Zool. J . Linn. SOC.,56: 91-129.With 6 figures
March 1975
A review of the genus Janua, including Dexiospira
(Polychaeta: Spirorbinae)
PHYLLIS KNIGHT-JONES, E. W. KNIGHT-JONES
Department of Zoology, University College of Swansea, U.K.
AND
TATUO KAWAHARA
Faculty of Fisheries,
Mie University, Edobashi, Tsu, Mie-Ken, 514 Japan
Accepted f o r publicarion July I974
Janua (Dexiospira) bradlien& and J. (0.)foraminosa are fully described for the first time and
compared by a pictorial key with other species of Dexiospim and Pillaiospira. The former is
very clox to J. (0.)pseudocorngara (= “SpirorbiS corngatus”) and particularly to J. (0.)
nipponica, whilst the latter occupies a central position in the group of five species (including J.
(0.)sreueri) which are characterized by having large four-lobed juvenile talons.
Janua (Janua) pagensrecheri is the most widespread species of the genus, whereas J.
(Leodora) knighrjonesi Seems to be confined to coral was. In all. 14 species of Janua are
recognized, whilst 19 names are here regarded as synonyms or varieties. Eleven other species
sometimes placed in Dexiospira or Leodora should rather constitute the genera Circeis,
Sinisfrella and Romanchella. These are re-defined and their affinities are studied.
The subgenus Dexiospira seems to be centred in the Indo-West Pacific, where it reaches
maximum species-diversity and extension into cooler latitudes. J. (0.)pseudocortugara var.
mannarensis is pantropical and overlaps in the Mediterranean with the west European form and
in the West Pacific with vars praeceps and f e n e m t o , and with subspecies a i m . which are
characterized by small modifications of the tube or abdominal setae.
CONTENTS
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92
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Introduction
Materials and methods
Genus Janua Saint-Joseph (1894)amended
Subgenus Janua
. . . . . . . . . . .
Subgenus DexiospiraCaullery&Mesnil(1897)amended
Janua (Dexiospira) brasiliensis (Grube. 1872)
Janua (Lkxiospira) foraminom (Bush, 1904)
Other subgenera of Janua
Subgenus Leodora St-Joseph (1894)amended
Subgenus Fauveldom Knight-Jones (1972)
SubgenusPiNoiospira Knight-Jones(1973)amended
7
91
92
P. KNIGHT-JONES ET AL.
Guide t o the species of Joncra
.
. . . . .
.
. . .
Some other taxa of Spirorbinae
Grnu.: Circeis St-Joseph (1894)
. . . .
Genus Rotnanchello Caullery & Mesnil ( 1 897)
Genus Sinisrrello ( = F'rorolcodora Pillai. 1970)
World tli.;trit)ution
. . . . . . . . . .
Summary
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Acknowledgments
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References
Appendix . . . . . . . . . . . . .
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100
117
118
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125
125
128
INTRODUCTION
The species that fall within the group Jaittta (Dexiospira) (see p. 4) are often
difficult to identify, for the differences separating some of them are very small.
Zibrowius (1968) suggested that a few of the old names are homonyms, but
other authors (Pillai, 1970; Uchida, 1971a; Knight-Jones, 1972, 1973; Vine,
1972a, b) propose further new species, some of which differ even less from
those previously described. The difficulties arise partly from the fact that
descriptions by early authors (Caullery & Mesnil, 1897; Bush, 1904; Sterzinger,
1909) were brief and seldom included comparisons with closely related species.
Comparative descriptions of a few Australian species have recently been
published (Knight-Jones, 1973;Knight-Jones, Knight-Jones 8c Llewellyn, 1974)
and further collections from Japan, north-west France, S. Africa and elsewhere
have now placed us in a position to review the genus Janua. In this we have had
invaluable assistance from the Museum authorities and others mentioned in the
acknowledgements (p. 125).
The classification into genera and subgenera used here is mostly a
compromise between the views of Chamberlain (1919), Bailey (1969b) and
Pillai (1970), with amendments by Knight-Jones (1972) and Vine, Bailey-Brock
& Straughan ( 1972). Descriptions of Janua (Dexiospira) brasifiensis (Grube)
and J. (D.1foraminosa (Bush) are given early in this account, for those species
have not been described adequately before. All species of Janua are later listed
in alphabetical order, with pictorial keys and notes on the world distribution,
so far as it is known.
MATERIALS AND METHODS
Our Japanese material was collected on Hokkaido, where Janua brasiliensis
(Grube) was abundant on a marine angiosperm at Akkeshi, whilst Janua
pseudocorrugata var. praeceps (p. 114)was abundant on stones at Hakodate; and
on Honshu, in Ago Bay, Mie Prefecture, where stones and shells bore abundant
J. pseudocorrugata var. a i m (p. 115) and a few Janua foraminosa (Bush) and
Jarzua nipponica (Okuda). The latter was more common on a bryozoan,
however. Our other sources of material may be understood from the details
given below for each species and from our later acknowledgements to
individuals and institutions (p. 125).
Our collections were preserved either in 5% formaldehyde or 70% alcohol
(both diluted with sea-water). The presence or absence of collar fusion
(Langerhans, 1880; Day, 1963; Pillai, 1970) was demonstrated by probing with
REVIEW OF THE GENUS JANUA
93
a mounted eyelash (Knight-Jones, 1972). Details of tubes and opercula were
distinguished in reflected light but, to examine setae, specimens with opercula
removed were mounted in polyvinyl-lactophenol and viewed by phase contrast,
with quartz-iodide illumination. Drawing5 were made on squared paper with
the aid of squared eyepiece graticules.
A guide to the setation of Spirorbinae has been given recently (Knight-Jones,
Knight-Jones & Llewellyn, 1974). In the descriptions which follow, the setae
and uncini selected for illustration were the largest in a given animal, but were
checked as being typical of the species whenever sufficient material was
available. The size of thoracic uncini (and the numbers of longitudinal rows of
teeth) vary within a single torus, whilst the blades of abdominal setae tend to
be shorter (but not usually narrower) in the posterior segments.
Genus Janua Saint-Joseph (1894) amended
Mostly with dextral coiling; six to seven tentacles; only two pairs of thoracic
tori; incubation in an opercular brood chamber, below which a secondary plate
(rudiment of next opercular plate) is formed soon after spawning; collar setae
obliquely geniculate and simple (without a toothed basal fin); those on the
concave side are never cross-striated; abdominal setae obliquely geniculate, as
big as or bigger than the collar setae and with elongated blades which are often
parallel-sided or taper very gradually; each fully developed abdominal seta
occurs singly, but may be accompanied by a single rudimentary seta (secondary
seta of Vine, 1972) which has the blade lying along the shaft of the main seta,
as though developing in the same follicle; thoracic uncini with thick anterior
pegs, which are pointed in surface view and usually single (rarely three or five),
followed by numerous teeth which are arranged in oblique rows and also in
between 5 and 8 longitudinal rows; distribution of abdominal tori more or less
bilaterally symmetrical, with largest tori in the anterior half of the setigerous
region; larvae have paired white attachment glands in the thoracic region. Italics
indicate characters not found in other Spirorbinae.
Subgenus Janua
Coiling usually dextral; brood chamber walls uncalcified and transparent;
sickle setae present in the third thoracic fascicles; thoracic collar folds not
fused dorsally.
Type. Janua (Janua) pagenstecheri (Quatrefages, 1865). See p. 111. This may
perhaps be the only species representing this subgenus.
Subgenus Dexiospira Caullery & Mesnil(l897) amended
Coiling usually dextral; distal part of brood chamber with calcified walls;
collar setae on the convex side may have crossstriated blades; no sickle setae in
thorax; collar folds usually fused dorsally, to form a tunnel over the faecal
groove.
Type. Janua (Dexiospira) pseudocorrugata Bush (1904); see p. 113.
At least eight other species are recognisable within this subgenus, viz. J. (D.1
brasiliensis (Grube, p. 94); J. (D.) foraminosa (Bush, p. 96); J. (0.)
formosa
94
P. KNIGHT-JONES €7-
AL.
(Bush, p. 106), J. ( D . ) lumellosu (Lamarck, p. 110);J. (D.)
nipponicu (Okuda,
p. 110);J. ( D . ) preucutu Vine (p. 113); J. ( D . ) steueri (Sterzinger, p. 115); and
J. (D.
) tumbalugumensis (Pillai, p. 116). Several other previously recorded species
Seem to be junior synonyms or varieties of these. We now offer full descriptions
of the first two listed above.
Junua (Dexiospiru) brusiliensis (Crube, 1872)
Type locality Florianopolis ( = Desterro) on algae.
The material was deposited in the Museum fur Naturkunde, HumboldtUniversitat, Berlin. Comparison shows that this (Fig. 4B, p. 104) is the same as
Spirorbis bellulus Bush (in Moore & Bush, 1904), which was dredged in Suruga
Ray, south-east of Honshu, in 63-75 fathoms on shells and stones. Most of the
following description is based on Bush’s type (Fig. 1K-S) and paratypes
(Fig. 1A-F), kindly loaned by the Smithsonian Institution, and on abundant
material from a marine angiosperm at Akkeshi, Hokkaido. It agrees with the
description of Dexiospiru oshoroensis Uchida (1971a), from Surgussum at
Oshoro, and was probably the form on Surgussttm at Asamushi, which Abe
(1943) suggested might be Spirorbis urgutus (p. 104). Straughan’s record (1969,
as S. bellulus) from under a rock at the Marshall Islands was indeed this species,
her note about egg-string incubation being a mistake. We have found the same
species at Cape Town and Port Elizabeth, on Cerurniurn plunum in shore pools.
This South African material was previously misidentified as J. ( D . )forurninosu
(Day, 1961) and J. (D.) nipponicu (Knight-Jones, 1973). Also in material from
Luminuriu pullidu holdfasts off Amsterdam Island (kindly sent by Dr Beurois);
on the walls of the aquarium at Leigh, near Auckland, New Zealand (Vine,
1975 as J. (0.)pseudocorrugutu); on a marine angiosperm from Biloxi,
Mississippi (Hartman, 195 1, as Spirorbis corrugutus);and in material from Port
Sudan (p. 104). Januu (Fuuveldoru) anti-corrugatu Vine (1972a) is best regarded
as a situs inversus form of this species.
Tube unremarkable, rounded in cross-section. Large tubes with up to four
equidistant longitudinal ridges, three easily seen from above (Fig. 1A), the
fourth (if present) being rather peripheral (Fig. 1B). Smaller tubes may have
only one ridge, for the inner and two outer ridges develop later.
Operculur plate slightly concave with a peripheral talon. Talons of Brazilian,
South African and Amsterdam Island populations show little variation. They
are somewhat triangular and bilobed terminally, like those usually found in
J. pseudocorrugutu (Fig. 4A, p. 101) and J. nipponicu Okuda (1934, Fig. 4C).
This shape was also found in Japanese material, but often the talon was broader
and sometimes serrated terminally (Fig. lD, E) or even axe-shaped (Fig. 1G).
An axe shape was also typical of the New Zealand material (Fig. 4B). Material
from Mississippi has talons somewhat axe-shaped, but bearing a terminal lobe
that is sometimes bifurcate, as in several other species (Fig. 4G-K). The base in
these broader forms of talon is curved, following the periphery of the plate
(Fig. 1F).
Brood chamber develops below the plate, with cylindrical walls which are
opaque and closely associated with the talon. Below the incubating embryos
REVIEW O F THE GENUS J A N U A
95
Q
F
c
.
0.5 mm
Figure 1. Janua (Deximpim) bmiliensts: A to F two paratypa and K to S holotype of
“Spirorbts bellufus Bush”; G . H. J and T specimens from Akkahi, Japan. A, tube viewed from
above; B, side view of A; C,side view of operculum; D,face view of C; E, face view of another
operculum; F, as E but viewed from below to show the peripheral curve of the talon; G,first
stage mature operculum partially dehisced. presumably to release embryos from the new
developing brood chamber below, which is without talon; H, a primary opsculum showing
another variation in talon; J. a fully mature operculum with well-developed embryos; K. collar
seta from the convex side, tide view; I, another collar scta from the same side, edge view; M.
collar sero from concave side, edge view; N, a m a typical of the second and third fascicles; P.
abdominal sap; Q, abdominal uncinus; R and S, thoracic uncini from each end of the same
torus; T, whole a n i d , dorsal view.
Scale: A and T as B; D to F as C;H and J as C ; L to S as K.
(up to 1 5 ) a secondary plate develops, which is the distal plate of the next
chamber. This and subsequent plates may lack talons (Fig. 1J).
Collar folds fused dorsally, forming a tunnel over the faecal groove. This
point could not be established in the museum material, however, which was
hardened and shrunken by alcohol.
96
P. KNIGHT-JONES
ET
AL.
Collar setae on convex side simple and obliquely geniculate, with fine
marginal serrations (about 7 teeth per 8 ~ m ) Cross
.
striations cannot be seen in
the usual side view (Fig. 1 K ) but may show faintly in the rarely seen edge view
(Fig. 1L). Those on the concave side are more finely serrated and without cross
striations even in edge view (Fig. 1M). Capillary setae are present in both collar
fascicles. The setae of the second and third fascicles are straight with almost
smooth margins and as slender as capillary setae for much of their length
(Fig. 1N).
Thoracic uncini each with a simple dense, pointed anterior peg bordered by
shorter, more transparent splayed “teeth”. The larger uncini have about four or
five longitudinal rows of teeth (Fig. 1s) but the smallest uncini (which are
found at the ventral ends of the tori) may have up to six such rows (Fig. 1R).
Abdominal setae obliquely geniculate, with long thin blades and coarse
recurved marginal teeth (Fig. iP). Uncini with splayed fluted anterior pegs
(Fig. 1Q).
Remarks. The name of this species is nearly as old as pagenstecheri and
would seem to be the oldest of its subgenus. The type material is in remarkably
good condition and the variations throughout the geographical range seem
unimportant. The New Zealand material has the margins of the collar setae not
quite as smooth as elsewhere. The Japanese material exactly resembles the
Brazilian in setation, but is more variable in the shape of the opercular talon.
Off South Africa and Amsterdam Island the anterior pegs of the thoracic uncini
have especially dense flanking teeth, resembling but not as long as the flanking
teeth in the trifurcate pegs of J. (0.) nipponica (p. 110). In Japanese
populations, however, these dense anterior pegs are conspicuously single,
indicating that no interbreeding takes place with J. (D.) nipponica.
Nevertheless, this is the only character separating J. ( D . ) nipponica. J. ( D . )
pseudocorrugata, the type species of Dexiospira, may likewise be distinguished
from J. ( D . ) brasiliensis only with difficulty (pp. 104 and 114).
Janua (Dexiospira) foraminosa (Bush) (Moore & Bush, 1904)
Type locality south-east of Honshu, on a red alga at 34 fathoms. Type and
paratype in Smithsonian Institution, compared by us and identified with
material (a) on stones and shells freshly collected from Ago Bay, Mie
Prefecture, (b) on an alga at 8 m from Oahu, Hawaii (Vine, Bailey-Brock &
Straughan, 1972 as J . ( D . ) steueri), (c) on an angiosperm from Lemon Bay,
Florida (Hartman, 1951, as Spirorbis spirillum), and (d) on Halimeda at 4 m on
Gurna Reef, Red Sea (Vine, 1972a as J. ( D . ) steueri var. heideri). Spirorbis
Jieideri Sterzinger ( 1909) from Suez is probably a junior synonym, as suggested
by Augener (1926) whose record from Ceylon seems reliable.
Other records from Japan (Okuda, 1938; Okuda & Yamada, 1954; Imajima
& Hartman, 1964; Kawahara, 1961, 1962, 1963, 1965), India, Ceylon and
Indonesia (Fauvel, 1932, 1953; Mesnil & Fauvel, 1939) are plausible but need
confirmation, as previous information then available was insufficient to
distinguish this species from closely related forms. Some under this name from
Vietnam (Fauvel, 1935, kindly loaned by the Paris Museum) proved to be
REVIEW OF THE GENUS JANUA
97
J. steueri. whilst other material from Krusadai Island, Madras (Fauvel, 1930)
was J. formosu. We failed to find J. foruminosus in South Africa and regard
Day's (1963) record as relating to J. brusiliensis. J. foruminosu may be defined
as follows:- Tubes rounded in cross section, with three equidistant
longitudinal ridges which may be interrupted and represented merely by rough
protuberances. Numerous small pits (hence the specific name) are arranged in a
diamond pattern on the periphery of the tube (between the outer ridge and the
substratum) and may also be found between ridges. The tube may coil in one
I
C
Figure 2. J a m (Dexiospiro) foramhaw: D and H to N holotype; P, paratype; A, B and G from
Tsu, Honshu, Japan; C, E and F, from Hawaii. A, B and C. variations in tube; D, the holotype
opaculum figured by Bush (Moore 8c Bush, 1904), but the talon is decalcified and no longer
visible; E, primary operculum face view; F as E. but side View; C, mature operculum without
talon; H, collar scta from the convex side; J, collar x t a e from the wncave side; K. seta typical
of the x w n d and third fascicles;,.I abdominal seta; M, abdominal uncinus; N. thoracic uncinus;
P, whole animal dorsal view.
Scales: B = A ; E s D ; P a s C ; J t o N a s H .
98
P. KNIGHT-JONES ET AL.
plane (Fig. 28,C) or the last whorl may ascend (Fig. 2A). Fully grown tubes
may develop a bell-shaped aperture (Fig. 2A,B).
Type operculum a narrow cylindrical brood chamber (Fig. 2D) with a
concave distal plate, the perimeter of which forms a remarkably wide collar.
The indistinct talon figured by Bush is not visible, .presumably through
decalcification. The chamber .contains embryos beneath which is a large
cup-shaped secondary plate, ready to develop into the distal flared “collar” of
the next brood chamber. The juvenile talon (found in material other than type)
is partially transparent, markedly bifurcate terminally and with angular lateral
wings (Figs 2E and 45). The walls of the brood chamber, which are slightly
opaque, develop in close association with this talon. Mature opercula may not
bear talons (Fig. 2G). The tentacles (about seven) may appear unusually long,
but this is probably due to the fact that the brood chamber (being much
narrower than the tube) does not compress the tentacles posteriorly when the
animal retracts (compare Fig. 2G & P with Fig. 1J & T).
Collar setue on convex side simple and obliquely geniculate, with fine
marginal serrations and indistinct cross striations (Fig. 2H). Those on the
concave side have almost smooth margins and no cross striations (Fig. 25).
Capillary setae present in both collar fascicles. Setae of the second and third
fascicles also have smooth margins, but are slender and almost straight
(Fig. 2K).
Thoracic uncini have a dense, pointed anterior peg bordered by less dense
splayed teeth. The larger uncini have about five or six longitudinal rows of
teeth (Fig. 2N).
AbdominuZ setue obliquely geniculate, with large fairly broad blades and fine
recurved marginal teeth (Fig. 2L). Uncini with a splayed fluted anterior peg
(Fig. 2M).
Remarks. See p. 106.
OTHER SUBGENERA OF JANUA
The original proposal of the genus Junua (St-Joseph, 1894)was so terse that
Caullery & Mesnil (1897) had every excuse for disregarding it when they
proposed the first reasoned scheme for a classification of the Spirorbinae. Their
scheme had an attractive simplicity and was followed by such authorities as
Pixell (1912), Fauvel (1927), Hartman (1959) and initially Rioja (1923).
Eventually their disregard for the rules of nomenclature brought retribution. As
Chamberlain (1919)remarked, if they were right in their grouping of dextral
forms, then Dexiospiru is a junior homonym of Janua. He saw that such a view
would be an oversimplification, however, and he proposed new genera and
redefined old ones (including Dexiospira), but many of these were separated
from others only by single characters.
Rioja (1942), Gee (1964), Harris (1969) and Zibrowius (1968)did not feel
ready to split the genus Spirorbis without more information about the
differences between the taxa proposed. Bailey (1969b)too used Chamberlain’s
taxa initially as mere subgenera, but started t o accumulate information about
the differences between them, including larval characters and the distribution
and form of uncini. Soon it became possible to offer more thorough definitions
REVIEW OF THE GENUS JANUA
99
of genera, including Janua (Knight-Jones, 1972, 1973; Vine, 1972 a & b; Vine,
Bailey-Brock & Straughan, 1972). At the same time it became clear that
Dexiospira as defined by Chamberlain should not be confused with Circeis
(p. 118), which is in fact quite close t o Paradexiospira (p. 118). Dexiospira,
moreover, is a senior homonym of Neodexiospira Pillai (1970-see
Knight-Jones, 1972), which is a useful taxon but nevertheless very close to
Janua. Of the characters used so far to help in generic diagnoses, more than a
dozen link Dexiospira and Janua and only two consistently separate them
(p. 93). Most of those which they have in common are found in few other
genera of Spirorbinae. It thus seems best to link them within the peculiarly
isolated genus Janua, whether or not they are also grouped as Januinae within
the family Spirorbidae proposed by Pillai (1970).
Although we thus regard Dexiospira (defined above, p. 93) as a mere
subgenus of Januu, it is nevertheless an important one, much more so than any
of the others defined below. We mention these here tentatively, for discussion,
but some might well be discarded and their species included within Dexiospira.
Subgenus Leodora St-Joseph (1894) amended
Coiling sinistral; sickle setae may or may not be present; collar folds separate
dorsally.
Type: Spirorbis laevis Quatrefages (1865), but this is a nomen dubium et
nudurn Its status has been fully discussed by Vine et al. (1972). Briefly,
Quatrefages’s original description (from Guettary, Bay of Biscay) was
inadequate and the beautiful figure of Claparede (1870) from Naples, which
illustrates “Spirorbis” in well-known textbooks of Zoology, may or may not
refer to the same species (p. 110). That figure showed a Janua species, which was
peculiar in having only two pairs of pinnate tentacles. Subsequent records of
Spirorbis laevis may well have referred t o abnormally sinistral individuals of
J. pagenstecheri or to the tropical species J. (L.) knightjonesi (de Silva,
1965-see Bailey, 1970) or to misidentifications (Zibrowius, 1973).
Leodora was used as a subgenus by Caullery & Mesnil(l897) and as a genus
by Chamberlain (1919). Chamberlain linked it with Romanchella (p. 119) but
this is a very distinctive genus which seems t o be confined to the Southern
Hemisphere (Knight-Jones, 1973; Knight-Jones, Knight-Jones & Llewellyn,
1974). Hartman (1959) linked Leodora with Sinistrella Chamberlain, which
comprises a number of northern cold-water species. Pillai (1970) and Uchida
(1971b) also included these with Leodora, but they are really related much
more closely toPileolaria than toJanua (see p. 119). They are not represented in
the Bay of Biscay or Mediterranean and are very different from Clapar6de’s
Spirorbis laevis. We must therefore remove from Leodora the species abnormis,
asperata. corona ta, evoluta, media, multiplata and perrieri listed by Uchida
(1971b). Spirorbis argutus Bush was probably a Eulaeospira (p. 104). The only
species remaining to represent Leodora, alongside the problematical type, is
J. (L.) knightjonesi. This is intermediate between J. pagenstecheri and the
Dexiospira group, in that it lacks sickle setae and calcified walls to its brood
chambers. It is thus quite convenient to retain Leodora in order to
accommodate it.
100
P. KNIGHT-JONES ET A L .
Subgenus Fauveldora Knight-Jones (1972)
Coiling sinistral; sickle setae absent; collar folds fused dorsally.
Type. Janua (Fauveldora) kayi Knight-Jones, 1972, which is perhaps the
only known species, if we regard J . (F.I ariticorrugata Vine (1972a) as based
upon an abnormally sinistral population of J. ( D . ) brasiliensis (see p. 104). I t
seems best to include the type with its close relatives in Dexiospira and to drop
this subgenus, now that situs inversus forms of Spirorbinae are better known.
Fauveldora was proposed to reduce the number of sinistral forms in Dexiospira.
but that seems unnecessary.
Subgenus Pillaiospira Knight-Jones ( 1973) amended
Coiling dextral; sickle setae absent; collar folds mostly separate dorsally.
Type. Janua (Pillaiospira) trifurcata Knight-Jones 1973 . These forms are very
close to Dexiospira, but the type species was separated because it showed no
collar fusion dorsally. We have recently found in South Africa, however, a
related but distinct species (p. 110) in which the collar folds are usually separate,
but may be partially fused (in young specimens remarkably enough). It seems
that this character alone is not very useful, but we are inclined to retain
Pillaiospira in order to avoid amending the definition of Dexiospira.
Finally, we should perhaps note that Dexiorbis Chamberlain 1919 (type
J. forarninosa) and Mera St-Joseph 1894 (type M. pusilla, p. 115) were based
upon misconceptions and have been dropped from taxonomy, whilst
Proroleodora Pillai (1970) is a junior synonym of Sinistrella (p. 119) and not
closely related to Janua.
GUIDE TO THE SPECIES OF J A N U A
A Janua species can be recognised as such in a microscopical preparation
(polyvinyl-lactophenol) either by the relatively large abdominal Setae or by the
pointed pegs of the thoracic uncini. Indeed, the genus can usually be
determined with a mere hand-lens, for it may be assumed to comprise all
opercular incubators that are dextrally coiling. If an opercular incubator is
sinistrally coiling and lives in warm seas, it can still be identified as a Janua if
the collar setae are simple, for with the possible exception of the peculiar
Anomalorbis (Vine, 1972a), all other opercular brooders with simple collar
setae belong to the genus Sinistrella (p. 119), which is mostly confined to the
cold northern seas of America, Russia and Japan, Sinistrella abnorrnis (Bush)
being the only representative known from California.
If a Janua is dextral and has the brood chamber cup-shaped with its lateral
walls completely transparent (Fig. 3 A-E), it may be identified tentatively as
J. (J.) pagenstecheri (p. 111). Possession of thoracic sickle setae (Fig. 3G)
positively identifies this very common and widespread form. The commonest
sinistral Janua is J. (Leodora) knightjonesi, which seems to be confined to coral
seas. This has brood chambers very like those of J. pagenstecheri, but more
subject to multiple stacking (Fig. 3J). Both these species have the collar folds
unfused dorsally.
The remaining Janua species (11 so far described and now deemed valid)
REVIEW OF THE GENUS JANUA
101
Figure 3. Junuu (Junuu) pugemrecheri: A to E, variations in form of opercular brood chamber
(p. 11 1); F, collar setae; G, sickle seta from third fascicles; H, abdominal seta. Junuu (LeodomJ
lolightjonesi: J, brooding operculum; K. collar seta; I. abdominal seta.
Scales: B to Emd J asA;G, H, K and L as F.
belong to the subgenera Dexiospira (fused collar folds) or Pillaiospira (unfused
folds). Fusion of the collar folds is particularly easy to determine in live
material. Most of these species have cylindrical brood chambers with heavily
calcified walls, but this calcification is thin and transparent in the most
widespread form J. (0.)pseudocorrugata (Fig. 4Ao). These species may be
identified from the pictorial key given in Fig. 4A-L and from the comparative
notes given in the list which follows, where the arrangement is in alphabetical
order of specific names.
ainu. For Dexiospira ainu Uchida (197 l a ) see Janua (Dexiospira)
pseudocorrugata subspecies ainu (p. 115).
alveolatus. Spirorbis alveolatus Zachs (1933) from the northern Sea of Japan
was inadequately described, without figures. I t was said to be similar t o
0.2 m m
..............-_.-...-._.
... ......
.......'
"
F
Figure 4. Dcxlacplm species and close relatives (for details r e p. 93). A, pseudocormguru;B,
bmsllfmsis; C, nfpponfca;D,trffuurcota;E,preacum; F, mmbalqgomensla
Figure 4 (continued-for details see p. 105). C,koyi; H, sreueri;J, forumfnosrr; K.fomato; L,
hmelloa
104
P. KNIGHT-JONES ET A L .
Spirorbis jbraiiiiriosiis in having simple setae and a tube with sharp ridges
flanked by depressions which are regularly located opposite one another and
occasionally penetrate the ridges. The tube of Janua riipponica (Pa 110) is
sometimes like that (Fig. 5A) which is presumably why lmajima & Hartman
( 1964) regarded tiipponica as synonymous with alveolata. However, perforated
ridges are seen in at least one other species of Jama (Knight-Jones, 1973-see
p. 114) and perhaps in other Spirorbinae. Uchida (1971b) rightly concluded that
nippoirica must be regarded as distinct. Zachs described a semitransparent tube
and incubation as probably occurring within it, which suggests a species of
Circcis (p. 118).
anticorrugata. Type material of Jama (Fauveldoral anticorrugata Vine
(1972a) from Port Sudan resembles J. ( D . ) brasiliensis in every detail except
direction of coiling. We have recently found normal J. ( D . ) brasiliensis in
material from Port Sudan kindly sent by Dr Vine and we consider
J. anricorrugafa a sirrts inversus form of this species. This is part of the case for
suppressing Fauveldora (see p. 100).
antillarum. Spirorbis antillarum Augener (1922), from the West Indies was
not figured and the brief inadequate description might be applied to most of
the nine species listed on pp. 93-4.
argutus. Spirorbis argutus sensu Abe (1943) was dextrally coiling, well
figured and probably J. ( D . ) brusiliensis (p. 94). The description of Spirorbis
argzctus Bush (Moore & Bush, 1904) from Japan was probably the first of what
has since been called Eulaeospira orientalis (Pillai, 1960). Pillai’s material has
collar fascicles with a mixture of simple and finned setae and it would have
been easy for Bush to have overlooked the latter. E. orientalis was fully
described by Vine (1972a) from the Red Sea and recently discovered in
collections from Hawaii which included other Japanese species of Spirorbinae
(pers. comm. from Dr Julie Bailey-Brock). This species has the anterior pegs of
the uncini distinctively multipronged, but this character could not be seen in
the microscopical preparation of Bush’s type specimen, which is fragmented
and in poor condition. It seems best to allow the name argutus to lapse.
Lulaeospira Pillai (1970-redefined by Knight-Jones, Knight-Jones &
Llewellyn, 1974) is a genus of tube incubators and its affinities lie not with
Junua but with various genera of the southern hemisphere (Knight-Jones,
Knight-Jones & Bregazzi, 1973).
bellulus. Spirorbis bellulus Bush (in Moore & Bush, 1904), is a junior
synonym of J. (D.) brasiliensis (p. 94). Japan.
benhami. Neodexiospira benhami Pillai ( 1970) is a junior synonym of
J. (D.)
pseudorrugata var. mannarensis (p. 114). Ceylon.
brasiliensis. Janua (Dexiospira) brasiliensis (Grube, 1872-see p. 94) = Spirorbis bellulus Bush = Spirorbis foraminosus sensu Day (1961) = Dexiospira
osltorensis Uchida (1971a) = J. (Elanticorrugata Vine (1972a). Distinguished
from J. (D.) pseudocorrugata only by the finer marginal striations on the collar
setae and the greater opacity of the brood-chamber walls, and from
J. ( D . )nipponica only because the teeth flanking the anterior pegs of the
thoracic uncini are shorter and do not form such a distinct and optically dense
trifurcation. The oldest Janua species of which type material remains.
Distribution (b in Fig. 6) described on p. 94 and discussed on pp. 123-4.
ceylonicus. Spirorbis ceylonicus Pillai ( 1960) is represented by a single
REVIEW OF THE GENUS J A N U A
105
lectotype, the tube of which was left unopened. This and the description
(which did not include the juvenile form) do not differ clearly from J. ( D . )
pseudocorrugata.
comptus. Spirorbis comptus Bush (1904) on red algae from California was
inadequately described, but seems to have been a Janua species. Hartman
(1942) was unable t o find the type material of this species in the Peabody
Museum, so it should be regarded as a nomen nudum.
cormgutus. Spirorbis corrugatus Montagu ( 1803) was an inadequately
described sinistral species, possibly Spirorbis rupestris Gee & Knight-Jones
(1962) to judge from Montagu’s ecological notes. S. corrugatus Montagu sensu
Langerhans (1880) was dextral and seems to have been a species of Janua
(Pillaiospira) (p. 116). The dextral S. corrugatus sensu Caullery & Mesnil and
most modern authors was renamed pseudocorrugatus by Bush (1904) who
pseudocornrgata, p. 113 ) .
realised the other name was preoccupied (see J. (D.)
epichysis. Spirorbis (Junua) epichysis Bailey (1970) is within the range of
variation shown by J. pagenstecheri (p. 111) in warm latitudes.
fuuveli. Neodexiospira fauveli Pillai ( 1970) was probably a descriptive
chimaera of J. (DJformosa (p. 106) and J. (D.)kayi (p. 107), for the paratypes
include both these species but none agreeing with the description.
fenestrutu. Janua (Dexiospira) fenestrata Knight-Jones (1973) is probably a
variety of J. ( D . )pseudocorrugata (p. 113).
foruminosu. Janua (Dexiospira) foraminosa (Bush in Moore & Bush,
1904) = Spirorbis heideri Sterzinger (1909) = Janua steueri var. heideri Vine
(1972a) = Januu steueri sensu Vine, Bailey-Brock & Straughan (1972); non
Figure 4. (see pp. 102 & 103). Comparison of the early opercula (0).abdominal setae (as),
convex side collar setae (a)
and thoracic uncini (u), in closely related species of Jonuo.
The fully mature opercula (as in Figs. 1J and 2C) have been omitted as they lack talons and are
therefore very similar and difficult to identify. All the first stage mature opercula, which have
talons, are figured with embryos in order to illustrate differences in opacity of the brood
chamber walls due to varying degrees of calcification. The opacula without embryos are of the
primary or juvenile type, before brood chambers are formed. All opercula are figured to the
same scale (top left) in face view and all serae to the scale on right, in the usual side view unless
otherwise stated. All species have thoracic collar folds fused except D and all are dextral except
C and occasionally B, H and J . Anterior pegs of thoracic uncini are as in Bu in all species except
Cand D.
A. Jonuo (Dexiospiro) pseudocorrugoto: 01, Plymouth U.K. (also N.W. France,
Mediterranean and Mozambique); 02, Roscoff. France and most world locations; 0 3 , as 1 and
a,Subspecies olnu, Japan; 04, Japan var. pmeceps (occasionally other locations and often in
subspecies oinu); as 2, same specimen as 0 4 and typical of most world locations.
B. J. (0.)bwsfliensis (see also Fig. 1) from type material in Berlin Museum, except 0 2 and
as2 which are from New Zealand.
C. J. (0.)nipponico. Japan.
D.J. (FWoiospiro)trifurcato, Australia. See also J. notdensis.
E. J. (0.)preocuto: 0 1 (primary operculum side view) and 0 2 figured from Hawaiian
material (see p. 113) and the setae from the Red Sea.holotype (a2shows edgc View).
F. J. (0.)tombologomensisparatype. Ceylon.
G . J. (Fauwldoro) koyi: 0 1 and setae from Kenyan specimen; 0 2 , Kuwait (similar concial
opercula seen in Mozambique and Ceylon).
H . J. (0.)steueri: 01. Australia; 0 2 , Kenya (also typical of material from Red Sea). 0 2 and
setae from same specimen.
J. J. (D.) forominom (see also Fig. 2), Red Sea.
K. J. (0.)fonnoso: 01, Ceylon (also typical of Bermuda, Kenya and Mozambique); 0 2 and
sctae. Australia.
L. J. (0.)l o m e l l ~Australia. The hidden terminal (bibbed) portion of the talon is
represented by a dotted line (see p. 110).
106
P. KNIGHT-JONES ET AL.
Figure 5. A. Janua (Dexiospira) nipponica tube on bryozoan, Tsu, Honshu, Japan. showing
paforaced ridges. Remainder J. (0.)p m d o c o m r g a t a : B. tubes on stone, Mannar lagoon,
Ceylon. round in cross-section, with indistinct ridges and a small peripheral flange on the last
whorl; C, tube of var. praeceps on stone, Hokodate, Hokkaido, Japan, rather square in
cross-section due to the prominent peripheral ridge and flange; D, tube of subspecies alnu. on
stone, Tsu, Honshu, Japan, showing single ridge and sloping peripheral flange.
foruminosus sensu Day (1961),which was brusiliensis, or sensu Mohammad
(1971),which was h y i . The tube (p. 96) bears fine pits (Fig. 2), hence the
name, but the depth of pitting varies and other Junuu species too may have
their tubes more or less pitted. The pits are arranged in a rather characteristic
mesh-like pattern but we have also Seen this in a few West Indies populations of
J. formosu and J. steueri.
The operculum is very similar to those of J. steueri and J. formosu, but the
brood chamber is narrower and less opaque whilst the distal rim is larger
(Fig. 4J).It is almost impossible to distinguish juveniles of these three forms by
their opercula (cf. Fig. 2E and Fig. 4H,J and K), whilst the setal characteristics
of J. foruminosu are intermediate between those of the other two. The collar
setae on the convex side (Jcs in Fig. 4) are more coarsely serrated than in
J. formosu but less so than in J. steueri. The abdominal setae are less broad
than in J. formosu but broader than in J. steueri. Most material of
J. foruminosu (not the Florida specimens) also differs from the other two
species in having the blades of the collar setae about the same size as those of
the abdominal setae. This character is found too in J. lamellosu, but the talon
of the latter and the very coarse serrations on its collar setae are highly
distinctive. The distribution (p. 96 and a in Fig. 6)is Indo-Pacific but always
north of the equator and discussed on p. 122.
formom. Junuu (Dexiospiru) formosu (Bush, 1904), which may include
REVIEW OF THE GENUS J A N U A
107
Spirorbis kuruitivensis Pillai ( 1960), was recently redescribed from Kenyan
material and Bermudan specimens identified by Bush (Knight-Jones, 1972). It
differs from J. (D.) foruminosu in that the brood chamber (Fig. 4K) is wider
and more opaque, with a less prominent distal rim, but it resembles
J. (0.)
sreueri in these respects. The juvenile (non-brooding) opercula of the
three latter species are very similar (Fig. 4H,J, K), though the opercular plates
(both juvenile and adult) of J. (D.)formosa are usually distinctive in that the
edge is sharply depressed on the side of the talon. It is, however, necessary to
examine the setae t o be certain of identification. The blades of the abdominal
setae of J. (D.) formosu seem to be uniquely wide compared with other Junuu
species, the length/breadth ratio being about 5 : 1, whilst the collar setae of
both fascicles have almost smooth margins and no cross striations, thus
differing from those of J. f D . )foruminosu and J. (0.)
steueri. Distribution
(p. 121 and f in Fig. 6) Bermuda, West Indies, Kenya, Ceylon (Knight-Jones,
1972) and Australia (Knight-Jones, Knight-Jones 8t Llewellyn, 1974).
gnomonicus. Spirorbis (Junuu) gnomonicus Bailey (1969a) is a variety of
J. pugenstecheri (p. 111).
heideri. Spirorbis heideri Sterzinger (1909) is almost certainly a junior
foruminosu (p. 1 0 5 ) .
synonym of J. (0.)
karaitivensis, Spirorbis kuruitivensis Pillai (1960) is represented by a single
paratype in the British Museum (Nat. Hist.). This is unmounted and the
operculum is rather decalcified, so detailed comparison with other Junuu
species could not easily be made. Pillai’s figures indicate that his material (from
Ceylon) resembled J. fD.)formosu Bush in having the rim of the opercular
plate sharply depressed in the region of the talon, but that the brood chamber
was less opaque, the abdominal setae blades not quite as wide, and the collar
setae margins not quite as smooth. I t may, however, be the same as
J. (D.)formosu, which has been found in Ceylon on the same substratum, a
marine angiosperm (Knight-Jones, 1972).
kayi. Junuu (Fuuveldoru) kayi Knight-Jones (1972) always showed sinistral
coiling in its three widely-separated Indian Ocean locations, but could not very
well be placed in Leodoru (p. 99), for that would have linked it with
J. knightjonesi, whereas its fused collar folds, opaque brood chamber and bifid
opercular talon show that it is nearer to Dexiospiru. I t then seemed justifiable
to erect Fuuveldoru, but we now favour dropping this taxon, since three species
of Dexiospiru are known to coil both ways (p. 122) and direction of coiling
seems generally of small taxonomic importance (Vine, 1972a; Knight-Jones and
Walker, 1973), whilst J. kuyi is very close to other Indian Ocean Dexiospiru
species. Its opercular talon is remarkably like those of J. (Dexiospiru) steueri,
J. ( D . ) foruminosu and J. (0.)
formosu (cf. Fig. 4 G,H, j and K), whilst its setae
are like those of J. (0.)brusifiensis, but it differs from all of these in having a
domed or conical opercular plate. In that respect it resembles J. (D.)
rumbalugurnensis (Pillai, 1970), but differs in shape of talon and lack of
cross-striations on collar setae (cf. Fig. 4F). From Kenya, Ceylon (amongst
paratypes of Neodexiospiru fuuveli Pillai, 1970) and Kuwait ( = Spirorbis
foruminosus sensu Mohammad, 1971). We have recently found this species in
Mozambique.
knightjonesi. Junuu (Leodoru) knightjonesi (de Silva, 1965) has unfused
collar folds, pin-like talon and transparent, uncalcified brood chambers, thus
8
108
P. KNIGHT-JONES ET
At.
Figure 6. Distribution of Janua species. Black triangles denote places where the genus has
appeared to be absent. Open circles indicate locations for Janua (Janua) pagensrecheri; open
squares J. (Leodora) knightjonesi; open triangles J. (Pillaiospira) rrifircata; triangle enclosing 1.
J. (P.) natalensis; open circles round letters denote species mostly belonging to the subgenus
Dexiospira, viz. a, foraminosa; 5, bmsiliensis: f, form-; k, kayi; I, lamellosa; n, nipponica; p.
preucuta; r, rugata; s, sreueri; t, tambalagamensls; 2. pseudocomcgota. For other details see text
f
44
a
41
. .
I
.-.__
REVIEW OF THE GENUS JANUA
109
40
0
40
110
P. KNIGHT-JONES ET AL.
differing from Dexiospira and resembling J. (Janua) pagenstecheri. I t differs
from the latter in its sinistral coiling, stacked opercula (Fig. 31, p. 100) and
absence of sickle setae. I t is peculiar in that all its mature brood chambers seem
to develop talons. Distribution (p. 121) Ceylon (type locality), West Indies
(Bailey, 1970), Great Barrier Reef and Hawaii (Vine, Bailey-Brock &
Straughan, 1972).
laevis. Spirorbis laevis Quatrefages ( 1 865) was inadequately described from
Guettary, near Biarritz, but Claparede (1870) soon applied the name to a
Naples form which was most unusual in having only two pairs of tentacles.
Otherwise it was a typical Janua, with simple collar setae, an opercular brood
chamber and the uncini of the thorax like those of the abdomen. The setae
seem to have been studied carefully, but sickle setae are not mentioned, so it
may perhaps be inferred that they were absent. The figure of the whole animal,
well-known from its appearance in text-books, looks like a dorsal view of a
dextrally coiling species, with the collar folds fused dorsally. It is in fact rather
like J. ( D . ) pseudocorrugata, except for the small number of tentacles. A small
figure of the tube, however, indicates sinistral coiling. There is no clear record
of anything like this having been seen since, and it cannot be regarded as a good
species without confirmation. It was used as the type for Leodora (p. 99) by
St-Joseph (1894), who did not define that genus distinctly. Chamberlain’s
(1919) statement “tube sinistral” was probably based on Claparede’s figure, for
there seems to have been no other previous reference t o the direction of
coiling. Vine et al. (1072) discuss this problem in more detail.
lamellosa. The original description of Janua (Dexiospira) lamellosa
(Lamarck, 1818) was based merely on the form of the tube which was later
figured by Chenu (1843), but is not very distinctive. Wisely (1962) gave the
first adequate description from Port Hacking, Sydney, basing his identification
on Chenu’s figure. A fuller description (Knight-Jones, Knight-Jones &
Llewellyn, 1974) has been based on material found near Adelaide and on
Wisely’s material kindly loaned by the British Museum (Nat. Hist.). This species
differs from other Janua (Dexiospira species in that the talon has a transverse
bar with bulbous ends, lying just below the deeply concave distal plate
(Fig. 4L). The bifid terminal part of the talon is not closely associated with the
opaque walls of the brood chamber and may be obscured by embryos. I t is
represented by a dotted line in Fig. 4L. J. ( D . ) lamellosa is close to J. (D.)
sreueri and J. ( D . ) foraminosa but the talon is distinctive and it also differs
from the former in having collar setae and abdominal setae of similar size (to
judge from a comparison of rather sparse material), whilst it differs from J. (D.)
foraminosa in that the serrations and cross-striations of the collar setae are
markedly more prominent. The species may be endemic to Australia.
mannarensis. Neodexiospira munnarensis Pillai ( 1970) is a synonym or
variety of Janua (Dexiospiral pseudocorruguta Bush ( 1 904) (p. 1 13).
natalensis. Janua (Pillaiospira) natalensis Knight-Jones & Knight-Jones
(1974) from Uvongo, southern Natal, has an operculum like that of J. (P.)
rriftcrcatu (Fig. 4D-see also p. 116) but differs in that (1) the brood-chamber
walls shatter irregularly; (2) the collar setae blades are coarsely toothed and
cross-striated; and (3) each thoracic uncinus has a single anterior peg.
nipponica. The original description of Janua (Dexiospira) nipponica (Okuda,
1934) from Akkeshi 8c Muroram, Japan, did not deal with the juvenile form.
REVIEW
OF THE GENUS JANUA
111
but this was described by Uchida (1971b). We examined material from Ago
Bay, Honshu, Japan, and found that the juvenile talon, seen also in early brood
chambers (Fig. 4C), is remarkably like that of J. (0.)pseudocorrugata and J.
(D.) brasiliensis. Some of the tubes from Ago Bay, which were attached to a
filamentous bryozoan, had longitudinal ridges perforated by transverse tunnels
(Fig. 5 , p. 114), similar to those in the Australian J. fenestrata (p. 114) which is
perhaps best regarded as a variety of J. pseudocorrugata.
J. nipponica differs from J. pseudocorrugata, however, in having collar setae
with almost smooth margins and no cross-striations, thoracic uncini with three
optically dense anterior pegs of similar length and a more opaque brood
chamber. The distinction between J. nipponica and J. brasiliensis is slight and
indeed we cannot rule out the possibility that the former is a variety of the
latter. In our collections from Japan, however, they formed distinct
populations (p. 122) and could always be distinguished by the difference
between the anterior ends of the thoracic uncini (p. 104). When compared with
material of J. (0.)brasiliensis from other parts of the world this difference may
not be so obvious. Material from South Africa, thought to be J. nipponica
(Knight-Jones, 1973), is best regarded as a variety of J. brasiliensis (p. 96). J.
nipponica has also been found in Hawaii (Vine, Bailey-Brock & Straughan,
1972).
oshomensis. We have not examined the type material of Dexiospira
oschoroensis Uchida (1971a) but judge from Uchida’s detailed description that
this Japanese form is J. (D.)brasiliensis (p. 94).
pagenstecheri. Janua (Janua) pagenstecheri (Quatrefages, 1865) is readily
distinguished from J. (D.) pseudocorrugata (and even more readily from other
Dexiospira species) in that the ampulla beneath the opercular plate is
cupshaped (not cylindrical), with transparent walls completely lacking in
calcification, whilst the talon when present is pin-like (Fig. 3A, p. 100). The
talon may be flattened in a radial direction, but never in a direction parallel to
the opercular wall. In these respects it resembles J. (Leodora) knightjonesi, but
differs in direction of coiling and in possessing sickle setae (Fig. 3G) in the
third thoracic fascicle, whilst the brood chambers differ in that they are usually
single, with opercular plates that often lack talons and may bear distal
processes.
These distal excrescences vary in development and the opercular plates thus
show a variety of shapes (Fig. 3A-E). We have found the full range of
opercular shapes in populations from warm latitudes (Malta, Chios, Cape Verde
Islands, Gulf of Mexico, Galapagos and the mid-Atlantic St Pauls Rocks) and
we agree with Vine, Bailey-Brock & Straughan (1972) that Spirorbis (Janua)
epichysis Bailey (1970) from the West Indies is to be regarded as a mere
variant, as is Spirorbis (Janua) gnomonicus Bailey (1969a) from the Aegean
(Knight-Jones, Knight-Jones & Llewellyn, 1974). We have likewise examined
paratypes of Spirorbis unicornis (Bailey & Harris, 1968) from the Galapagos
and can find no difference between them and the more exaggerated forms of J.
pgenstechen (Fig. 3D) from various warm localities. On the other hand, we
have not found prominent distal opercular processes (e.g. Fig. 3C, D or E) in
the more temperate waters of north-west Europe and southern Australia.
The collar setae of this species have been rightly described as simple but,
rather misleadingly, as having no gap between fin and blade. It should not be
112
p, KNIGHT-JONES ET A L
inferred that there is any distinct basal fin. Stereoscan pictures show that the
blade is rounded, as usual in Spirorbinae, and that the basal region is swollen,
with teeth that are slightly enlarged, but the transition between this and the
rest of the blade is gradual (Knight-Jones, Knight-Jones & Llewellyn, 1974), SO
we agree with Zibrowius (1968) that no fin can be distinguished. From certain
viewpoints the basal swelling may appear in a light microscope t o have a fairly
abrupt boundary and this appearance, though not often seen, has probably
been the basis of figures which show these blades as distinctly differentiated
into proximal and distal parts (Caullery & Mesnil, 1897; Borg, 1917; Rioja,
1923; Bergan, 1953; Gee, 1964; Bailey, 1969b; Vine, Bailey-Brock &
Straughan, 1972).
Gee (1964) observed two colour forms of J. pagenstecheri in South Wales,
an orange and brown form from rocky shores and a pale sublittoral form,
which he called var. incoloris. We find that the highly pigmented form is
particularly characteristic of rock pools on British west coasts and in Brittany.
Since the type locality for J. pagenstecheri is the Golfe du Lion, and since all
the material of this species which we have observed from there and other
Mediterranean localities has been pale, it seems inappropriate to refer to var.
incoloris as though it were in that respect atypical. The orange form is further
from the type and may be called forma aurantiaca.
We have found these forms in different zones of the same rocky shores,
particularly at Dale Point, Pembrokeshire, South Wales, and Spiddal, near
Galway, Eire. Their colours were matched against those in the Methuen
Handbook (Kornerup & Wanscher, 1963) with the following results:-
forma aurantiaca (a) base of tentacles and opercular stalk orange 5A6-6A7;
(b) thorax deep orange 6A8-7A8; (c) stomach dark brown 7E4, with trace of
dull violet 15E5; (d) abdomen orange 5A6-6A7; (e) embryos grey-orange 5B6,
but changing during development:
forma incoloris (a) colourless; (b) pale grey-orange 5B6 or grey-yellow 4B3;
(c) brown 5E5 or pale brown 4C5; (d) pale grey-orange 5B6 or grey-yellow
4B3; (e) embryos red-grey 10B2 developing to green-grey 2C3-29B2.
On these shores forma aurantiaca is abundant, but only in shallow pools high
in the tidal zone. The tube is often evolute, without longitudinal ridges, but we
could find no other morphological peculiarities. F o r m incoloris is widespread
throughout the lower shore and sublittoral. There is some overlap and some
individuals have intermediate colouration, but the general zonation is clear. The
pigmentation and uncoiling of the tube may, however, be ecophenotypic, at
least in part. Similar pigmentation may be seen on South African shores in
Paralaeospira levinseni and Paralaeospira patagonica, which are red-orange in
sunlit situations, but relatively colourless in shaded crevices. The pigment may
perhaps be developed as a protection against solar radiation.
Zibrowius ( 1968) has discussed the problem of “Spirorbis” pusilloides Bush
(1904) nom. nov. pro Mera pusilla Saint-Joseph (1894) and agrees with most
authors that this is a synonym of J. pagenstecheri. This species may show
inverse coiling. Two specimens of J. pagenstecheri from the north east coast of
Costa Rica were sinistral and the Mediterranean material called Spirorbis hevis
by Zibrowius (1968) may well have been this species (p. 99).
The distribution (open circles in Fig. 6) is almost world-wide in warm and
REVIEW OF THE GENUS JANUA
113
temperate latitudes, probably as a result of transport from a centre in western
Europe (P. 123), sometimes no doubt by shipping (p. 124). Remarkably enough,
it has not been recorded from Japan, northeast America or the tropical east
coasts of Africa. Although Pixel1 (1912) recorded it (as pusilloides) from near
Vancouver, Potswald (1965) was unable to find it in that area.
pantulus. We have examined two paratypes of Spirorbis (Janua) parvulus
Bailey (1969a), kindly made available by Dr Bailey-Brock, and found them
remarkably like small specimens of J. (0.)pseudocorrugata (p. 114). Each had
the typical coarsely serrated collar setae, long thin abdominal setae and a
mature stage operculum (without talon) with lightly calcified walls and a very
small distal rim. One of the opercula had two embryos as figured in the original
description. We have sometimes found specimens of J. (0.)pseudocorrugata
with as few as three embryos, and feel that the marginally smaller number of
embryos cannot be regarded as sufficient grounds for separating parvulus from
J. (D.1 pseudocorrugata, which was recorded from the same collecting stations
at Chios (Bailey, 1969a).
preacuta. The type mount and paratypes of Janua (Dexiospira) preacuta
Vine (1972a) from the Red Sea were kindly loaned by the British Museum
(Nat. Hist.). Unfortunately the opercula were decalcified. We also examined
turrita Vine (1972b) from Hawaii, kindly loaned by Dr
paratypes of J. (0.1
Vine. The latter species was separated from the former because it bore rough
protuberances on the ridges of the tube. Such ornamentation can vary within a
single species however (see pp. 110 8c 114), and a close comparison revealed no
other differences, so we consider J. (D.) turrita t o be a junior synonym of J.
(D.1 preacuta.
This species has an extraordinary long talon, sometimes so long that it is
curved in side view (Fig. 4E, 01) to conform with the curve of the tube.
Judging from the general morphology, its closest relatives seem to be J. (0.)
pseudocorrugata and J. (0.)steueri. The talons of the former, though variable
in shape, differ in being quite short. Those of the latter are always bifid
terminally and bear large angular lateral wings. J. (0.)
preacuta (both Red Sea
and Hawaiian material) differs from both species in having a remarkably long
asetigerous region, whilst the abdominal setae and collar setae seem to be much
smaller relative to the size of the operculum than in J. (D.
1 pseudocorrugata, to
judge from three specimens in which this character was checked.
pseudocormgata. The first useful descriptions of Janua (Dexiospira)
pseudocorrugata (Bush, 1904) nom. nov. pro Spirorbis corrugatus Caullery &
Mesnil (1897) non Montagu non Langerhans, were given by Sterzinger (1910)
and Rioja (1923). These relate to a form which extends, according t o our
observations, from the type locality (north-west France) to the Mediterranean
and as far north as Gulland Rock (Padstow, Devon) and Galway Bay (Eire). It
is always attached to marine plants and has a rounded tube without conspicious
ridges. The juvenile talon is usually thin and almost peg-like, so this form may
at first glance be confused with J. pagenstecheri, until its collar fusion and lack
of sickles are established. The talon is somewhat broadened tangentially,
however, whereas that of J. pagenstecheri is rounded or broadened radially.
Sometimes the talon is considerably broadened, so we cannot distinguish this
form specifically from those which follow.
With this in the Mediterranean there is another form which is common in
114
P. KNIGHT-JONES ET A L .
most warm seas. We propose to call it var. mannarensis, for comparison with
paratypes shows that this is the same as that named Neodexiospira mannarensis
Pillai (1970), before it was made clear that “Spirorbis corrugatus” had fused
collar folds (Knight-Jones, 1972). In fact this is the most widespread form of
the species and the best known, following the accounts of Zibrowius (1968),
Harris (1968), Bailey (1969a, 1970), Vine (1972a) and Vine, Bailey-Brock &
Straughan (1972). Like the other varieties which follow, this is usually attached
to stones or shells and much less frequently to plants. The tube usually has
three longitudinal ridges and a peripheral sloping flange. The talon is
comparatively broad, often bilobed and somewhat triangular (Fig. 4A, 0 2 and
03), thus resembling the talons of J. (D.) brasiliensis and J. ( D . ) nipponica.
Both the latter, however, have finer marginal serrations on the collar setae and
more opaque brood-chamber walls. We therefore regard J. pseudocorrugata as
distinct, rather than as a junior synonym of J. brasiliensis (p. 00), whilst J.
rzipponica differs from both in having trifurcate anterior pegs to the thoracic
uncini.
Neodexiospira benhami Pillai (1970) is also indistinguishable from this form.
The original description suggested that N. benhami lacks a juvenile form, but
we found two amongst the paratypes, with talons typical of J. pseudocorrugata
var. mannarensis. We have examined material of this variety from the
Mediterranean, West Indies, Mozambique, Ceylon, Hawaii and Catalina Island
(off California).
Pending further information, we are inclined to regard Spirorbis (Janua)
parvulus Bailey (1969a) as another variety of this species (p. 113) exemplifying
the “nanisme oriental” noted by Perks & Picard (1958) in other Aegean forms.
The juvenile operculum is unknown. The tube is smooth, with the mouth
relatively broad and overlapping, so that only a single whorl is visible.
Some Indo-West Pacific populations show another trend in tube
modification which seems worth characterizing. We designate as var. praeceps
our material from Hakodate, Hokkaido, Japan, which invariably has the
outermost of the three longitudinal ridges steeply projecting and almost
overhanging the peripheral slope (Fig. 5C). The only other differences between
this and var. mannarensis were that the talon seemed always t o be short
(Fig. lA, 04) and the uncini somewhat more numerous in all the thoracic tori
except the first on the convex side. The single paratype of Spirorbis
(Dexiospira) ceylonicus Pillai (1960) has a tube showing the characters of var.
praeceps, though the original description does not make this feature clear. It
showed a rather transparent mature operculum, as in J. pseudocorrugata, but
did not describe the important juvenile form with talon (and we were reluctant
to open the only tube available). It also showed an abdominal seta and collar
seta which were similar in size, but it is possible that an atypical abdominal seta
was figured, for it must be admitted that an atypical abdominal seta was
figured for J. ( D . )fenestrata (Knight-Jones, 1973) from South Australia.
The latter is best regarded as var. fenestrata and very close to var. praeceps.
Its tube is similar, but has numerous holes perforating the longitudinal ridges.
The holes are no more than deep extensions of the transverse intercostal
grooves, which are already seen in var. mannarensis but become more obvious
in var. praeceps. for they make deeper indentations as the longitudinal ridges
become more prominent. A similar trend leading to perforated longitudinal
REVIEW OF THE GENUS JANUA
115
ridges is to be seen in J. (D.) nipponica (Fig. 5A). This is one reason why we
feel that tube differences must be regarded as of no more than varietal
importance, unless supported by other evidence. Certainly tubes are unlikely to
be as taxonomically important as setae, for differences in setal structure must
be determined very closely by the genetic control of their development, since
this takes place within the protection of the tube, whereas tube differences
must result from complex interactions between the structure of the collar
region, the behaviour of the whole animal, and the nature of the substratum
and general environment. The setae, in fact, must reflect peculiarities of the
gene complex rather directly, whereas tube development is at least one step
further removed from genetic control.
All these varieties seem to have similar setae, but another form, which should
perhaps be regarded as subspecies ainu, has the shaft of each abdominal seta
remaining peculiarly narrow at its junction with the blade (Fig. 4A, as 1). This
agrees with Dexiospira ainu Uchida (1971a), which is otherwise a typical J.
pseudocorrugata. The tube is unremarkable, like that of var. mannarensis.
Sometimes the median ridge is much more prominent that the others (Fig. 5D).
We have found this form abundantly in our collections from Honshu and also
from Sydney in Australia. In Australian material from Kangaroo Island, near
Adelaide, it w a s found on the same shore as var. fenestrata and on similar stony
substrate. Nevertheless the two populations seemed to remain distinct; further
studies in shore ecology may well promote some of these varieties t o specific
rank.
The present picture of distribution, which is still very sketchy, therefore
shows this species t o be represented in the warm waters of all oceans by var.
munnarensis, with the other varieties branching out in cooler latitudes. The
taxonomic type is confined to Europe and is characteristically Lusitanian. In
the temperate west Pacific the species extends north to Honshu as var.
praeceps, south t o Adelaide as var. fenestrata and nearly as far in both
directions as subspecies ainu.
pusilloides. Spirorbis pusilloides Bush (1904) nom. nov. pro. Mera pusilla
Saint-Joseph (1894) cannot be distinguished from Janua pagenstecheri (p. 11 1).
rugatus. The brief description of Spirorbis rugatus Bush (1904) indicates the
genus Janua and is reminiscent of J. (0.)pseudocorrugata. Although one figure
shows the wall of the cylindrical brood chamber t o be rather shallow (and
probably transparent), this might depict an incomplete stage of development.
The locality (southern Alaska) is far north for a record of J. pseudocorrugafa,
but receives the warm current from the west Pacific (see Fig. 6). Moore (1923)
records this species off California. We have not examined his material, but we
have found J. pseudocorrugata sparsely on stones collected from Californian
shores by Dr Dale Straughan and Prof. R. P. Dales. If these two east Pacific
forms are the same, J. rugata would be senior synonym (page priority), with
the further advantage of brevity, but it might alternatively be regarded as a
nomen nudum, the type material consisting of a minute fragment of decalcified
tube (Hartman, 1942).
stared We have not examined type material of Janua (Dexiospira) steueri
(Sterzinger, 1909) from Suez, but other Red Sea material kindly loaned by Dr
Vine (Vine, 1972a) agrees well with Sterzinger's description and with our
material on plants, stones and ships' hulls from Kenya (Knight-Jones, 1972),
116
Y. KNIGHT-JONES ET
AL.
Australia (Knight-Jones, Knight-Jones & Llewellyn, 1974), West lndies (Bailey,
1970), New Zealand (Vine, 1975), Vietnam (Fauvel, 1935, as foruminosu),
Mozambique (lnhaca) and Tenerife (Playa de Teresitas, near Santa Cruz).
Material from Hawaii, which was thought to be J. ( D . ) steueri (Vine,
Bailey-Brock & Straughan, 1972), we found to be J. fD.)foruminosu. These
two species are indeed easily confused, but J. ( D . ) steueri differs in having a
wider more opaque brood chamber, with a smaller distal rim, and collar setae
which are much more coarsely serrated and cross-striated on the convex side
and are usually smaller than the long slender abdominal setae (cf. Fig. 4H,J).
The operculum of J. ( D . ) steueri is even more similar to that of J. ( D . )formosu,
but the setae are quite different. The collar setae of J . (D.)formosa have almost
smooth margins and no cross striations, and the abdominal setae are
characteristically wide (compare Fig. 41, K). It is interesting to note that both
these closely related species can be found together in the West Indies, Kenya
and Australia.
S. treudwelli Pillai (1965) from the Philippines is almost certainly a junior
synonym of J. ( D . )steueri (see below).
tambalagamunsis. Paratypes of Juiiiia (Dexiospird tumbulagumensis (Pillai,
1970) from Ceylon were kindly loaned by the British Museum (Nat. Hist.) and
one specimen was mounted to show the setae. J. ( D . ) pseudocorrugutu is
probably its closest relative. Both species have collar setae with fairly coarse
serrations and cross-striations, very long slender abdominal setae and a
somewhat similar talon. The talon of J . ( D . ) tumbulugumensis, however, is more
bulbous and the marginal teeth of the abdominal setae seem to be finer
(compare Fig. 4A and F). The most characteristic feature of J . (D.)
tumbulugurnensis is the opercular plate which is domed, very thick and glassy. It
closely resembles the opercular plate of the sinistral species J. (Fuuveldoru)
kayi. but this species differs in having a markedly different talon (long and
bilobed with angular lateral wings), collar setae with almost smooth margins
and no cross-striations (compare Fig. 4 F and G).
treadwelli. A paratype of Spirorbis treudwelli Pillai (1965), from the
Phillipines kindly loaned by the British Museum (Nat. Hist.) was mounted and
examination showed the collar setae to be very coarsely serrated (and cross
striated) and the abdominal setae to be narrow and longer than the collar setae.
In these and other respects it was indistinguishable from J. (0.)steueri.
trihrcata. Januu (Pilluiospira) trifurcutu Knight-Jones (1973) from
south-west Australia differs from all other Januu species in having the following
aggregate of characters. The anterior pegs of the thoracic uncini are strongly
trifurcate, the lateral teeth being as large and as optically dense as the central
one; the collar setae on the convex side have almost smooth margins and lack
aoss-striations; the juvenile operculum is bilaterally asymmetrical (Fig. 4D);
the mature opercular brood chamber (without talon) smashes into pieces like
the staves of a barrel (see original description); no specimen examined had a
fused thoracic membrane over the dorsal groove.
I t resembles J. (0.1
nipponicu in having similar uncini and collar setae, but it
differs in the other characters mentioned above.
The mature operculum and lack of collar fusion recalls a species from
Madeira tentatively but mistakenly identified as Spirorbis corrugatus by
REVIEW OF THE GENUS JANUA
117
Langerhans (1880). The details of the juvenile operculum and uncini are not
known but the collar setae seem to have more coarsely serrated margins.
J. (P.1 natalensis (p. 110) from South Africa has a similar juvenile operculum
and usually has the thoracic membrane split dorsally (a few juvenile specimens
had it fused) but differs in the other three respects. For a discussion of this
subgenus see p. 100.
turrita. For Janua (Dexiospira) turrita Vine (1972b) see J. preacuta (p. 113).
unicornis. For Spirorbis unicornis Bailey (Bailey & Harris, 1968) see J.
pagenstecheri (p. 111).
SOME OTHER TAXA OF SPIRORBINAE
It may be useful to deal now with other groups which possess simple collar
setae and have therefore been linked from time to time with those defined
above. Circeis St Joseph seems particularly worthy of attention. Following
Chamberlain (1919) and Pillai (1970), this would appear to be the best generic
name for the well-known and widespread species “Spirorbis” spirillum (L.),
yet other authorities (Hartmann, 1959) have regarded this as the type species
of Dexiospira. Circeis was in fact based upon Circeis armoricana St-Joseph
(1894), which is another well-known form. Caullery & Mesnil (1897) pointed
out that it might reasonably be regarded as a mere variety of Spirorbis spirillum
and is almost certainly the same as Spirorbis sinistrorsus Montagu (1803),
which was however inadequately described. Chamberlain therefore cited the
Linnaean species as type when he reinstated the genus Circeis, He gave no
formal definition, so we propose one here, augmenting those of Rioja (1942)
and Pillai (1970) and using information from Bergan (1953), Gee (1964),
Bailey (1969b) and our own studies of material from many localities in Europe,
America and Japan, including the brooding arrangements (Knight-Jones,
Knight-Jones & Vine, 1973) and more recent stereoscan studies of the setae.
We find that Circeis armoricana is distinct from C. spirillum so should
presumably be regarded as the type. It lacks collar seta crossstriations, which
are present in the Linnean species.
We must also consider the sinistral forms which are still associated with the
name Leodora by some authorities (p. 99), but should rather be distributed
between the genera Romanchella and Sinistrella (see below). Romanchella was
redefined recently (Knight Jones, 1973) but was then thought to include
several northern hemisphere forms which were nevertheless different in
important respects from the Southern Hemisphere type species. For these the
subgenus Bushiella was proposed, with Spirorbis evolutus Bush as type, t o
include Spirorbis asperatus Bush, Spirorbis coronatus Zachs and Spirorbis
medius Pixell. On examining further material however, we have found that this
group should undoubtedly include Spirorbis validus Verrill and Spirorbis
abnormis Bush (=Leodora multipluta Uchida, 1971b) and that these two
species are opercular incubators. Bush (1904) could find “no difference . . . in
the essential characters” between S. validus and Spirorbis verruca Fabricius,
whilst Chamberlain (1919) designated the latter type species of his genus
Sinistrella, in which he also placed S. media. The first good definition of the
118
P. KNIGHT-JONES ET
AL.
group was given by Pillai (1970) under the name Protoleodora, but the name
Sinistrella has priority.
Since we had also accumulated material of all other well-established genera
and most species of Spirorbinae, through the kindness of several individuals and
institutions (p. 125), we thought ‘it useful to try “numerical taxonomy”
comparisons with the taxa redefined here. We used the admittedly small
number of characters ( 1 to 16) indicated below in the first diagnosis (of
Circeis), without any weighting but with appropriate value judgements (e.g.
when considering tube translucency, and what proportion of individuals and/or
species show dextral or sinistral coiling). The calculated percentage similarities
to other taxa are given after diagnosis. Diagnoses of most of these other taxa
are given in Knight-Jones (1972), Knight-Jones & Walker (1972), Vine (1972),
Knight-Jones (19731, Knight-Jones, Knight-Jones & Bregazzi (1973), and
Knight-Jones, Knight-Jones & Llewellyn (1974).
Genus Circeis St-Joseph 1894
Coiling (1) usually dextral; material of tube (2) usually porcellaneous and
more or less translucent; collar folds ( 3 ) not fused dorsally; collar setae
flat-sided and lacking any fin (4) of larger teeth, more or less sharply
geniculate, with a few rows of small teeth extending along the blade; these are
fairly regular, and may show cross-striations (5); sickle setae (6) absent; only
two pairs of thoracic tori (7); thoracic uncini with a broad somewhat square
anterior peg (8) and with numerous teeth which are arranged in transverse rows
(9) and also in at least four longitudinal rows (10); blades of abdominal setae
( 1 1 ) and collar setae somewhat similar in size and shape; abdominal uncini
numerous on concave side and sparse on convex side (12); embryos incubated
within the tube ( 1 3 ) , adhering individually to the tube wall (14) and to their
neighbours in about two irregular rows; larvae without any white attachment
glands (15); there is no tendency t o retain earlier opercular plates (16).
Type. Circeis armoricana St-Joseph. The percentage similarities of Circeis to
other taxa were calculated to be:
Paradexiospira s.s . . . . . . 85 Janua(Pil1oiospira) . . . . . . 46
Paradexiospira (Spirorbid es) . . 76 Janua (Dexiospira) . . . . . . 43
Eiclaeospira . . . . . . . . . 66 Janua s.s . . . . . . . . . . 41
Anomalorbis Vine (1972) . . . 60 Pileolaria (Simplicaria) . . . . 40
Spirorbis including Spirorbella . 58 Protolaeospira s.s . . . . . . 39
Romanchella . . . . . . . . 52 Janua(Leodora) . . . . . . 38
Helicosiphon . . . . . . . * 5 1 Sinistrella . . . . . . . . . 3 6
Protolaeospira (Dextralia) . . . 49 Pileolaria s.s . . . . . . . . 34
Me talaeospira . . . . . . . * 49 Pileolaria (Ouplicaria) . . . . 28
Paralaeospira . . . . . . . . 49 Amplaria Knight-Jones (1973) . 21
Circeis is thus remarkably close to Paradexiospira violacea (Levinsen), which
was designated type species of Paradexiospira Caullery & Mesnil (1897) by
Chamberlain (1919), and is sufficiently distinct from its only two known
relatives Paradexiospira cancellata (Fabricius, 1780) and Paradexiospira vitrea
(Fabricius) to justify their separation under the name Spirorbides Chamberlain
(lYl9), which is here regarded as a subgenus. This disposition of the three
REVIEW OF THE GENUS JANUA
119
species has priority over that proposed by Pillai (1970). All three have subarctic
distributions, like Circeis.
Genus Romanchella Caullery & Mesnil 1897
Coiling mostly sinistral; tube often porcellaneous but opaque white; collar
folds fused to form a tunnel over the mid-dorsal groove; collar setae almost
straight (not strongly geniculate), lacking fins and with blades bearing small
teeth and no cross-striations; sickle setae present in third fascicles; only two
pairs of thoracic tori; thoracic uncini with a broad somewhat s uare anterior
peg and with numerous teeth arranged in oblique rows (about 45% ); abdominal
setae less than a third of the size of the collar setae, strongly geniculate and
with short, rapidly tapering blades, often accompanied by hooked capillary
setae; abdominal uncini numerous on concave side and sparse on convex;
embryos incubated within tube, forming an oval egg mass which is attached
anteriorly to the body, by a stalk which arises within the mid-dorsal groove but
eccentrically, on its concave side; larvae probably lack any white attachment
glands; opercular plates tend to be retained after moulting in some species.
Type. Spirorbis (Romanchella) perrieri Caullery & Mesnil 1897. The
percentage similarities of Romanchella were t o
Metalaeospira . . . . . . . 69 Janua(Leodora) . . . . . . 49
Helicosiphon
. . . . . . . 68 Circeis . . . . . . . . . . 48
?+o tolaeospira (Dextralia) . . . 67 Pileolaria(Dup1icaria) . . . . 47
Eulaeospira . . . . . . . . 63 Janua S.S. . . . . . . . . . 44
Protolaeospira s.s . . . . . . 61 Janua (Dexiospira) . . . . . . 43
. . . . . . . 43
Paralaeospira . . . . . . . . 61 Pileolaria s.s
Paradexiospira
s.s
. . . . . . 43
.
.
.
58
Anomalorbis . . . . .
Janua(Pil1aiospira)
. . . . . 41
Amplaria . . . . . . . . . 57
Pileolaria
(Simplicaria)
. . . . 41
Spirorbis + Spirorbella . . . . 52
Sinistrella . . . . . . . . . 51 Paradexiospira (Spirorbides) . . 3 7
Evidently the genus is a distinctive one but has affinities with the other six
groups of Southern Hemisphere tube-incubators (the first six listed above). Five
species, all sinistrally coiling, have been described by Harris (1969),
Knight-Jones (1973) and Vine (1975), and we have recently encountered
another, which is dextral. All seem to be restricted to cool southern seas.
Genus Sinistrella Chamberlain 1919
(=Protoleodora P illai, 1970)
Coiling always sinistral; tube opaque white; collar folds unfused; collar setae
slender and almost straight, the blades bearing fine teeth and no cross-striations
or separate fins, though the wider basal part may show a gradual differentiation
aver a small area, which may include a slight indentation not separating it
clearly from the rest of the blade; sickle setae present in third fascicles; only
two pairs of thoracic tori; thoracic uncini with a broad gouge-shaped anterior
peg and very slender, most of the length of the largest bearing only one or two
rows of comparatively large teeth ; abdominal setae obliquely geniculate,
tapering uniformly like a pennant, and about half the size of the thoracic setae;
120
P. KNIGHT-JONES€7-
AL.
numbers of abdominal uncini more or less similar on left and right sides;
incubation in an opercular brood chamber which often retains the protection
of one or more older opercular plates, always lacks any secondary plate below
the embryos, and has an opening on the side facing the tentacles; larva with a
single white attachment gland in the abdomen.
Type. Spirorbis verruca Fabricius (March, 1863 ; this may =Spirorbis validus
Verrill in Smith & Harger, 1874).
The percentage similarities of Sinistrella were to
Pileolaria (Duplicaria) . . . . 87 Paralaeospira . . . . . . . . 5 5
A’leolaria 5.s . . . . . . . . 83 Protolaeospira(Dextra1ia) . . . 5 3
Pileolaria (Simplicaria) . . . . 81 Protolaeospira s.s . . . . . . 5 1
Amplaria . . . . . . . . . 79 Anomalorbis . . . . . . . . 5 1
Spirorbis + Spirorbella . . . . 68 Eulaeospira . . . . . . . . 49
Janua (Leodoral . . . . . . . 65 Janua(Pil1aiospira) . . . . . 47
Romancltella
. . . . . . . 61 Janua(0exiospira) . . . . . 44
Metalaeospira . . . . . . . 61 Circeis . . . . . . . . . . 41
Janua s.s . . . . . . . . . 60 Paradexiospira . . . . . . . 3 7
Helicosiphon . . . . . . . . 56 Paradexiospira (Spirorbidesl . . 3 3
Sinistrella is thus so close to Pileolaria that it might be regarded as a mere
subgenus, particularly as the figures above are not influenced by a resemblance
of the brood chamber of S. valida and S. abnormis to those of Pileolaria
granulata (L.) and Pileolaria quadrangularis Stimpson. The two latter have a
subarctic distribution similar to that of Sinisrrella.
We are reviewing the genus Sinistrella and think that some of the seven
species mentioned above (p. 117) may be synonyms. We have not yet found
brooding opercula in Sinistrella species other than S. valida and S. abnormis.
Instead we find egg masses distending the thin skin and initially believed,
probably wrongly, that these were incubated within the tube. Indeed, we have
not yet ruled out the possibility that some of these species have evolved
coelomic incubation, as an adaptation for larger brood size in high latitudes.
The group seems to be confined to the north Pacific and north-west Atlantic.
Finally, the percentage similarities of Janua sensu strict0 to other taxa are:
Janua (Pillaiospira) . . . . . 91 Pileolaria (Simplicaria) . . . . 48
Janua(Dexiospira)
. . . . . 84 Protolaeospira(Dextra1ia) . . . 48
Janua(Leodora) . . . . . . 83 Metalaeospira
. . . . . . . 47
Anomalorbis . . . . . . . . 72 Circeis . . . . . . . . . . 47
Sinistrella . . . . . . . . . 63 Paradexiospira . . . . . . . 44
Spirorbis + Spirorbella . . . . 56 Romanchella
. . . . . . . 48
Pileolaria s.s . . . . . . . . 5 5 Paradexiospira (Spirorbides) . . 43
.4mplaria . . . . . . . . . 5 3 Helicosiphon . . . . . . . . 41
Eulaeospira . . . . . . . . 5 2 Paralaeospira . . . . . . . . 3 9
A’leolaria (Duplicaria) . . . . 5 1 Protolaeospira s.s . . . . . . 32
The only species which would appear to be fairly closely related to Janua is
Anomalorbis (Vine, 1972a), which is so little known that only ten characters
could be used in the comparisons involving it.
REVIEW OF T H E GENUS JANUA
121
WORLD DISTRIBUTION
The foregoing data on distribution of Janua species (pp. 93-117)are
summarized in Fig. 6,which also shows the surface circulation of the oceans.
Warm currents are indicated by complete arrows, cold currents by interrupted
ones. An equatorial belt of unshaded water, where the surface temperature
never falls below 20°, is bounded by two shaded belts, where it never falls
below 15". These isotherms are taken from Sverdrup et al. (1942).Minimum
temperatures are chosen as likely to be particularly critical for this
predominantly tropical genus. Black triangles are evidence that the genus may
be absent, each indicating a locality where collections of Spirorbinae have been
made without discovering any species of Janua. A small black triangle indicates
that from 1 to 3 species of other genera were present, whilst a large one
indicates that more than three were collected, from an area where the
Spirorbinae are thus quite well known.
The most widely distributed species of Janua is J. pagenstecheri (empty
circles). It is found in all oceans and extends from the equator to (in Norway)
latitude 70". Warm water populations often include the variants gnomonicus.
epichysis and unicornis (p. 111). In the Southern Hemisphere it seems to be
uncommon and inconspicuous, but in Britain it is extraordinarily common and
widespread, settling abundantly not only on stones and shells, but on certain
algae (e.g Corallina and Laminaria). I t occurs at high shore levels and
considerable depths, and it is more resistant than any other of the British
Spirorbinae to turbidity, insolation and freezing (Gee & Knight-Jones, 1962;de
Silva, 1967). In the Aegean, however, its percentage contribution to
Spirorbinae populations in very shallow water was found to be comparatively
small (Bailey, 1969a),whilst its scarcity in Galapagos collections led to the
suggestion that it may perhaps become more sublittoral in the tropics (Bailey &
Harris, 1968). At the Cape Verde Islands and South Africa we have found it
only on the undersides of large stones bedded in gravel, whilst in New Zealand
it has been found so far only at depths greater than 30 m (Vine, 1975).We
have never found it on plants in warm latitudes, but always on stones and
shells. All this suggests that it is primarily a boreal form, which has crossed the
tropics by becoming strongly photonegative and has not re-emerged to face the
sunshine of the south, perhaps because it has not yet had time to evolve new
behaviour patterns, but more probably because of competition from the
successful and strongly photopositive southern species Paralaeospira levinseni
Caullery & Mesnil. Recent immigrants do not necessarily dominate a native
fauna. Amongst Spirorbinae, one of the most likely candidates for immigrant
status is Paralaeospira malardi Caullery & Mesnil, but this is one of the least
conspicuous of British species, though it is quite widespread in Britain.
Considering the other small subgenera, J. (Leodora) knightjonesi,
represented in Fig. 6 by open squares, is perhaps the mos-t restricted of all these
forms to warm seas and indeed t o coral or mangrove areas, but in such areas it
Seems to be practically world-wide. The other monotypic and somewhat
dubious subgenus Fauveldora (circle enclosing k), is known only from the
Indian Ocean, whilst F'ilkiospira (represented by white triangles and two
species) is known so far only from Africa and Australia.
In the major subgenus Dexiospira the species problem is particularly
122
P. KNIGHT-JONLS E7’ AL.
difficult. There is of course the possibility that several or all of the forms here
recognized represent one or more polymorphic supcrspecies. One point in
favour of such a view is the evidence for parallel mutations to be seen, for
instance, in (a) the tube-sculpturing of some J. nipponica (p. 110), which
resembles that of J. pseudocomgata var. fenestmta (p. 114), and in (b) the situs
inversus mutations of the Red Sea region, which have affected at least three
apparently distinct forms there. Stcrzinger (1909) first recognized this
phenomenon at Suez in “heideri” (which we must now call foraminosa) and in
steueri. Vine (1972) confirmed it for steueri and recognized another sinistral
form anticorrugata (here regarded as a situs inversus main of bresflfensis).
Sinistral forms are otherwise unknown in Dexiospim and indeed in Janua, apart
from the Indian Ocean J. (Fauveldora) kayi, the circumtropical J. (Leodora)
knightionesi and occasional individuals of J. pagenstecheri. It does not
necessarily mean, however, that the Red Sea forms constitute a single
interbreeding complex, for they may well be separate species responding
similarly to warm conditions, for of course parallel mutations may occur in
forms which are related but distinct.
Another kind of evidence, that may suggest interbreeding, is the existence of
forms that are in some respects intermediate between two other t a n . The
forms J. (0.)formosa and J. (0.l steueri, for instance, are closely similar in
distribution, not only geographically, but sometimes ecologically. Both occur
together on Cymodocea off Kenya. They are particularly distinct from one
another in setal characters, but the setal characters of J. (0.)fominosa are
intermediate between them. It is scarcely conceivable, however, that this could
be due to crossing, for the three forms do not form a continuum. Each seems
to be distributed independently of the others. It is noteworthy, moreover, that
in distribution fomm’nom is markedly different from the other two. Its
extension to cooler latitudes is greatest in the Northern Hemisphere (i.e. Japan
and Hawaii), whilst the reverse is true of the others. It rarely overlaps with the
other two, so we are reminded of the rule that closely related species are never
sympatric (Mayr, 1942). but this has exceptions, of course, and fomminosa
overlaps with sfeueri at Suez, with formom off Ceylon and with both in the
Caribbean.
Another form which is in some respects intermediate between formow and
steueri is h y i . All three of these occur in the Mida Creek, Kenya, yet once
w i n Mayr’s rule seemed to apply to our collections, but on a smaller
ecological scale rather than a geographical one, for the intermediate form was
found on a separate angiosperm EnhaZus.
The pseudocomgata-brasiliensbnipponica group is particularly compact
taxonomically (A, B, C in Fig. 4). yet all three members are found in Japan (2,
b, n in Fig. 6). whilst the former occurs with nipponiuz in Hawaii and with
brasiliensis in South Africa and the Caribbean. Where they overlap these forms
seem to remain ecologically distinct. J. pseudocomgata occurs on stones, as it
d o a in most warm seas, whereas nipponicrr in Japan and Hawaii generally
occurs on algae, whilst brariuensis occurs in a marine angiosperm at Hokkaido,
but on algae in South Africa (where nipponica is not found) and in Brazil. The
only region w h m we have found pseudomrrugata occurring commonly on
algae is Europe, which is outside the temperature range of other Dexiospira
species. Our material from the Atlantic coast (Portugal to Ireland) has been
REVIEW OF THE GENUS JANWA
123
exclusively on algae. It seems that where physical conditions become difficult
for the species, towards the northern limit of its distribution, it is able t o
survive only as a result of larval skill in selecting algal substrata, for these
characterize suitable habitats. In warm waters there is less need for this and less
advantage, since such substrata are also favoured by more exclusively tropical
species of Dexiospira. In setal characters pseudocorrugata is almost
indistinguishable from steueri, and where these overlap they are separated
ecologically by preference of the latter for plant substrata. In favouring plants
of temperate regions, but shunning those of the tropics, pseudocorrugata
exactly parallels pagenstecheri. Probably both have temperature optima which
are warm-temperate rather than tropical, so they must avoid the intense
insolation of many tropical plants.
Considering how these patterns of distribution may have arisen, we have seen
that even a widely distributed species such as J. pagenstecheri may show
evidence of its centre of origin, for off Europe it reaches its maximum
extension into cold latitudes, depth ranges and diverse habitats (p. 121),
suggesting that prolonged occupation of the north-east Atlantic has allowed
time for appropriate adaptations to occur. The Dexiospira group, on the other
hand, displays its greatest taxonomic diversity in the Indo-West Pacific. In the
West Pacific region, moreover, several otherwise tropical species display their
maximum adaptation to cooler latitudes, foraminosa to the north, formosa and
steueri to the south, brasiliensis in both directions.
As for methods of dispersal, most Janua may attach themselves to algae,
including some of those algae which float when detached. In Europe
pseudocorrugata is often found on Cystoseira, whilst pagenstecheri sometimes
occurs sparsely on Fucus vesiculosus, but we have seen that these two most
widely distributed of Spirorbinae are never found on algae in tropical waters.
Probably their wide distribution has been brought about by long distance
transport of shells lodged in the holdfasts of large floating algae such as
Ecklonia and Macrocystis. Near Sydney (Australia) pseudocorrugata is often
found on the algal part of such holdfasts. Self-fertilisation is possible, in
pagenstecheri at least (Gee & Williams, 1965), so even a single adult thus
transported to a littoral pool may establish the species in a new locality. Other
widely distributed species (J. brasiziensis and steueri) build up large breeding
populations on Surgassum in shore pools at, for instance, Mossel Bay (South
Africa) and Kangaroo Island (Australia), whilst J. formom is the common
species on Sargassum drifting near Bermuda. The last three species may perhaps
have rounded Africa and crossed the Atlantic in that way, though transport by
the old sailing ships is another possibility.
It seems that remote transport through some agency must have enabled
formosa, foraminosa and steueri to by-pass Africa, for they could scarcely have
rounded the Cape by gradual extension of their ranges, in conditions like those
of today. Careful surveys of South African shores failed to find them anywhere
south of Mozambique, not even in pools or sheltered places, like Knysna or
Saldanha Bay. In Africa their adaptation to cooler latitudes is much less than in
Australia, perhaps because of lack of time, or scarcity of sheltered
embayments, or competition with Paralaeospira, a genus which is very
abundant on the shores of South Africa, South America and New Zealand, but
seems to be absent from Australia. This was the only genus (represented by
9
124
P. KNIGHT-JONES ET AL.
Paralaeospira patagonica) that could be found on shores on South-West Africa,
visited by one of us (P. K. J.).
Figure 6 shows that very few Janua species occur on the western shores of
continents, probably because the prevailing winds and currents are such that
they have never been transported there. As Bailey (1970) remarked, the great
difference between the two sides of the isthmus of Panama indicates that much
of the evolution of modern Spirorbinae has occurred since the early Pliocene,
when that ocean link was finally closed. The difference is less likely to have
resulted merely from the old sailing vessels bringing Janua species to the
Caribbean, because there are several Spirorbinae (of other genera) present on
the west, but absent from the east side of the isthmus. Although ship’s fouling
may include abundant J. steueri (p. 96), J. pseudocorrugata and J.
pagenstecheri, it seems unlikely that ship-borne transport has played a major
part in producing the patterns shown in Fig. 6, for otherwise more links would
be expected between Hawaii and California, and between Australia and New
Zealand. Considering the Hawaiian species, J. foraminosa and J. nipponica
probably drifted from the west on buoyant algae, but further transport to the
west coast of America was impossible without going outside the temperature
range of these species. J. knightjonesi and J . preacuta seem t o need particularly
warm conditions, so may have reached Hawaii via the equatorial
counter-current. J. pseudocorrugata, however, may have got there from
California and J. pagenstecheri very probably did so, if it is indeed absent from
Japan. Both these species occur amongst the fouling of small ships in the
Aegean (Bailey, 1960a) and such habits may indeed help to account for their
remarkably wide distributions, but the transport of J. pagenstecheri to St Paul’s
Rocks, in mid-Atlantic, was more probably due t o flotsam or shipwreck than t o
ordinary shipping. I t is possible too that this cold-tolerant species extends deep
enough to be able to migrate along the ocean ridges. We have dredged it
commonly at 130 m in the Hurd Deep (English Channel). It seems to have been
the only Janua species to reach the Galapagos and Easter Island, perhaps
because it was the only one established on the nearest mainland.
SUMMARY
Two widespread species of the subgenus Dexiospira are fully described for
the first time and compared with other Janua species by means of a pictorial
key. Janua (Dexiospira) brasiliensis (Grube, 1872 =Spirorbis bellulus Bush) is
pseudocorrugata by its almost unstriated collar setae
distinguished from J. (0.)
and more calcified brood chamber and from J. (D.1 nipponica by its thoracic
uncini bearing single pegs. I t seems still more distinct from J. (D.)
tambalagamensis, J. (D.) preacuta, Janua (Pillaiospira) trifurcata and J, (P.)
natalensis.
Janua (Dexiospira) foraminosa (Bush, 1904 =Spirorbis heiden Sterzinger) is
characterized by a pitted tube and narrow brood chamber with a wide distal
collar. In setal characters it is intermediate between J. (DJformosa and J. (D.)
steueri. J. (0.1lamellosa and J. ( 0 . ) kayi also fit into this group of five
species, all of which have long juvenile talons, terminally bifid with wide lateral
lobes.
Janua (Janua) pagenstechen is thought t o include Spirorbis pusilloides, S.
REVIEW OF THE GENUS JANUA
125
momonicus, S. epichysis and S.unicornis. The deeply pigmented form in rock
pools of Britain and Brittany is named forma aurantiaca. In the tropics and
Southern Hemisphere this species (forma incoloris) is generally restricted to
deeper water, interstitial environments or other shaded situations.
Janua (Leodora) knightjonesi, found only in coral areas, is perhaps the sole
representative of its subgenus.
In an alphabetical review 19 names are regarded as junior synonyms or mere
varieties and six others as relating to inadequate descriptions. Eleven other
species which have sometimes been placed in Dexiospira or Leodora clearly
belong to Circeis, Sinistrella and Romanchella. These genera, which are
redefined, are not closely related to Janua (to judge from a brief exercise in
numerical taxonomy), but respectively resemble Paradexiospira, Pileolaria and
various Southern Hemisphere forms which brood within their tubes.
J. pagenstecheri seems to be the most widespread Janua species, though it
has not yet been found on the north-west sides of the Atlantic and Pacific
Oceans. J. (D.) pseudocornrgata is almost equally widespread. The west
European form is found only on algae and this overlaps in the Mediterranean
with var. mnnarensis, which occurs on stones and shells in most warm seas.
The latter also overlaps with the west Pacific vars praeceps and fenestrata,
characterized by tube modifications, and subspecies ainu, with slightly
modified abdominal setae. The subgenus Dexiospira probably originated in the
Indo-west Pacific, for there it attains its greatest species diversity and extension
into cooler latitudes. Though not found on the west coast of Africa, four of its
species have nevertheless got past Africa to the West Indies and Brazil,
presumably transported by flotsam and/or shipping.
ACKNOWLEDGEMENTS
Particularly important collections of type material were obtained from
national museums through the kindness of Dr J. D. George (London), Dr Gerda
Hartwich (Berlin), Dr M. Imajima (Tokyo), Dr Marion H. Pettibone
(Washington) and Dr J. Reynaud-Morant (Paris). We are also much indebted,
for further material, facilities and/or advice, to Prof. Julie H.Bailey-Brock, Dr
J. Beurois, Prof. X. da Cunha, Prof. R. P. Dales, Prof. J. H. Day, Dr D. M.
Devaney, Dr K. Fauchald, Dr L. C. Llewellyn, Dr Margaret C. Lloyd, Prof.
Maria Teresa Lopez, Dr T. G. Pillai, Dr Dale Straughan, Dr H.ten Hove, Dr H.
Uchida, Dr P. J. Vine and Dr H.Zibrowius.
REFERENCES
ABE, N., 1943. The ecological observation on Spirorbis, especially on the post-larval development of
Spirorbisugutus Bush. Sci. Rep. mhoku Univ. (Ser. 4, Biol.), 17: 327-51.
AUCENER, H., 1922. Ueber litorale Polychaeten von Westindien. Sber. Ges. narurf. Freunde Bed., 1922:
338-52.
AUCENER, K, 1926. Ceylon-PolychLten. Jeno. 2.~ t u r w 62:
. 435-72.
BAILEY, J. H., 1969a. Spirorbinae (Polychaeta : Serpulidae) from Chios (Aegean Sea). Zoo1 J, Linn.
Soc., 48: 363-85.
BAILEY, J. H., 1969b. Methods of brood protection as a basis for the reclassification of the Spirorbinne
(Serpulidae). Zool. J. Linn. Soc., 48: 387-407.
BAILEY, J . H., 1970. Spirorbinae (Polychaeta) from the West India. S W . FUUMcUm$uo, 3 2 58-81.
BAILEY, JULIE H. & HARRIS, M. P.. 1968. Spirorbinae (Polychaeta : Scrpulidae) of the Galapagos
Islands. J. Zool., Lond., 155: 161-84.
126
P. KNIGHT-JONES ET AL.
BERGAN. P., 1953,The Nonv-ian species of Spirorbis (Daudin). Nytt Mug. 2001.. I : 27-48.
BORG. F.. 1917. Uber die Spirorbisanen Schwedens nebst einem Venuch zu einer neuen Einteilung der
Cattung Spirorbis. 2001.Biidr. Upps.. 5: 15-38.
BUSH, K. J.. 1904.Tubicolous annelids of the tribes Sabellidae and Serpulidae from the Pacific Ocean.
Hornmon AIPtko Exped., 12: 169-355;
CAULLERY. M. & MESNIL, F.. 1897. Etudes sur la morphologie comparCe et la phyl&nie d a a p i c e s
Jes Spirorbes. Bull. x i . Cr. Belg.. 30: 185-233.
CHAMBERLAIN. R. V., 1919.The Annclida Polychaeta. Mem. Mus. comp. 2001.Horv.. 48: 472-9.
CHENU. M.. 1843.IIlustrotions Conchyliologiques. 1: Paris.
CUPAREDE. E.. 1870. Les a n n t l i d a chktopodes du golfe de Naples (Supplement). M b . Soc. Phys.
Hist. not. Cknkve, 20: 365-542.
DAY, 1. H., 1961.The polychaete fauna of S. Africa. Part 6.Sedentary specia dredged off Cape coasts,
with a few new records from this shore. J. Linn. Soc. (20ol.J.. 44: 463-560.
DAY, J. H.. 1963. The polychaete fauna of S. Africa. Part 8.New species and records from grab samples
and dredgings. Bull. Br. Mw. not. Hist, (Zool.)., 10: 381-445.
DE SILVA, P. H. D. H.. 1961.Contribution to the knowledge of the polychaete fauna of Ceylon. Spolh
zeylon.. 29: 164-94.
DE SILVA. P. H. D. H.. 1965. New species and records of Polychaeta from Ceylon. &'roc. zool. SOC.
LO&., 144: 537-63.
DE SILVA, P. H. D. H.. 1967. Studies on the biology of Spirorbinae (Polychaeta). J . Zool., Lond.. 152:
269-79.
FABRICIUS. 0..1780.Founa Crornlondico. Copenhagen & Leipzig.
FAUVEL, P.. 1919. Anntlides p o l y c h t t a des iles Gambier et Touamotou. Bull. Mus. Hist. nut..
Poris, 25: 3 36-43.
FAUVEL. P., 1927. Polychhes stdentaires. Foune Fr.. 16: 1-494.
FAUVEL. P., 1930. Anne'Tida Polychaeta of the Madras Government Museum. Bull. Modras Covt Mus.
(Nor. Hist. Ser.), I : 1-72.
FAUVEL. P.. 1932.Annclida Polychaeta of the Indian Museum Calcutta. Mem. Indion Mus. 12: 1-262.
FAUVEL, P.. 1935. Annilides polychites de L'Annam. Memorie Accod. pont. Nuovi Lincei. (Ser. 3). 2:
279-358.
FAUVEL. P., 1953. Annelida Polychaeta. In R. B. Seymour-Sewell. (Ed.), The founo oflndiu. including
Pokistoa Ceylon, Burmo ond Moloyo. Allahabad: Indian Press.
GEE, J. M., 1964. The British Spirorbinae (Polychaeta : Serpulidae) with a description of Spirorbis
cuneatus sp.n. and a review of the genus spirorbis. Proc. zool. Soc. Lond.. 143: 405-41.
GEE, J. M. & KNIGHT-JONES, E. W.. 1962. The morphology and larval behaviour of a new species of
Spirorbis (Serpulidae). J. mor. biol. Ass. U.K.. 42: 641-54.
GEE, J. M. & WILLIAMS, G. B.. 1965. Self and cross fertiliution in Spirorbis boreolis and S.
pagenstecheri. J. mor. biol. Ass. U.K.. 45: 275-85.
GRUBE, A. E., 1872. Uebersicht der b i s h a beschriebenen Terebellen und uber T.onguicomus und einige
Serpulacean. Jber. schles. Ces. voterl. Kult., 49: 48-53.
HARRIS. T.. 1968. Spirorbis species (Polychaeta : Spapulidae) from the Bay of Naples with the
description of a new species. Pubbl. Stoz. zool. Nopoli, 36: 1-20.
HARRIS. T., 1969. Spirorbis species (Polychacta : Serpulidae) from the South Atlantic. Discovery Rep.,
35: 135-78.
HARTMAN, 0..1942.A review of the types of polychaetous annelids at the Peabody Museum of Natural
History, Yale University. Binghom Occonog. Coll. Bull., 8: 1-98.
HARTMAN, 0..1951. The littoral marine annelids of the Gulf of Mexico. Publ. Mor. Inst. Sci. Univ.
Texas, 2: 7-124.
HARTMAN. 0.. 1959. Catalogue of the polychaetous annelids of the world. Occ. Pop. AI&n Honcock
Fdn, 23: 628pp.
HARTMAN. 0.. 1966. Polychaeta Myzostomidae and Sedentaria of Antarctica. Antarctic Res.. 7: 1-1 58.
IMAJIMA, M. & HARTMAN. O.,1964.Polychaetous annelids of Japan II. Occ. Pop. Allon Honcock Fdn,
26: 239-452.
KAWAHARA, T., 1961. Regional differences in the composition of fouling communities in Ago Bay.
Rep. Foe. Fish prefect. Univ. Uie, 4: 65-80.
KAWAHARA. T.. 1962. S r u d i a on the marine fouling communities 1. Development of a fouling
community. Rep. F a . Fish. prefect. Univ. M e , 4: 27-41.
KAWAHARA. T.,1963. Studies o n the marine fouling cornmunitis. 11. Differences in the development
of the test block communities with reference to the chronological difference of their initiation. Rep.
Foe. Fish. prefect. Univ. Uie, 4: 391-418.
KAWAHARA, T., 1965. Studies on the marine fouling communities. 111. Seasonal changes in the initial
development of t a t block communities. Rep. F a . Fish. prefect. Univ. Mie, 5: 319-64.
KIRKEGAARD. J. B., 1959.The Polychana of W a t Africa. Sci. Res. Danish Exped. Trop. W. Afr.
1945-46.Ailonride Rep., 5: 7-117.
REVIEW OF THE GENUS JANUA
127
KNIGHT-JONES. E. W.. KNIGHT-JONES, PHYLLIS & BREGAZZI. P., 1973. Helicosiphon biscoensis
Gravier and its relationship with other Spirorbinae. 2001.J. Linn. Soc..52: 9-21.
KNIGHT-JONES, E. W., KNIGHT-JONES, PHYLLIS & VINE, P.. 1972. Anchorage of embryos in
Spirorbinae (Polychaeta). Mur. Biol.. 12: 289-94.
KNIGHT-JONES, E. W.. KNIGHT-JONES, PHYLLIS & LLEWELLYN, L C., 1974. Spirorbinae from
south eastern Australia Part 2. N o r a on taxonomy, ecology and distribution. Rec. Ausr. Mus.. 29:
107-51.
KNIGHT-JONES. PHYLLIS, 1972. New species and a new subgenus of Spirorbinae (Serpulidae : Polychaeta) from Kenya. J. Zool.. Lond., 166: 1-18.
KNIGHT-JONES, PHYLLIS, 1973. Spirorbinae (Sapulidae : Polychaeta) from south eastern Australia. A
new genus and seven new species. Bull. Br. M u . nut. Hisr.. 24: 230-59.
KNIGHT-JONES, P. & KNIGHT-JONES, E. W., 1974. Spirorbinae (Serpulidae : Polychaeta) from
southern Africa, including three new species. Mur. Biol., 25: 253-61.
KNIGHT-JONES, PHYLLIS & WALKER, A. J. M., 1972. Spirorbinae (Serpulidae : Polychaeta) on
limpets from the South Orkney Islands. Br. Anrurcr. Surv. Bull.. 31: 33-40.
KORNERUP, A. & WANSCHER, J. M.. 1967. Methuen handbook of colour. London: Methuen.
LAMARCK, J. B. DE. 1818. Historie Nuturelle des Animuuxsuns Vertkbres, 5: 1-612.
LANGERHANS. P., 1880. Die Wurmfauna von Madeira 111. Z. wfss. Zool., 34: 86-143.
LANGERHANS, P.. 1881. Uber einige canarische Anneliden. Nova Actu kgl. Leop.-Curol. Deursch. Akad.
Nuturf., 42: 93-124.
MAYR. E., 1942. Sysremarics und the origin of species. New York: Columbia University Prcss.
MESNIL, F. & FAUVEL. P.. 1939. Polychites sidemaires de I'expCdition du "Siboga". Maldanidae,
Cirratulidae, Capirellidae, Sabellidae et Serpulidae. Sibogu Exped., 24: 142.
MOHAMMAD, M-B. M.. 1971. Intertidal polychaetes from Kuwait, Arabian Gulf, with descriptions of
three new species. J. Zool., Lond., 163: 285-303.
MONTAGU, G.. 1803. Tesraceu Brimnnicu. Romsey: Hollis.
MOORE. J. P., 1923. Polychaetous Annelids dredged by U.S.S. "Albatross" off the coast at Southern
California in 1904. IV. Spionidae to Sabellaridae. Proc. Acud. nut Sci. Philad.. 75: 179-259.
MOORE, J. P. & BUSH, J., 1904. Sabellidae & Serpulidae from Japan with descriptions of new species of
Spirorbis. Proc. Acad. nat. Sci. Philad.. 56: 157-79.
MaRCH, 0. A. L., 1863. Revisio critica Serpulidarum et bidrag tie roromena Naturhistorie. Nunwhist.
Henrik Kr#yer, K4benhuven. (Ser. 3). I : 347-470.
OKUDA, S.. 1934. Some tubicolous annelids from Hokkaido. J. Fuc. Sci. Hokkaido Univ. (Ser. 6). 3:
2 3 3-46.
OKUDA, S., 1938. Polychaetous annelids from the vicinity of the Mitsui Institute of Marine Biology. Jup.
J. ZOO^.. 8: 103-5.
OKUDA, S. & YAMADA. M.. 1954. Polychaetous annelids from Matsushima Bay. J. Fuc. Sci. Hokkaido
yniv. (Ser. 6 Zool.). 12: 175-99.
PBRES, J. -M. & PICARD, J.. 1958. Rechercha sur l
a peuplements benthiques de la Meditcrranke
nord-orientale. Ann. Inst. ockunogr. Monuco. 34: 21 3-91.
PILLAI, T. G., 1960. Some marine and brackish-water serpulid Polychaeta from Ceylon, including new
genera and species. Ceylon J. Sci. biol. Sci., 3: 1-40.
PILLAI, T. G.. 1965. Annelida Polychaeta from the Phillipina and Indonesia. Ceylon J. Sci. biol. Sci.. 5:
110-77.
PILLAI, T. G., 1970. S t u d i a on a collection of Spirorbis from Ceylon. together with a critical review and
revision of Spirorbid sysrematia and an account of their phylogeny and zoogeography. Ceylon J. Sci.
biol. Sci., 8: 100.72.
PIXELL, H. L. M.1912. Polychaeta from the Pacific coast of North America 1. Scrpulidae. with a revised
table of classification of the genus Spirorbis. Prm. 2001. Soc. Lond., 65: 784-805.
POTSWAW, H. E., 1965. Reproductive biology und development of Spirorbk Ph.D. Thesis, University
of Washington.
QUATREFAGES, M. A. DE. 1865. Histoire nuturelle des Annilides murins er d'euu douce. Annilides et
Cephyriens, 2. Sedenruriu. Paris: Roret.
RlOJ A, E,1923. Ertudio sistemitico de las especis ibccicas dd suborden Sabelhformia. n a b . MUS.nuc.
Cien. nut., Mudr. (Ser. Zool.), 48: 1-143.
RIOJA, E., 1942. Estudios anilidologicos V. Observaciones acerca de *nos
especies del gknero
SplrorMS Daudin de 10s costas Mexicans del Pacifico. An. 1-t. BiOL Univ. Mix.. 13: 137-53.
RIOJA, E., 1959. Estudios anilidologicos XXII. D a t a para el conmimiento dc la fauna dc an6lidos
poliquetos dc I
scostas orientals de MCxico. An, Inn. Biol. Univ. Mix.. 29: 219-301.
RULLIER, F., 1964. Annilides P o l y c U t s (campape de la "Calyp~o" 1959 aux Dles du Cap V e d .
Annals Inst. ocianogr.. MOMCO, 41: 113-218.
SAINT-JOSEPH, BARON DE, 1894. Les annilides p o l y c h h s d a cbtes de Dinar4 3. Annls Sci. nut.
(Zod.), 17: 1-395.
SMITH, S. 1. & HARGER. 0..1874. Report on the dredgings of the region of George's Bank in 1872.
Annelid.. by A. E. Verrill. ?Vans. Conn. Acud, ArWSci., 3: 1-57.
0. KNIGHT-JONES ET AL.
128
SOURIS, R., 1954. conltibptioa & l'ttudc kdoeiCpe d a c6tcs mcheusu du s
w . M b n . Inrt. ft. Aft.
38: 7-342.
STGRZHCER, I., 1909.
m e Spirorbk-Ancn aus Suez. S k r . A M . wlrr. Wkn Wh~wrWw.
Kl.), 118: 1U1-19.
WERZINCER. I.. 1910. ihm die Spfmwbis-Artender akdkhcn Adria. A b h wok -hot.
5:
1-13.
STRAUCHAN. D.. 1969. Spirorbinlc (Annelid8 : Polychma) from Eniwctok, M.nhdl Id.ndr.
Mkr@t&m, (I: 111-3.
SVERDRUP, H. U.. JOHNSON, M W. & FLeMINC. R. H.. 1942. Tha octheir phyrler, ch.nfrtry
~ n d
b i d q y . Ncw Yorkt Pmuice-)t.ll.
UCHlDA, H.. 1971.. S p W i e (Polychaco. %?pdidsc) from H d r l u i b , I.J. Fr. Scl Ho&Lotdo Unlu.
(Ser. 6), 17: 628-62.
UCHIDA. H, 1971b. Spimrbinac(hlychacta. Serpulidrc) from Hokk.id0. II.1. Fac. Sd. Holrlutdo Udv.
(Se. 6. ZOOL). 18: 193-226.
VINE, P. J.. 1972.. Spitotbilue (Pdychaeta : Serpulidae) from the Red Sa. iadudillg descriptionrafa
ntw genus and four new @a zod J . Wnn. SOC.. 51: 177-201.
VINE, P, J., 1972b. Spirorbinac (Rdychaeu : Sapdidae) of thc hdhn chin. Part 1. hkW rpcda
pbclf: scl.. 26: 1409.
VINE, P. J.. 1975. S p i r O h i n ~(pdycb.cu :sCrpuW.e) from Ncw ZaLnd
ineluding W P h m
of clcrnn new speck Bull. N.Z. lkp. sdmt. ind. Re& (in pterr).
VINE, P. 1.. BAILEY-BROCK, J. H.& STRAUCHAN, 0,1972. S p i d i n r e (Polychwta :Scrpulidac) of
thc Hawinn chain. Rtt 11. pM/. Scl. 26: 150-82.
WESfiNBERELUND, E, 1952. S c r p u l i (Polyehaeta) collected by C Dons along the Norwqian coast.
Skr. K. nor& Vlcknrlt.S d k . 6: 1-22.
WISELY, B.. 1962. Two spirorbid tubcwornrr (Sapulidae. Polychaua) from 01tcfll Australia. Rcr. Awt.
Mu., 25: 342-8.
ZACHS. J. M., 1933. polycb.rp of thc North Japanese Sa (in Rori.n). Inn. HydrlbloL Expkw. Men
URSS. 19: 125-37.
ZIBROWIUS, H, 1968. &udc maphdogiquc, sydmatique ct b b g i q u e da sapulidrc (Mi
Fdychaua) & la w n de M n d k . Red. 7hv. St. mar. h d . WClL, 43: 81-252.
ZIBROWIUS, H.. 1969. -qua
bQp*cucI r&oltcr dc S e r p u l i i (Polychacta rsdcntui.) drnr k W e
& Cab& ct en Tripokanic. Bull. I N t . oc&nogr. P?&q Sdhmbo, 1: 123-37.
PBROWIUS, H, 1970. comribuno
' n iI'cNdc da Sapulidse (pdyebrrrr scdwnum
* 1 du BtcliLBolrn
ItW
SRttlO, 19: 1-32.
UBROUIUS, H., 1973. scrpulibc (Annelid.. pdychacta) da
ouclc dc I'Afriqw u da archipels
VOiriw Annlr Mw. r. Afi. cent (Sn. 8,). 207: 1-93.
**.,
APPENDIX
Sources from which the distributions of Janua species have been plotted
(Fig. 6). Asterisks denote that collections from the area have been examined by
one of us (P.K.-J.). An asterisk following an author's name indicates that that
author's material has been so checked.
Northeast Atlantic. Norway, Bergan, 1953; Wesenburg-Lund, 1952; Britain.,
Gee, 1964; N.W. France*, Caulluy & Mesnil, 1897; Spain, Rioja, 1923.
Mediterranean. France & N. Africa, Zibrowius, 1968,1969; Italy,Harris 1968;
Adriatic, Stetzinger, 1910;Aegean, Bailey, 196%*; Malta..
East Atlantic. Madeira' and CanarkP, Langerhans, 1880,1881; Cape V d e .
Is.*, Kirkcgaard, 1959; Rullier, 1964; Senegal, Sourie, 1954; Angola. and
S.W. Africa..
West Atlantic. West Indies., Bailey 1970*; Gulf of Mexico, Hartman 1951,
Rioja 1959; Costa Rica*; Brazil, Gmbe 1872*, Zibrowius 1970;
Newfoundland*.
Indian Ocean. South Africa.;
Mozambique*, Mndagasclr.;
Kenya,
Knight-Jones 1972*; Red Sea, Stazinger 1909,Vine 1972a.; Persian Gulf*;
caullery & Mesnill897; Ceylon, Augena 1926, de Silva 1961, Pillai, 1970.;
India, Fauvel1930*; Amsterdam Is.*.
Australasia South Australia and New South Wales*, Knight-Jones, 1973;
REVIEW O F THE GENUS J A N U A
129
Knight-Jones, Knight-Jones & Llewellyn, 1974*; Queensland, Vine,
Bailey-Brock & Straughan, 1972; New Zealand, Vine, 1974*.
West and Central Pacific. Indonesia & Vietnam, Mesnil& Fauvel, 1939, 1935*;
Phillipines, Pillai, 1965; Japan*, Moore & Bush, 1904*; Okuda, 1934;
Uchida, 1971a, b; Marshall Islands, Straughan, 1969*; Touamotou, Fauvel,
1919; Hawaii, Vine, Bailey-Brock & Straughan, 1972*; Easter Island*.
East Pacific. Alaska, Bush, 1904; British Columbia, Pixell, 1912; California*,
Bush, 1904; Mexico, Rioja, 1942; Costa Rica*; Chile*; Galapagos, Bailey &
Harris, 1968*.
Southern Ocean and Antarctic. Harris, 1969; Hartman, 1966; Knight-Jones,
Knight-Jones & Bregazzi, 1973*; Vine, 1975.