ZoologicalJournal ofthe Linnean Society, 64: 201-240. With 18 figures
November 1978
New Spirorbidae (Polychaeta: Sedentaria)from
the East Pacific, Atlantic, Indian and
Southern Oceans
PHYLLIS KNIGHT-JONES
Department of Zoology, University College of Swansea,
SA2 8PP, U.K.
Acceptedforpublication February 1977
Spirorbis b$matus, Spirorbis rothlisbergi, Spirorbis spatulatus, Pileolaria lateralis, Pileolaria margznata,
Pileolaria spinfer, Pileolaria tiarata and Pileolaria potswaldi are described from California, Spirorbis
strigatus frofn Madeira, Pileolaria dakaremis from W. Africa, Pileolaria (Duplicaria) zibrowii from
Madagascar, Pileolaria data and Pileolaria (Nidularia) nidica from Curacao, Pileolaria (Nidularia)
palliata from S . Africa, Pileolaria (Jugaria) atlantica from north of the Azores, Rumanchella bicava from
Amsterdam Island, Romanchella pustdata from Chile and Helicosiphon platyspira from Marion Island.
Of the new subgenera, Jugaria is proposed for forms having compound opercula (as in Pileolaria
granulata L. and most Sinistrella species)and Nidularia for those with opercula that do not completely
protect the embryos.
Some of the Californian forms have probably resulted from sympatric speciation in both the
genera concerned. The morphology and zoogeography of spirorbids suggest division into the
subfamilies Spirorbinae, Circeinae, Pileolariinae, Januinae, Romanchellinae and Paralaeospirinae.
KEY WORDS :-Polychaeta-Serpulidae-Spirorbidae-speciation-~onomy-zoogeography.
CONTENTS
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Introduction
Methods
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Taxonomy of Spirorbidae
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Subfamily Spirorbinae
Genus Spirorbis Daudin (1800)
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Spirorbish$rcattlrr sp. nov.
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Spirorbis rothlisbergisp. nov.
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Spizorbisspotulahrc sp. nov.
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Spirorbis strigatus sp. nov. . . . . . . . . . . . .
Subfamily Pileolariinaenov. . . . . . . . . . . . . . .
Genus Pileolaria Clapari.de (1868) . . . . . . . . . . . .
Subgenus Pileohria
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Pile&& a h sp. nov. . . . . . . . . . . . . .
Pileokaria a’akarcnsis ap. nov. . . . . . . . . . . . .
Pileokaria lateralis sp. nov. . . . . . . . . . . . .
Pile&
margincrta sp. nov. . . . . . . . . . . . .
Pikolariu rpiniii sp. nov. . . . . . . . . . . . .
Pilcokuiatiarata sp. nov.
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Subgenus Simplicaria Knight-Jones (1973)
Pile&riu(S.)potswaldinom.nov. . . . . . . . . . .
0024-4082/78/110201-40/502.00/0
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0 1978TheLinneanSocietyofLondon
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P. KNIGHT-JONES
Subgenus Niduloria subgen. nov. .
Pileohria ( N . ) nidua sp. nov. .
Pilrdaria (N.)pallioto sp. nov. .
SubgenusDuplicariaVine(1972a)
Pilcolmia (0.)zibrowu sp. nov. .
Subgenus Jugaria subgen. nov. . .
Piledoria (J.) atlantua sp. nov. .
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SubfarnilyRomanchellinaenov. . . . . . . . . .
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Genus Rumanchdla Caullery & Mesnil(1897)
Romanchella &ma sp. nov.
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Rnnrmchelloputhrlnto sp. nov. . . . . . . .
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GenusHelicosiphon Gravier(l907)
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Helicosiphonj&tyspirasp. nov. . . . . . . . . . . . . . . . 233
Specific characters, speciation and higher taxa in Spirorbidae
Summary
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Acknowledgements
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References
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INTRODUCTION
Most of the material for these studies was collected and kindly provided by Dr
J. Beurois, Prof. R. P. Dales, Mr R. Dubois, Dr P. Freon, Dr Gilchrist, Dr M.
Jones, Prof. E. W. Knight-Jones, Prof. Maria Lopez, Dr L. M. Marhue, Dr R.
Rowe, Dr P. C. Rothlisberg, Dr J. W. Scoggen, Mr W. M. Shepherd, Mr J. F.
Shisko, Dr Dale Straughan, Dr H. ten Hove, Dr J. Vacelet, Dr A. F. de Villiers
and Dr H. Zibrowius. Further specimens occurred in material kindly loaned by
the Smithsonian Institution (U.S.N.M.); the Hancock Museum (University of
Southern California); the British Museum (Nat. Hist.); the National Museum of
Canada; the Universitets Zoologiske Museum, Copenhagen; and Zool. Mus.
Humboldt-Universitat, Berlin.
Half these new species are from California, for Hartman ( 1969)had included
no original work on spirorbid taxonomy, but relied for identifications of species
on studies by Caullery & Mesnil (18971, Bush (1904) and Berkeley & Berkeley
i19521, which were based on material from Panama, Alaska and Canada
respectively. She placed most of her dextrally coiling species, moreover, under
the generic name Dexiospira Caullery & Mesnil, although that had been redefined
more precisely by Chamberlain ( 1919) according to the rules of nomenclature.
The dextrally coiling Spirorbis of California might possibly be grouped in
Spirorbella Chamberlain, following Rioja ( 1942),but in fact direction of coiling is
of little use in spirorbid taxonomy, 'as is amply confirmed by the species of
Spirorbis and Romanchella described here.
METH 0DS
The material was preserved in formalin or 70%alcohol and examined between
1971 and 1976. The animals were extracted from their tubes with dissecting
needles and the general morphology observed with a dissecting microscope in
reflected light whilst in preservative. The setae were viewed by phase contrast with
quartz iodide transmitted light after mounting the animal (minus operculum) in
polyvinyl - lactophenol.
Drawings were made on squared paper with the help of squared graticules in
the eyepiece, the largest setae and uncini of each type and species being selected
for illustration. Further details of techniques have been given earlier (KnightJones, Knight-Jones & Llewellyn, 1974).
NEW SPIRORBIDS FROM FOUR OCEANS
209
TAXONOMY OF SPIRORBIDAE
The classification used here is derived mainly from Chamberlain (19191, with
appropriate modifications (Knight-Jones, 1973; Knight-Jones, Knight-Jones &
Llewellyn, 1974; Knight-Jones, Knight-Jones 8c Kawahara, 1975). Genera are
defined on the balance of a considerable number of characters in combination,
for reasons previously discussed by Vine (1972a). The group is regarded as a
family distinct from Serpulidae, as suggested by Pillai (19701, which allows
related genera to be grouped into subfamilies. Several of these are here proposed
and it should be noted that the Spirorbinae are thus defined much more
narrowly than before.
FAMILY SPIRORBIDAE
Body asymmetrical and tube coiled in a spiral; thorax with setigerous rudiments of 3, 4, or 5 “segments” (Vine 1972a; Knight-Jones 1973); embryos
incubated in the tube or in an opercular brood chamber.
SUBFAMILY SPIRORBINAE
Embryos in an egg “string” which is attached posteriorly by a thread to the
inside of the tube wall; larvae with a single attachment gland; thoracic uncini
slender, usually with 3 or no more than 4 longitudinal rows of teeth for most of
their length and with a blunt anterior peg; numbers of abdominal uncini similar
on both sides of the body; abdominal setae pennant-shaped, usually with a thick
(optically dense) projecting “heel”.
Genus Spirorbis Daudin ( 1800)includes Spirorbella Chamberlain ( 19 19)
Coiling usually sinistral (p. 202); margins of thoracic membrane not fused
dorsaIly over thoracic groove; collar setae with gap (usually distinct) between
distal blade and proximal fin and blade with o r without cross striations; sickle
setae in third thoracic fascicles; only two pairs of thoracic tori ; largest abdominal
tori usually in anterior half of abdomen.
Type. Serpula spirorbis L. =Spirorbis borealis Daudin ( 1800).
Spirorbis bfurcatus sp. nov.
(Fig. 1)
Holotype. B.M. (N.H.) No. Z.B. 1976: 633aand 633.
Paratypes. B.M. (N.H.) No. Z.B. 1976: 634-50andU.S.N.M. No. 54358.
Tube about 1.75 mm in coil diameter, either sinistral or dextral, with the last
whorl frequently coiling upon the previous one. There are usually two
longitudinal ridges, one toward the periphery and one on the inner edge of the
whorl, making the tube somewhat quadrangular in cross section, but younger
specimens may lack these ridges.
Tentacles about seven, fairly long. Material examined a few weeks after
preservation was rather colourless, with a brown ochre stomach. Most specimens
were breeding, with up to twenty-four embryos in each egg-string.
Opercular plate slightly concave. Talon thick, blunt and with a proximal
bifurcate projection on the side away from the tentacles (Fig. 1D) in all but the
youngest specimens.
NEW SPIRORBIDS FROM FOUR OCEANS
205
This species is close to the mainly dextral species Spirorbis (Spirorbella) marioni
Caullery & Mesnil (1897), particularly to the “bu5hi” stage of opercular
development (Vine, Bailey-Brock & Straughan, 197 2). Such opercula lacking any
distal ornamentation are found in mature S. marioni at La Paz (Mexico), Canary
Islands and St Helena (pers. obs.), but the talon in these populations and in more
typical material (with distal projections) from Hawaii (Vine, Bailey-Brock &
Straughan, 19721, Galapagos (Bailey 8c Harris, 1968), Easter Island (Kohn &
Lloyd, 19731, Costa Rica (kindly loaned by Dr Straughan) and Panama
(U.S.N.M.) has no terminal bifurcation but only shallow irregularities. The
clearest difference between S. marioni and S. b$matus, however, involves the
abdominal setae, particularly the relative thickness of shafts and blades (cf. Fig.
lK,J). S. marioni, moreover, is a much smaller species (Fig. lM), with fewer
abdominal segments (18 compared with 24 in mature specimens). It has not yet
been found north of La Paz and may not overlap with S. bfurcatus
geographically.
S . cuneatus Gee (1964) from Europe is the only other Spirorbis species with a
large talon and cross-striated setae. I t differs from S. bfurcatus in that the talon
has a pointed terminal projection and two angular lateral flanges; the collar setae
fins are more streamlined; the abdominal segments are fewer (about 14, even
though the animal is almost as large); and the coiling never seems to be dextral.
Spirorbis rothlisbergi sp. nov.
(Fig. 2)
Holotype. B.M. (M.H.) N o . Z.B. 1976: 651aand 651.
Paratypes. B.M. (N.H.) No. Z.B. 1976: 652-77,&U.S.N.M. No. 54359.
Tube about 2 mm in coil diameter, usually dextral, coiled in one plane,
without longitudinal ridges and with each whorl round in cross section except
for a small peripheral flange adjoining the substratum (Fig. 2A).
Tentacles about 9, fairly short. Material received a few weeks after preservation
had thorax and abdomen pale green, stomach pale ochre, and ovaries and
embryos cream. Egg strings contain up to 70 embryos in about two rows.
Opercular plate slightly concave; talon large and tapering with no projections
other than shallow irregular bulges (Fig. 2C). Collar setae of both fascicles have
basal fin of coarse and medium sized teeth and distal blade with fine marginal
serrations. A few very indistinct oblique cross-striations are sometimes associated
with the most proximal marginal teeth. Capillary setae also present, making
about 10 setae in each collar fascicle. Second and third fascicles have the usual
simple setae, with sickle setae also present in the third. There are two pairs of
thoracic tori, the largest with up to 70 uncini. Longest uncini have about 4
longitudinal rows of teeth, the smallest about 5. These teeth may also be
arranged in transverse rows, particularly in the smaller uncini at the ventral end
of each torus. The anterior peg is blunt and gouge-shaped, which gives a
bidentate appearance viewed from certain angles.
Asetigerous region short, about three times the distance between the first and
second abdominal tori. These tori total about 20 pairs. The largest tori, each with
about 25 uncini, occur at about the 6th setiger. Abdominal setae have a thick
(optically dense) “heel” (Fig. 25) which projects beyond the line of the shaft, and
tapered blades (with coarse marginal serrations) that are unusually large for a
Spirorbis species, being longer and wider than the collar setae blades. These setae
206
P . KNICHT-JONES
Figurc 2. A - j . Spirorbrs rothlisbergi: A, tube; 8, whole animal dorsal view; C, D, operculum (face and
side view!; E, collar seta; F. sickle seta; G , largest thoracic uncinus; H, smallest thoracic uncinus; J ,
dxiorninal seta. K. Spirorbis bidentalus abdominal seta with accompanying hooked scta.
are either single or in pairs. Hooked capillary setae occur only in the terminal
<<
segments’’.
Remarks. This description is from material intermixed with Circeis armoricana St
Joseph (1894) on Egregia laevzgata cast ashore at La Jolla and referred to as S.
marioni by Rothlisberg (1974). It differs from S. marioni and S . bEfurcatus in lacking
distinct cross-striations on the collar setae and any ornamentation on the tube
and talon.
Spirorbis tridentatus (Levinsen, 1883) has similar collar setae on the convex side,
but those on the concave side differ in not having a gap between fin and blade.
The talon, moreover, is broad bearing distinct lateral protuberances.
Spirorbis bidentatus Bailey from the Galapagos (Bailey & Harris, 1968) differs in
being a much smaller species; its short abdominal setae blades have a very large
projecting heel and there are hooked capillaries in each abdominal fascicle (Fig.
2 K ) whilst the collar setae in both fascicles often have minimal gaps between fin
and blade.
Spirorbis spatulutus sp. nov.
(Fig. 3)
Holotype. B.M. (N.H.)No. Z.B. 1976: 678aand 678.
Paratypes. B.M. ( N . H . )No. Z.B. 1976: 679-99,&U.S.N.M. No. 54360.
Tube up to 2 mm in coil diameter, usually dextral, usually coiled in one plane,
round in cross section but with a narrow median ridge (Fig. 3A).
Tentacles about 8 , very short. Any colour that the material might have had was
NEW SPIRORBIDS FROM FOUR OCEANS
20 7
Figure 3. A-K. Spirorbis spatulatus: A, tube; B, whole animal dorsal view; C, egg string attached to
tube wall fragment; D, E, operculum (face and side view); F, collar seta; G, sickle seta; H , largest
thoracic uncinus; J , smallest thoracic uncinus; K, abdominal seta. M . Spirorbis inornutus abdominal
seta.
lost by long immersion in preservative. Embryos up to 50, in the usual “string”
attached to the tube (Fig. 3C). Mature embryos have a particularly opaque
attachment gland.
Opercular plate strongly concave, bearing an eccentric somewhat spatulate
talon with the terminal edge a little longer on the side nearest to the substratum.
Setae and uncini remarkably like those of S. rothlisbergz in form and distribution, but the abdominal setae blades are more slender (cf. Fig. 3K and 25). The
abdomen too is more attenuated (though the numbers of tori are similar), the
tentacles are shorter, whilst the talon and probably the tube appear to be fairly
distinctive.
Remarks. This material was collected by Prof. R. P. Dales (February 1951) on
Macrocystis pyrzfera from Punta Barda near Ensenada, Baja California, and from
help at La Jolla. A few specimens on a kelp fragment were also found by Dr
Straughan (September 1968) at Catalina Island and four sinistral specimens on a
small piece of algae at Santa Cruz Island by Mr Shisko. The species was
intermixed with Circeis armoricana and the new species of Pileolaria (p. 2 14) at most
locations.
The concave opercular plate and the outline of the talon are somewhat like
those of Spirorbis inornutus L’Hardy & Quihreux ( 1962) and S. corallinae de Silva 8c
Knight-Jones ( 1962), which are sibling species from northwest Europe (KnightJones), Knight-Jones & Al-Ogily, 197.5). The talons of those species, however,
differ from S. spatulatus in being thicker in side view and curved when viewed
from below, whilst the abdominal setae are of very different proportions (cf. Fig.
3M, K).
208
P. KNIGHT-JONES
Figurr 4 . .Spcrorbu xtrtgufus: A, tube; B, whole animal dorsal view; C, D, operculurn iside and face
\ i e w ~ l E,
; collar seta; F. largest thoracic uncinus; G, smallest thoiacic uncinus; H, sickle seta (blade
unnaturallv folded);J , abdominal seta.
Spirorbis sttigutus sp. nov.
(Fig. 4)
Holotype. B.M. (N.H.) No. Z.B. 1976: 700aand 700.
Paratypes. B.M. (N.H.) No. Z.B. 1976: 701-10.
'Tube sinistral, coil diameter about 1.5 mm, with two longitudinal ridges, one
nearly peripheral and one toward the inner edge of the whorl, giving the tube a
somewhat quadrangular cross-section (Fig. 4A).
Tentacles about seven, fairly long. Live material rather colourless, except for a
yellow ochre or light brown stomach. Only one specimen (of 10 extracted from
their tubes) was breeding, with 9 embryos in an egg string attached posteriorly to
the tube.
Opercular plate flat. Talon thick, pointed, unusually long and curving (in side
view) to conform with the tube (Fig. 4B, C).
Collar setae have an angular basal fin of large and small teeth, and a distal
blade in which the marginal serrations are mostly very fine but a little coarser
basally. At this position fine, very indistinct cross-striations may be seen with a
good microscope. Thoracic uncini have surface teeth arranged in rows which are
both longitudinal (about 4 in the largest) and transverse. Largest torus has about
70 of these uncini.
Asetigerous region 3 to 4 times distance between first and second abdominal
tori. There are about 11 pairs of such tori, the largest (with about 17 uncini)
occurring at about the 8th segment. The abdominal setae have shafts gently
widening distally, almost covering the proximal part of the blade. The blade is
tapered with medium sized marginal serrations. Other details in form and
distribution of setae and uncini similar to those of S . bfuricatus (p. 204).
Remarks. About 30 specimens were collected by Prof. Knight-Jones and myself
in February 1976 from rocks at depths of 25 and 30 m on the seaward side of
NEW SPIRORBIDS FROM FOUR OCEANS
209
Funchal Harbour wall. There were also 3 specimens at 15 m off Punta do
Espinbaco 2 k m west of Funchal. The first collection also contained one
specimen of Pileolaria clavus (Harris, 1968) and the latter abundant material of
Janua pagelwtecheri (Quatrefages, 1865). Competition with hydroids, bryozoans.
sponges and Filograna (Serpulidae)seemed to be intense.
S. strigatus differs from S . marioni, S. cuneatus and S . bfurcatus in not having
coarsely serrated and distinctly cross-striated collar setae. S. tridentatus differs in
that its collar setae on the concave side lack gaps between fin and blade, whilst
the surface teeth of its thoracic uncini are arranged in oblique and not strictly
transverse rows. Both S. bidentatus and S . rothlisbergi differ in having abdominal
setae with large bulbous heels and S. bidentatus further differs in usually having
collar setae (on both sides) with minimal gaps between fin and blade, and in
having hooked capillary setae in each abdominal fascicle.
S. strigatus differs from all the above species in that the largest abdominal torus
lies posteriorly, whilst the talon is very pointed and markedly longer than it is
wide (hence the specific name).
SUBFAMILY PILEOLARIINAE NOV
Incubation in an opercular brood chamber which is an invagination from the
exterior, so that the embryos are morphologically distal to the cuticle (seeThorp,
1975).The chamber can be used for more than one brood; larval attachment
gland single; thoracic uncini with a blunt anterior peg; numbers of abdominal
uncini similar on both sides of body; abdominal setae pennant-shaped, usually
with a thin (optically transparent) projecting “heel”.
Genus Pileolaria Claparede ( 1868)
Coiling sinistral; margins of thoracic membrane not fused over dorsal groove;
collar setae with a gap between distal blade and proximal fin; only two pairs of
thoracic tori.
Subgenus Pileolaria
Brood chamber almost enclosed, with embryos released by dilation of an
epithelial pore on the side next to the tentacles; rest of chamber wall is calcified
arid shaped like a helmet or cap; non-brooding (“primary”) opercula not
retained after brood chamber is fully formed; they have talons but these are not
peripheral; collar setae blades have coarse marginal serrations and distinct cross
striations; sickle setae present; largest thoracic uncini usually have single row of
teeth posteriorly; largest abdominal tori occur in posterior half of abdomen.
Type. Pileolaria militaris Clapari.de (1868).
Pileolaria alata sp. nov.
(Fig. 5)
Holotype. B.M. (N.H.) N0.Z.B. 1976: 711aand 711.
Paratypes. B.M. (N.H.). No. Z.B. 1976: 7 12-13 and in Rijksuniversiteit Utrecht.
Tube up to 1.8 mm in coil diameter. Whorls round in cross section, with
indistinct transverse growth rings and even vaguer suggestions of longitudinal
ridges.
The material had been long preserved, but crystalline patches in the thoracic
dorsal groove still showed some pigment (faded orange). These patches were
210
P. KNIGHT-JONES
Figuw ,5. /'rlcoiarra d a t a . A, tube; B, whole animal dorsal view; C, D, primary operculum (side and
h c v iieivsi: E. F, H , variable Ornamentation o n brooding opercula; G , diagram of approximate
ot Fviewed from above, dotted lines represent the probable direction of fusion of
rpine ;~i-i-angeinent
piired spiiirs to produce the distal wings in H ; J, side view of H ; K, collar seta; M , aickle seta; N,
el)doiriinal wta; P, lai-gest thoracic uncinus.
paired and diffuse. Abdominal cloak (see p. 221) lacking as in most spirorbids.
Tentacles short, about 7 .
Primary operculum seems to be unique among Pileolaria species in being
bilaterally asymmetrical, for the talon is eccentric and bears a single wing-like
blade on the side away from the substratum (Fig. 5 C , D). Distal plate concave
with a rim that dips sharply near the talon and also on the opposite side. Brood
chamber helmet shaped with variable ornamentation. There may be a double
row of tubercles o r spines encircling the distal area with sometimes additional
spines in the centre of this (Fig. 5 G ) . Radially adjacent pairs o f spines, one from
the inner and one from the outer circle may fuse together to form one (Fig. 5E)
o r more (Fig. 5H, J ) radiating wings. I t is possible that the tuberculate stage is an
early growth form and the radially winged crown a later stage.
Collar setae have basal toothed fins and long distal blades with coarse
serrations (and associated cross-striations), about 12 in the proximal half of each
blade. Capillary setae are associated with these making about 10 setae in each
fascicle. Second and third fascicles have each up to 13 setae, most of them of the
usual simple type, with sickle setae also present in the third fascicles. The largest
thoracic torus has about 70 uncini, each uncinus with a blunt somewhat gougeshaped anterior peg and a single row of teeth for most of its length (Fig. 5P).
NEW SPIRORBIDS FROM FOUR OCEANS
211
Asetigerous region about 3 times the distance between the first and second
abdominal tori. Largest such torus with about 15 uncini occurs at about the 13th
abdominal setiger. In all there are about 16 such setigers. Abdominal setae have
large blades with fine marginal serrations and thin (optically transparent) winglike angular heels, each projecting considerably beyond the line of the shaft
(Fig. 5N).
Remarks. Eight specimens (3 adult and 5 juvenile) were found on a vertical coral
reef 600 m west of Klein Curacao Lighthouse at a depth of 30 m (Stn 2103B
October 1970) intermixed with another new Pileolaria species (p. 222). P. alata
also occurs at Curacao. Four specimens were found at 10m depth on a
submarine iron buoy at Caracas Baai (Stn 1334 November 1955); a single
specimen at 13-16m on a reef at Playa Kalki (Stn 2037B May 1970); two
specimens at 12- 15 m on a vertical reef near the swimming pool at Bullenbai (Stn
2048D April 1970). At Bonaire there were four specimens at 13-15m on a
vertical reef at Santa Barbara near Hato (Stn 21 12GeJuly 1970).All this material
was collected and kindly forwarded by Dr ten Hove.
Remarks. The only species in this subgenus that has a talon other than peg-like
is Pileolaria militaris. This differs from P. alata in that the talon is bilaterally
symmetrical, with paired “wings” and a median spur projecting towards the
branchial crown. P. militaris further differs in having a peripheral rim to the
brood chamber, a single dense thoracic crystalline patch which turns black when
preserved, and very coarse teeth on the abdominal setae.
If juvenile specimens are not present, other species of Pileolaria sensu stricto that
have brood chambers with multiple distal spines may be distinguished from P .
data as follows :P. grandis Pillai (1970) from Ceylon has coarser marginal serrations on the
collar and abdominal setae and unusually numerous uncini (about 200) in the
largest thoracic torus, even though it is similar in size (Fig. 9M).
P. quusimilitaris (Bailey, 1970), which also occurs in the West Indies, is a much
smaller species with setae blades half the size. The collar setae have serrations that
are fewer, though just as large, and the pattern of opercular spines is random.
P. pseudoclavus Vine (1972a=P. semimilitaris Vine, 1972b, see p. 217) is also a
much smaller species occurring in the West Indies, but its abdominal setae have
“heels” that do not project beyond the line of the shaft (Fig. 10s) and the small
distal spines on its brood chamber are always peripheral (Fig. 6K).
The juveniles in each of these three species have the primary operculum with
the usual simple eccentric talon, vestigial in P. grandis (pers. obs.) and pin-like in
the other two.
Pileolaria dakarensis sp. nov.
(Fig. 6)
Holotype. B.M. (N.H.) No. Z.B. 1976: 714a.
Paratypes. B.M. (N.H.) No. Z.B. 1976: 715-20.
Tube very small (coil diameter up to 1.2 mm including a peripheral sloping
flange), with meandering growth rings and incipient longitudinal ridges.
Live material brightly coloured, tentacles (about 7 ) orange and thorax bright
red, with paired bright orange crystalline “patches” dorsally. Patch on concave
side large and oval and that on convex thin and diagonally elongated, following
the faecal groove around towards the ventral side and ending, like the thoracic
S3NOr-LH3INX ‘d
NEW SPIRORBIDS FROM FOUR OCEANS
213
rim, except that they are somewhat flattened distally. Those brood chambers are
also deeper (helmet-shaped)and the animals larger and more attenuated.
P. pseudoclauw Vine 1972 usually has a distal rim, but this is always papillate
and once again the chamber is deep (Fig. 6K). The abdominal setae differ in that
the heel does not project beyond the line of the shaft (Fig. 10s).
P.dakarensis seems to be closest to a new species from California (p. 214).
Pileolaria lateralis sp. nov.
(Fig. 7 )
Holotype. B.M. (N.H.) No. Z.B. 1976: 721aand 721.
Paratypes. B.M. (N.H.) No. Z.B. 1976: 722-30 8cU.S.N.M. No. 54361.
Tube sinistrally coiled, usually in one plane, up to 1.5 mm in coil diameter,
without longitudinal ridges, each whorl round in cross section and widening
gradually towards the aperture.
After 6 weeks in formalin a single small crystalline thoracic patch could be distinguished dorsally (Fig. 7B), but it was not conspicuously different from the
underlying orange pigment of the stomach. These pigments are probably
brighter in live material and the rest of the animal may also be pigmented. There
are about seven tentacles.
Primary opercular plate concave, with a well developed eccentric talon. Brood
chamber develops beneath, quickly forming a central dome, with a prominent
rim on the side away from the substratum. When fully developed the calcified
part bears from one to five spines randomly placed on the dome and is somewhat
helmet-shaped, but always bilaterally asymmetrical because of the lateral rim.
G\
H
Figure 7 . A-M. Pileolaria lateralis: A, tube; B, whole animal dorsal view; C, D, primary opercula still
attached to developing brood chambers; E, F, brooding operculum (face and side views); G , another
brood chamber; H, collar seta; J, largest thoracic uncinus; K, sickle seta; M, abdominal seta; N,
Pileolaria berkeleyanu brooding operculum (faceview).
214
P. KNIGHT-JONES
Collar setae have basal fins and blades with coarse serrations and associated
cross-striations. Abdominal setae blades also have a coarsely serrated margin
and an angular heel projecting beyond the line of the shaft (Fig. 7 m). Form and
distribution of other setae and uncini like those in P. alata, except that the
asetigerous region is a little shorter.
Remarks. More than 40 specimens were found intermixed with Janua
pugenstecheri on Sargassum at a location 15 miles east of La Paz, Baja California.
They were collected by Mr Shepherd in August 1975.
Two other species with bilaterally asymmetrical opercula are P. heteropoma
(Zibrowius, 1968) from the Mediterranean and P. berkeleyana (Rioja, 1942) from
Mexico. P . heteropoma differs in having only a single spine on the operculum, but
P. berkeleyana is usually similar in having a lateral rim. Rioja did not describe
sickle setae but as these are easily overlooked, and the type material no longer
exists it seems best to regard Bailey’s description of material with sickle setae
(Bailey & Harris, 1968) as representing Rioja’s species. Certainly her material
from the Galapagos agrees perfectly with material from Acapulco, Mexico,
examined by me.
P. berkeleyana differs from P. lateralis in that the rim (when present) forms the
most distal part of the operculum (cf. Fig. 7E, N ) ; the brood chamber never bears
spines; sickle setae are longer relative to the blades of the largest collar setae; and
the favoured substrata are rocks o r shells and never algae. Both species are found
at La Paz, distinctly separated by their different choice of substrata.
Pileolaria marginata sp. nov.
(Fig. 8 )
Holotype. B.M. (N.H.) No. Z.B. 1976: 731aand 731.
Paratypes. B.M. (N.H.) No. Z.B. 1976: 732-40, & U.S.N.M. No. 54362.
Tube up to 2 mm in coil diameter, usually coiled in one plane, either in a tight
whorl or evolute. The surface bears growth rings that may be faint or distinct,
meandering in such a way as to suggest incipient longitudinal ridges.
Tentacles about seven. N o distinct pigments could be seen in the preserved
material.
Primary operculum concave, with a small eccentric talon (Fig. 8D). Brood
chamber large, with up to 16 embryos, but the calcified part is no more than a
shallow rimmed cap. This is usually symmetrical although the rim is sometimes
vestigial on the side nearest to the sub-stratum (Fig. 8 G ) .
Collar setae blades (with marginal serrations and cross-striations) are
unusually small compared with the size of the animal. Abdominal setae blades
have very coarse marginal serrations and a thin obtuse wing-like heel that
extends an unusually long way down the shaft (Fig. 8M). The form and
distribution of other setae and uncini are similar to those of P. alata (p. 209) but
the asetigerous region is a little shorter.
Remarks. This species is described from material collected on an unidentified
piece of kelp from La Jolla. It agrees with abundant material on Macrocystis
pyr@ra from Todos Santos (near Ensenada, Baja California), both samples
collected by Prof. R. P. Dales, October 1950 and February 1951. Other material
was found at Catalina Island (California), Playas del COCOS
and Rincon (Costa
Rica) on pieces of kelp collected by Dr Dale Straughan (September 1969 and
November 1968). It also occurs on Dictyoneuropsis reticulata in Monterey Bay
NEW SPIRORBIDS FROM FOUR OCEANS
215
/
Figure 8 . Pzleolaria marginata: A, tube; B, whole animal dorsal view; C, D, primary operculum (face
and side views); E, F, typical brooding operculum (side and face views); G, atypical brooding
operculum showing more bilateral asymmetry; H, collar seta; J , largest thoracic uncinus; K, sickle
seta; M , abdominal seta.
(Hancock Museum 002547) and had previously been misidentified as Spirorbis
borealis. The species seems to be abundant at most locations and is frequently
intermixed with Circeis armoricana and Spirorbis spatulatus (p. 2 14).
P. margznata differs from P. moerchi sensu Rioja, P. moerchi semu Harris and P.
Pseudoclaws in the same characters as listed for P. dakarensis (p. 21 1).
P. dakarensis is very close to P. marginata. The former is a much smaller species,
however, but the collar setae blades are nevertheless bigger, the sickle setae are
more attenuated, and the asetigerous region is proportionately twice as long.
Pileolaria spinijii sp. nov.
(Fig. 9)
Holotype. B.M. (N.H.) No. ZB. 1976: 741aand 741.
Paratype. B.M. (N.H.) No. ZB. 1976: 742.
Tube up to 2 mm across coils, usually coiled in one plane with outer whorl
(round in cross section) widening rapidly toward the aperture. Surface
roughened by indistinct growth ringsand incipient ridging.
Tentacles about 6, fairly long. Material had been long preserved but a
crystalline dorsal patch in the posterior part of the thorax still showed a rich rust
pigment. This patch is rather dense and sausage-shaped on the concave side,
more diffuse in the centre of the dorsal groove and virtually absent on the convex
side. The colour in life is likely to be brighter (perhaps orange or red) and other
parts of the animal may also be pigmented.
Primary opercular plate very concave, with a short eccentric talon (Fig. 9C).
P. KNJGHT-JONES
216
I
D
,
/’
Figure 9. A-J. Pileoluriu spimfr: A, tube; B, whole animal dorsal view; C, primary operculuni; D, E,
brooding operculum (face and side views); F, collar seta; G, thoracic uncinus; H, sickle seta (from a
dissected paratvpe);J, abdominal seta; K, Pileoloria quasimilitaris abdominal seta; M , Pileolaria grandic
whole animal dorsal view.
Calcified part of brood chamber deep and helmet-shaped, with a random
distribution of distal spines. It may shelter up to 12 embryos.
Collar setae have basal fins, coarsely serrated blades and associated cross
striations. Sickle setae unremarkable except that the distal toothed part is
unusually thin and difficult to see. Abdominal setae blades (with coarse marginal
serrations) are carried on a shaft that widens distally but the “heel” projects only
slightly beyond the line of the shaft (Fig. 9J). Form and distribution of other setae
and uncini as in P . alata (p. 2 10).
Remarks. Four adult and three juvenile specimens were found intertidally on
limpets and stones 15 miles east of La P a , Baja California, by Mr Shepherd in
the summer of 1972.
P . spinver is similar in size to P . militaris but the latter difkrs in having a
complex juvenile talon (p. 2 1 1), a thoracic crystalline patch that turns black when
preserved and a distinct rim around the brood chamber.
P . grandis from Ceylon has a similar brood chamber and primary operculum,
but differs in having nearly four times the numbers of uncini (p. 000) though the
animal is a little smaller. The calcified area of the brood chamber is smaller; the
tentacles and opercular stalk are shorter (Fig. 9M) and the collar setae have
unusually coarse marginal serrations.
P . quasimilitaris from the West Indies has a similar brood chamber with random
spines, but it is a much smaller species and its small abdominal setae have a
relatively large thin “heel” projecting beyond the line of the shaft (Fig. 9K).
NEW SPIRORBIDS FROM FOUR OCEANS
217
P. alata markedly differs in having a winged primary talon and abdominal
setae blades with much finer marginal serrations.
P . lateralis is a sibling species, differing mainly in having an asymmetrical
operculum and abdominal setae with a large thin heel projecting beyond the line
of the shaft (Fig. 7M). P. lateralis and P. spinfer occur on different substrata on the
same shore. Specific separation seems justified, but further study is desirable.
P . spinrer was found intermixed with abundant P. berkeleyana, but separation of
both young and adult forms was made easy by the characteristic rust-coloured
“patch” of P. spinrefer. The pigments of P. berkeleyana had completely faded. To
judge from live material from the Canary Islands the patches of P. berkeleyana are
small, paired and glistening orange.
Pileohria tiarata sp. nov.
(Fig. 10)
Holotype. B.M. (N.H.) No. ZB. 1976: 743a and 743.
Paratypes. B.M. (N.H.) NO.ZB. 1976: 744-48.
Tube about 1.5 mm in coil diameter, sinistral, with the last whorl sometimes
coiled upon the previous one, or evolute, or ascending, with two to four
longitudinal ridges (Fig. 10A) positioned in such a way as to give a rather
quadrangular shape to the cross section of each whorl.
The material was very hardened by alcohol and the numbers of tentacles etc.
could not be ascertained.
Primary operculum more or less flat, with a blunt eccentric talon (Fig. 10H).
Brood chamber helmet-shaped, broadest in “face” view (Fig. 10E, G), with a
transverse undulating rim or row of spines forming a “tiara” on the side
adjoining the tentacles. Other distal spines randomly arranged and variable in
number. Embryos up to ten.
Collar setae with basal fin, coarse marginal serrations and associated crossstriations. Sickle setae attenuated, with long basal portions. Abdominal setae
blades have coarse marginal serrations and a thin heel projecting beyond the line
of the shaft (Fig. 1OP). Form and distribution of other setae and uncini as in P.
alata, except that asetigerous region is nearly twice as long.
Remarks. Five adults and two juveniles were found by Dr R. Rowe on a coralline
alga, Lithothrix aspergillum, intertidally at San Clemente Island, California.
P. militaris differs in several ways, the most diagnostic characters being its
complex primary talon (p. 2 1 1) and thoracic crystalline “patch” that turns black
in preservative.
P. pseudoclavus differs in usually having an encircling rim of spines and the
differences separating it from P. lateralis also separate it from P. tiarata. The
holotype of P. semimilitaris from Hawaii was closely compared with P. tiarata. The
setae, particularly those of the abdomen, are quite different (Fig. 10% R, S), but
are so characteristic of P. pseudoclavus that P. semimilitaris is likely to be a junior
synonym, particularly as P. pseudoclavus is now known to be widespread in warm
and temperate waters (pers. obs.).
P. grandis differs in several ways. It is characterized by having unusually large
thoracic tori (p. 2 11) and vely coarsely serrated collar setae.
P. dakarensis and P. margznata differ in having a shallow distal rim encircling the
brood chamber.
SBNO[-IH3INI ‘d
812
NEW SPIRORBIDS FROM FOUR OCEANS
219
Figure 11. A-M. PiLeolaria (Simplzcuria) potswaldi: A, aggregated tubes; B, whole animal dorsal view;
C, primary operculum; D, F, variation in brooding opercula; E, view from above of D; G, collar
seta; H, simple seta typical o f second and third fascicles; K, largest thoracic uncinus; M, abdominal
seta; N, P . PileolaTia rnoerchi collar seta and brooding operculum.
Tentacles about seven, bright red in a field photograph. Some preserved
material retained a little colour, most of the animal being pale salmon pink with
two crystalline thoracic patches of reddish brown. One patch is large, ovoid and
diffuse on the dorso-concave side and the other slender on the dorso-convex. In
other samples all colouration had disappeared. The anterior margin of the
thoracic collar covers about half the operculum (the opercular stalk being very
short) and the dorso-convex margin extends posteriorly to form an “abdominal
cloak”. This is attached to the abdomen along the convex side of the faecal
groove and it reaches to about the sixth abdominal segment. It would be best
seen in live material or animals preserved out of their tubes (Fig. 1 lB), but it can
still be demonstrated by probing with a mounted eyelash in animals that have
compressed themselves into their tubes.
15
“0
P. KNIGHT-JONES
Prirnarv operculum asymmetrically concave, with a small tubercle beneath
representing an eccentric talon. Calcified part of brood chamber helmet-shaped
hut relativelv shallow. The periphery of the operculum may be either smooth, or
marked by a’row of small tubercles (Fig. 1 ID) which may be reduced or absent on
the sides nearest the substratum and the tentacles. Any bilateral asymmetry is
slight. Centre of “helmet” smooth, but may have a shallow median groove giving
a bilobed appearance (Fig. 11F). Chamber may contain up to 30 embryos.
Collar setae have basal fins of large and fine teeth and distal blades with fairly
coarse serrations and associated cross-striations. Capillary setae also present,
rnaking up to 1 7 setae in each fascicle. Sickle setae absent. Largest thoracic torus
has about 75 uncini, each with a single row of unusually large teeth for most of
their length.
Asetigerous region very long, about seven times distance between first and
second abdominal tori. There are about 28 pairs of such tori, the largest, with
about 23 uncini, occurring between the tenth and eighteenth “segment”.
Abdominal setae have a thin “heel” (with a well-defined margin) projecting only
slightly beyond line of shaft (Fig. 11M).
Remarks. This description is based on specimens from a small population at
Piiios Boint, Monterey, which was intermixed with Spirorbis bfurcatus (p. 2041,
Paradexiospira vitrea and Sinistrella abnomis on the shaded sides of stones. The
material was collected by Prof. Dales (March and May 1951) and Dr Gilchrist
( J u h 1974). I t was also found by Prof. Dales in dense aggregations at nearby
Pacific Grove. Other material from the Monterey Peninsula in the British
Museum (1958 3:1:62 collected by Dr G. F. Parker) is also this species. I t is also
found at False Narrows, British Columbia (U.S.N.M. No. 41 129, 1938 identified
as Spirorbis moerchi Levinsen by E. & C. Berkeley, 1952) and it is clearly the species
referred to as S . moerchi by Pixel1 ( 19 12) from Departure Bay, Vancouver Island,
and Potswald (1968) from San Juan Island, Washington, for both those authors
referred to the remarkable posterior extension of the thoracic membrane.
Potswald (1967, 1968) also described various aspects of the biology of this
species.
The presence of P. moerchi (Levinsen 1883) at Alaska and Queen Charlotte
Island, as inferred by Bush (19041, cannot be discounted, but this species has not
occurred in collections examined by me. Levinsen’s material contains one
complete specimen from Greenland (now mounted and returned to the
Copenhagen Museum). Levinsen’s species basically differs from P. potswaldi in
having sickle setae. The brood chamber moreover, is deeper (Fig. 11P) and large
numbers of specimens from Iceland and Newfoundland show that it is usually
bilobed distally (Bush 1904 and pers. obs.); there is no posterior extension of the
thoracic membrane; the collar setae differ in having a smaller basal fin and larger
distal blades with remarkably well-defined cross-striations (Fig. 15N); the setae
are even more numerous in each collar fascicle (up to 201, but the thoracic uncini
are fewer and the asetigerous region shorter.
If we can regard P. berkeleyana (Rioja, 1942) as having sickle setae (see p. 214)
there are only two species beside P. potswaldi in the subgenus Simplicaria. The
best-known of these, P. pseudomilitaris, differs from P. potswaldi in having a
smaller brood chamber with a very pronounced lateral rim (or a lateral row of
spines); abdominal setae with smaller blades and coarser marginal serrations;
and no abdominal cloak extending posteriorly from the thoracic membrane. The
remaining species in the subgenus is P. (S.) ovata Vine (1977). It differs from P.
NEW SPIRORBIDS FROM FOUR OCEANS
32 1
potswaldi in having a remarkably deep, bilaterally symmetrical, distal rim nearly
encompassing the brood-chamber, very small collar setae and no abdominal
setae.
The “cloak” of P. potswaldi is unusual amongst Spirorbids, but a similar
structure occurs in two species of Sinistrella, a genus closely related to Pileolaria
(Knight-Jones, Knight-Jones & Kawahara, 1975).These are S. media Pixel1 (1912)
and S. coronata Zachs (1933; Imajima & Hartman, 1964).A “cloak” that is even
more extensive is described below (p. 223).
Subgenus Nidularia subgen. nov.
As Pileolaria sensu strict0 but the brood chamber is nest-shaped, either forming a
simple bowl (e.g. Fig. 13F) or like an American Hang-nest, with a roof and sideopening (Fig. 12D). The opening is not restricted by a soft-walled annulus, as in
Pileolaria, Simplicaria and a related subgenus (p. 227), nor covered by a specialized
plate as in Duplicaria. The embryos may protrude from it, but seem to be retained
securely by an adhesive matrix.
Type. Spirorbis clauus (Harris, 1968. For a fuller description see Bailey (1969a,
b); Mediterranean, Madeira & Canary Islands.
Pileolaria (Nidularia) nidica sp. nov.
(Fig. 12)
Holotype. B.M. (N.H.) No. ZB. 1976: 766a and 766.
Paratypes. B.M. (N.H.) No. ZB. 1976: 767 and in Rijkuniversiteit Utrecht.
Tube very small, with meandering growth rings and about 2 incipient ridges;
coil diameter up to 1.2 mm, including peripheral flange.
No colours could be seen in the preserved material but there is a single small
crystalline dorsal area in the thorax (with a dotted outline in Fig. 12B) which
appeared distinctly white and may never have been pigmented. Tentacles about
6.
Primary operculum slightly concave, with a pin talon. The entire brooding
operculum is densely calcified, but fragile, except for a large opening on the side
towards the branchial crown (Fig. 12E). There is a small proximal projection
near the stalk (Fig. 12D). One of the chambers contained two embryos but there
seemed to be room for about four.
Collar setae have basal fins and distal blades with coarse marginal serrations
and associated cross-striations. There are about 3 of these setae and 3 capillary
setae in each collar fascicle. The other thoracic fascicles are also unusually small
for a Pileolaria species. The sickle setae blades are twice the length of those of the
collar setae and the distal part has extremely fine teeth. The thoracic uncini
(largest torus with about 40) have blunt anterior pegs and are very thin with
extremely fine surface teeth at or beyond the limits of microscopical resolution.
There are, however, probably no more than two longitudinal rows of such teeth.
Asetigerous region about 4 times the distance between the first and second
abdominal tori. About 11 pairs of such tori, the largest with about 8 uncini
occurring at the 1st and 10th “segment”. Abdominal setae single. Their blades
have very fine marginal serrations and the heel does not project beyond the line
of the shaft (Fig. 12H).
Remarks. The above material (one adult and one juvenile) was found 600m
222
P. KNIGHT-JONES
1 2 . A-J. Prirolarin /.Vidularia) nidica: A, tube; B, whole animal dorbal view; C, primary
mature operculum; F, collar seta; G , outline of largest thoracic- uncinus; H ,
, i l ~ ( I o i i i i ~ i creta:
d
J . iickle seta; K-N. P. pseudoclavw; K, sickle sera; M , brooding operculum; N , whole
I i i i i n i r i l d o i sal view; P, Q. P. pseudoclouus var. lulparensis; P, prima? operculum still attached t o
1)i-oocI chain be^-: Q, brood chamber with a new chamber developing below.
I.ipii-c
~ p ~ i ~ i l ~ iD.
i i E,
i ;
west of Klein Curacao lighthouse on a vertical reef of dead and living coral at a
depth of 40 m (Stn 2103B October 1970), associated with Pileolaria alata. P. nidica
is also found at Curacao. Three specimens occurred on a vertical coral reef near
the swimming pool at Bullen baii, at 12-15 m, and one at 35-38 m (Stns 2048D
arid 2048Fa respectively April 19701, and three specimens at Salinja Fuik near
Ceru Preekstul from a vertical reef at 30 m (Stn 2088A September 1970).All this
material was collected by Dr H. ten Hove.
Pileolaria nidica is closely related to P. clauus (Harris, 1968; see also Bailey,
1969) but the latter differs in that the brood chamber is flattened distally, has no
basal projection and is nearly transparent with an opening about 0.75 the depth
of the operculum (Fig. 12M), whilst the sickle setae blades are smaller (similar in
size to those of the collar setae) with coarser serrations on the distal parts (Fig.
12K).
Material from Ilot Passerhina off Sio Vicente in the Cape Verde Islands
collected at 12-15 m by Dr J. Laborel (1974) may well be this species judging
from the operculum, but the animals were very hardened by preservative and the
setae (particularly the diagnostic sickle setae) could not be observed. One
operculum had about 3 embryos but there was room in the chamber for about 8.
Material from Tulear, Madagascar (collected by Dr J . Vacelet from coral
tunnels, Grand Recife), resembles P . nidica in opacity of the brood chamber and
smallness of size, but is closer to P . clavus in having ( 1) the operculum flattened
distally, (2) a relatively large brood chamber ‘‘0 ening” and (3) short sickle setae
with coarsely toothed distal margins. The Ma agascan material differs from P.
clauus in that the distal rim to the operculum is sometimes more clearly defined,
whilst the parapodial rudiments and overall size are smaller. These differences
seem unimportant, and it seems best at this stage to refer to this form as P.clavus
B
NEW SPIRORBIDS FROM FOUR OCEANS
223
vur. tuleurensis JB.M.(N.H.)ZB. 1976: 768a & 769-80). Several developmental
stages were observed in the Madagascan material which are likely to occur in the
closely related P. nidicu. The brood chamber develops beneath, and prior to the
loss of, the primary operculum (Fig. 12P). The distal part of the brood chamber
may have a transverse median scar, where the side of the primary talon was
recently attached. The primary operculum is probably soon lost, judging from
the fact that only one out of about 50 specimens had it attached to a brood
chamber. Subsequent chambers may develop below (without producing another
primary” operculum) and the early formation is cup-like (Fig. 12Q) as in other
species of the subgenus (see below). One specimen of the South African form P.
(P.) duyi Knight-Jones & Knight-Jones (1974) showed a similar cup-shaped stage
to the developing brood chamber and it must be admitted that its mature
operculum too is quite like those described here. P. duyi, however, is placed in
Pileolariu sensu strict0 because the non-calcified area of the operculum (an ovoid
“
opening’’ as wide and almost as long as the chamber) is covered by soft-walled
epithelium, with a central contractile opening. P. (P.)duyi thus links the two
subgenera Pileoluriu and Niduluriu but it has two characters atypical of either
subgenus, ( 1) the thoracic uncini have three or four rows of teeth for most of their
lengths and (2) the largest abdominal tori occur anteriorly.
(<
Pileolaria (Nidularia) palliata sp. nov.
(Fig. 13)
Holotype. B.M. (N.H.) No. ZB. 1976: 781a and 781.
Paratypes. B.M. (N.H.) No. ZB. 1976: 782-99 and in Zoology Department,
University of Cape Town.
Tube thick walled, coil diameter up to 2 mm but coiling may be irregular.
Often found in aggregations so that last whorl is forced to ascend. Surface of
tube rather uneven and without distinct ridges, though whorls may sometimes be
subquadrangular in cross section.
During life stomach is orange-brown, embryos orange and rest of animal
(including epithelium of operculum) orange-red. Two sausage-shaped crystalline
patches lie on each side of the thoracic dorsal groove. These are vivid orange in
life, vivid red when first preserved and rust-coloured after some months preservation, by which time other colours disappeared. The most characteristic
feature of this species is that the margin of the thoracic collar on the convex side
extends throughout the abdomen like a “cloak” (Fig. 13B & C), hence specific
name proposed. Only one specimen (a small one) was found with a reduced
“cloak”, and that reached as far as the third abdominal “segment”. This
character is not very obvious in animals which have been preserved in their tubes,
but it can still be demonstrated by probing with a mounted eyelash. Tentacles
(about 7 ) and opercular stalk are very short. The latter is about one third the
depth of the operculum.
In young specimens opercuium is concave, with a small eccentric talon.
Mature operculum is funnel-shaped, with walls that are thin distally but thick
and opaque at proximal apex. Embryos (10-20) incubated within the funnel,
secured by what seems to be hardened mucus. They lie distally to the cuticle,
within the opercular funnel (Thorp, 1975: fig. 5). One specimen had one
operculum inside another, the apex of the distal funnel being partially attached
P. KNIGHT-JONES
I.igtiw 13. &I. Pdrolorui (,\du/arm) palhafa; A, tube; B , C, whole anirnal ventral and dorsal views
\liowiiig ”< I o A ” n n d paired pigmented patches: D. E , young operculum; F, G , brooding opercula;
t1, i i e ~
o p ( ~ ~ c u l u \\.irIi
n i the old operculurn “delaminating” distal[!; J, collar seta, with a dotted line
iiitilcariiig \trap(. ot t i n i i i P (.\./ porillator (see PI; K, sickle seta; M , largest thoracic uncinus; N,
, i l ) ( l o i i i i r r ~ i l x u : P, [’. \’.i p o n N d o r tloi-sal vie\\ ot wholr animal showing ringlc pignrwt patch rind
i i o
“(
IOdk“.
t o the one beneath (Fig. 13Hi. This seems to have resulted from a moulting
process like that described for Spirorbis spirorbis (Thorp & Segrove, 1975).
Collar setae have fairly narrow basal fins, and blades with a -coarsely serrated
niargin and associated cross-striations. Capillary setae are associated with these,
making about 10 setae per fascicle. Sickle setae have basal and distal portions of
siiiiilar length (Fig. 1 3 K ) . Each of the larger thoracic uncini has a blunt anterior
peg and a single r o ~of’ teeth for most of length. Largest torus contains about 55
of these uncini.
Asetigerous region about 4 times distance between first and second torus.
Largest abdominal tori (about 15 uncinii may occur within a tairl
bet\veen fifth and twelfth “segments”. About 18 abdoniinal setige
setae usually paired and rather coarsely serrated, with thin angular wing-like
hc.els projeciing beyond line of shaft (Fig. 13N).
Remarks. The type locality Sunny Cove, False Bay, South Africa, where it occurs
abundantly on Fasciolaria at 13 m depth, in association wirh Protolaeospira capensis
iDav, 1961). I t is also found intertidally in crevice situations formed by stones
and‘ mussels, where it is associated with Paralaeospira patagonica and Paralaeospira
leuinseni (both Caullery & Mesnil, 1897-see also Knight-Jones, Knight-Jones &
Al-Ogilv, 1974). It occurs on most rocky shores from Saldanha Bay (north of
NEW SPIRORBIDS FROM FOUR OCEANS
225
Cape Town) to Shoemakerskop near Port Elizabeth (as species 9, Knight-Jones 8c
Knight-Jones, 1974). The material was collected by E. W. Knight-Jones and
myself in 1971. This species was also found by the Galathea Expedition at
Simonstown, False Bay (Stn 172, 1951, material in Zool. Museum, Copenhagen).
P. palliata is similar in size and most characters to P. pocillator Vine ( 1977). The
latter, however, differs in not having an abdominal “cloak” (Fig. 13P); the
operculum seems always to bear a small protuberance near the apex of the
funnel; the collar setae have much broader basal fins (indicated by the dotted line
in Fig. 135); the sickle setae have a longer basal untoothed portion; the
abdominal setae have a more rounded projecting heel and the thorax has a
single, large dense, milky-orange crystalline patch (Fig. 13P-colours observed
ten days after preservation, in material collected from Kaikoura, New Zealand,
by Dr M. Jones).
P. dalestraughanae Vine (1972b) differs in being a much smaller species with
most parapodial rudiments smaller in numbers and size. The small operculum is
more deeply cup-shaped and is borne on a very long opercular stalk, its length
two or three times the depth of the brood chamber.
The abyssal species P. levensteinae Bailey-Brock 8c Knight-Jones ( 197 7 ) is much
larger than P . palliata, though the parapodial rudiments are similar in size and
form. The operculum is also more cup-shaped and the opercular stalk more than
twice the depth of the operculum.
Material of “Spirorbis” langerhansi Caullery 8c Mesnil (1897) no longer exists.
The description could apply to any of the fore-going species and this should
probably be regarded as a nomen nudum (Bailey-Brock 8c Knight-Jones, 1977).
The abdominal “cloak” of P. palliata is more extensive than those described
for P. potswaldi, Sinistrella media and S . coronata (p. 2211, which end at about the
6th abdominal setment.
Subgenus Duplicaria Vine (1972a)
As Pileolaria sensu stricto, except that the brood chamber consists of two parts,
an upper unit with a thin proximal wall (somewhat helmet-shaped) and a lower
cup-shaped unit, the junction of the two being the line of dehiscence for embryo
release; one or more earlier opercular plates, each with a peripheral talon, may
be retained distal to the brood chamber; the thoracic uncini have more than a
single row of teeth (2 to 6) for most of their length; and collar setae blades lack
cross striations.
Type. Spirorbis koehleri Caullery 8c Mesnil( 18971, a species which is widespread in
warm seas.
Pileolaria (Duplicaria) zibrowii sp. nov.
(Fig. 14)
Holotype. B.M. (N.H.) No. ZB. 1976: 800a.
Paratypes. B.M. (N.H.) No. ZB. 1976: 801-25.
Adult tube one of the smallest found in spirorbids, up to 0.7 mm in coil
diameter. There are usually three longitudinal ridges that may be distinct
(roughened with small protuberances) or vestigial. The median ridge is the most
prominent, sometimes giving the whorl a subtriangular cross-section which is
emphasised by a sloping peripheral flange. Colours and numbers of tentacles
could not be determined in these very small preserved specimens. Opercular
stalk quite long, exceeding length of operculum itself.
226
P KhICHT-JONES
1
'
-~
/
i
I
/
1.igur.r 11. X-P. hlf*olann(Duplican'a) ribmm'i: A, tube; B, whole animal dorsal view; C, priinarv
o p e n u l u l n ; D, E, br-ood chamber without embryos; F, brood chamber with embiyos; G. inside of
tlic (op half of'an einpt) brood chamber viewed partly from below (after Zibrowius); H, collar seta
11-omcoiivcx \ide ofaiiirnal;J , K, collar seta fr-om the concave side; M , sickle seta; N , largrst thoracic
uiicitiu\; P. dbdoininal seta; Q. R. P. (D.)
koehleri primary and mature operrula rcsprctivrly; S, T. P.
i l j . i r r h c m m s r ~priinar-\ and tnatut-e opercula respectively.
Primary operculum a flat and relatively thick plate, with a small peripheral
protuberance (talon)on the side away from the branchial crown. This operculum
is retained after the formation of the brood chamber and probably throughout
the brooding cycle. I t is keyed into the top edge of the chamber but is easily
broken away. The top half of the chamber is somewhat helmet-shaped and keys
similarly into a basal unit which is like a spoon or rounded shovel. This has a
thickened protuberance at its proximal apex. These two units hold two to four
embryos and the line of dehiscence between them is oblique but curved. Collar
setae o n convex side have a basal "fin" differentiated from the distal blade by its
slightly larger teeth (Fig. 14H) but there is virtually no gap between the basal and
distal parts. Collar setae blades on the concave side have no such differentiation
arid can certainly be regarded as simple setae (Fig. 145 & K). Neither type of seta
has cross-striations, There are only two or three such setae in each collar fascicle,
accompanied by two or three capillary setae. Second and third fascicles have the
usual simple setae, but these are remarkably short and geniculate like the collar
setae on the concave side (e.g. Fig. 14K).Sickle setae in the third fascicles are also
unusually short, but otherwise typical, with coarse marginal teeth distally (Fig.
14Mi. Thoracic tori small, the largest with about 25 uncini, each uncinus being
unusuallv broad (for a Pileoluria species) with 5 or 6 longitudinal rows of teeth
arid a blunt anterior peg (Fig. 14N).
Asetigerous region appears to be very long compared with the length of the
animal but this is associated with the unusual shortness of the abdomen, which
has only about 4 setigers. Numbers of uncini per torus in one specimen are
4,4,9and 12 on the concave side and 2 , 3 , 7 and 9 on the convex. The largest
tori are therefore unusual in occurring at the posterior end. Abdominal setae,
which occur singly, have a small heel scarcely projecting beyond the line of the
shaft.
NEW SPIRORBIDS FROM FOUR OCEANS
227
Remarks. Found commonly at Grand Recif, Tulear, south-west Madagascar, in
coral tunnels at 3-5m. The material was collected by Dr J. Vacelet and
recognized as being an unusual species by Dr H. Zibrowius, who kindly supplied
the material, with notes and preliminary drawings.
P . zibrowii is closest to P. koehleri=P. bernardi Caullery 8c Mesnil, 1897=P.
gelasinfera Pillai, 1970 (pers. obs.)=P. poljoperculata (Straughan, 1969-see Vine,
Bailey-Brock and Straughan, 1972). P. koehleri differs in that the primary
operculurn has a large talon and a concave plate (Fig. 14Q); there are normally
more than one of these opercula retained above the brood chamber, and the
latter’s line of dehiscence is not markedly curved (Fig. 14R); collar setae of both
fascicles have a gap between a coarsely-toothed basal fin and an elongated distal
blade; thoracic uncini are narrower with only 2-3 rows of teeth; sickle setae are
twice as long but with smaller marginal teeth; and abdominal setigers are nearly
three times as many.
P. endoumensis (Zibrowius, 1968) differs from P. zibrowii (and P. koehleri) in
having a primary operculum with a brood but fairly shallow talon (Fig. 14s)
which does not seem to be retained after the full development of the brood
chamber. The upper brood chamber unit, moreover, has a deeper proximal wall
with the lower margin transverse to the axis of the chamber (Fig. 14T).
SubgenusJugaria subgen. nov.
As Pileolaria sensu strict0 except that the “primary” operculum has a peripheral
flattened talon and is not naturally shed after the brood chamber is formed
beneath it, but remains as an additional protection. Thoracic uncini have more
than a single row of teeth for most of their length; collar setae lack distinct cross
striations; largest abdominal tori can occur either anteriorly o r posteriorly.
Type. Spirorbis quadrangularis Stimpson ( 1854)= Spirorbis affinis Levinsen and
Spirorbis carinatus semu Levinsen 1883 (pers. obs.)= Laeospira pacifica Uchida
(1971). This is holarctic and north temperate, but not recorded from Europe,
except Trondheim (pers. obs.) Spirorbis granulatus L. in the sense used by most
authors is a close relative but distinct from the above (pers. obs.) and confined to
Europe. It has suffered some nomenclatural confusion, so is not used as the type.
Pileolaria (Jugaria) atlantica sp. nov.
(Fig. 15)
Holotype. B.M. (N.H.) No. ZB. 1976: 826a and 826.
Paratypes. B.M. 1N.H.) No. ZB. 1976: 827-50 and in the National Museum of
Canada.
Coil diameter up to 1.8 mm, including a peripheral flange. Tube may coil in
one plane or with last whorl ascending and may have a median ridge.
Material was preserved, hardened and colourless. Body is very attenuated with
an opercular stalk about 3 times as long as the operculum and proportionately
long tentacles (Fig. 15B).
Primary operculum with a flat opercular plate and a long flat peripheral talon
with rounded lateral “wings”. This operculum becomes wholly fused to the
brood chamber which develops beneath. Behind the talon the chamber has a
helmet-shaped calcification that is so transparent that it can be distinguished
only at certain angles of illumination and by its relative stiffness compared with
P KNIGHT-JONES
the soft epithelium of the remainder of the chamber. The chamber is very
narrow, holding up to three embryos one beneath the other (Fig. 15D).
Collar setae have large streamlined fins and distal blade with fine marginal
serrations and very close cross-striations (Fig. 15F). Capillaty setae are associated
with these itiaking about 8 setae per fascicle. Sickle setae have basal and distal
parts of similar length (Fig. 15C). Thoracic uncini have gouge-shaped anterior
pegs arid about two rows of teeth for most of their length (Fig. 15H). The largest
torus has about 55 uncini.
Asetigerous region long, about 5 times distance between first and second
abdotninal tori. There are 11 pairs of these tori, the largest with about 10 uncini
occurring at about the 8th “segment”. The abdominal setae (one or two per
fascicle) have blades with fine serrations. The heel does not project beyond the
line of the shaft (Fig. 15J).
Remarks. Over 40 specimens were found o n coral (Solenosmilia uariabilis) dredged
from the mid Atlantic ridge 4 0 0 miles n o r t h of the Azores (Stn
A266-16 45’12, 2”; 28O54, 7’W). The depth was not recorded but S. uariabilis
was also found at a nearby station (45O 14, 2”; 28O55, 2’W) at a depth of 1800 m.
The material (Nat. Mus. of Canada) was collected by J . W. Scoggen and L. M.
Marhue in August 1966 abroad “C.S.S. Hudson” from the Bedford
Oceanographic Institute, Nova Scotia and kindly forwarded by Dr Zibrowius.
P. atlantica is much smaller and more attenuated than any other member of
this subgenus, and it seems to be the only one with cross-striations o n the collar
setae blades. Cross-striations as fine as these can be difficult to see with most
microscopes, however, so P. atlantica is further compared with the other species
as follows :P. quadrangulan‘s (holarctic) has a similar “primary” talon but differs in that the
distal plate is usually domed and the brood chamber with which it is wholly fused
has a much larger opaque calcified area; the largest abdominal tori are towards
NEW SPIRORBIDS FROM FOUR OCEANS
229
the anterior end and the thoracic uncini are unusually large, longer than the
blades of the collar setae.
P. granulata, from the northeast Atlantic, is similar to P. atlantica in having the
largest abdominal tori towards the posterior end, but the primary operculum
differs in being attached to the brood chamber only at the end of the talon,
which is small and bilobed or fan-shaped (Fig. 15K).
P. stimpsoni (Verrill, 1879, north-west Atlantic) differs in having the largest
abdominal tori towards the anterior end; collar setae with virtually no gap
between fin and blade; and the “primary” operculum, like that of P . granulata,
attached to the brood chamber by a short fan-shaped talon. The brood chamber
is elongated with a shallower area of opaque calcification.
P. similis (Bush, 1904, north-east Pacific) is similar in having the primary
operculum wholly fused to the brood chamber but differs in that the talon is fanshaped; the brood chamber is very wide and the calcification opaque; hooked
capillary setae occur in each abdominal fascicle; and the largest abdominal tori
are towards the anterior end.
SUBFAMILY ROMANCHELLINAE NOV.
Embryos incubated within the tube in an egg “sac” attached anteriorly to a
specialised process arising at the posterior end of thoracic dorsal groove; larvae
usually without a white attachment gland; thoracic uncini with a blunt anterior
peg; abdominal tori asymmetrical in size and distribution (vestigial on the
convex side).
Genus Romanchella Caullery & Mesnil(1897)
Coiling usually sinistral; margins of thoracic membrane fused dorsally to form
a tunnel over dorsal groove; collar setae simple (without a basal fin); sickle setae
present in third thoracic fascicles; only two pairs of thoracic tori.
Type. Spirorbis (Romanchella) perrieri Caullery 8c Mesnil (18971, which is
widespread in the Southern Ocean.
Romanchella bicava sp. nov.
(Fig. 16)
Holotype. B.M. (N.H.) No. ZB. 1976: 85laand 851.
Paratypes. B.M. (N.H.) No. ZB. 1976: 852-70.
Tube dextral, up to 1.8 mm in coil diameter, thick-walled and with three
distinct longitudinal ridges.
Colours could not be determined in the preserved material; tentacles about 6.
None of the specimens are with embryos but, a long embryo attachment “stalk”
is clearly visible (Fig. 16B).
Opercular plate lies almost at right-angles to stalk and frequently bears
previous partially delaminated flat plates which can give the operculum a stacked
or domed appearance (Fig. 16E).Talon beneath is thick and rather tooth-like in
side view, but on the side away from the branchial crown it bears two very
distinct depressions, like eye sockets separated by a nose (Fig. 16D).
Collar setae blades are simple and small. Their shafts are about half the length
of those of the other thoracic fascicles. Capillary setae are also present making
about 10 in each fascicle. The second and third fascicles each contain about 25
and 20 setae respectively. The third fascicle includes some sickle setae which have
230
P. KNIGHT-JONES
Figure 16. A-J. Romancheh bicava. A, tube; B, whole animal, dorsal view showing e n i h r y attiirhnieiit
process; C, D, nvo views of same operculurn; E, operculum with a thickened distal plate (dotted line
iiidicates niaxiinuin doming observed); F, collar seta; G, largest thoracic unrinus; H , sickle seta
Ilkon1 a paratype);J , abdominal seta with accompanying hooked capillan'; K-N. R. rcorestpi; K, M,
two views of operculum; N,largest thoracic uncinus.
the coarsely toothed portion longer than the unserrated basal part. Thoracic
uncini long (the largest about the same size as the collar setae blades) and
slender, with about 3 to 4 longitudinal rows of teeth for most of the length.
Smallest more ventral uncini have about 5 rows of teeth. Anterior peg is blunt
and gouge-shaped, which gives a bidentate appearance at certain angles. Largest
torus contains about 80 uncini.
Asetigerous region short, about three times distance between first and second
abdominal tori. The largest of these, with about 35 uncini, occurs at about the
sixth setiger, there being about 16 in all. Each abdominal seta has a blade with
coarse marginal serrations and a shaft that widens to give an optically dense
bulbous heel. These setae occur mostly in groups of two and three (sometimes
singly), but anteriorly o n the convex side the setae can number 10 in one fascicle.
Each fascicle contains a single hooked capillary.
Remarks. Most of the tubes had been bored by predators and were empty
except for opercula, but they seemed to be very abundant. They were found on
small dredged stones from Amsterdam Island (30 m Station B1) and St. Pauls
Island (50rn Station D3) in the Indian Ocean by Dr J . Beurois between
December, 197 1 and January, 1972 (Beurois, 1975).
All the specimens were dextral, which is the first record of such coiling within
this genus. As there are several instances of sztus inuersus coiling in other genera,
close comparisons were made with other sinistral relatives, particularly R.
scoresbyi (Harris, 1968) which also has a substantial talon. The operculum of the
latter differs in having a very oblique slightly concave distal plate. The talon is
less thick (in side view) and bears several protuberances o n each side (Fig. 16K,
MI. Although there are subterminal depressions, these are vestigial and separated
NEW SPIRORBIDS FROM FOUR OCEANS
23 I
by a broad ridge. The parapodial rudiments are similar except that the
abdominal setae do not exceed 2 per fascicle, the thoracic uncini have twice the
numbers of longitudinal rows of teeth (Fig. 16N)and the asetigerous region is up
to 3 times as long.
Other species of Romanchella differ in having either a small peripheral talon, as
in R. inuentis (Harris 1969) and R. quadricostalis Knight-Jones (19731, or a long
thin talon as in R. perrieri (Caullery & Mesnil, 1897).
R. bicaua is superficially similar to Paradexiospira uiolacea (Levinsen, 1883),
particularly in opercular form and direction of coiling. Young specimens of the
latter frequently have only two pairs of thoracic tori and if the method of
incubation cannot be established the two forms could be confused. P. uiolacea
however lacks simple setae in the third fascicles and is not known outside boreal
waters.
Romanchella pustulata sp. nov.
(Fig. 17)
Holotype. B.M. (N.H.) No. ZB. 1976: 871aand 871.
Paratypes. B.M. (N.H.) No. ZB. 1976: 872-82.
Tube up to 4 mm in coil diameter with thick walls, frequently bearing a welldefined median ridge or even three ridges, but may be fairly smooth except for
indistinct transverse growth rings.
Natural colouring could not be judged from preserved material. Anterior
margin of the thoracic membrane covers most of the lengths of the tentacles,
which are about 6 in number. Dorsal membrane thin, easily torn and fused for
only about half the length of the thorax. In brooding specimens, the egg “sac”,
of about 160 embryos, is held anteriorly by a very short attachment stalk.
Figure 1 7 . A-J. Romanchella pustulata: A, tube; B, C, D, different views of same operculum; E, collar
m a ; F, sickle seta; G, largest thoracic uncinus; H, abdominal seta with accompanying hooked
capillary; J, whole animal dorsal view showing very small embryo attachment process with embryo
sac detached (arrows denote position of attachment); K. R . scoresbyi whole animal dorsal view.
932
P. KNIGHT-JONES
Operculum with a very oblique concave plate. Talon thick, with two
depressions o n the side away from the branchial crown bordered by many
pustules that are seen clearly only in clove oil. These depressions are separated by
a ridge which bears a cone-shaped terminal hole (Fig. 17C). This forms the
attachment area for the fibres which pass through the opercular stalk, anchoring
the operculum to the thoracic uncini.
Thoracic setae similar in shape, size and number to those of R . bicava, except
that the sickle setae have longer basal portions and shorter distal parts. Thoracic
uncini very large (largest considerably longer than the blades of the collar setae)
with up to 4 longitudinal rows of teeth and a blunt gouge-shaped anterior peg.
Largest torus contains about 100 uncini.
Asetigerous region short and abdominal uncini similar in distribution to those
of R. bicaua except that the largest torus (the sixth) has 50 to 60 uncini and there
are about 24 “segments” in all. Abdominal setae occur in groups of not more
than 3 ; the blades are larger and the “heel” thin but optically dense at the outer
edge (Fig. 17H).
Remarks. Five specimens occurred in the type locality Caleta Leandro,
Talcahuano, Concepcion, Chile, three of them on the decapod Taliepus dentatus
and one each on Aulacomya ater and Semimytilus algosus. Twenty more were Sound
at Castro, Chiloe also o n Aulacomya ater; two were o n the gastropod Concholepas
concholepa at Isla Quiriquina, Talcahuano; two on a limb of Homalaspis plana
from San Vincente. This material was collected between 1972 and 1974 and
forwarded by Dr Maria Lopez. Twelve more were found by Mr Random Dubois
in the Bay of Herradura near Coquimbo (U.S.N.M. Reg. No. 53672). R . pustulata
may be found in association with Paralaeospira levinseni Caullery 8c Mesnil.
R . scoresbyi Harris has an operculum like that of R . pustulata but it is more
slender in side view, with a wider median “rib”. The shallow depressions on each
side o f the “rib” are not bordered with pustules. R . scoresbyi furthermore has the
asetiger-ous region up to three times as long (even though it is a much smaller
species); the embryo attachment stalk is longer (Fig. 17K);most of the parapodial
rudiments have half as many uncini though similar numbers of setae; the
thoracic uncini are shorter and wider with about eight longitudinal rows of
teeth; arid the sickle setae are smaller but with proportionately longer distal
toothed portions.
K. bicaua is also closely related, but the sickle setae have different proportions;
the opercular plate is fiat and sometimes stacked; the talon is without pustules or
kriobs; the overall size is much less; and the coiling is dextral.
The very reduced size of the embryo attachment stalk recalls that of R .
quadricostalis. A section of the latter showed that this process was the extended
edge of ail oviducal pore (Knight-Jones, Knight-Jones & Vine, 1972). R.
quadricostalis differs however in having a vestigial peripheral talon and thoracic
uncini with numerous longitudinal rows of teeth.
Sjirorbis chilensis Gay ( 1849), recorded from San Carlos, Chiloe, Chile on shells
a n t i stones, was clearly a very different species, for the brief description mentions
that the opercul um was funnel - shaped.
Genus Helicosiphon Gravier 1907
Coiling sinistral; incubation in an egg “sac” attached anteriorly to a
bpecialised process arising in the thoracic dorsal groove; larvae seem to be
NEW SPIRORBIDS FROM FOUR OCEANS
23.3
without a white attachment gland; margins of thoracic membrane not fused
dorsally; collar setae simple (without a basal fin); sickle setae absent; three
thoracic tori on the concave side and two on the convex; largest abdominal tori
are in anterior half of the abdomen.
Type. Helicosiphon biscoensis Gravier 1907. Graham Land, South Shetlands
(Monro, 19301, South Georgia (Hartman, 1953) and South Orkneys (KnightJones, Knight-Jones & Bregazzi, 1973).
Helicosiphonplatyspira sp. nov.
(Fig. 18)
Holotype. B.M. (N.H.) No. ZB. 1976: 883a and 883.
Paratypes. B.M. (N.H.)No. ZB. 1976: 884-99.
Tube very large, about 5 mm in coil diameter when coiled in one plane, but
frequently with the last whorl coiled upon the previous one or irregularly coiled
against the substratum. No characteristic markings other than the usual growth
rings.
Colours during life not recorded. Tentacles about 15, fairly long. Egg “sac”
which may contain about 90 embryos, is attached anteriorly in the thoracic
dorsal groove near the bases of the second and third fascicles on the convex side.
The attachment process is very small.
Opercular plate very oblique and slightly concave, with a peripheral subquadrangular talon that is curved and thin in side view (Fig. 18C).
All the thoracic fascicles large, comprising only straight, slender, simple setae
with accompanying capillary setae. In the first, second and third “segments”
each fascicle contains about 25,35 and 35 setae respectively. Thoracic tori have
Figure 18. Helicosiphon platypirat A, B, tubes; C, D, operculum from different viewpoints; E, whole
animal dorsal view showing vestigial embryo attachment process and embryo “sac” detached; F,
collar seta; G, largest thoracic uncinus; H, detail of anterior part of uncinus; J, abdominal seta; K,
surface details of J .
P. K S I C H T - J O N E S
234
nunierous uncini, the largest (with about 165) being the middle of’the three o n
the concave side. Each uncinus bears two longitudinal tows of‘teeth for most of
its length and has a blunt gouged-shaped anterior peg.
Asctigerous region remarkably short, less than half distance between first and
second abdominal tori. These tori are well spaced anteriorly but crowded
tobxm-ds the posterior end, there being about 40 setigers. Most of the tori are very
large, the largest with up to 160 uncini occurring at approximately the fifth
segment o n the concave side. Tori on the convex side vestigial and very few.
Al,tioininal setae difficult to see as their shafts are short compared with girth 0 1
aI)tloiiien, and setae blades tend to show an edge \riew rather than the usual
prolilc. There are at least 5 of these setae in some anterior fascicles. The 1)lacles
arc rrlativelv short but with very thick marginal teeth. Hooked capillary setae
j Fig. 1 8 K ) also occur in the last 8 “segments”.
Remarks. This material was found at Marion Island (one of the Prince Edward
Islands, southeast of Port Elizabeth) in Februan and March 1973. I t occurs at
low water springs O R sheltered rocky surfaces. There were about 40 specimens at
Ships Cove (SC. 1) and about 30 at Transvaal Cove (Z. 36). Thc specimens
occurred in collections from “The third biological expedition” to the Island
rnadc b\- Mr A. F. de Villiers (in press). The species was also obtained at
Observatorv Bay, Kerguelen, but was wrongly identified as Spirorbls perrieri bv
Ehlei-s (19 13) and as Helicosiphon biscoensis by Knight-Jones, Knight-Jones &
Brcgazzi (1972). This inaterial (now Berlin Museum Reg. no. 5940) comprises
about I 5 animals, mostly out of their tube, manv incomplete and all without
attached einbrvos, but with two loose egg “sacs” amongst them.
H. biscoensis‘Gravier is the only other species recorded within the genus. I t
differs from H . platyspira in being nearly twice the size; in having relatively short
tentacles but a long embryo-attachment process; the opercular plate more or less
at right angles to the peduncle; the talon larger and set in a less peripheral
position; setae fewer in the thoracic fascicles particularly at the collar ( I 5 each);
and tori markedly smaller in the abdomen (thoracic tori are similar). The tube
also differs in being helically coiled away from the substratum, but to judge from
Pileolariu leuensteinue Bailey-Brack & Knight-Jones (1977) this is likely to be an
ecophenotypic response rather than a specific character. It is likely that H.
hiscoensis does not occur on the shore as all records have been between 60 and
200 111.
The verv small process for embryo-attachment in H . platyspira recalls the
similarly vestigial structure in Romanchella quadricostalis and R. pustulata (p. 23 1).
5PECIFIC CHARACTERS. SPECIATIOX AND H I G H E R TAXA I N SPIRORBIDAE
The ornamentation of Spirorbid tubes was regarded as a useful specific
bv most early authors and some recent ones, initially including myself,
proved to be basically unreliable. Tubes that usually bear longitudinal
ridges commonly lack them in the younger stages of growth (Crisp, Bailey 8c
Knight-Jones, 1967 ; Knight-Jones, Knight-Jones & Llewellyn, 1974) and may
lack them throughout life in areas of reduced water movement (Gee, 1964). High
perforated ridges may be produced by certain populations, yet the animals wiII
not differ in any way from others where the ridges are normal o r incipient
(Knight-Jones, Knight-Jones & Kawahara, 1975).The terminal coils of tubes may
ascend o r become evolute, when in dense aggregations (p. 218) o r covered by
c hnracter
but it has
NEW SPIRORBIDS FROM FOUR OCEANS
235
algal filaments or encrustations (Gee & Knight-Jones, 1962). In deep silty areas
most of the tube can become erect (Knight-Jones, Knight-Jones & Bregazzi, 1973;
Bailey-Brock & Knight-Jones, 197 7 ) .
Opercular characteristics are more useful in taxonomy but specimens of all
ages should be examined. Not only is there dimorphism involving primary and
brooding opercula of Janua, Pileolaria and Sinistrella (Zibrowius, 1968, 1969;
Knight-Jones, Knight-Jones & Kawahara, 19751, but the brood chambers too
may vary during growth and in different localities. Those o f J a n u a pagenstecheri
can vary from flat to domed and may even bear a distal projection. Distal
ornamentations which normally develop on older brood chambers may be
absent from young ones and may never develop at all in populations of P.
berkeleyana at La Paz, Mexico and P. pseudoclavus in some parts of the West Indies.
Similarly, in tube incubating species, the opercular excrescences of Spirorbis
marioni which develop during ontogeny (Vine, Bailey-Brock & Straughan, 19 7 2 )
may be suppressed altogether in populations at La Paz, St Helena and Canary
Islands (pers. obs.), whilst the opercular plates of Romanchella bicava can vary from
flat to domed within a single population.
Fine differences in proportion and ornamentation of abdominal and collar
setae can be more diagnostic than differences in opercula (Knight-Jones, KnightJones & Kawahara, 1975) and this certainly applies to the separation of s. marioni,
S. bfurcatus and S. rothlisbergi (pp. 204 and 205). On the other hand it does not
necessarily follow that forms with similar setae are conspecific, as is illustrated by
the sibling pair Spirorbis corallinae and Spirorbis inornatus (Knight-Jones, KnightJones & A1 Ogily, 1975). It is on the basis of experience with those two British
species that Spirorbis spatulatus is here regarded as distinct from S. rothlisbergi, and
Pileolaria tiarata from P . lateralis. The members of the former pair and probably
the latter too have overlapping distributions but are separated ecologically
having different algae as substrata. Careful field studies will be needed to
establish the status of these forms and it should be noted that choice of specific
algae is not in itself good evidence for specific status. Local populations of
Spirorbis spirorbis in Canada (Doyle, 1974) and S. inornatus in Britain are adapted to
choose different dominant algae, but can be regarded as no more than the
earliest stages in speciation through that process.
It has often been supposed that most speciation is allopatric, and several of the
forms here (viz. Spirorbis strigatus, Pileolaria dakarensis, P. palliata, P . zibrowii and
Romanchella bicava) are indeed found in rather isolated areas and may have
diverged for that reason. In general, however, the spirorbids of isolated islands
are amongst the most widely distributed species of the family (Knight-Jones,
Knight-Jones & Kawahara, 1975) and as Scudder (1974) has remarked for other
animals (mainly insects) there seem to be “as many examples of sympatric
speciation as there are geographical isolates”. California in particular would
appear to have been a centre of evolution for Pileolaria and Spirorbis sensu stricto.
The genus Spirorbis now contains 14 species and these tend to fall into two
morphological and geographical groups. Those having small talons, collar setae
blades without cross-striations and thoracic uncini with oblique rows of teeth are
mostly European, whereas those having large tooth-like talons, collar setae
blades with cross-striations and thoracic uncini with transverse rows of teeth are
mainly centred in the east Pacific. There are, however, also species with
intermediate characteristics and some which are typical of one of the groups but
nevertheless occur far from the main geographical area. Situs inversus coiling in
2%
P. K Z I G H T - J O S E S
this genus is iiiostly confined to the east Pacific and several of‘ the species
concerned are usually dextral, whereas 9 of the 14 Spirorbis species arc always
siiiistral, most of these being European. The genus Spirorbella Chamlm-lain was
suggested for dextral forins with the characteristics of SpirorbiJ, but dii-ection of
cx)iling is not sufficient to characterize even a subgenus, especially considering the
UI-iabilitv of coil direction in several species described abovc. I t is difficult to
rci+.r.itettie diagnosis of Spirorbella to cover other characteristics of‘ the Pacific
gi.oup, because of the existence of intermediate forms. I t ther-cfbrcsecrtis best to
drop ttie taxon altogether and regard the genus Spirorbis as a fairly unified group
\+4tli 110subgenus.
The genus Pileolaria is the largest within Spirorbidae, now comprising 35
specie,. Of‘ these 18 resemble the type quite closely, but the remainder niay be
divided between several small subgenera. Simplicaria Knight-Joiies ( 1973) with 3
species is thr closest to Pileolaria sensu stricto, differing only in the absence of sickle
sctae. Duplicaria Vine i 1972a) is the most distant. Duplicaria was established for
species with a brood chamber formed of two interlocking parts (e.g. P. ( D . )
korhlerii, but P. dalestraughanae (Vine 1972b) and P. leuemteinae Bailey-Brock 8c
Knight-Jones ( 197 71 were later included, because their cup-shaped opercula
resc.nible the lower unit of the brood chamber in species like the tvpe P. ( D . )
koehleri. P. dalestraughanae and P. palliala (p. 223) however have to rely on an
adhesive matrix to secure their embryos ( P . leuemteinae was not breeding),
whereas the tvpe of Duplicaria has them between the two opercula. I t thus seems
best t o separate from Duplicaria (now 3 species) the new subgenus Nidularia (6
spccies) comprising those with a single brood chamber unit.
.l‘idularia moreover is closer to Pileolaria sensu stricto than to Duplicaria. As
Ttiorp ( 19751 suggests, the brooding operculum typical of Pileolaria sensu stricto
rnav have developed from the simple form represented in P. (A’.) dalestraughanae,
by &a\ of stages like those in P. (A1.) claw3 and P. ( P . ) dayi. The primary opercula
of both subgenera are closely similar.
,/ugaria (4 species) is also close to Pileolaria sensu stricto. The characteristic
granulata throughout
retention of the “priinaq” operculum goes on in P . 0.)
several broods (Thorp, 1975). The peripheral flattened talon is connected to
strengthening fibres, which are easily seen by low power microscopy in larger
species such as P. 0.) quadrangularis. These fibres lead from the talon to the
thoracic uncini on the concave side. They are so strong that both tori are often
torn free if the operculum is pulled ’away from the animal, for separate
examination. In P. (J.) granulata a strand-like zone of cuticular sclerotinization is
embedded within the talon and fused proximally with the cuticle (Thorp, 1975).
This secure anchorage of the primary operculum presumably gives added
strength to the brood-chamber and enables it to be withdrawn decisively when
the animal is endangered by a browsing predator, such as a fish (Qasim, 1957).
Such strands are also noticeable in species of the genus Sinistrella, which have an
opercular structure like that ofJugaria. Both these taxa occur mainly in arctic and
north temperate waters where species of Pileolaria sensu stricto are rare. Probably
this form of opercular strengthening is one of the adaptations for increased size
in cool waters. Similar fibres are easily seen in Antarctic and Southern Ocean
genera such as Romanchella and Helicosiphon (Figs 16-18), but with a few
exceptions (Knight-Jones and Knight-Jones, 1974) they are not visible in the
numerous tropical species o f Pileolaria sensu stricto andJanua.
The other diagnostic characters of Jugaria (p. 227) are also those of Sinistrella,
NEW SPIRORBIDS FROM FOUR OCEANS
23 7
except that the latter has simple collar setae, lacking fins. In Spirorbidae it has
been traditional to separate taxa on this character, but it is not always clear.
Some collar setae in Sinistrella bear an obscure marginal indentation which could
be said to differentiate a proto-fin, whilst Pileolaria (J.) stimpsoni has a very
reduced gap between fin and blade, reminiscent of that of P. (0.)zibrowii (Fig.
14H). Evidently there is a close relationship here and Jugaria might well
become a subgenus of Sinistrella, or Sinistrella could be a subgenus of Pileolaria. If
either of these arrangements were used, however, it would be desirable for the
sake of uniformity to lump other genera. Protolaeospira, for instance, would
become a subgenus of Helicosiphon. The latter however, has only recently been
recognized as a spirorbid (Knight-Jones, Knight-Jones & Bregazzi, 1973) and
lumping of the two would cause confusion and would be contrary to modern
tendencies which are to split rather than lump.
The close relationship between Helicosiphon, Protolaeospira and Romanchella is
best acknowledged by associating them in the Romanchellinae, as proposed here
(p. 229), the most important unifying character being the epithelial process
attaching the embryo sac to the thorax. Other sinistral tube incubating genera
lack such processes and may be grouped in another new subfamily, the
Paralaeospirinae, comprising Paralaeospira Caullery & Mesnil, Eulaeospira Pillai
and Metalaeospira Pillai. These two subfamilies are alike in being centred in
southern or warm seas and in not attaching their embryos to their tube walls as
do the northern tube incubators (Knight-Jones, Knight-Jones & Vine, 1972) viz.
Spirorbinae and Circeinae, the latter accommodating Circeis St. Joseph and
Paradexiospira Caullery & Mesnil. The affinities of Anomalorbis Vine ( 1972a) and
Neomicrorbis Rovereto (see Zibrowius, 197 2) are obscure, but Amplaria KnightJones ( 1973)has an opercular blood chamber very like Duplicaria and could well be
accommodated in the Pileolariinae. The latter would certainly include Sinistrella
(see above) but the predominantly dextral opercular incubators might well be
separated as the Januinae, for these would appear to be a rather isolated group.
SUMMARY
Eighteen species are described, eight from S. California. Of the latter Spirorbis
bfurcatus sp. nov., Spirorbis rothlisbergi sp. nov. and Spirorbis spatulatus sp. nov. are
close to Spirorbis marioni Caullery & Mesnil, and could be placed in “Spirorbella”;
but this taxon is rejected since situs inversus coiling is not infrequent. Pileolaria
lateralis sp. nov., Pileolaria marginata sp. nov., Pileolaria spinqer sp. nov. and
Pileolaria tiarata sp. nov. are remarkably close to each other and to Pileolaria
berkeleyana (Rioja), but Pileolaria (Simplicaria) potswaldi nom. nov. pro. Spirorbis
moerchi sensu Potswald is more distinctive and not closely related to Pileolaria
moerchi (Levinsen).
Madeira yields Spirorbis strigatus sp. nov. which is also close to Spirorbis marioni.
Pileolaria alata sp. nov. from Curacao has a primary talon with a lateral wing
whilst Pileolaria dakarensis sp. nov. from West Africa resembles P. margznata (see
above) but is smaller, with a longer asetigerous region.
A new subgenus Nidularia is proposed for forms with opercula that do not
completely protect the embryos. This includes Pileolaria (Nidularia) nidica sp. nov.
from Curacao in which the embryos are partially exposed laterally, and Pileolaria
(Nidularia) palliata sp. nov. from S. Africa in which they are very exposed distally.
238
P. KNIGHT-JONES
The latter has a longitudinal skin-fold forming an abdominal cloak, like that
found in P. potswaldi (see above) and in two Sinistrella species. Pileolaria
(Duplicaria) zibrowii sp. nov. from Madagascar is like Pileolaria (Duplicaria) koehleri
(C. 8c M.) but much smaller and with several of its collar setae of the simple type,
thus fonning another link between Pileolaria and Sinistrella. Pileolaria atlantica sp.
nov. from north of the Azores is placed with Pileolaria granulata L. in a new
subgenusJugaria, having compound opercula as in most Sinistrella species.
Romanchella bicava sp. nov. from Amsterdam Island is the first dextral member
of this southern genus. Romanchella pustulata sp. nov. from Chile has an
operculurn with lateral knobs as in several species of Protolaeospira. Helicosiphon
platyspira sp. nov. from Marion Island has flat coiling and a vestigial brood stalk.
Several of these island species may well be vicariant but some of the
Californian forins have probably resulted from sympatric speciation in both
genera concerned. From morphological and zoogeographical considerations the
spirorbids may conveniently be divided into the subfamiles Spirorbinae,
C i r ce i n ae , P i 1e o 1a r i i nae , J a nu i na e, R o m a n ch el 1in a e a n d perhaps
Paralaeospirinae.
ACKNOWLEDGEMEKTS
I a m much indebted to all those (listed o n page 202) who provided this
mnterial and to the Royal Society for a grant towards the cost of my visit to
Madeira and Dakar. At Madeira a launch was kindly provided by Dr G. E. Maul,
Curator of the Museu Municipal d o Funchal, and diving equipment by Mr
Manuel Biscoito. At Dakar similar invaluable help was given by Dr Pierre Freon,
through the kindness of Dr J . Chabanne, Director of the Centre de recherches
oceanographiques de Dakar-Thiaroye. Dr H . A. ten Hove’s work at Curacao was
supported by the Netherlands Foundation for the Advancement of Tropical
Research.
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