---------------- - - - - -
Spirorbinae (Polychaeta, Serpulidae) of the
Hawaiian Chain
Part 2, Hawaiian Spirorbinae 1
PETER
J. VINE,2 JULIE H.
BAILEY-BROCK,3 and DALE STRAUGHAN4
No TAXONOMIC STUDY of tubeworms in the
water formalin or 70 percent ethyl alcohol. It
was found that those preserved in formalin were
easier to study in detail than the others, which
soon became hardened and colorless. Details of
setation were studied in polyvinyl-Iactophenol
mounts, with the phase microscope, using quartziodide illumination. The presence of dorsally
fused collar folds in some species was confirmed
by probing with an eyelash (Knight-Jones,
1972) .
subfamily Spirorbinae has been made previously
in Hawaii except for a recent description of
three new species (Vine, 1972). This second
paper deals with 10 other species, previously
described from elsewhere, which were collected
around Oahu and Hawaii.
Collections have been made by each of the
authors and preliminary studies were made independently. Settlement plates have been used by
two of us (Bailey-Brock and Straughan) and it
was found that several species settled very
readily on these. Most of the collections were
littoral or from shallow water and it is likely
that other species will be found at greater
depths.
The descriptions and drawings presented here
are the work of one of us (Vine) although each
author has contributed to the ecological data.
Spirorbinae were usually removed from their
substrates by slicing them off with a sharp scalpel and were then preserved in 5 percent sea-
SPECIES ENCOUNTERED
Taxonomy of Spirorbinae has been recently
reviewed by Gee (1964), Bailey (1969b), and
Pillai (1970). Important advances are now being made, however (Knight-Jones, 1972, in
press; Vine, in press), and the arrangement
used here follows that adopted by Phyllis
Knight-Jones (1972), which represents a slight
modification of that proposed by Bailey
(1969b).
KEY TO SPECIES OBSERVED
1. Eggs incubated in tube
2
3
Eggs incubated in operculum
2. Tube dextral (mouth faces anticlockwise) .. . Spirorbis (Spirorbella) marioni (p. 151)
Tube sinistral (mouth faces clockwise)
Protolaeospira species A (p. 155)
3. Collar setae have a distinct fin and blade (tubes all sinistral)
Collar setae lack a distinct proximal fin (all except one are dextral)
4
7
4. Operculum has spines
Operculum without spines
5
6
5. Sickle setae present in third thoracic fascicle
Tube lacks longitudinal ridges
Tube has three longitudinal ridges
----Manuscript received 25 June 1971.
University College of Swansea, Department of
Zoology, Swansea, Wales, United Kingdom. Present
address: University College, Department of Zoology,
Galway, Republic of Ireland.
1
2
Pileolaria (Pileolaria) militaris (p. 157)
Pileolaria (Pileolaria) semimilitaris (p. 140)
3 University of Hawaii, Department of Zoology,
Honolulu, Hawaii 96822.
4 University of Southern California, Allan Hancock
Foundation, Los Angeles, California 90007.
150
Hawaiian Spirorbinae-VINE, BAILEy-BROCK, AND STRAUGHAN
151
Sickle setae absent from third thoracic fascicle
.
·
Pileolaria (Simplicaria) pseudomilitaris (p. 158)
6. Tube has two or three irregular longitudinal ridges
.
·
Pileolaria (Duplicaria) koehleri (p. 161)
Tube has four narrow longitudinal ridges
.
·
Pileolaria (Duplicaria) dalestrattghani (p. 143)
7. Tube dextral (mouth faces anticlockwise)
8
11
Tube sinistral (mouth faces clockwise)
8. Tube has three longitudinal ridges and lacks any other distinctive pattern
.
Janua (Janua) pagenstecheri (p. 163)
9
·
Tube has transverse as well as longitudinal ridges
9. Collar setae are not cross-striated
Collar setae are
Operculum
·
Operculum
·
Operculum
·
10
cross-striated
has transparent walls and short spade-shaped talon
.
Janua (Dexiospira) pseudocorrugata (p. 166)
has calcified walls and bifid talon with lateral wings
.
Janua (Dexiospira) steueri (p. 168)
has transparent walls and long, asymmetrical, pointed talon
.
Janua (Dexiospira) turrita (p. 145)
10. Operculum has transparent walls and a short spade-shaped talon; thoracic uncini have
Janua (Dexiospira) nipponica (p. 170)
trifurcate anterior pegs
11. Opercular chambers may be stacked and each one has a short talon
.
" Janua (Leodora) knightjonesi (p. 172)
·
Genus Spirorbis Daudin, 1800 (amended)
The genus is described as follows: coiling
usually sinistral; thorax with two pairs of tori;
collar folds normally unfused; collar setae with
basal fin and distal blade which may be crossstriated; sickle setae present in third fascicles;
thoracic uncini with several longitudinal rows
of teeth and broad anterior pegs; abdominal
tori without any marked bilateral asymmetry;
abdominal setae smaller than collar setae; embryos incubated in an egg string attached posteriorly to the tube; larvae usually have a single
attachment gland.
Type
Spirorbis spirorbis (Linnaeus)
Daudin.
= S.
borealis
Subgenus Spirorbella (Chamberlin, 1919)
This subgenus may be described as above, except that it is coiled dextrally.
Type
Spirorbis (Spirorbella) marioni (Caullery
Mesnil, 1897).
&
The genus Spirorbella was originally separated by Chamberlin and has recently been recognized by Pillai (1970). Here it is given
subgeneric status as the only feature dividing
it from the subgenus Spirorbis is its direction
of coiling (and, in certain Spirorbinae, this may
vary, even within the same species; see Bailey,
1969b, and Vine, in press).
Spirorbis (Spirorbella) marioni
Caullery & Mesnil
Fig. 1, 2a, 3a
Material
Twenty-four specimens.
Location
Nanakuli, Oahu, Hawaii.
-- ------------------------ --- --------- -------- ------ ------------------------------------------------------1
PACIFIC SCIENCE, Volume 26, April 1972
152
~
b
Q'Smm
~1
o.t)
d
C'l
6
E
E
k
m
n
C'l
o
6
FIG. 1. Spirorbis (Spirorbe/la) marioni. a, Tube viewed from above; b, side view of tube; c, dorsal view
of whole animal showing eggstring attached posteriorly to a fragment of the tube (the specimen was hardened
and collar folds could not be distinguished); d, dorsal view of juvenile operculum; e, side view of juvenile
operculum; f, later stage of operculum; g, dorsal view of fully developed operculum; h, side view of fully
developed operculum; ;, collar seta; k, thoracic uncinus; m, simple seta from second thoracic fascicle; n, sickle
seta from third thoracic fascicle; 0, abdominal seta; p, abdominal uncinus.
SCALE:
band c as
(I;
e,
f,
g, and h as d; k, m, n,
0,
and p as ;.
Hawaiian Spirorbinae-VINE, BAILEy-BROCK, AND STRAUGHAN
Habitat
The species is found in the intertidal zone on
igneous rocks.
Description of Species
TUBE: Tube is dextral and characteristically
flattened against the substrate. An outer longitudinal ridge demarcates the indented flat top
from the oblique sides. It is tightly coiled with
the terminal portion and mouth overlapping the
previous whorl, making the tube almost round
when viewed from above. In adult specimens
two whorls can be seen on the underside of the
tube (Fig. la). Coil diameter is 1.5 to 2.0 mm.
OPERCULUM: As Berkeley and Berkeley
(1941) have commented, the operculum has
a variable structure. In the past this has given
rise to descriptions of three separate species:
S. marioni Caullery & Mesnil, 1897; S. bttshi
Rioja, 1942; and S. tricornigert/s Rioja, 1942.
Study of opercula of S. marioni collected in Hawaii shows that, as they grow, they develop an
increasingly complex structure. The plate and
talon are always lightly calcified and more or
less transparent. The earliest operculum (Figs.
Id, e) has a slightly concave plate and a massive
cone-shaped talon. At this stage the operculum
is reminiscent of an ice-cream cone with the
dome licked off. The next stage is for the concavity to deepen, whilst a central peg develops
from the plate and a slight depression appears
on the dorsal edge of the plate (Fig. If). Finally, the edge of the plate thickens on each side
of this depression (Fig. Ig).
THORAX: The thorax has two pairs of tori.
The dorsal folds of the collar do not appear to
be fused, but the specimens are hardened and
the collar is difficult to distinguish. Collar setae
have rather broad blades with coarse, pointed
teeth and fine cross-striations. A proximal fin
has several large teeth centrally and smaller
teeth on each side of these. Capillary setae are
associated with the collar setae on the convex
side. The second and third fascicles have simple, finely serrated setae and the third fascicles
also have sickle setae. The blades of the latter
have their proximal and distal portions about
equal in length. Uncini have fine teeth arranged
153
in approximately four longitudinal rows and
many transverse rows, which are unusual in being mostly perpendicular to the long axis. Each
uncinus has a broad peg at the anterior end,
which appears bifid.
ASETIGEROUS REGION: This is long, being
approximately equivalent to eight times the distance between the first two abdominal tori.
ABDOMEN: This structure has approximately
nine segments and the largest tori are toward
the middle segments of the abdomen. Setae are
geniculate with blunt-ended teeth; they appear
to have a fan of more pointed teeth, but stereomicroscope study shows that this effect is due
to sculpturing of the gaps between the larger
teeth (Phyllis Knight-Jones, personal communication). The first tooth of the blade is recurved and prominent. Some fascicles contain
paired setae. Uncini have approximately nine
longitudinal rows of fine teeth (illustrated by
cross-hatching in Fig. IP) and an apparently
bifid anterior end similar to that of the thoracic
uncini.
INCUBATION: Incubation occurs in an egg
string attached posteriorly to the tube (Fig. lc).
Distribution Elsewhere
Species has been found in Panama (Caullery
and Mesnil, 1897); Mexico (Rioja, 1942);
Galapagos (Bailey and Harris, 1968); southern
California (Berkeley and Berkeley, 1941).
Remarks
Specimens intermixed on the same substrate
and having similar tubes and setation showed
enough opercular variation to indicate clearly
that the two species described by Rioja (1942)
as S. bushi and S. tricornigerus are the same as
S. marioni. In having cross-striated collar setae
and thoracic uncini with straight transverse
rows of teeth, this species resembles Spirorbis
(Spirorbis) cuneattlS Gee, but differs from most
species of Spirorbis sensu stricto. This arrangement of teeth is indeed very unusual amongst
Spirorbinae in general (Phyllis Knight-Jones,
personal communication).
_._-_._------------- - - - -
a
10
+~
b
+- +++
c
~+
H-++
+
d
+
~
H+
e
FIG. 2. Chart showing distribution of setae and uncini in: a, Spil'ol'bis (Spil'ol'bella) marioni; b, Pl'otolaeospil'a sp. A; c, Pileolaria (Pileolaria) militaris (abdominal setae could not be counted on the single slide of
this species); d, Pileolaria (Simplicaria) p.reudomilitaris; e, Pileolaria (Duplical'ia) koehleri.
The histograms indicate the number of uncini per torus on the concave and convex sides of the animal.
Continuous lines within these indicate the number of setae in each fascicle and symbols represent various kinds
of ancillary setae, viz.:
+ = long thoracic capillaries and shorter abdominal capillaries or hooked setae
= sickle setae
. = secondary setae with rudimentary shafts
X
Species are shown as they appear in situ. Thus, in sinistral species the concave side is on the reader's right,
whereas in dextral ones the concave side is on the reader's left.
The length of the asetigerous region is represented as a multiple of the distance between the first and
second abdominal tori on the concave side. The distance between tori on the histograms is standardized and,
thus, the histograms do not necessarily give a true indication of the length of the asetigerous region in relation to that of the whole animal. In some species the anterior abdominal tori are themselves unusually far
apart and this will result in the asetigerous region appearing relatively shorter in the histogram,
The figures represent single typical specimens but, where sufficient material was available, three counts were
made of each species and the variations were small. The exact distribution of paired setae and capillary setae
in the abdomen was not constant. Generally speaking, it is the overall pattern of the histogram which characterizes a particular species rather than separate details which vary with age and show slight variation within
a species, even between individuals of the same size,
Hawaiian Spirorbinae-VINE, BAILEy-BROCK, AND STRAUGHAN
I
II
/
a
/
I
L((
II ((('
/
/
155
on the concave side; thoracic uncini are usually
very long and slender, with a blunt anterior peg.
Abdominal setae are less than a quarter the
length of the collar setae and have vestigial
brushlike blades. The operrular talon is massive
and usually bears lateral projections. A single
white larval attachment gland may be present.
Type
Protolaeospira ambilateralis (Pixell, 1912).
b
Protolaeospira sp. A
Fig. 2b, 3b, 4
Material
A single specimen has been preserved as a
polyvinyl-Iactophenol mount. Fragments of the
tube are preserved in alcohol and the operrulum
is in clove oil.
d
e
FIG. 3. Relative sizes of setal shafts and blades
in: a, Spirorbis (Spirorbella) marioni; b, Protolaeospira sp. A; c, Pileolaria (Pileolaria) militaris;
d, Pileolaria (Duplicaria) koehleri; e, Pileolaria (Simplica1'ia) pseudomilitaris.
The stick diagrams represent relative lengths of setal
shafts and blades. The numbers in Fig. 3a represent
the arrangement of setae in all the diagrams. The
central seta is abdominal. Numbers 1, 2, and 3 represent setae from the collar, second and third fascicles
on each side. Spirorbis (Spit'01'bella) marioni (Fig. 3a)
is dextral and in this case the concave side is on the
observer's left, whereas all the other species in Fig. 3
are sinistral and the concave side is represented by
the stick setae on the observer's right.
Genus Protolaeospira Pixell (1912) redefined
(= Marsipospira Bailey, 1969b; includes
Pixellia Pill ai, 1970)
Coiling is usually sinistral. Three rows of tori
ocrur on the concave side of the thorax; other
traces of a fourth thoracic segment mayor may
not be present. Embryos are attached to a stalk
which arises dorsally from the floor of the fecal
groove, approximately level with the first thoracic torus. Collar setae have separate fins and
broad cross-striated blades; sickle setae are present in the third and fourth fascicles. Thoracic
and abdominal uncini are much more numerous
Location
Coconut Island, Oahu, Hawaii.
Habitat
The animal was found on a settlement plate
in shallow water.
Description of Species
TUBE: Tube is sinistral with thick bulging
transverse ridges, translucent, and vitreous. In
some gaps between the ridges the tube is transparent. Terminal section of tube widens out to
form a broad mouth. Coil diameter is approximately 3.5 mm.
OPERCULUM: The operculum is translucent
except for a median densely calcified rod on the
dorsal side of the asymmetrically lobed talon,
which lacks a central spur. The plate is slightly
concave. The whole operrulum is notable for its
extremely light calcification.
THORAX: The thorax has three tori on each
side, but the third torus on the convex side has
only two uncini. There are no setae associated
with the third tori, Collar setae are large, partirularly those on the convex side. The longer
shafts of the latter originate behind the first
torus on the convex side and behind the origin
of the second thoracic fascicle on that side. They
-.------.-------.------------------.-.----------..---------------1
PACIFIC SCIENCE, Volume 26, April 1972
156
,O·5m.m
a
c
f
E
E
9
h
N
o
o
FIG. 4, Protolaeospira sp. A. a, Tube; b, dorsal view of operculum; c, side view of operculum; d, collar
seta; e, seta from second thoracic fascicle; j, seta associated with collar setae; g, abdominal uncinus; h, sickle
seta from third thoracic fascicle; j, abdominal seta; k, thoracic uncinus.
SCALE:
c as b; e, j, g, h, j, and k as d.
-----------------------------
Hawaiian Spirorbinae-VINE, BAlLEy-BROCK, AND STRAUGHAN
have a proximal fin with three or four large
teeth and a broad distal blade which has coarse
teeth and cross-striations across the narrow width
of the blade. The section between the fin and
blade is unusually broad. Capillary setae associated with the collar setae bear minute teeth.
Simple setae in the second and third fascicles are
numerous and have smooth blades lacking any
distinct striations. Capillary setae are also present
in these fascicles except for the third fascicle on
the convex side. The third fascicles also have
sickle setae with a proximal smooth blade and
a distal portion bearing relatively large recurved
teeth. Uncini have three or four longitudinal
rows of teeth and a blunt gouged anterior end
lacking teeth. In some cases the anterior end
appears to have an irregularly indented edge,
possibly a result of convolutions.
ASETIGEROUS REGION: This region is short,
approximately 1.5 times the length between the
first and second abdominal tori.
ABDOMEN: The abdomen has approximately
14 segments. Setae have short, brushlike blades
with pointed teeth. Anteriorly they may be in
groups of three and, in one fascicle on the
convex side, five were counted. More posteriorly
they are paired or single. The most posterior
segments also have simple finely serrated nongeniculate setae. Uncini are unusually widely
spaced within tori on the concave side. They have
approximately seven longitudinal rows of small
teeth and are short and broad with blunt anterior
ends. Many segments on the convex side lack
uncini but have setae. Thus the distribution of
uncini is very asymmetrical.
INCUBATION: Embryo mass is born on a stalk
originating from the dorsal side of the thorax
close to the first thoracic torus.
157
culum and that of S. translttcem. This specimen
resembles S. tramlttcens in having three or four
rows of teeth on each thoracic uncinus; most
Protolaeospira species have single rows of teeth
on their thoracic uncini. It also bears some resemblance to a species from southern Australia
(Knight-Jones, in press), but that species has a
densely calcified opaque tube, a slightly different
operculum, and thoracic uncini with single longitudinal rows of teeth.
Genus Pileolaria Claparede, 1868
(amended Knight-Jones, 1972)
The genus is described as follows: sinistral
coiling; incubation in the operculum; only two
pairs of thoracic tori; fin and blade collar setae,
with the blades usually bearing large teeth and
distinct cross-striations and greatly exceeding in
length the blades of abdominal setae; sickle setae
mayor may not be present; thoracic uncini very
slender, often with only one longitudinal row of
teeth; pegs of thoracic uncini about as broad as
rest of uncinus and not pointed in surface view;
arrangement of abdominal tori almost bilaterally
symmetrical and with the largest tori lying
toward the posterior end; larvae with single,
white, middorsal attachment glands.
Subgenus Pileolttria
Opercula of juvenile and adult are morphologically different from one another; the opercular plate is single with embryos brooded under
it. Collar setae are cross-striated. Thoracic uncini are usually found with single rows of teeth.
Type
Pileolaria
1868.
(Pileolaria)
militaris Claparede,
Remttrks
Inasmuch as only a single specimen has been
found, it is not yet possible to state whether this
should be regarded as a new species. It appears
to be closely related to, or perhaps the same as,
S. translttcem Bailey & Harris (1968) from
the Galapagos Islands. The opercular structure
of other Protolaeospira species tends to be somewhat variable, and it is difficult to be sure how
significant are the differences between this oper-
Pileolaria (Pileolaria) militaris
Claparede, 1868
Fig. 2c, 3c, 5
Material
One specimen.
Location
Coconut Island, Oahu, Hawaii.
158
PACIFIC SCIENCE, Volume 26, April 1972
Habitat
Animal was found in the intertidal zone, on
an oyster.
(Vine, in press), and off southern Australia
(Phyllis Knight-Jones, in press). It thus appears
to show a worldwide distribution in tropical,
subtropical, and warm temperate waters.
Description of Species
Remarks
TUBE: Tube is sinistral, with indistinct wavy
transverse growth thickenings, the terminal portion ascending. Coil diameter is 3.0 mm.
The description agrees closely with those of
previous authors. The operculum of the species
described here differs slightly from that of specimens recently described from the Red Sea (Vine,
in press) in that the rim of the plate has a
definite depression on the dorsal side and lacks
spines there. Zibrowius (1967) studied this species in the Mediterranean and showed that its
operculum exhibits quite considerable variation
in structural detail. Recent research tends to suggest that some widely distributed Spirorbinae
such as P. (P.) militaris should be treated as
superspecies, but there is not yet enough information to characterize geographical races.
OPERCULUM: The operculum formed a brood
chamber containing eggs. The plate is slightly
convex and covered with spines. A rim of spines
almost surrounds it, but is discontinuous on the
dorsal side where the rim dips. Walls of the
chamber are densely calcified and opaque. On
the ventral side at the base of the wall are five
short spines; and a number of eggs can be seen
clearly from this side, where the wall does not
extend as far down as it does on the dorsal side.
THORAX: The thorax has two pairs of tori and
the collar folds are not fused. A dark granular
region partially obscures the tori on the concave
side and extends into the asetigerous region.
Collar setae have fins and distinctly cross-striated
blades. Capillary setae are associated with all the
thoracic fascicles. Second and third segments
have simple setae, and sickle setae are found in
the third fascicles. Uncini usually have a single
longitudinal row of teeth and a bifid anterior
end.
This is rather long and
equivalent to approximately four times the distance between the anterior abdominal tori.
ASETIGEROUS REGION:
The abdomen has about 12 segments and the longest tori are toward the posterior end. Setae are geniculate with rounded
teeth. Uncini have about six longitudinal rows
of teeth and a bifid anterior end.
ABDOMEN:
INCUBATION:
Incubation occurs in operculum.
Distribution Elsewhere
The distribution of this species has been reviewed in considerable detail by Zibrowius
(1968). It is a widely distributed species with
many records in the Mediterranean Sea and
Atlantic and Pacific oceans. Recently it was
found near Plymouth, England (Phyllis KnightJones, personal communication), in the Red Sea
Subgenus Simplicaria Knight-Jones, in press
This subgenus resembles Simplicaria, but lacks
sickle setae.
Type
Pileolaria (Simplicaria) pseudomilitaris (Thiriot-Quievreux, 1965).
Pileolaria (Simplicaria) pseudomilitaris
Fig. 2d, 3e, 6
Material
Thirty specimens.
Locations
Oahu, Hawaii: Sand Island in Kaneohe Bay
and Koko Head.
Habitat
Animals were taken in the intertidal zone, to
3 m, on stones.
Description of Species
TUBE: Tube is sinistral, with transverse
growth thickenings; terminal section often ascends and coiling may be helical. Coil diameter
is 2.0 mm.
159
Hawaiian Spirorbinae--VINE, BAILEy-BROCK, AND STRAUGHAN
a
E
E
0·5 mm
l<")
~
~l
1
III
! !I,
Ii,iI~ i
~
, ,I
~
I'
i
9
e
h
~
~
~
o
~G~
k
m
FIG. 5. Pi/eo/aria (Pi/eolaria) militaris. a, Tube; b, dorsal view of operculum; c, side view of operculum;
d, ventral view of operculum, showing eggs; e, collar setae; f, seta associated with collar setae; g, seta from
second thoracic fascicle; h, sickle seta from third thoracic fascicle; j, thoracic uncinus; k, abdominal seta;
m, abdominal uncinus.
SCALE:
c and d as bj
f,
g, h, j, k, and m as e.
._----------------- --- - - - - -------- -----
PACIFIC SCIENCE, Volume 26, April 1972
160
E
E
N
9
0
d
c
e
f
9
h
Iii
FIG. 6. Pileolaria (Simplicaria) pseudomilitaris. a, Tube; b, mature operculum; c, collar seta; d, thoracic
uncinus; e, simple seta from second thoracic fascicle; f, capillary seta associated with collar setae; g, side view
of abdominal uncinus; h, edge-on view of abdominal uncinus; ;, abdominal seta.
SCALE:
d, e,
f,
g, h, and
i as
c.
OPERCULUM: Juveniles had a slightly concave
plate and an eccentric fingerlike talon. Adults
have a helmet-shaped calcified chamber with a
slightly convex plate bearing several spines and
an incomplete rim bearing short spines.
THORAX: The thorax bears two pairs of tori.
Each collar seta has a proximal fin and a distal
toothed blade, with fairly distinct cross-striations. Capillary setae are associated with these
and are present in all the thoracic fascicles. The
second and third fascicles "'ear simple setae and
lack sickle setae. Uncini have a somewhat bifid
gouged anterior end and a single row of teeth.
ASETIGEROUS REGION: This region is fairly
long, equivalent to four times the distance between the first two abdominal tori.
ABDOMEN: The abdomen has approximately
17 segments. Setae are geniculate with blunt
recurved teeth. Uncini have four or five longitudinal rows of distinct teeth and a blunt anterior end lacking teeth.
Hawaiian Spirorbinae-VINE, BAILEy-BROCK, AND STRAUGHAN
INCUBATION: Incubation takes place in opercular brood chamber.
BODY COLORS: Tentacles are pale pink or
orange; gut is nut brown; eggs in ovary are
green-blue.
Distribution Elsewhere
The species has been taken from the Mediterranean Sea (Thiriot-Quievreux, 1965; Zibrowius,
1968; Harris, 1968, as S. berkeleyanus); Aegean
Sea (Bailey, 1969a); Galapagos Islands (Bailey
and Harris, 1968, as S. rega/is); West Indies
(Bailey, 1970); southern Australia (KnightJones, in press); northeastern Australia (personal observations by P.J.V.); New Zealand
(unpublished, P.J.V.).
Remarks
S. regalis from the Galapagos Islands is almost
certainly the same as Pileolaria (Simplicaria)
pseudomilitaris. Bailey and Harris (1968) separated these two species on the basis of the former's having a domed opercular plate with
several spines and a smooth tube lacking longitudinal ridges. More recent studies have shown
that P. (S.) psettdomilitaris often has a relatively smooth tube and the opercular plate may
be flat or slightly domed.
Subgenus Duplicaria Vine, in press
Subgenus resembles Pileolaria, except that
adult operculum in Duplicaria develops from
juvenile form without sharp dimorphism and
has two or more opercular plates stacked one
above the other; embryos are brooded below
them in a chamber with lightly calcified, rather
delicate walls; collar setae not distinctly crossstriated; thoracic uncini have several longitudinal
rows of teeth; anterior abdominal torus on concave side is usually split into two unequal portions.
Type
Pileolaria (Duplicaria) koehleri Caullery &
Mesnil (1897).
Pileolaria (Duplicaria) koehleri
Fig. 2e, 3d, 7
161
Material
One hundred fifty-one specimens were preserved in tubes and five were mounted in polyvinyl-Iactophenol.
Location
Maili Point, Oahu, Hawaii; Hilo, Hawaii.
Habitat
Animals were found off Oahu at 8 m on
stones. They were taken off the island of Hawaii
from the intertidal zone and shallows and were
found to be abundant on lava rocks.
Description of Species
TUBE: Tube is sinistral, somewhat porce1aneous, and tightly coiled. Outer longitudinal ridge
projects almost horizontally from the top outer
edge of tube. Mouth is often wider than rest of
tube. Irregular vertical ridges occur on sides.
It is often colored brown by algal growth. Coil
diameter is 1.5 mm.
OPERCULUM: This structure has two or three
slightly concave plates interlocking by peg and
socket arrangement and is also supported by
lateral wings on talon. Underneath these plates
brood chambers develop with lightly calcified,
rather delicate, walls, through which eggs can be
distinguished.
THORAX: The thorax has two pairs of tori.
Collar folds are not fused. Collar setae have a
broad fin with many narrow teeth at the outer
edge and approximately five larger teeth toward
the opposite side. There is hardly any gap between this and the distal blade, which is fairly
coarsely serrated but lacks cross-striations.
Smooth-edged, flexible, capillary setae are associated with ~hes
and are also present in the
second and third fascicles. Simple setae are the
main setae in the second fascicles and are also
present in the third, although sickle setae tend
to be more numerous there. The proximal portion of each sickle blade is only a quarter of the
length of the whole blade. Uncini have two or
three longitudinal rows of teeth and a broad
anterior end which lacks teeth.
ASETIGEROUS REGION: This is rather short,
approximately 1.5 times the distance between
the first two abdominal tori.
------.------ .-----1
162
PACIFIC SCIENCE, Volume 26, April 1972
a
0·5
mm
d
c
e
FIG. 7. Pileolaria (Duplicaria) koehleri. a, Tube; b, mature opercular brood chamber; c, collar seta;
d, thoracic uncinus; e, capillary seta associated with collar setae; f, sickle seta from third thoracic fascicle;
g, simple seta from second thoracic fascicle; h, abdominal seta; j, abdominal uncinus.
SCALE: d,
e,
f,
g, h, and j as c.
ABDOMEN: The abdomen consists of about 14
segments. Setae are geniculate with rather blunt,
deeply gouged teeth. The first tooth of the blade
is large and slightly recurved, partially overlying the next tooth. Uncini have about nine
longitudinal rows of fine teeth and a broad anterior end lacking teeth. The anterior torus on
the concave side is split into two unequal sections, of which the one nearest the concave edge
bears more uncini than the other and is out of
line with the other tori on this side.
INCUBATION:
chamber.
Incubation occurs in opercular
Distriblltion Elsewhere
This species has been taken from the Mediterranean Sea (Caullery and Mesnil, 1897; Zibrowius, 1968); Aegean Sea (Bailey, 1969a);
West Indies (Bailey, 1970); Red Sea (Vine, in
press); New Zealand (unpublished, P.J.V.);
Australia (personal observations, P.J.V.).
Remarks
It is possible that Spirorbis
opermlata Straughan (1969)
the Marshall Islands is the
koehleri. Straughan separated
(Pileolaria) polyfrom Eniwetok in
same as P. (D. )
the two species on
Hawaiian Spirorbinae-VINE, BAILEy-BROCK, AND STRAUGHAN
the basis of S. (P.) polyoperculata having a
smooth tube and lacking wings to its talon. The
tube of P. (D.) koehleri is variable, however,
and the wings of the talon are often almost
transparent and may be partly obscured by secondary plates. Recent studies which have extended the distribution of P. (D.) koehleri to
the Red Sea, New Zealand, and Australia (as
well as to Hawaii) tend to support this suggestion. Further studies are needed to show if
these two species are synonymous.
Genus Janua de Saint-Joseph, 1894
(redefined Knight-Jones, 1972)
Most species of this genus have dextral coiling. Incubation is in an opercular brood chamber, below which a secondary plate (rudiment
of next opercular plate) is formed soon after
spawning. Only two pairs of thoracic tori are
present. Collar setae are without a toothed fin;
abdominal setae have blades as big as, or bigger
than, those of the collar setae, and often are
accompanied by secondary setae with rudimentary shafts. Thoracic uncini have anterior pegs
narrow and more or less pointed in surface view.
The largest abdominal tori lie in the anterior
half of the setigerous region. Larvae have paired
white attachment glands in thoracic region.
Subgenus Jamla sensu stricto
This subgenus exhibits dextral coiling. The
talon when present is a simple peg. Sickle setae
are present in the third thoracic fascicles. The
collar does not form a tunnel dorsally.
Type
Ja11ua (Janua)
1865.
pagenstecheri Quatrefages,
Janua (Janua) pagenstecheri
Fig. 8a, 9
163
Habitat
J. pagenstecheri was found
zone, on stones.
III
the intertidal
Desc1'iption of Species
TUBE: The tube shows dextral coiling with
three longitudinal ridges. Terminal section usually partly overlies previous whorl and is sometimes upturned. Coil diameter is 1.3 mm.
OPERCULUM: The operculum has a lightly
calcified, partly concave, opercular plate which is
somewhat convex but has a central concavity like
an inverted saucer. The walls of the chamber
are more or less transparent and two or three
embryos, with white attachment glands, can be
distinguished inside the cup-shaped operculum.
A secondary plate forms the basal plate and this
gives rise to the terminal plate of a later chamber. The opercula studied lacked talons, probably because the chambers were not primary
ones.
THORAX: The thorax has two pairs of tori.
Collar folds are not fused dorsally. Collar setae
have a proximal "fin" of fine teeth and this is
followed immediately by the distal blade with
slightly narrower teeth. The splayed "fin" is
quite distinct on the two slides prepared of these
Hawaiian specimens but has proved less clear on
specimens from some other areas. The fin is not
like those of most other fin and blade collar
setae, however, and it appears to be due to
slightly larger proximal teeth tending to splay
when squashed whereas the finer distal teeth lie
flat. The fin is less distinct on the smaller collar
setae of the concave side. Several capillary setae
are associated with the collar setae. The second
and third fascicles have simple, finely serrated
setae and the third fascicle also possesses sickle
setae. Uncini have two or three longitudinal
rows of teeth and a tapering peg at the anterior
end around which several teeth are splayed.
Material
Three specimens are preserved in tubes and
two are mounted in polyvinyl-Iactophenol.
ASETIGEROUS REGION: The asetigerous region
is fairly long, approximately five times the distance between the first and second abdominal
tori.
Location
Sand Island, Oahu, Hawaii.
ABDOMEN: This structure has about six segments. Setae are geniculate with slightly re-
PACIFIC SCIENCE, Volume 26, April 1972
164
a
b
10
II (
++
c
d
(
++++
FIG. 8. Distribution of setae and uncini and comparative sizes of setal shafts and blades in: a, Janua
(Janua) pagenstecheri; b, Janua (Dexiospira) pseudocorrugata; c, Janua (Dexiospira) steueri; d, Janua
(Dexiospim) nipponica; e, Janua (Leodora) knightjonesi.
For explanation of histograms and setal stick diagrams see legend for Figs. 2 and 3.
Hawaiian Spirorbinae---VINE, BAILEy-BROCK, AND STRAUGHAN
165
,I'
:1\,
'10.
,1:1
E
E
d
1'1
e
N
0
0
('r
f
,I\i
FIG. 9. Janua (Janua) pagenstechej·i. a, Tube; b, side view of adult operculum; c, dorsal view of adult
operculum; d, collar seta with rudimentary fin; e, simple seta from second thoracic fascicle; f, sickle seta from
third thoracic fascicle; g, abdominal seta; h, thoracic uncinus; j, abdominal uncinus.
SCALE:
c as b,. e,
f,
g, h, and
i as
d.
curved teeth. Uncini have approximately 10
longitudinal rows of teeth and a rather broad
anterior fan lacking teeth.
INCUBATION: This occurs in the operculum.
Distribution Elsewhere
Zibrowius (1968, p. 201-203) reviewed in
some detail the taxonomic history and present
zoogeographic knowledge concerning this species. It is widely distributed in the Mediterranean Sea and Atlantic Ocean and, in addition
to the locations listed by Zibrowius, has been
found off the Galapagos Islands (Bailey and
Harris, 1968) and off Chios in the Aegean Sea
(Bailey, 1969a). S. epichysis Bailey, 1970, from
the West Indies is probably conspecific, because
the main difference noted by Bailey was the
absence of a proximal fin in the collar setae.
This feature has recently been shown to be frequently absent from J. (J.) pagenstecheri (see
Zibrowius, 1968, p. 202). J. (J.) pagenstecheri
has recently been found in southern Australia
(Phyllis Knight-Jones, in press) and in New
Zealand (unpublished, P.J.V. ) .
Other records from the Pacific show that the
species is found off North America as S. pttsil-
---------------------------------- - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -
166
loides (Pixell, 1912; see Zibrowius, 1968, for
discussion); from Mexico (Rioja, 1941, 1942);
from the Tuamotu atolls (Fauvel, 1919, 1947).
In the Indian Ocean it is known from Ceylon
(de Silva, 1961).
I. (J.) pagenstecheri thus appears to have a
worldwide distribution. The recent records from
southern Australia and New Zealand, together
with the present one from Hawaii, tend to support the record from Ceylon by de Silva (which
has been questioned by Pillai) and also that
from the Tuamotu atolls by Fauvel. Thus the
species appears to display an Indo-Pacific, as
well as Atlantic, Caribbean, and Mediterranean,
distribution. Zibrowius (1968) stated that S.
pusilloides and S. pagenstecheri are conspecific,
and this extends its distribution to the west
coasts of Mexico, the United States, and Canada.
Remarks
J. (J.) pagenstecheri differs slightly from all
other species in the genus by having the rudiments of a proximal fin in the collar setae. The
rudimentary fin has been observed by some authors and not by others. It is not a prominent
feature and its appearance partially depends on
how the setae are orientated and to what extent
they are squashed, in a polyvinyl-lactophenol
mount.
This species may also be unique amongst
lanua species in possessing sickle setae. It seems
possible that the sinistral forms with sickle setae,
described by Zibrowius (1968) as Spirorbis
laevis, may have been situs inversus of J. pagenstecheri. The puzzling status of Spirorbis laevis
is discussed later.
Subgenus Dexiospira Caullery & Mesnil, 1897
(= Neodexiospira Pillai, 1970)
Subgenus is like lanua, with coiling usually
dextral, but sickle setae are absent and the margins of the collar usually are fused to form a
tunnel over the middorsal thoracic groove.
Type
lanua (Dexiospira) pseudocorrugata (Bush,
1904).
PACIFIC SCIENCE, Volwne 26, April 1972
lanua (Dexiospira) pseudocorrugata
Fig. 8b, 10
Material
More than 100 specimens comprise the material.
Location
Coconut Island, Oahu, Hawaii.
Habitat
Species was found on settlement plates
shallows.
In
Description of Species
TUBE: Tube is dextral, with three longitudinal
ridges formed by rows of calcified "knobs" on
top of the tube. The central ridge is the most
prominent. Rather narrow transverse ridges pass
through these "knobs." The sides of the tube
slant outward and are vertically ridged. Two
whorls can be distinguished from above, and the
terminal section of the tube partially overlies the
previous whorl.
OPERCULUM: The operculum has a bifid calcified talon which is almost peripheral. In some
cases it dips under the rim of the lightly calcified plate whereas, in others, it overlaps the
edge of the plate. Walls are more or less transparent. A basal plate is present in some forms
and secondary (or later) opercula lack talons.
THORAX: Thorax has two pairs of tori. The
collar folds are fused to form a tunnel dorsally.
Collar setae on the convex side lack a proximal
fin and are coarsely toothed with indistinct crossstriations. Collar setae in the fascicle on the concave side lack cross-striations and are like the
simple setae of the second and third fascicles.
Uncini have a single anterior peg and approximately three longitudinal rows of teeth.
ASETIGEROUS REGION: This region is rather
long, approximately six times the distance between the first two abdominal tori.
ABDOMEN: This has approximately six segments. Setae are geniculate with long, rather
narrow blades and recurved teeth. Secondary
setae are present in some fascicles (see Vine, in
Hawaiian Spirorbinae-VINE, BAILEy-BROCK, AND STRAUGHAN
167
!:{!/
O,'~:
p .....
~
E
E
It)
N
"7'
o
a
E
E
9
f
e No
o
d
h
FIG. 10. Janua (Dexiospira) pseudocorrugata. a, Tube; b, side view of tube; 1:, juvenile form of operculum; d, mature form of operculum without eggs; e, collar seta; f, thoracic uncinus; g, simple seta from
second thoracic fascicle; h, abdominal uncinus; j, abdominal seta.
SCALE:
b as
aj
d as
I:j
f,
g, h, and j as
e.
press). U ncini have five to seven longitudinal
rows of fine teeth and have an anterior, somewhat convoluted, fan.
INCUBATION: This takes place in the operculum.
Distribution Elsewhere
Zibrowius (1968) reviewed the distribution
of this species in detail (under the name Spirorbis corrugatus), but he considered that several
other species were synonymous with it (see remarks below).
It seems likely that some of the species listed
by Zibrowius, though closely related, are not
conspecific and the distribution discussed here
relates only to J. (D.) pseudocorrugata.
It is widely distributed in the Atlantic Ocean
and Mediterranean Sea: English Channel (Caullery and Mesnil, 1897); Roscoff (i'Hardy and
Quievreux, 1964); northwestern Spain (Rioja,
1923); Morocc;o (Fauvel, 1936); Madeira
168
PACIFIC SCIENCE, Volume 26, April 1972
(Langerhans, 18806 ); Azores (Fauvel, 1909,
1914); Sargasso Sea (Fauvel, 1909, 1914);
Mexico (Rioja, 1959); Mediterranean Sea
(Caullery and Mesnil, 1897; Sterzinger, 1909,
1910; Fauvel, 1911; Casanova, 1954; Bellan,
1959; Zibrowius, 1968; Bailey, 1969a); West
Indies (Bailey, 1970). The only previous record
from the Pacific of which we are aware is that
of Knight-Jones (in press) from southern Australia.
J. (D.) foraminosa is an acceptable species with
a distribution that may be very limited.
Phyllis Knight-Jones (1971) discussed the
taxonomic history of J. (D.) pseudocorrugata
and established that Bush was correct in renaming, as S. pseudocorrugata Bush (1904), the
species called S. corrttgatus Montagu by Caullery
and Mesnil (1897), since the latter name had
been first applied to a sinistral form.
Remarks
Janua (Dexiospira) steueri Sterzinger, 1909
(= J. (D.) foraminosus)
There are a number of Janua species which
show slight variations in structure of tubes,
opercula, and details of setation. The mature
operculum of J. (D.) pseudocorrugata, which
lacks a talon, is difficult to distinguish from
a number of other species. Zibrowius (1968)
considered S. steueri Sterzinger (1909), S.
heideri Sterzinger (1909), S. foraminosa Moore
& Bush (1904), and S. treadwelli Pillai
(1965) to be conspecific with J. (D.) pseudocorrugata. Recent study in the Red Sea (Vine,
in press) suggests that J. (D.) steueri may be a
distinct species and that S. heideri should perhaps be considered as a subspecies of the latter.
The tube of J. (D.) stetteri usually has three
longitudinal ridges with deep indentations between them, forming a crisscross network of
ridges and indentations. Its juvenile operculum
has a bifid talon with wings (which are sometimes transparent) and the mature chamber has
rather heavily calcified, almost opaque, walls.
J. (D.) pseudocorrugata has a less distinctive
crisscross pattern on its tube and the talon of
the juvenile operculum has its broadest section
distally. In addition, the walls of the brood
chamber are more or less transparent. Pillai's
description of S. treadwelli, in details of tube
and operculum, resembles J. (D.) steueri rather
than J. (D.) pseudocorrugata, but he failed to
compare his material with either of these closely
related species.
The situation with regard to J. (D.) foraminosa is slightly more confusing since a recent
description by Day (1961) probably referred
to another species. This problem is discussed
more fully elsewhere, but it seems likely that
OPERCULUM: In the juveniles, a slightly concave plate has a bifid talon with calcified wings.
The bifid ends bend sharply and project ventrally. The talon persists in the primary brood
chamber of the mature worm and the wings
tend to be lightly calcified or even completely
transparent. The chamber walls are rather lightly
calcified and developing embryos can be distinguished, but not as clearly as in J. (D.)
pseudocofrttgata. Secondary brood chambers lack
a talon and are then more difficult to distinguish
from those of J. (D.) pseudocorrugata.
(; These records probably do not refer to the same
species as that described above.
THORAX: Thorax has two pairs of tori. Collar
folds are fused to form a tunnel dorsally. Collar
Fig. 8c, 11
Material
Three specimens were preserved in tubes and
one mounted in polyvinyl-Iactophenol.
Location
Maili Point, Oahu, Hawaii.
Habitat
Animals were found on an alga, at approximately 8 m deep.
Description of Species
TUBE: Tube is dextral with three longitudinal
ridges on the top and one on the side. The
outer ridge on top of tube has outwardly projecting blunt processes. There are deep indentations between the ridges and the two outer
ridges are actually perforated where indentations
on opposite sides connect. Tube has one or two
whorls. Coil diameter is 1.5 to 2.0 mm.
Hawaiian Spirorbinae-VINE, BAILEy-BROCK, AND STRAUGHAN
169
O'5mm
d
9
'i
;
1
h
!
FIG. 11. Janua (Dexiospira) steueri. a, Tube; b, juvenile operculum; c, mature operculum; d, thoracic
seta from second fascicle on convex side; e, collar seta from convex side; !, collar seta from concave side;
g, abdominal seta; h, abdominal uncinus; j, thoracic uncinus.
SCALE:
c as bj e,
f,
g, h, and j as d.
setae on the convex side have simple blades,
with coarse teeth and rather faint cross-striations,
whereas those on the concave side have smaller
blades, narrower teeth, and no cross-striations.
Second and third fascicles have finely striated
simple setae and there are no sickle setae in the
third fascicles. Uncini have three or four longitudinal rows of teeth and these splay out
around a triangular peg at the anterior end.
ASETIGEROUS REGION: This area is long, approximately 10 times the distance between the
first two abdominal tori.
ABDOMEN: The abdomen has about 14 segments. Setae are geniculate with a rather broad
blade and small teeth. Uncini have about six
longitudinal rows of teeth and a broad fan at
the anterior end.
INCUBAnON: This occurs in operculum.
Distribution Elsewhere
J. (D.) steueri, of which this is probably a
variety, is rather widely distributed. It was first
recorded by Sterzinger (1909) in the Red Sea,
at Suez, and has recently been found farther
170
PACIFIC SCIENCE, Volume 26, April 1972
south in the Red Sea, near Port Sudan (Vine,
in press). Phyllis Knight-Jones (1972) has recorded it from Kenya and in discussing this
species she tentatively suggested, after studying
original material, that S. treadwelli is synonymous. This has been recorded from the Philippines (Pillai, 1965) and at Heron Island on
the Great Barrier Reef of Australia (Straughan,
1967). J. (D.) stetteri was also found in southern Australia by Phyllis Knight-Jones (in
press), and off northeastern Australia and South
Island, New Zealand, by Vine (personal observation). In addition to these records in the
Indo-Pacific, Bailey (1970) recorded J. (D.)
steueri in the Caribbean Sea. It is, therefore,
rather remarkable that it has not been discovered
elsewhere in the Atlantic or in the Mediterranean.
setae are not so clearly cross-striated as those of
the Red Sea form. The asetigerous region is
longer in the Hawaiian form and the blades of
the abdominal setae are broader, approaching
in width those of J. (D.) foraminosa or J. (D.)
formosa.
The Hawaiian variety of J. (D.) steueri bears
some striking resemblances to J. D. foraminosa
Moore & Bush, 1904, from the Sea of Japan.
The original material has since been reexamined
by Phyllis Knight-Jones who finds that the tentacles are particularly long, their ends reaching
the outer rim of the opercular brood chamber.
The species also differs from J. (D.) steueri in
that the blades of the abdominal setae are shorter
and broader, approaching those of J. (D.) formosa. The species differs from the latter, however, in having cross-striated collar setae. It
seems best at present to regard J. (D.) steueri
as being distinct from J. (D.) foraminosa.
It now seems clear that this Hawaiian material is really J. (D.) foraminosa (Bush, in
Moore and Bush, 1904). This species was inadequately described, but Phyllis Knight-Jones
(personal communication) has examined the
type from the Smithsonian Institution and fresh
material from Japan. She found that it usually
differs from the true J. (D.) steueri in the pattern of pitting between ridges on the tube, and
always differs in having more transparent walls
to the brood chamber and wider blades on the
abdominal setae. (In the last character, which
is easily seen, it is intermediate between J. [D.]
steueri and J. [D.] formosa.)
Remarks
J. (D.) steueri has, until recently, been a
rather doubtful species. Zibrowius (1968) considered it to be synonymous with J. (D.) pseudocorrugata but material studied from Kenya
(Phyllis Knight-Jones, 1972) and the Red Sea
(Vine, in press) has indicated several differences between these closely related species. The
tube of J. (D.) steueri is usually more intricately
patterned with three or four longitudinal ridges
and deep transverse indentations. The juvenile
opercula differ, that of J. (D.) steueri having
asymmetrical lateral wings arising from the
middle of a long, bifid, densely calcified, central talon. The other species has a more fanshaped talon with the broadest section distally,
and the walls of the mature opercular brood
chamber are less densely calcified and almost
transparent. J. (D.) steueri varies in different
areas, but there is considerable variation in
tubes and talons of specimens from the same
atea, so it seems sensible for the present to
regard these slight geographical variants as belonging to the same species.
The Hawaiian variety described here differs
in several points from the forms found in the
Red Sea area (Vine, in press). Only dextrally
coiling specimens were found. The tube has
four longitudinal ridges (compared with three
on the Red Sea species) and perforations between the ridges are often complete. Collar
Jantta (Dexiospira) nipponica Okuda, 1934
Fig. 8d, 12
Material
Eight specimens.
Location
Nanakuli, Oahu, Hawaii.
Habitat
J. nipponica was found in the littoral zone on
a red alga.
171
Hawaiian Spirorbinae-VINE, BAILEy-BROCK, AND STRAUGHAN
a
b
O'5mm
d
E
E
f
e
N
c
9
o
o
FIG. 12. Janua (Dexiospira) nipponica. a, Tube; h, juvenile operculum; c, adult operculum; d, collar
seta from convex side; e, collar seta from concave side; f, simple seta from second thoracic fascicle; g, abdominal seta; h, abdominal uncinus; j, thoracic uncinus.
SCALE:
cas h; e,
f,
g,
h, and j as d.
Description of Species
TUBE: Tube is dextral, with three longitudinal ridges on top and one on side of tube.
Transverse bars between these ridges create an
appearance of deep oblong indentations in the
surface of the tube. Two whorls are present.
Terminal section sometimes ascends. Coil diameter is 1.5 to 2.0 mm.
OPERCULUM; The operculum is small and
has a flat or slightly concave calcareous plate
with a wedge-shaped talon. An indentation may
be present creating a slightly bifid appearance.
Brood chamber has a narrow transparent rim
around a more or less flat calcified plate. The
walls are almost transparent and the calcified
talon is usually less bifid than in Janua pseudocOt'rugata Bush (1904).
172
PACIFIC SCIENCE, Volume 26, April 1972
THORAX: Thorax has two pairs of tori. lJorsal collar folds are fused. Collar setae are simple, finely serrated blades, not coarsely toothed
or cross-striated on either side. Second and third
fascicles have similar simple setae and there
are no sickle setae in the third segment. Uncini
have at the anterior end a broad peg and two
large teeth, one on each side, creating a distinctly trifid appearance. Teeth of the uncini
are arranged rather irregularly, in approximately four longitudinal rows.
of J. (D.) alveolata. Their description clearly
refers to J. (D.) nipponica rather than to
J. (D.) alveolata.
Day (1961) described a species from South
Africa which he identified as J. (D.) foraminosa
and kindly made available for examination. It
clearly has trifurcate thoracic uncini (as Day
described) and is apparently the same as the
species described above as J. (D.) nipponica.
In type material of J. (D.) foraminosa Bush,
kindly supplied by the Smithsonian Institution,
the thoracic uncini have single pointed pegs.
The shape of the talon suggests close affinity
with J. (D.) psettdocorrttgata (also described
above), but it can be easily distinguished from
this species because it lacks cross-striations on
the collar setae and has trifid pegs on the
thoracic uncini.
ASETIGEROUS REGION: Region is long, approximately equivalent to 10 times the distance between the first two abdominal tori.
ABDOMEN: This structure has six rows of
uncini on each side. Setae are geniculate and
have their blades relatively shorter than in many
lantta species. Secondary setae are present in
some segments. Uncini are broad and have approximately six longitudinal rows of fine teeth.
The anterior end lacks teeth and appears somewhat fluted.
INCUBATION:
This occurs in the operculum.
Distribtttion Elsewhere
Species is known in Japan as S. nipponictts
(Okuda, 1934, 1937) and as S. aiveolattts (Imajima and Hartman, 1964). In South Africa it
is known as S. foraminostts.
Remarks
It has been suggested by Ushakov ( 1955) ,
Imajima and Hartman (1964), and Pillai
(1965) that J. (D.) nipponica is synonymous
with J. (D.) alveolata Zachs (1933). Although
the description by Zachs lacks diagrams and
does not satisfactorily describe the operculum
or details of setation, it nevertheless seems to
refer to a species quite different from J. (D.)
nipponica. The tube of J. (D.) alveolata is semitransparent and Zachs mentions that incubation
probably takes place in the tube. Although he
drew a loose comparison between it and J. (D.)
joraminosa based on both having smooth collar
setae and foramina in their tubes, he dismissed
the possibility that they could be the same because the opercula were not similar.
Imajima and Hartman follow the suggestion
of Ushakov and list S. nipponica as a homonym
Subgenus Leodora de Saint-Joseph
(1894), amended
Sinistral coiling is present; collar folds are
unfused; collar setae are not coarsely toothed or
cross-striated; sickle setae are absent.
Type
lantta (Leodora) laevis (Quatrefages, 1865).
lantta (Leodora) knightjonesi (de Silva, 1965)
Fig. 8e, 13
Material
Eight specimens are preserved in tubes and
three mounted in polyvinyl-lactophenol.
Location
Sand Island in Kaneohe Bay, Oahu, Hawaii.
Habitat
Species was found
stones.
10
the intertidal zone on
Dem·iption of Species
TUBE: This is sinistral, with three well-defined
longitudinal ridges. Mouth of tube is sometimes
upturned away from the substrate. Coil diameter
is 1.5 mm.
OPERCULUM: Operculum often bears one,
two, or occasionally three brood chambers in se-
Hawaiian Spirorbinae-VINE, BAILEy-BROCK, AND STRAUGHAN
173
E
E
I/')
N
o
Q'5mm
m
[~
f
I
'.
E
E
9
f
N
0
0
h
l
k
e
FIG. 13. Janua (Leodora) knightjonesi.
d, side view of adult operculum; e, collar
concave side; g, simple seta from third
j, thoracic uncinus; k, abdominal uncinus;
SCALE:
b as a; d as c;
f,
a, Tube; b, side view of tube; c, dorsal view of adult operculum;
seta from fascicle on convex side; f, collar seta from fascicle on
thoracic fascicle; h, capillary seta from third thoracic fascicle;
m, primary and secondary abdominal setae.
g, h, j, and k as e.
ries, each with very lightly calcified, transparent
walls, and a more or less flat plate bearing a
white calcified talon, which is approximately
round in cross section. Each chamber contains
developing embryos, those in the distal chamber being more developed. The basal plate of
the distal chamber is also the terminal plate of
the chamber immediately underneath.
THORAX: This structure has two pairs of tori.
Collar setae are rather geniculate and broad
proximally. Those on the convex side have
blades almost twice as long as those of the
concave side, but the basic structure of the collar
setae on both sides is similar, each having blades
with jagged edges, the teeth being somewhat
coarser proximally than distally. The blade is
---- . _ - - - - - - 174
constructed of several layers of teeth and in
some cases these have peeled back (on the polyvinyl-lactophenol mounts) to reveal the multilayered structure of the blade. There are no distinct cross-striations, however, inasmuch as the
rows of teeth on the edge of the blade are too
few and too irregularly arranged. Associated
with the collar setae on each side are several
capillary setae. Setae of the second and third
fascicles have extremely narrow, finely striated
blades and are accompanied by some capillary
setae. There are no sickle setae in the third
thoracic fascicles. Each uncinus has a blunt anterior peg and on each side of this there are
several teeth which tend to fan out, giving the
anterior end a shape reminiscent of a tapering
spade. This is rather characteristic of this species
and has not been mentioned by previous authors.
ASETIGEROUS REGION: This is long, approximately seven times the distance between the first
and second abdominal tori.
ABDOMEN: The abdomen has approximately
six segments. Setae are geniculate with rather
long narrow blades and slightly recurved teeth.
Secondary setae are present in several abdominal
segments. These have already been noted in
other members of the genus Janua (Vine, in
press; Knight-Jones, 1972). They have rudimentary shafts and blades approximately the
same length as those of the primary abdominal
setae. Uncini have approximately 10 longitudinal rows of fine teeth and a broad, somewhat
convoluted fan.
INCUBATION: Incubation in Janua knightjonesi takes place in opercular brood chambers.
Distribtttion Elsewhere
Species is found in Ceylon (de Silva, 1965);
West Indies (Bailey, 1970); Australia (personal
observation by one of us, P.J.V.). As S. laevis,
it is found in CuraC;ao (Augener, 1936).
Remarks
De Silva (1965) described Janua (Leodora)
knightjonesi from Ceylon. He recognized its
affinities with the subgenus Leodora and the
type "S. laevis" Quatrefages (1865). He listed
other species which he believed should be included in this subgenus. They were: S. verrttca
PACIFIC SCIENCE, Volume 26, April 1972
Fabricius (1780); S. valida Verrill (1874); S.
perrieri Caullery & Mesnil (1897); S. abnormis
Bush (1904), and S. coronatus Zachs (1933).
These were grouped on the basis of their sinistral coiling and simple collar setae and, in several cases, the mode of brood protection had not
been recorded.
Bailey (1969b) reviewed the taxonomy of
Spirorbinae and emphasized the importance of
methods of brood protection. On this basis. the
subgenus Leodora is separated from the genus
Romanchella Caullery & Mesnil (1897) (see
Knight-Jones, in press). Recent studies have
shown that the latter genus includes R. perriefi
and probably R. coronata. Of the remaining
little-known species which de Silva grouped in
the subgenus Leodora, only S. laevis has opercular details somewhat similar to those of J. (L.)
knightjonesi.
The status of J. (L.) laevis was recently discussed by Bailey (1970) who found that material identified by Augener (1936) as S. laevis
was the same species as that described by de
Silva (1965) from Ceylon as S. knightjonesi.
In view of the taxonomic history of J. (L.)
laevis it was correct of de Silva to apply a new
name to the species which had already been
described by Augener because it was clearly
different from that first described by Quatrefages as S. laevis (see below). J. (L.) knightjonesi differs from every adequate description
of material identified as S. laevis (except for
that of Augener) in at least some of the following characters: its sinistral tube with three
distinct longitudinal ridges; its stacked opercula, every chamber of which possesses a talon;
unfused collar; broad geniculate collar setae
which lack a proximal fin or cross-striations; the
shape of the anterior end of the thoracic uncini;
and the absence of sickle setae from the third
thoracic fascicles.
J. (L.) knightjonesi has now been found in
Ceylon (de Silva, 1965); West Indies (Augener, 1936; Bailey, 1970); Australia (personal
observation, P.J.V.), and Hawaii. It thus appears to have an Indo-Pacific and Caribbean
distribution. It has not been recorded on the
west coast of the Americas or in Europe, and
Bailey (1970) offered a possible explanation for
these apparent gaps in its distribution. It is pos-
Hawaiian Spirorbinae-VINE, BAILEy-BROCK, AND STRAUGHAN
sible, however, that some records of S. laevis
in Europe have referred to this species and further research may show that these gaps are more
apparent than real.
THE PROBLEM OF "SPIRORBIS LAEVIS"
The original description by Quatrefages
(1865) of a species from Guettary was inadequate. It did not refer even to the direction of
coiling, but some later authors appear to have
regarded the name as indicating sinistral coiling,
because the Latin laevtts means on the left side.
In fact, the Latin laevis means smooth and it is
uncertain whether the species which Quatrefages
described was sinistral or dextral. He was clearly
referring to smoothness of the tube in the name
laevis, because he placed his species under the
heading "Spirorbe lisse." His description covered several other important characters. The
animal had only four branchiae and the cylindrical operculum had a narrow rim around the
distal plate. He described the operculum as containing red cells which were presumably eggs.
He completed this short description (which did
not include the setae) with the comment: "Je
regret de n'avoir pas pris de notes etendues sur
cette petite Annelide."
Claparede (1870) provided a much more
complete description of a sinistral species from
Naples which he considered fitted the important
points in Quatrefages' description of S. laevis.
At that time taxonomists were not aware of the
large number of species of Spirorbinae present
in the Mediterranean and N. Atlantic and Claparede was probably justified in guessing that
his Naples species tallied with Quatrefages'
completely inadequate description of a species
from the Bay of Biscay.
Claparede described a sinistral smooth tube
and illustrated a rather more cylindrical operculum than that figured by Quatrefages. He described the setae in some detail but made no
mention of sickle setae. In fact, he inferred that
these were absent: "Les soies dorsales des deux
autres segments thoraciques sont filiformes,
lanceolees a l'extremite." He gave drawings of
all the setae and his study appears to have been
a careful one, so there is no reason to suppose
that the species he described had sickle setae in
the third thoracic fascicles. His drawing of the
175
whole animal is perhaps the best drawing from
life of any of the Spirorbinae and the best
known, having been reproduced in several major
textbooks of invertebrate zoology. It seems
doubtful, however, if there has been any record
since that time of a mature form with only two
pairs of tentacles.
Since the work of these two early authors,
almost every mention in the literature of S.
laevis has been either (1) a reference to these
first descriptions, without any new material (de
Saint-Joseph, 1894; Caullery and Mesnil, 1897;
Rioja, 1923; Fauvel, 1927; Prenant, 1927;
Cornet and Rullier, 1951; Laubier and Paris,
1962; L'Hardy and Quievreux, 1964; Cabioch,
L'Hardy, and Rullier, 1968) or (2) records of
its occurrence usually comprising just a single
sentence, not supported by drawings or a description or any note of material deposited in
a museum (Pruvot, 1897). Pixell (1913) recorded this species at Suez but her brief description referred to a form with three longitudinal
ridges. That may well have been the sinistral
form of Jantta stetteri, which was recorded there
by Sterzinger (1909) and was recently rediscovered in the Red Sea (Vine, in press). Ehlers
(1913) described a species from South Africa
as S. laevis but this had a characteristically ridged
tube with deep indentations. and his drawing
is also somewhat reminiscent of the sinistral
form of J. (D.) stetteri found in the Red Sea
(Sterzinger, 1909; Vine, in press) and of a
sinistral species on pearl oysters in the Tuamotu
atolls (Fauvel, 1919).
Augener (1936) was the first taxonomist
since Claparede to provide an adequate description of material from the Caribbean Sea, which
he called S. laevis. His specimens differed from
earlier descriptions in that they had three distinct longitudinal ridges and often had multiple
opercular chambers, each with a talon (see
Bailey, 1970). De Silva (1965) discovered the
same species as this in Ceylon and, recognizing
that it differed from S. laevis, called it S. knightjonesi. Although Augener had already provided
an almost adequate description of de Silva's
species there was good reason for de Silva to
rename it inasmuch as it was clearly different
from S. laevis Quatrefages, and there is a possibility that later authors may discover a species
176
which tallies closely with that recognized by
Claparede as S. laevis.
Zibrowius (1968) described a few sinistral
opercular-incubating specimens which he obtained from a submarine cave near Marseille;
he mentioned a tube with three longitudinal
ridges, an operculum like that of J. (J.) pagenstecheri, and sickle setae in the third thoracic
fascicles. In fact, his description differs from
that of J. (J.) pagenstecheri only in direction
of coiling. In three important details it differs
from the species described by Quatrefages and
Claparede. Its tube was not smooth, its operculum was more cup-shaped than cylindrical
(like Quatrefages' drawing), and there were
sickle setae in the third fascicles (not found by
Claparede, Fauvel, Pixell, Augener, or any other
author).
Direction of coiling may be reversed both
within a single species (see Sterzinger, 1909;
Bock, 1953; Potswald, 1965; and also Vine, in
press) and within a genus (see Knight-Jones,
1971 and Vine, in press), so that it appears
possible that the few specimens described by
Zibrowius and identified as S. laevis were, in
fact, sinistral mutants of J. (J.) pagenstecheri
which he recorded from the same locality and
habitat.
If one discounts these records by Augener
(1936) and Zibrowius (1968), no record of
S. laevis has been supported by an adequate description or drawings, since that of Claparede
(1870).
In view of the recent rediscovery of the
sinistral variety of J. (D.) stetteri in the Red
Sea (Vine, in press) and of the sinistral species
within the subgenus Fattveldora Knight-Jones
(1972), it seems quite probable that some
records of sinistral opercular incubators with
simple collar setae may have been attributed
to S. laevis without close investigation of details
of tube structure, opercular structure, or setation.
In view of the inadequacy of the original description by Quatrefages and the subsequent
failures to find anything that is comparable in
detail with the figures of Claparede, it seems
that S. laevis should be regarded as a nomen
dttbittm. It is unfortunate that de Saint-Joseph
(1894) selected such a doubtful species when
he established the genus Leodora.
PACIFIC SCIENCE, Volume 26, April 1972
ECOLOGICAL OBSERVATIONS
The majority of species collected were present
in the intertidal zone settling on lava rock, dead
corals, and molluscan valves, especially oysters
(Iso gnommn sp.), or algae (see Table 1). Only
Pileolaria (Pileolaria) semimilitaris, Pileolaria
(Duplicaria) dalestrattghani, and the Protolaeospira sp. were absent from the intertidal collections.
Shallow rock pools in the upper shore and
splash zone contained the tube-incubating species
Spirorbis (Spirorbella) marioni which is presumably better adapted to avoid desiccation and
salinity fluctuations than are most of the opercular-brooding species. The other species which
can apparently tolerate the conditions in the
upper shore is the opercular incubator Jamta
(Dexiospira) psettdocorrugata which often settles on valves of the oyster (Isognomttm sp.)
and on lava rock.
Most other species could be found in shallow pools of the middle and lower shore under
stones and On algae.
The rather large species Pileolaria (Pileolaria)
militaris settles on a calcareous red alga which
encrusts the undersides of platelike coral structures. Pileolaria (Simplicaria) psettdomilitaris
is an abundant intertidal species usually found
on the undersides of stones on the lower shore
and in the shallow sublittoral Zone. Pileolaria
(Dttplicaria) koehleri was found intertidally on
stones in rock pools but appears to favor settlement below tidemarks and can live in the intertidal zone only when it is permanently covered
by water. Pileolaria (Dttplicaria) dalestrattghani
settles on bryozoa encrusting dead corals in
deeper water (such as McVey's artificial reef off
Oahu, 15 m).
Jantta (Jamta) pagenstecheri occurred sparsely
on the underside of stones in the intertidal zone
and Bailey and Harris (1968) suggested that it
may be more sublittoral in tropical conditions.
Jantta (Dexiospira) psettdocorrttgata, J. (D.)
stetteri, and J. (Leodora) knightjonesi all settle
on hard substrate on the middle and lower shore.
J. (D.) stetteri also settles on algae whilst J.
(D.) nipponica and J. (D.) tttrrita were found
exclusively on algae.
Settlement plates lowered at various times into
shallow water at Coconut Island, off Oahu, col-
Hawaiian Spirorbinae--VINE, BAILEy-BROCK, AND STRAUGHAN
177
TABLE 1
SPECIES AND NUMBERS OF SPJRORBINAE COLLECTED, BY INTERTIDAL AND SUBTIDAL ZONES
INTERTIDAL
SPECIES
Spirorbis (Spiro, bella)
marioni
Protolaeospira sp. A
Pileolat'ia (Pileolaria)
militaris
Pileolat'ia (Pileolaria)
semimilitaris
Pileolaria (Simplicaria)
pseudomilitar'is
Pileolaria (Duplicaria)
koehleri
Pileolaria (Duplicat·ia)
dalestraughani
Janua (Janua)
{Jagenstecheri
Janua (Dexiospira)
pseudocor'rugata
Janua (Dexiospira)
steueri
Janua (Dexiospira)
tunita
Janua (Dexiospira)
nipponica
Janua (Leodora)
knightjonesi
ON STONES,
DEAD CORALS,
OR SHELLS
SUBTIDAL
ON ALGAE
ON STONES,
DEAD CORALS,
OR SHELLS
ON ALGAE
abundant
(24)
rare (l)
uncommon
(l)
uncommon
(2)
abundant
(11)
abundant
(+)
probably
common (7)
abundant
(19)
abundant
(156 +)
possibly
locally
common (3)
abundant
(100 +)
abundant
in shallows
present
probably
common (3)
abundant
(14)
probably
common (8)
probably
present
common (11)
NOTE: Numbers in parentheses indicate number of specimens collected.
quantitative. Comments are results of observations by authors.
lected many of the species described here.
Pileolaria pseNdomilitaris and JanNa (Dexiospira) psetldocormgata were particularly abundant on the plates. After approximately a
month's settlement, plates had many fully mature adults of these two species, with welldeveloped embryos in their opercula.
The paucity of species collected on the island
of Hawaii (in comparison with Oahu) and the
absence of species from the upper littoral zone,
such as Spirorbis (Spirobella) marioni, may
perhaps be explained by active volcanic action,
for a massive lava flow had occurred in the collecting region several years prior to our study.
Frequent rises in coastal sea temperature asso-
+
indicates large numbers of individuals-not
ciated with these flows must have resulted in
repeated reconstruction of the littoral zone. The
only possibly endemic species which was found
in the littoral zone, Jamla (Dexiospira) tttrrita,
came from this region, suggesting that the disruptive influence of lava flows may have promoted invasion of the littoral zone by an
endemic, but normally sublittoral, species,
through the removal of more successful and
widely distributed competitors. It must be admitted that this species was not found sublittorally, but then the sublittoral collections
were not very extensive.
There is considerable scope for the study of
the ecology of Hawaiian Spirorbinae, and it is
------ ---------------------------------------
178
PACIFIC SCIENCE, Volume 26, April 1972
hoped that the present taxonomic study will
provide a good basis for further work in this
area.
also show close affinities with the Hawaiian
species. Although geographically closest to Hawaii, the west coast of America shows least
similarity in its assemblage of Spirorbinae (only
three out of nine species in common with Hawaii), and the east Atlantic has a similarly low
number of species in common with Hawaii.
Ekman (1967, p. 19) has drawn attention to
the high percentage of endemic species around
Hawaii, citing molluscs, crustaceans, echinoderms, and fish, but Spirorbinae differ from
many of these forms in being easily transported
as adults and, indeed, as breeding colonies. Such
transport of adults is likely to be much more
effective in dispersal than the transport by currents of comparatively ephemeral planktonic larvae.
Considering the ease with which many adult
Spirorbinae species may be transported on floating algae and the stones or shell fragments
associated with these, and on crustacean carapaces, turtle shells, driftwood, ships' bottoms,
and perhaps on algae or barnacles attached to
cetaceans, the differences in distribution of species is unlikely to be explained by differential
transport to various regions. It seems more likely
that there are a number of species (approxi-
ZOOGEOGRAPHICAL DISCUSSION
The species described here show several characteristics typical of tropical and subtropical
Spirorbinae. The two dominant genera, Pileolaria and JanNa, are both opercular incubators.
Only two out of 13 Hawaiian species incubate
their embryos in their tubes. It appears that
opercular incubators are generally favored in
warm waters, to judge from recent research elsewhere in the tropics (Bailey, 1970; Pillai, 1970;
Knight-Jones, 1972; and Vine, in press). Although the opercular brood chambers seem more
exposed to predation, they must help embryonic
respiration, which may present problems to tube
incubators in warm waters.
The affinities between Spirorbinae of the
Hawaiian chain and those from other regions
at similar latitudes where adequate studies have
been carried out are summarized in Table 2.
Closest affinity is with the western Pacific (eight
out of nine spp.). Spirorbinae from the Indian
Ocean (six out of nine), Mediterranean (five
out of nine), and Caribbean (six out of nine)
TABLE 2
SPECIES OF SPIflORBINAE AND AREAS IN WHICH THEY WERE COLLECTED
HAWAiiAN
SPECIES
Spirorbis marioni
Pileolaria militaris
Pileolaria
pseudomilitaris
Pileolaria koehlel'i
Janua pagenstecheri
Janua
pseudocol'rugata
Janua steueri
Janua nipponica
Janua knightjonesi
Percentage similarity
of species
assemblage with
that found
around Hawaii
AREA 2
AREA ,3
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
89%
67%
AREA 1
AREA
4
AREA 5
AREA
6
X
33%
X
X
X
X
X
X
X
X
X
X
X
X
X
X
33%
56%
67%
KEY: X, species has been recorded; -, species has not been recorded.
AREAS: 1, Eastern Pacific; coast of U.S.A.; 2, western Pacific; Australia, and New Zealand; 3. Indian Ocean and Red
Sea; 4, western Atlantic; Spain and North Africa; 5, Mediterranean Sea; 6, Caribbean Sea.
Hawaiian Spirorbinae-VINE, BAILEy-BROCK, AND STRAUGHAN
mately eight) which show a widespread distribution throughout regions with water warm
enough for growth of hermatypic corals. Although at similar latitudes to Hawaii, the west
coasts of California and of North Africa do
not have coral reefs. Cold coastal currents lower
surface sea temperatures in these regions. The
most widespread warm-water Spirorbinae appear
to be Pi/eolaria militaris, P. pseudomilitaris, P.
koehleri, Jamta pagenstecheri, J. pseudocorrugata, J. stetteri, J. nipponica, and J. knightjonesi.
Of these eight species, Pileolaria militaris, Janua
pagenstecheri, and J. pseudocorrttgata appear to
be eurythermal, also occurring widely in temperate regions.
Not enough Spirorbinae from deeper water
localities elsewhere in the Pacific have been
studied to establish whether the three new species from the Hawaiian Islands are endemic.
The most likely endemic species is perhaps J.
(D.) turrita, which was abundant on red alga
in the intertidal zone on Hawaii but not found
on Oahu. It is remarkable that the Hawaiian
list of Spirorbinae does not contain more endemic species. Evidently there have been continuous invasions of species from other regions.
SUMMARY
Collections from the Hawaiian chain included
13 Spirorbinae, three of which are new. Eleven
species belong to the opercular-incubating genera
Pileolaria or Janua, whereas two tube-incubating
genera, Spirorbis and Protolaeospira, are each
represented by single species. Most of the species
are widely distributed in coral reef areas and
close links exist between Hawaii, the western
Pacific, Indian Ocean, Mediterranean, and the
Caribbean, whereas links with the west coast of
America seem to be slighter. The low numbers
of endemic Spirorbinae in the Hawaiian chain
and the widespread distribution of species can
perhaps be explained by the ease with which
adults may be transported. Distribution of species appears to reflect hydrological conditions
and physiological tolerances of species rather
than greatly different transport.
ACKNOWLEDGMENTS
We wish to thank Rotary International and
the University of Wales for supporting the
179
travel and research of one of us (P.J.V.) and
the American Association of University Women
for a fellowship enabling one of us (D.S.) to
collect Spirorbinae off Oahu. For assistance in
diving and collecting we are grateful to Professor A. H. Banner and Dr. J. H. McVey of the
University of Hawaii. We are grateful to Mrs.
Phyllis Knight-Jones for discussions on the
taxonomy of Spirorbinae and for prepublication
copies of her papers on Spirorbinae from southern Australia and Kenya, and to Professor E. W.
Knight-Jones for advice on presentation. The
work was carried out by one of us (P.J.V.) as
part of the requirement for the Ph.D. degree.
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