014
Fig. 1. Illustrations of Pyrgopsella annandalei modified from Gruvel (1907), plate II. A, general view; B, view from above; C, fragment of shell; D, scutum
and tergum; E, mandible; F, maxilla II; G, apex of penis; H, enlargement of shell surface, outer view.
‘Echantillons’ can be translated as samples or specimens.
However, Gruvel (1906) remarked, without explicitly stating the number of specimens examined, “Quand on examine
un des échantillons par sa face superieure, . . . ” [When examining one of the “échantillons” from its upper side], and
in this context an “échantillon” is unambiguously a specimen, not a sample. Gruvel was aware that three specimens
were collected by HMS Investigator, but we cannot say how
many he had at hand. The fact remains, though, that Gruvel
(1907) presented the dimensions of a single specimen and
not a range of sizes from two or three specimens we therefore assume that he examines a single specimen.
Recently, a jar was found in the cirripede collection of
the Natural History Museum (NHM), London, with one
barnacle inside and a typed label on the outside reading
“Syrgopsis [sic] annandalei Gruv. off Blair Reef. Pres by
(I)ndia M.” The label inside the jar reads, “Pyrgopsis
annandalei Gruv. off Blair Reef. J. Wood Mason cotype. 1907.7.23.16.” Concerning this sample, the register
of the NHM indicates, “present by Dr. Annandale, Indian
Museum Calcutta.” According to recommendation 73E of
the ICZN the use of the term cotype should be avoided. The
labeling of the NHM specimen “co-type” and the location
of its collection “off Blair Reef” (11°41 00 N, 92°45 00 E),
indicate that it is the same place of origin as that of the
nominal species, clarify that the present specimen is part
of the material collected together with Gruvel’s material
and according to Article 73.2.1 and 73.2.3 it should be
designated as syntypes.
A detailed study of the morphology of the syntype of
P. annandalei, together with a comparison to the material
described by Achituv and Simon-Blecher (2006) as P.
youngi, shows that the diagnosis of P. youngi was based
on morphological differences that exist as well among
specimens of P. annandalei, and that there are no other
significant morphological differences. We conclude that
Pyrgopsella is a monospecific genus, and the specific name
youngi should be abandoned as a junior subjective synonym.
M ATERIALS AND M ETHODS
Sample
1907.7.23.16 Natural History Museum, London (NHM): Pyrgopsis annandalei “co-type,” labeled as noted above.
AR27804 previously described as P. youngi, Zoological Museum,
Steinhardt National Collections of Natural History, Tel Aviv University
(TAU), Israel. Host: Co32349 (TAU). Symphyllia radians from Bali,
Indonesia.
NHMUK 2014.396: Pyrgopsella annandalei. Natural History Museum,
London, four specimens. Host coral: NHMUK 2014.3, Symphyllia radians,
Bali, Indonesia.
The morphology of 1907.7.23.16 (NHM) was described and compared
to specimens previously described as P. youngi (AR27804 (TAU)) and to
NHMUK 2014.396 (NHM), the specimens of these samples were used in
both morphological and molecular studies.
After more than a century of preservation, the physical condition of the
specimen from the NHM, marked as “co-type,” was very delicate. Its hard
parts were fragile, soft parts very pliant, and photographs could not show
all the details. In order to reduce damage, cirri were removed from only one
side of the body. Our efforts to isolate and sequence DNA from its muscles
proved futile.
ACHITUV AND SIMON-BLECHER: REDISCOVERY OF PYRGOPSELLA ANNANDALEI
Wall plates and opercular valves were immersed for about an hour in
household bleach, rinsed in tap water followed by distilled water, and then
dried. The specimens were examined under a dissecting microscope, and
adhering chitin was removed using needles and fine forceps. Dried samples
were mounted on brass stubs, coated with gold, and examined with a
QUANTA 200F (FEI, Hillsboro, OR, USA) scanning electron microscope
at 25 kV.
The cirri and mouthparts were mounted in glycerin jelly on microscope
slides and the slides sealed with nail varnish. The slides were examined and
photographed using an Olympus Vanox microscope.
DNA was extracted from the four specimens used for morphological
study using the High Pure PCR Template Kit (Roche, Mannheim, Germany). REDTaq ReadyMix R2523 (Sigma-Aldrich, St. Louis, MO, USA)
was used for amplification by the polymerase chain reaction (PCR) (Saiki
et al., 1988), with 50 ng DNA per reaction.
For amplification and sequencing of the 12S subunit of mitochondrial rDNA and cytochrome oxidase subunit 1 (CO1), we followed the
protocol of Brickner et al. (2010). The new sequences were deposited
in GenBank under accession numbers KJ397966-KJ397974. In addition
to the newly generated sequences two sequences of 12S rRNA of Pyrgopsella sp. were retrieved from GenBank and 10 sequences of 12S
rRNA and CO1 Galkinia Ross and Newman, 1995 (see Table S1 in the
Supplementary Material in the online edition of this journal, which can
be accessed via http://booksandjournals.brillonline.com/content/journals/
1937240x). Sequences were then aligned using MUSCLE (Edgar, 2004a,
b) embedded in MEGA6. Distances were calculated using the Kimura
2-parameter model (Kimura, 1980) embeded in MEGA6 (Tamura et al.,
2013).
S YSTEMATICS
Superorder Thoracica Darwin, 1854
Order Sessilia Lamarck, 1818
Suborder Balanomorpha Pilsbry, 1916
Superfamily Balanoidea Leach, 1817
Pyrgomatidae Gray, 1825
Pyrgomatinae Gray, 1825
Pyrgopsellini Ross and Newman, 1995
Pyrgopsella Zullo, 1967
Pyrgopsella annandalei Zullo, 1967
Pyrgopsis annandalei Gruvel, 1906
Pyrgopsella annandalei Zullo, 1967
Pyrgopsella youngi Achituv and Simon-Blecher, 2006
Specimen.— 1907.7.23.16 NHM London.
Diagnosis.— Balanoid with elliptical, concrescent shell
carino-rostral elongated. separated opercular valves, scutum
transversely elongated, basis membranous.
Description.— Shell (Fig. 2A) white, concrescent, thin,
oval, low-conical, carino-rostral diameter 8 mm, lateral
diameter 4 mm. Outer surface with concentric growth lines
consisting of series of small projections (Fig. 2C). Shell
tubiferous with lateral septa terminating as small spines
at sheath perimeter; inner surface of shell white; sheath
white, with concentric growth lines, reaching to margins
of shell (Fig. 2A). Orifice oval, located at carinal end of
shell; aperture carino-rostral diameter 1/3 of shell carinorostral diameter. Basis membranous, basal part elongated
and peduncle-like.
Scutum and tergum white, separate. Scutum (Fig. 2B, D,
G, H) transversally elongated, thin, total length (including
tergal tooth) about five times maximal width; basal margins
slightly sinusoidal; adductor muscle pit shallow, distinct;
adductor ridge small, low; lateral depressor muscle pit
indistinct; width of tergal tooth about 1/2 width of tergal
margin, located closer to occludent margin than to basal
665
margin; with growth lines on outer surface; narrow, oblique
furrow beginning at tergal margin halfway between tergal
tooth and occludent margin, ending near occludent basal
angle. Tergum (Fig. 2E, F) triangular, with growth lines on
outer surface; spur short, barely distinct; external groove
running from middle of scutal margin to basi-carinal apex;
basi-scutal angle pointed; inner surface of scutal margin with
recess accommodating tergal tooth, growth lines present
within this recess, pointed inward-projecting tooth present
on spur.
Trophi: Labrum bilobed (Fig. 3B, outline indicated by
arrow), lobes oval, notch between lobes narrow and deep.
Palpus rounded, club-like, with long setae on upper and
distal margins.
Mandible (Fig. 3C) with four teeth along cutting edge,
distances between teeth unequal. Upper three teeth occupying about 3/4 of cutting edge, gap between first and second
teeth smaller than half length of cutting edge. Inferior angle
with short spines. Surface of mandible bearing rows of short,
sharp setae.
Maxillae II (Fig. 3A) with proximal lobe in too poor
condition to allow detailed description. Distal lobe round;
setae on both lobes, those on distal lobe simple.
Maxillae I (Fig. 3D) with 13 strong setae of equal size on
cutting edge, one short seta at inferior angle, other setae on
upper and lower margins. Surface of maxillule with rows of
short, sharp setae.
Cirrus I (Fig. 4A) with unequal rami, anterior ramus
antenniform, long, slender, with 12 articles, posterior ramus
2/3 length of anterior ramus with 6 articles. Pinnate setae
on apical article of both rami, simple and pinate setae on
proximal articles.
Cirrus II (Fig. 4F) with anterior and posterior rami
subequal, with 10 and 8 articles respectively, articles of
both rami with protuberant anterior margins. Pinate setae
on apical segment of anterior ramus; pinnate and bipectinate
setae on apical segment of posterior ramus.
Cirrus III (Fig. 4C) with anterior and posterior rami
subequal, with 12 and 10 articles respectively. Apical article
of anterior ramus with simple, pinnate, and pectinate setae,
posterior ramus with simple and pectinate setae on apical
article.
Cirri IV-VI (Fig. 4D) with number of articles as follows:
cirrus IV, 20 and 24; cirrus V, 18+ and 22; cirrus VI, 23 and
25. Three to 4 pairs of setae on anterior side of each article,
of which distal pair longest. Apical and other long setae of
each article pinnate.
Penis (Fig. 4E) long, pointed, annulated, with short,
simple setae scattered along it.
Molecular Results
Four sequences of CO1 and of 12S rRNA of Pyrgopsella
were generated; in addition, we used two sequences of 12S
rRNA retrieved from GenBank. Six sequences of 12S rRNA
and six of CO1 of the pyrgomatid Galkinia Ross and Newman, 1995 were added to the analyses. Accession numbers
of sequences used in the analyses are given in Table S1
in the supplementary material in the online edition of this
journal, which can be accessed via http://booksandjournals.
brillonline.com/content/journals/1937240x.
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JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 34, NO. 5, 2014
Fig. 2. Pyrgopsella annandalei sample 1907.7.23.16, NHM, London. A, shell, outside and inside view; B, inside view of shell with opercular valves in situ;
C, enlargement of shell surface showing growth lines consisting of series of small projections; D, inside and outside view of scutum; E, tergum, outer view;
F, tergum, inner view showing internal tooth; G, side view of scutum and tergum attached showing (arrow) tergal tooth; H, attached tergum and scutum,
inner view.
ACHITUV AND SIMON-BLECHER: REDISCOVERY OF PYRGOPSELLA ANNANDALEI
667
Fig. 3. Trophi of Pyrgopsella annandalei sample 1907.7.23.16, NHM, London. A, maxillae II (P : proximal lobes); B, mandibular palpus and labrum
(indicated by L and an arrow); C, mandible; D, maxillae I. Scale bar in C applies to all figures.
Table 1 presents the differences in evolutionary distance
beween specimens of Pyrgopsella, three species of Galkinia
and four specimens of G. altipiculus Chan et al. (2013)
based on sequences of 12S rRNA and CO1. Specimen 5
exhibited a truncated tergal spine, while the others exhibited
a pointed spine. Analyses were conducted using the Kimura
2-parameter model.
D ISCUSSION
There are slight differences in the number of articles of
the cirri, those of the syntype being a bit higher than
those of Gruvel’s described specimen. However, in barnacles
there is variability in size and the number of articles
among specimens of the same species (Marchinko, 2003;
Chan and Hung, 2005; López et al., 2007), and even
differences between the two sides of the same specimen have
been reported. The NHM specimen differs from Gruvel’s
described specimen in the presence of a pointed internal
tergal spine (Fig. 2F) on the spur instead of a truncate spine
(Fig. 1D). A second difference concerns the absence of the
pear-shaped distal part of the penis ornamented with short
spines, described by Gruvel (1907). In the syntype the end
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JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 34, NO. 5, 2014
Fig. 4. Cirri of Pyrgopsella annandalei sample 1907.7.23.16, NHM, London. A, cirrus I; B, cirrus II; C, cirrus II; D, cirrus IV; E, tip of penis. Scale bar in
C applies also to A and B.
of the penis is elongated, bearing a few setae. These features
agree with the description of P. youngi, and are the very
features upon which the latter species was established. Their
presence in the rediscovered syntype casts doubt on the
validity of P. youngi.
Examination of the terga of six specimens of Pyrgopsella
retrieved from of Symphyllia radians (NHMUK 2014.396),
four of them used for the molecular analyses, showed that
the internal tergal spine is pointed in four of them, as in the
syntype, while in the other two, the tergal spine is truncate,
as in the description by Gruvel (1907; cf., Fig. 5). We
conclude, therefore, that the shape of the internal spine is
variable within the species and is not diagnostic.
The description of the distal end of the penis by Gruvel
(1907) is enigmatic; this part of the penis of the five
specimens examined by us, including the syntype, does not
exhibit the pear-shaped structure described by Gruvel. There
are now over 40 species of pyrgomatid coral barnacles for
which the soft parts have been described and the penis
illustrated (Brickner et al., 2010; Chan et al., 2013a, b), but
not one was found to resemble Gruvel’s (1907) description
and drawing. We suspect that what Gruvel described as a
Table 1. Pairwise distance between specimens of Pyrgopsella and based on mitochondrial sequences of CO1 (below the diagnonal line) and 12S rDNA
(above the diagonal line) genes. Specimen 5 exhibits a truncated inward-projecting internal tooth on the spur of the tergum. The sequences of Pyrgopsella
are compared to sequences of three species of the Galkinia. Analyses were conducted using the Kimura 2-parameter model (Kimura, 1980). Analyses were
conducted in MEGA6 (Tamura et al., 2013). n.a. = not available. GenBank accession numbers are given in the Supplementary material in the online edition
of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1937240x.
Sample
1
2
3
4
5
6
7
8
9
10
11
12
Pyrgopsella specimen 5
Pyrgopsella specimen Y
Pyrgopsella specimen 4
Pyrgopsella specimen 1
Pyrgopsella specimen 3
Pyrgopsella sp.
Galkinia tabulaus
Galkinia depressa
Galkinia altipiculus 1
Galkinia altipiculus 2
Galkinia altipiculus 3
Galkinia altipiculus 4
1
2
3
4
5
6
7
8
9
10
11
12
–
0.000
0.004
0.011
n.a.
n.a.
0.163
0.145
0.164
0.162
0.167
0.159
0.003
–
0.004
0.011
n.a.
n.a.
0.163
0.148
0.163
0.162
0.167
0.159
0.006
0.003
–
0.011
n.a.
n.a.
0.160
0.148
0.162
0.159
0.165
0.148
0.003
0.006
0.003
–
n.a.
n.a.
0.163
0.164
0.162
0.167
0.159
0.145
0.003
0.006
0.003
0.006
–
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
0.003
0.000
0.003
0.000
0.003
–
n.a.
n.a.
n.a.
n.a.
n.a.
n.a.
0.114
0.111
0.111
0.111
0.118
0.118
–
0.073
0.071
0.069
0.071
0.073
0.128
0.124
0.125
0.124
0.125
0.128
0.046
–
0.065
0.063
0.067
0.069
0.135
0.131
0.132
0.131
0.139
0.131
0.043
0.027
–
0.002
0.002
0.014
0.135
0.131
0.132
0.131
0.139
0.131
0.043
0.027
0.006
–
0.004
0.002
0.131
0.132
0.135
0.135
0.142
0.138
0.046
0.046
0.003
0.003
–
0.012
0.135
0.135
0.142
0.138
0.138
0.135
0.030
0.046
0.003
0.003
0.000
–
ACHITUV AND SIMON-BLECHER: REDISCOVERY OF PYRGOPSELLA ANNANDALEI
669
Fig. 5. Terga of Pyrgopsella extracted from Symphyllia radians (sample NHMUK 2014.396, NHM, London). A, outer view; B, inner view with pointed
internal tooth; C, inner view with truncated internal tooth.
penis was really a misinterpretation of an organism that
was dredged along with the barnacles, with a pear-shaped
structure resembling the proboscis (introvert) of a priapulid.
The short spines might then be scalids.
Table 1 presents the pairwise distances beween specimens
of Pyrgopsella based on nucleotide sequences of the 12S
rRNA and CO1 genes. The table include sequences of three
species of the pyrgomatid Galkinia, randomly selected four
sequences from one of this species, G. altipiculus (Chan et
al., 2013a, b), are also included in the analyses. The distence
between specimens of Pyrgopsella range between 0.000 and
0.006 for 12S rRNA and between 0.000 and 0.011 for CO1.
This values represented no more than the level of variation
found within the population of G. altipiculus. Specimen 5
of Pyrgopsella had a truncate tergal spine, while the others
exhibited pointed spines, the distances between specimen 5
and the other specimens range between 0.003 and 0.006 for
12S rRNA. Moreover, there was no difference in the CO1
sequence between the specimen with a truncate spine on
the tergal spur and specimen Y, which originated from the
original sample described by us as P. youngi. The differences
between the three species of Galkinia that cluster in the
maximum-likelihood tree (Chan et al., 2013) range berween
0.063 to 0.073 for CO1 and between 0.027 to 0.046 for 12S
rRNA. The molecular data thus do not support the separation
of P. youngi from P. annandalei on the basis of internal
tergal spine shape alone. Regrettably, it proved impossible
to amplify genes from the 125-year-old specimen.
Based on the morphology of the “co-type” of Pyrgopsella
annandalei and the molecular analyses of specimens hitherto assigned to P. youngi, we conclude that the name P.
youngi is a junior subjective synonym of P. annandalei and
should be abandoned.
ACKNOWLEDGEMENTS
The visit of Y.A. to the Natural History Museum (London, UK) was
supported by the SYNTHESYS Project (http://www.synthesys.info) (Grant
GB-TAF-2892), which is financed by the European Community Research
Infrastructure Action under the FP7 Integrating Activities Programme. The
study was suported by ISF grant 574/10. The help of Ms. Miranda Lowe
and Mr. Andrew Cabrinovic of the Natural History Museum is greatly
appreciated. We thank Mr. Meir Moial for donation of the colony of
Symphyllia radians carrying Pyrgopsella. The authors thank Ms. T. GuyHaim for comments on the manuscript. Dr. Yaakov Langzam of the Mina
and Everard Goodman Faculty of Life Sciences, Bar Ilan University, helped
with the SEM work.
R EFERENCES
Achituv, Y., and N. Simon-Blecher. 2006. Pyrgopsella (Cirripedia: Balanomorpha: Pyrgomatidae) is not a sponge-inhabiting barnacle. Zootaxa
1319: 29-42.
Alcock, A. 1902. A Naturalist in Indian Seas, or, Four Years with the Royal
Indian Marine Survey Ship ‘Investigator.’ John Murray, London.
Brickner, I., N. Simon-Blecher, and Y. Achituv. 2010. Darwin’s Pyrgoma
(Cirripedia) revisited: revision of the Savignium group, molecular analysis and description of new species. Journal of Crustacean Biology 30:
266-291.
Chan, B. K. K., Y.-Y. Chen, and Y. Achituv. 2013b. Crustacea Fauna
of Taiwan, Vol. II, Cirripedia: Thoracica: Pyrgomatidae. Biodiversity
Center Academia Sinica, Taipei, 364 pp.
,
, and H.-C. Lin. 2013a. Biodiversity and host specificity of
coral barnacles of Galkinia (Cirripedia: Pyrgomatidae) in Taiwan, with
descriptions of six new species. Journal of Crustacean Biology 33: 329331.
, and O. S. Hung. 2005. Cirral length of the acorn barnacle Tetraclita
japonica: effect of wave exposure and tidal height. Journal of Crustacean
Biology 25: 329-332.
Darwin, C. 1854. A Monograph on the Sub-class Cirripedia, with Figures
of All the Species. The Balanidae, (or Sessile Cirripedes); the Verrucidae,
etc., ect., etc. The Ray Society, London, viii + 684 pp.
Edgar, R. C. 2004a. MUSCLE: multiple sequence alignment with high
accuracy and high throughput. Nucleic Acids Research 32: 1792-1797.
. 2004b. MUSCLE: a multiple sequence alignment method with
reduced time and space complexity. BMC Bioinformatics 5: 113.
Gray, J. E. 1825. A synopsis of the genera of Cirripedes arrangedin natural
families, with a description of some new species. Annals of Philosophy
n.s. 10: 97-107.
Gruvel, J. A. 1906. Sur une forme nouvelle de Cirrhipède operculé
(Pyrgopsis annandalei n. g., n. sp.). Comptes Rendus Hebdomadaires
des Séances de l’Académie des Sciences Paris 142: 1558-1559.
. 1907. Cirrhipèdes operculés de l’Indian Museum de Calcutta.
Memoirs of the Asiatic Society of Bengal 2: 1-10.
Kimura, M. 1980. A simple method for estimating evolutionary rate of
base substitutions through comparative studies of nucleotide sequences.
Journal of Molecular Evolution 16: 111-120.
Lamarck, J. B. P. 1818. Histoire naturelle des animaux sans vertèbres,
présentant les caractères généraux et particuliers de ces animaux, leur
distribution, leurs classes, leurs familles, leurs genres. Verdière, Paris 5:
1-612.
Leach, W. E. 1817. Distribution systématique de la classe des Cirripèdes.
Journal de Physique, de Chimie et d’Histoire Naturelle Élémentaire 85:
67-69.
López, B. A., S. Y. Guaitro, and D. A. López. 2007. Density-dependent
effects on the cirral structure of the barnacle, Jehlius cirratus (Darwin,
1854) (Cirripedia). Crustaceana 80: 793-801.
Marchinko, K. B. 2003. Dramatic phenotypic plasticity in barnacle legs
(Balanus glandula Darwin): magnitude, age dependence and speed of
response. Evolution 57: 1281-1290.
Milne Edwards, H., and J. Haime. 1849. Mémoire sur les polypiers appartenant à la famille des Oculinides, au groupe intermediate des Pseudoastréides et à la famille des Fongides. Comptes Rendus Hebdomadaires des
Séances de l’Académie des Sciences 29: 67-73.
670
JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 34, NO. 5, 2014
Nilsson-Cantell, C. A. 1938. Cirripedes from the Indian Ocean in the
collection of the Indian Museum, Calcutta. Memoirs of the Indian
Museum 13: 1-81.
Pilsbry, H. A. 1916. The sessile barnacles (Cirripedia) contained in the
collections of the U.S. National Museum; including a monograph of the
American species. Bulletin of the United States National Museum 93:
xi + 366 pp.
Rosell, N. C. 1973. On two less well-known balanids (Cirripedia Thoracica)
from the Sulu Archipelago, Philippines. U.P. Natural Science Research
Center Technical Report, 4: 1-12.
. 1975. On two noteworthy balanids (Cirripedia Thoracica) from the
Sulu Archipelago, Philippines. Crustaceana 29: 206-214.
Ross, A., and W. A. Newman. 1995. A coral-eating barnacle, revisited
(Cirripedia; Pyrgomatidae). Contributions to Zoology (Amsterdam) 653:
129-175.
Saiki, R. K., D. H. Gelfand, S. Stoffel, S. J. Scharf, R. Higuchi, G. T.
Horn, K. B. Mullis, and H. A. Erlich. 1988. Primer-directed enzymatic
amplification of DNA with a thermostable DNA polymerase. Science
239: 487-491.
Simon-Blecher, N., D. Huchon, and Y. Achituv. 2007. Phylogeny of coralinhabiting barnacles (Cirripedia; Thoracica; Pyrgomatidae) based on
12S, 16S and 18S rDNA analysis. Molecular Phylogeny and Evolution
44: 1333-1341.
Tamura, K., G. Stecher, D. Peterson, A. Filipski, and S. Kumar. 2013.
MEGA6: molecular evolutionary genetics analysis, version 6.0. Molecular Biology and Evolution 30: 2725-2729.
Zullo, V. A. 1967. Pyrgopsella, new name for Pyrgopsis Gruvel, 1907
(Cirripedia, Thoracica) non Pyrgopsis de Rochebrune, 1884. Crustaceana
13: 123.
R ECEIVED: 12 March 2014.
ACCEPTED: 2 June 2014.
AVAILABLE ONLINE: 15 July 2014.