<oological Journal of the Linnean Society (1983), 79: 245-295. With 21 figyres
Contributions to the taxonomy of
Sabellidae (Polychaeta)
PHYLLIS KNIGHT-JONES
<oology Department, University College of Swansea, Swansea
Received April 1983, accepted for publication June 1983
The genus Potamilla Malmgren is particularly characterized by dorsal lips which lack any radiolar
midrib; Pseudopotamilla Bush by compound radiolar eyes, dorsal collar lappets and flanges alongside
the bases of the most dorsal radioles; Demonax Kinberg by a wide dorsal collar gap and companion
se:ae with big heads and narrow blades; and Potamethus Chamberlin by an elongated first segment,
enlarged ventral sacs, very long shafts to the thoracic uncini, and inferior abdominal setae much
shorter than the superior ones. Perkinsiana gen. nov. lacks most of these characters, but the type
species ‘Potamilla’ rubra Langerhans is remarkable in having red blood. It bores into limestone
abundantly off S Wales and is here redescribed. Oriopsis hyncnsis sp. nov. from SW Ireland has
a smooth collar set low on the first segment, with its ventral margin entire and free from the apex of
the peristome. The new name Demonax langerhnsi is proposed for Sabella (Potamilla) incerta
Langerhans non Demonax incertus Kinberg.
K E Y WORDS:-Sabellidae
taxonomy.
- Potamilla - Demonax - Potamethus -
Oriopsis - Perkinsiana gen. nov.
-
CONTENTS
Introduction . . . . . . . . .
Materials and methods
. . . . . .
Order Sabellida . . . . . . . .
Family Sabellidae . . . . . . . .
Subfamily Sabellinae Rioja (1923) . . . .
Genus Potamilla Malmgren (1865) amended
Genus Pseudopotamilla Bush (1905) . . .
Genus Demonax Kinberg (1867) amended .
Genus Potamethus Chamberlin (1919) .
.
Genus Perkinsiana gen nov.. . . .
Subfamily Fabriciinae . . . . . . .
Genus Oriopsi~Caullery and Mesnil (1896) .
Summary . . . . . . . . . .
Acknowledgements
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References
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INTRODUCTION
Whilst studying British sabellids it became necessary to examine the type
species of Potamilla, P. neglecta (Sars, 1851). This species lacks the radiolar midrib
supporting the dorsal lips as described in “Potamilla” renformis by Orrhage
(1980). Potamilla torelli Malmgren is also like P. neglecta in this respect, but it has
on one hand been wrongly synonymized with P. neglecta (Lukasch, 1910;
245
0024--4082/83/011245+51 $03.00/0
0 1983 The Linnean Society of London
246
P. KNIGHTdONES
Johansson, 1927) and, on the other hand, the name has been commonly given
to southern forms which must now be placed in a different genus. Langerhans
(1880) was probably the first to use P. torelli as a name for a southern
(Madeiran) form but his description does not agree with that of Malmgren.
Marenzeller ( 1885), following Langerhans’s concept of P. torelli, identified
material from the Adriatic and Japan as that species and regarded Sabella
brachychona (1870, Claparkde from Naples) as a junior synonym. Most authors
have followed Marenzeller using the name Potamilla torelli for both warm water
and Arctic forms, often suggesting a cosmopolitan distribution, but in fact
Claparede’s species should be brought back into use and, judging from the
original description, placed in the genus Demonax. Fauvel’s material from
Cherbourg ( M H N P A40), labelled Potamilla torelli and Potamilla incerta (large
and small specimens respectively), is best referred to as Demonax brachychona and
his suppression (1906) of the specific name incerta was unfortunate, as the
description fits a species that is not uncommon in southern Britain and Ireland;
it too is a species of Demonax but not Demonax incertus Kinberg (1867) from Chile.
A search was therefore made amongst known types of Demonax and Potamilla to
ensure that a form agreeing with incerta Langerhans did not exist already under
another name. Species that could be placed in the genus Pseudopotamilla Bush
(1905, see Hartman, 1938) were not examined as these are quite distinct, in spite
of McIntosh’s (1916, 1922) remarks that P. reniformis (the type species of Pseudopotamilla) is a variable species. He regarded as a variety of ‘Potamilla’ rengormis a
form that is clearly ‘incerta’ and also synonymized with it a red-blooded species
from South Wales (Watson, 1906), in spite of the fact that P. reniformis has green
blood, compound eyes on the radioles and several other differences. Watson’s
species is almost certainly Sabella (Potamilla) rubra Langerhans (1880), but its
genus is neither Sabella, Potamilla nor Demonax. Potamethus seemed to be a
possibility following Hartman’s (1967) concept of that genus, but a review of
the species reveals that only one, Potamethus littoralis Hartman ( 1967), is closely
related to the British form. Eventually it became clear that a new genus would
have to be created to accommodate both these two forms and also a number of
species that had been inappropriately associated with the genus Potamilla.
Thomas H . Perkins (Department of Natural Resources, St. Petersburg, Florida),
who had kindly made available to me his pre-publication studies of Demonax
species mainly from North America, suspected that a new genus was needed and
it is named Perkinsiana in recognition of his expertise in this and other related
fields (Perkins, 1979; 1980; 1981; in press). A species of Oriopsis that seems to be
new to science is also described.
New diagnoses of the foregoing genera of Sabellidae are given here basing
them on multiple numbered characters. For brevity only the diagnosis of the
first genus of each subfamily and that of the new genus are given in full. In other
genera, the only characters that are listed are those that differ from the full
diagnosis and these are similarly numbered. Characters, or combinations of
characters, that are especially useful are emphasized by italics.
MATERIALS A N D METHODS
A few of the following species were obtained by collecting from the shores of
S Wales, Plymouth, Madeira and Chile, and the remainder were received from
TAXONOMY OF SOME SABELLIDAE
247
various institutions and individuals. Donors are indicated by initials in the text
but named in full in the acknowledgements. Live material was usually
narcotized with menthol crystals, fixed and preserved in 5% seawater
formaldehyde and later transferred (after washing) to 70% alcohol.
Several drawings of the head and lower crown were made from preliminary
camera lucida outlines, with details added from the usual examination under a
dissecting microscope with bright incident illumination. Studies of the crown
were made in two ways. Museum material was laid on a dark wax-lined dish
under alcohol with the radioles splayed out by pinning between them with
entomological pins, to display the dorsal and ventral lips. I n the larger
specimens of personally collected material one half of the crown was removed
prior to examination. A sharp scalpel was used for this and for making
transverse sections of the base of one of the second dorsal radioles to study the
axial cartilage. Fine hand-cut sections could be seen adequately in glycerol
under a cover slip. Setae and uncini were studied in polyvinyl-lactophenol
mounts, of the whole body with worms as small as Oriopsis, but otherwise a
whole torus or a whole fascicle was teased apart by fine needles. Sufficient
details could be seen with phase-contrast microscopy and quartz-iodide lighting,
but stereoscan electron micrographs were also made of some setae (techniques
described in Knight-Jones & Fordy, 1979). All species examined have been
drawn in detail. Drawings not offered for publication have been kept in
personal files and copies are available on request. Permanent mounts of the
setae and uncini of type material have been, or will be, returned to museums
that hold the types.
ORDER SABELLIDA
Abdominal tori dorsal to the fascicles of setae, an arrangement the reverse of
that found in the thorax and in sedentary polychaetes generally; usually with a
distal crown of radioles that have both filter-feeding and respiratory functions;
faeces conveyed along a tract which runs ventrally along the abdomen, round
the right side of the body and dorsally along the thorax, to be ejected out of the
mouth of the tube.
FAMILY SABELLIDAE
No operculum; thoracic membrane only distal or absent; tube membranous
(or gelatinous), never calcareous, but often including particles of mud, sand or
shells; body bilaterally symmetrical except for faecal tract.
SUBFAMILY SABELLINAE
Each radiole of the crown with two rows of pinnules; companion setae
adjacent to thoracic uncini in most genera; thoracic uncini avicular, each with a
distal toothed crest adjacent to the anterior peg or beak, the ‘shaft’ strongly
curved and swollen anteriorly beneath the ‘neck’ to form a breast (Fig. lL),
posterior portion varying from long to truncated.
P. KNIGHT-JONES
248
Y
0.04rnm
..
H
J
K
Figurc 1. f'o'olamilla neglecta holotype: A, schematic view of the inside of the left half' of the crown,
dorsal lip on the left, crosshatched area representing the position of the 'bridge' between the two
halves of the crown, the lower dotted line representing the base of the thoracic collar; B, terminal
part of a radiole; C, cross-section of the base of a dorsal radiole, just above web; D & E, anterior
thorax and base of crown; D, dorsal view; E, ventral view; F, superior thoracic seta; G & H, inferior
hooded thoracic seta; G, three-quarter view; H, 'face' view; J, showing the position of the two types
of thoracic seta in an epithelial sleeve; K , abdominal seta side view; L, thoracic uneinus and M,
ad.jacent companion scta, both side view; N, detail of straightened companion seta bladc. Scales: D
& E as A; G , H, K & N as F.
Genus Potamilla Malmgren, 1865 amended
Radioles ( I ) without eyes, (2) without longitudinal ridges, (,3)
without appendages, other than the pinnules, and (4) without flanges but with
(5) the lower parts finely webbed, sometimes vestigially, and (6) with the more
basal parts strongly fused to their neighbours; the axial skeleton of each radiole
(7) appears in a cross-section just distal to the webbed region as a rounded
group of cells (Fig. 1C), these axial supports arise from the cartilaginous bases of
the crown which forms two semicircles, one each side, separated ventrally and
connected mediodorsally (8) without flanges or notches (Fig. 1D); (9) the paired
DIAGNOSIS:
TAXONOMY OF SOME SABELLIDAE
249
dorsal lips which have often been termed ‘tentacles’ or ‘palps’ are without a radiolar
midrib and thus not tapered or grooved, each being a simple blunt lamella
arising from the mouth and (10) attached to the adjacent pinnule (Fig. 1) of the
most dorsal radiole; ( 1 1) the length of the first segment, from the annulus below
the collar setae to the base of the crown and including the peristomium, is about
13 times the length of the next thoracic segment; (12) the ventral sacs are
vestigial and adjoin (13) a ventral cleft between the ventral lappets of the collar;
(14) the collar is in two parts, one on each side, each extending from the ventral
cleft to the dorsal side, (15) without lateral notches and (16) without dorsal
lappets or lamellae; ( 17) the dorsal gap in the collar is either narrow (Fig. 1 ) or
a mere slit (Fig. 2B), its dorsal margins are partly fused to the floor of the faecal
groove; (18) the collar setae are arranged in oblique oval tufts, and (19) the
superior setae of the other fascicles in the thorax form sublongitudinal arcs; (20)
both superior and collar setae are slender with a fine or vestigial border
(Fig. 2G); (21) the inferior setae are numerous and form oblique clusters
comprising ill-defined subtransverse rows within the arc of the superior setae;
(22) each inferior seta is slightly curved distally, with tapered wings like the
hood of a cobra (Fig. lG, H) and with a very long shaft more than 13 times the
length of the hood; (23) the avicular thoracic uncini (see subfamily diagnosis)
arranged in a tight row (torus), the length of each shaft being similar to, or a
little longer than, the distance between the breast and the crest; (24) companion
setae (sometimes known as pick-axe or pennoned setae or soies en pioche) with
blades shaped like leaves that are broad proximally (Fig. l M , N); (25)
abdominal setigers more numerous than those of the thorax; (26) abdominal
setae slender and geniculate, the inferior ones differing from the superior ones in
being slightly wider at the ‘knee’ and slightly shorter between the ‘knee’ and tip;
(27) abdominal setae arranged in a transversely elongated fascicle; (28)
abdominal uncini like those ofbthe thorax, but with shafts shorter than the
distance from the breast to the’ crest; (29) glandular areas most conspicuous
ventrally, forming prominent shields (scutes or cushions); (30) no glandular
girdle around the second setiger.
Type species: Potamilla neglecta (Sars, 1851) ZMHUB 6796
Designated type species by Bush ( 1905), collected Hammerfest and Tromso a t
about 80 m depth.
A single specimen dredged off northern Norway is well preserved except for a
damaged posterior. I t has about 85 segments of which 8 are thoracic. The
thorax is oval in cross-section, 1.6 mm being the widest diameter. The body is
25 mm in length, the crown being a further 10 mm with 13 radioles on each
side, each with long, widely spaced pinnules along most of the length. The
terminal part of the radiole is very short and the adjacent pinnules fairly long
(Fig. lB), indicating that the shortness of the tip is probably normal and not the
result of predation. Each rachis has an axial skeleton of cartilaginous cells
forming about 3 parallel rows in side view and, in cross-section, a circular group
of 6-7 cells. These are surrounded by a substantial sheath, outside which is a
thin, even layer of epithelium with no thickened corners or ridges on the outer
surface (Fig. 1C bottom). The waisted outline each side of the dark central
blood vessel may be characteristic.
The dorsal lips are blunt, supported a t the side only by a pinnule and not
G
H
Figure 2. Potamilla lorellz: A-C, anterior thorax and base of crown; A, lateral view; B, dorsal view;
C:, ventral view; D, schematic view of the inside of the right half of the crown, dorsal lip and
webbed pinnules on the right; E, a cross-section of the base of a dorsal radiole just above the
vestigial web; F, optical section (longitudinal) of the lower part of a radiole (dorsal) showing
pinnule junction with the axial core; G , superior thoracic seta; H, inferior hooded thoracic seta,
same fascicle as G ;J, thoracic uncinus side view; K, blade of an adjacent companion seta; L & M,
abdominal seta; L, inferior; M, superior from the same fascicle as L. Scales: B & C as A: H-M as G.
TAXONOMY OF SOME SABELLIDAE
25 1
higher than the curved ventral lips (Fig. 1A). T h e absence of midrib with a
distal taper led to the remark in Sars’s later paper (1862), that the species was
without tentacles.
Th e collar is oblique in side view (interrupted line Fig. lA), shallow
dorsolaterally and set low on the first segment, showing the peristomium above.
The dorsal margins are separated by a narrow gap (Fig. 1D). Ventrally the
collar extends to form narrow, obliquely elongated lappets (Fig. lA, E), the tips
of which cross when forced back against the body. These are equal in length to
one and a half thoracic shields. The latter (except the first) are 3 times as broad,
transverse to the body axis, as long, separated by a deep transverse groove and
traversed by a shallow one. T h e first shield has the upper half extended laterally
and anteriorly to form a bulbous corner on each side, with an indentation
between and below the collar cleft and small ventral sacs (Fig. IE).
The fifth thoracic fascicle (mounted) is composed of about 6 superior setae
that are very fine and virtually without a border (Fig. 1F). The inferior setae
(about 26) are arranged in a tight group each with wings forming a hood that
tapers to a point and is nearly 3 times as long as broad (Fig. IG, H). The
convex surfaces of the hoods are staggered, forming an oval aggregation (e.g.
Fig. 1J) which gives a maximum surface to grip the tube wall. The shafts are
very long, more than 13 times the length of the hood and partially encased in an
epithelial sleeve. The avicular uncini have a shaft, 1+ times the distance between
the maximum curvature of the ‘breast’ and the distal crest (Fig. 1L). The
companion setae have fairly substantial leaf-shaped blades (Fig. 1N) which are,
when not distorted, held a t right angles to a long shaft (Fig. 1M). The
abdominal uncini (sixth abdominal fascicle mounted) have a less swollen
‘breast’ and a shorter shaft. The abdominal setae are slightly geniculate. The
superior ones (about 15) are slender, whilst the inferior ones are wider at the
knee, shorter distally (Fig. 1K) and more numerous (about 20).
The tube, about 3 mm in diameter, has a hardened mucous lining covered
with tightly packed sand grains. Basally there are several Bryozoa, Foraminifera
and an empty spirorbid tube, probably Circeis.
Brood protection of embryos within the mouth of the tube has been described
in material dredged off Shetlands labelled P. torelli (BMNH. 1931.10.7.32)
which turned out on examination to be P. neglecta (Knight-Jones & Bowden, in
press). Similar brooding of juveniles with rudimentary crowns was described by
Wesenberg-Lund (1950) in material from Iceland identified as P. neglecta, but it
was probably P. torelli as she noted that the radioles were “without any palmar
membrane” and such webbing is a t least obscure in P. torelli (see below). The
distribution of P. neglecta is difficult to ascertain. I t may have been frequently
mistaken for P. torelli as Sars’s Latin descriptions (1851, 1862) were without
figures and’,thoseof Malmgren (1865) depict a species that agrees better with
P. torelli. Misidentifications throughout the literature are therefore
understandable.
Potamilla torelli Malmgren (1865, later figured 1867)
Malmgren’s Icelandic material has not been found but some (ZMUB)
collected by the Norwegian North-Atlantic Expedition 1876-1878 at a depth of
547 m between Faroes and Iceland was identified by Hansen as P. torelli and the
material agrees well with Malmgren’s description. Five specimens are present,
252
P. KNIGHT-JONES
one wholly out of its tube, with four others protruding from damaged tubes. The
former has a few fascicles missing and is damaged posteriorly but 50 abdominal
segments remain. The body is 32 mm long and 2 mm wide plus a crown 11 mm
long with 13 radioles on each side. Malmgren’s material may have been more
complete and/or more mature as he describes the body as 63 mm long, 3.5 mm
wide, with a crown 8-12 mm long, with 14 to 16 radioles on each side. He
describes the “tentacles” (=dorsal lips) as being short, broad and almost
rounded (e.g. Fig. 2D) agreeing with Potamilla in the restricted sense defined
above. Each dorsal radiole (Hansen’s material) has three basal pinnules webbed
to each other, the proximal one being also fused to the adjacent dorsal lip. This
species differs from P. neglecta in that the webbing between the radioles is
vestigial (Fig. 2A) and a cross-section at the base of a dorsal radiole near to the
webbing is less waisted in outline, with a thicker epithelium, thinner skeletal
sheath and a more irregular arrangement of axial cells in both cross-section and
longitudinal view (Fig. 2E, F). The tip of each radiole however, is similar to
that of P. neglecta in being short compared with the adjacent pinnules (e.g.
Fig. IB).
The collar reaches the base of the crown as shown in Fig. 2A (and fig. 76,
Malmgren, 1867) and the dorsal margins are very close forming a mere slit
(Fig. 2B). Ventrally the two collar lappets are less acute that those of P. neglectu,
overlapping slightly a t the median cleft. The first thoracic shield is not indented
distally and is deeper than the others which are about twice as broad as long
(Fig. 2C).
The thoracic fascicles have a similar arrangement to P. neglectu (Fig. 1J). The
superior setae (about 14) are, however, much more numerous than in neglectu.
The setae (mounted 7th fascicle) are a little wider than in Potamilla neglectu (cf.
Figs lF, 2G) but the inferior setae are similar, with a hood nearly 3 times as
long as broad, including the taper which is more narrow distally (Fig. 2H) and
in agreement with that in Malmgren’s figures. The shafts of the longest inferior
setae are about 12 times the length of the distal hood. The thoracic uncini have
a shaft no longer than the distance between breast and crest (Fig. 25) and the
companion setae have leaf-shaped blades that are difficult to see by transmitted
light, and are doubtless very thin (Fig. 2K). The abdominal setae are geniculate
(Fig. 2L, M ) and the inferior ones (5-9) have shorter blades and more bulbous
knees than the superior ones, which number about 4-5 in each fascicle. The
abdominal uncini have shorter shafts than those of the thorax.
The construction of the tube is very similar to that of P. neglectu. One empty
tube about 2.5 mm across has a curved basal part fixed to small anchoring
stones. Just above the curve, presumably at substratum level, two tubes are
fused along its side, one with a diameter of 1 mm and, in the angle between it
and the larger tube, another just 0.3 mm across containing a tiny sabellid with
the crown exposed. The lower part of this tube is hardened mucus with sandy
detritus distally. The embedded sand grains, typical of the adult are not present.
If embryos are developed without a pelagic stage (see P . neglectu above)
settlement on the tube of the parent might be necessary if firm substrata were
scarce.
Material from the Bay of Fundy (labelled P . neglectu USNM 44786) also has
these characters, to judge from drawings kindly loaned by Thomas H . Perkins,
and should be regarded as P. torelli. The species is therefore fairly widespread in
TAXONOMY OF SOME SABELLIDAE
253
the NW Atlantic but other records from southern locations are not reliable (see
below).
Potamilla leptochaeta Southern (192 1)
Collected from Port Canning, Lower Bengal amongst Entoprocta from a
brackish pool, not examined.
This species may indeed belong to Potamilla in the above restricted sense.
According to Southern’s description (1921) the dorsal margins of the collar are
near the median line and the setae seemed to be typical of the genus with the
possible exception that the abdominal superior setae are very much longer than
the inferior ones (hence the specific name), and reminiscent of these in the genus
Potamethus. This species cannot however, be put into Potamethus as the first
segment seems to be shallow according to Southern’s figure (29A), but further
details are uncertain as the figure seems to be a pictorial chimera of both ventral
and dorsal views, to judge from his lateral view of the worm (fig. 28B). The
species is small, 4.5 mm long, of which the crown is 1.3 mm.
Genus Pseudopotamilla Bush (1905)
As Potamilla except that, ( 1 ) most dorsal radioles bear compound eyes on
their outer surfaces (Fig. 3A, B) and (2) have indistinct ridges but (5) are without
webbing although (6) proximally fused; (8) each free dorsal margin of the
cartilaginous base of the crown with a thin narrow flange surmounted by a
notch level with the base of the radioles (Fig. 3A); (9) the paired dorsal lips
have a radiolar midrib (grooved and tapered) and (10) each dorsal lamella is
fused to one or more pinnules at the base of the adjacent radiole (Fig. 3C); (12)
ventral sacs are small (but larger than in Potamilla) and above the ventral
lappets of the collar; ( 14) the collar is in four parts, two extending from the ventral
cleft to deep indentations a t the laterodorsal quarters (there are no truly lateral
notches), and the other two forming (16) tall dorsal lappets reaching above the
DIAGNOSIS:
9
I[
![
0
A
8
B
C
D
Figure 3. Pseudopotamilla renifrmis: A, anterior thorax and base of crown, dorsal view; B, crosssection of dorsal radiole (2nd on right) through the proximal compound eye which is situated on
the corner (representing a longitudinal ridge) furthest from the dorsal midline of the animal (see A);
C , camera lucida drawing of the basal part of the inside of the left half of the crown, dorsal lip with
radiolar midrib on the left, cross hatching representing the cut bridge between the two halves of the
crown, circles representing pinnules on the radioles furthest from view; D, inferior thoracic seta with
a spoon-shaped hood and distal point ‘face’ view. Scales: C as A.
25.1
P. KNIGHT-JONES
base of the crown; the nearly mid-dorsal margins of these lie close to one
another (1 7) the lower part being fused to the floor of the faecal groove; (2 1) the
inferior setae are strongly curved distally in the region of paired tapered wings
which form spoon-shaped hoods (Fig. 3D), each having a shaft about 6 times
the length of the hood.
Type species: Pseudopotamilla reniformis (Bruguiere, 1789)
Amphitrite rentformis Bruguiere ( 1789) “Die nieren formige Amphitrite”
Muller = Potamilla reniformis Malmgren (1867).
Species other than Pseudopotamilla reniformis (British material) have not been
examined during the present studies. Hartman (1938) made a start in reviewing
the genus, suggesting some synonyms, and Banse & Hobson (1968) point out
further specific problems. It is nevertheless probable that P. renzformis has a
world-wide distribution in warm and temperate waters.
Genus Demonax Kinberg (1867)
As Potamilla except that (3) radioles have outer epithelium usually
with ridges (subquadrangular in cross-section) ; (5) radioles are without webs;
( 7 ) in cross-section the skeletal cells are arranged in a reniform, oval or circular
shape; (9) paired dorsal lips have a radiolar midrib with flanking lamellae
(grooved and tapered) with the outer margin (as in Potamilla) fused to an
adjacent dorsal pinnule; (17) dorsal margins of the two part collar with wide
separation; (22) inferior setae have shafts only about five times the length of the
hood; (23) thoracic avicular uncini have shafts not much longer than the
distance between breast and crest; (24) companion setae with bulbous toothed heads
(hooked in side view) and a very narrow tapered blade orjilament arising from the centre at
right angles to each shaft.
DIAGNOSIS:
Type species: Demonax krusensterni Kinberg (1867)
Holotype NRS 577. Collected at Honolulu, Hawaii, amongst dead littoral
coral. Subsequent designation by Bush (1905).
The crown of the single specimen (separate from the body) is 7 mm long,
including the cartilaginous base and radioles (about 20 each side) which are
fused basally (Fig. 4A). The tapered terminal filament is about 0.8 mm long and
the outer surface of the base of some radioles is ridged (probably
subquadrangular in cross-section but sections were not made). Their exact
shape, however, is uncertain as most of the epithelium had sloughed off, but the
crown seems to be mature with no sign of recent regeneration. The radiolar
midribs of the dorsal lips scarcely project above the flanking lamellae or the
ventral lips.
The body is 20 mm long and 3 mm wide with 8 thoracic setigers and 43
abdominal ones. The collar is set high on the first segment and very shallow,
especially dorsally where it is nearly level with the peristome (Fig. 4C). The
shallow dorsal margins are widely separate with a gap ofjust over one-third of
the width of the thorax. Ventrally there is a small cleft (in front of damaged
ventral sacs) between shallow angular ventral margins of the collar, which are
scarcely as long as the average ventral shield (Fig. 4B). These shields are 4-5
times as long as broad and indented laterally by the ventral ends of each torus.
TAXONOMY O F SOME SABELLIDAE
a
Figure 4. A-G Dnnonax krusenstemi holotype: A, complete detached crown ventral view; B & C,
anterior thorax; B, lateroventral view; C, dorsal view, D, thoracic uncinus side view, shaft slightly
bent; E, adjacent companion seta three-quarter view; F, superior thoracic seta; G, inferior thoracic
seta with tapered hood. H-M Demonax leucuspis holotype (except for H, N and P from fresh
material): H, camera lucida drawing of the inside of the lower left half of the crown showing the
dorsal lip on the left; J, lateral view of detached crown; K-M, anterior thorax; K, lateral view; L,
dorsal view; M, ventral view, dotted lines representing the position of ventral lappets when not
curved outwards; N, inferior thoracic hooded seta; P, superior thoracic seta from same fascicle; 0 &
R, companion seta; Q three-quarter view; R, ‘face’ view with bent filament. Scales: C & J-M as B;
E-G & N-R as D.
14
255
256
P. KNIGHT-JONES
The first shield is longer, and indented medially below the collar cleft. The
cuticle, on the left side of the thorax, has lifted bringing with it the long tori.
One of these (5th) about 1 mm long has been mounted showing 37 uncini and
companion setae (Fig. 4D, E). The adjacent fascicle which seemed to be
complete has 9 slender superior setae (Fig. 4F) and inferior setae with a tapered
hood (Fig. 4G) about 4+ times as long as broad. These are not in very obvious
transverse rows (prior to mounting) as they are so crowded, about 24 in all.
The abdominal setae are slightly geniculate and numerous, about 22 in a
tight oval cluster with a transverse axis. Both inferior and superior setae are
slender with a narrow border differing only in that the inferior ones are slightly
wider at the knee and shorter distally. The pygidium is blunt without lobes or
eyes, although presence or absence of the latter can not be a reliable specific
character when material is scarce. Eye spot clusters may not be re-established
after posterior regeneration.
Johansson ( 1927) synonymizes this species with Demonax leucaspis Kinberg but
it has priority and should be treated as separate (following Johansson, 1925) at
least until the two small differences below can be confirmed (or otherwise) by
the study of fresh Hawaiian material. In Demonax leucaspis the collar seems to be
deeper throughout with the dorsolateral margin covering the base of the crown
and part of the fused area of the radioles. The superior thoracic setae of
D. leucaspis are much more numerous than is evident on the type of
D. krusensterni.
The syntypes of Demonax cooki Kinberg (1867, figured 1910) also collected at
Honolulu, Hawaii on littoral dead coral (Holotype NRS 579), could well be
younger specimens of D. krusensterni showing regeneration after damage. Several
radioles of the attached crown of both specimens have tips that are mere stubs
whilst the adjacent pinnules are represented only by papillae. T h e anterior
thorax is also relatively narrow but whether these characters are a result of
incomplete reorganization or are of specific importance it is difficult to say. In
spite of the fact that the dorsolateral margins of the collar are relatively, but
slightly, higher than those figured (4B, C) it seems best to regard the species as
synonymous with D. krusensterni until such time as fresh material is examined.
It has been suggested by Johansson (1927) that D. cooki too is synonymous
with D. leucaspis (as well as D. krusensterni). Unfortunately representative
parapodia were not taken whilst the material was under loan and the relative
difference in numbers between the thoracic inferior and superior setae could not
be estimated. This may be a character separating D. leucaspis from D. krusensterni
(see below).
Demonax leucaspis Kinberg ( 1867, figured 1910)
Collected off San Lorenzo Island near Callao, Peru, intertidal. Holotype
NRS 576. = Demonax incertus Kinberg (1867, figured 1910) collected from
Valparaiso, Chile amongst Fucus (holdfasts). Holotype NRS 579.
Kinberg separated D. leucaspis from D. incertus on account of the different
proportion of the crown to the body. He seemed to be unaware that sabellids
frequently have their crowns damaged by predators but can easily regenerate
the missing parts. The crown of the holotype of D . leucaspis is just 4 mm long and
the ends of the radioles show incipient growth into pinnules. T h e normal crown
as typified by the material labelled ‘incertus’ is 6.5 mm long from cartilaginous
TAXONOMY OF SOME SABELLIDAE
257
base through the areas of basal radiolar fusion and the free parts of the radioles
to the terminal filaments. The latter are gently tapering, about 20 each side.
The dorsal lips at the base of the detached crowns are triangular, scarcely
higher than the ventral lips or about the length of two thoracic segments. Fresh
material from Coquimbo (at 5 m depth) shows more clearly the radiolar midrib
and the attachment of the outer lamellae to the basal dorsal pinnule on each
side (Fig. 4H). The body of the type is in two pieces totalling 20 mm long
( 3 mm wide) with 8 thoracic and 57 abdominal segments.
The collar is deeper dorsolaterally than that of D. krusensterni. I n side view the
distal margin is at right-angles to the body but the base of the collar is set
obliquely on the first segment (Fig. 4K) so that the dorsal part of the collar is
half below and half above the position of the base of the crown (Fig. 4L).
Ventrally the paired lappets are scarcely longer than the length of the average
thoracic shield (Fig. 4M) which is about 4 times as wide as long, each indented
by the ventral ends of the long tori. A 5th torus was mounted showing 61 uncini
and companion setae (Fig. 4% R ) similar to those of D. krusensterni (e.g.
Fig. 4D). A thoracic fascicle was not mounted from the damaged type material
as Johansson’s (1925) figures agreed well with those from live material (Fig. 41v,
P). The hoods of the inferior setae are nearly 4 times as long as broad, a little
smaller than those of D . krusensterni, but the relative numbers of inferior setae
and superior setae (slender and bordered) are very different. For instance there
are 14 superior and 15 inferior setae in D. leucaspis compared with 9 and 24
respectively in D. krusensterni. The abdominal setae are similar to those of
D. krusensterni in shape but fewer, 17 judging from the 21st segment.
Demonax saxicola (Grube, 186 1) new combination
Syntype as Sabella ZMHUB 5198, collected by Grube from Cherso, Italian
Adriatic. Numbers in brackets are those given by Grube showing differences in
specimens other than that selected as type.
The body is 30 mm long (1.3 mm wide) comprising about 90 (100) segments
of which 8 (10, 11 & 24) are thoracic (Fig. 5A). T h e detached crown (8 mm
long) has 12 (9-15) radioles each side which are fused basally. The epithelium
of the radioles has mostly sloughed off. A cross-section of a dorsal radiole at the
base shows a substantial skeletal sheath (the outer epithelium is lost),
surrounding a subcircular group of about 11 irregular cells (Fig. 5B). A sagittal
view of the base shows it to be 4 cells wide which are arranged regularly like
bricks in a wall (Fig. 5C) but halfway along the radiole it is just 2 cells wide,
with a single row towards the terminal filament. This seems to be tapered and
fairly long, but most of the tips and pinnules are in poor condition. Grube
(1861) describes the latter as mostly long, those nearest the tip becoming
suddenly shorter. He also records that the radioles are decorated with a simple
series of 3-8 eyes along their backs. Distally, where some epithelium has
remained, two radioles show diffuse pigment patches that could be punctate
ocelli. They occur more on the sides than on the back of the radioles. O n one
radiole two patches occur together, one each side, but on the other there is no
juxtaposition between the two remaining patches (Fig. 5D). The basal pinnule
on each dorsal radiole is webbed to the dorsal lip which is supported by a long
radiolar midrib (Fig. 5E) about 53 times the length of the average thoracic
segment. These are the white “tentacles” referred to by Grube.
P. KNIGHT-JONES
258
-B
0.1mm
G
![
8
H
K
Figure 5. Demonux suxicolu holotype: A, thorax and anterior abdomen lateral view; B, cross-section of
base of dorsal radiole; C, optical section (longitudinal) of the lower part of a dorsal radiole; D,
detail of a pigment patch near the distal end of radiole, dotted circle represents similar patch lower
on thr other side of rachis; E, lower left half of crown, inner view showing tapered dorsal lip; F & G,
anterior thorax; F, dorsal view; G , ventral view; H, hooded seta from the thorax with distal part bent;
J, as H showing full length of shaft and relative position to K, the adjacent superior seta; L,
companion seta 'face' view (filament either broken or bent out of view); M, thoracic uncinus side
view; N, companion seta side view, detail; P, as N with whole shaft figured; Q abdominal seta.
Scales: F & G as A; C as B K & Q a s J; M as L, N as P.
TAXONOMY OF SOME SABELLIDAE
259
The dorsal margins of the collar arise close to the first fascicles and thus have
a wide separation (Fig. 5F) but the collar is shallow, set high on the first
segment. In side view (Fig. 5A) the distal margin, which would cover the base of
the crown, is nearly at right-angles to the body. The ventral lappets are small
and triangular, the separating V-shaped cleft indenting the first thoracic shield
(Fig. 5G) The following shields are indented laterally by the ventral ends of the
tori. These shields are 2 or 23 times as broad as long. The cuticle of the body has
lifted, bringing with it some fascicles of setae and uncigerous tori. Under the
dissecting microscope the companion setae seemed to be particularly prominent.
A fascicle and torus on the right side of the 4th segment is now mounted. The
superior setae (about 6) are long, slightly geniculate and bordered above a
narrow knee (Fig. 5K) whilst the accompanying inferior setae (9) have distinct
hoods at least 33 times as long as broad (Fig. 55). The tips of most of these are
damaged but one seemed to be complete though bent distally (Fig. 5H). The
uncini (35) have fairly short shafts, no longer than the distance between breast
and crest (Fig. 5M) whilst the companion setae have the usual bulbous head as
in other Demonax and very long, fine blades (Fig. 5L, N, P).
Grube described the body as being yellowish grey with white ventral shields
and a dark bordered collar with additional ‘violet’ colouring in the thoracic
region. The crown, basically pale, had 4 or 5 bands variously pigmented with
violet-brown, white or grey-brown. The tube was described as being semihorny
and pellucid, lining burrows of calcareous rock at Martinsica, Portore and
Cherso.
This species clearly differs from D . krusensterni and D . leucuspis in that the
dorsal lips are so prominent. A single smaller specimen from a ‘lithothamnion’ pool
at Carnsore Point, County Wexford, Ireland (kindly loaned by Brendan
O’Connor) seems to agree with this description.
Demonax brachychona (Claparcde, 1870) new combination, Naples
The whereabouts of the type is not known.
Claparkde’s figures (1870) of companion setae indicate that his species
belongs to Demonax. It differs from D . krusensterni and D. leucuspis in having two
‘tentacles’ ( = tapered dorsal lips) and differs from all three Demonax species
described above, in that it has just 6 thoracic segments. Although a single
specimen may have few thoracic segments as a result of incomplete
regeneration, each of eight sabellids from ‘lithothamnion’ pools near Cherbourg
(misidentified by Fauvel, 1906, 1927, as Potamilla torelli MHNP A470) has a
short thorax. Three have 6, four have 5 and one has just 4 thoracic segments. As
this material agrees with Claparkde’s description it is here fully described as
D . brachychona. In cross-section the cartilaginous cells of the radiolar skeleton are
numerous (more than lo), but as they have collapsed (through some chemical
action of fixative or preservative) their group arrangement is uncertain but
then, so is Claparkdes. If his drawing (plate 14, Fig. 5A) is in true side view a
cross-section of his material would suggest a circular grouping. If it is not, the
cross-section could be reniform, made up of numerous cells, in at least two rows
of five. In Fauvel’s material the outer epithelium (in cross-section) is
subquadrangular in outline with a substantial epithelium, thickened at the
outer corners denoting two longitudinal ridges along the outer rachis. The
specimens are smaller than that described by Clapartde, the largest being
260
P. KNIGHT-JONES
20 mm long (60 mm Claparkde) and 2 mm (3 mm) wide. The larger specimens
have between 100 and 130 abdominal segments posterior to the short thorax. In
Fauvel’s material the largest crown was 5 mm long and does indeed appear to
be short relative to the body, a character which gives brachychona its name. Some
had been regenerating their tips but typical terminal filaments are fairly long
and gently tapered (Fig. 6B). There are about 9 radioles on each side (more in
the larger specimens described from Naples) and these are fused basally. The
pinnules are fairly short, the longest occurring midway along and being about 2
to 3 times the width of the rachis. The basal ones on the dorsal radioles are
enlarged and webbed to the dorsal lips which are supported by a radiolar
midrib, 5 times the length of the average thoracic segment.
The collar is very oblique in side view (Fig. 6Ej. The dorsal margins are
separated by a fairly narrow gap arising low on the first segment, midway
between the first fascicles and the dorsal midline (Fig. 6C). The ventral lappets
are long and acutely pointed, separated by a narrow, V-shaped, deep cleft
which indents the first thoracic shield medially (Fig. 6D). The other shields are
about twice as broad as long, indented laterally by the ventral ends of long
thoracic tori. The uncini have short shafts no longer than the distance between
breast and crest (Fig. 6G) and the usual (for Demonaxj companion setae have
swollen heads and filamentous blades (Fig. 6G, H ) . The superior setae (5 to 6,
have slender bordered blades (easily broken) and the inferior setae (8-10) have
a winged hood about 3 times as long as broad (Fig. 6F). The tips of these are
also easily damaged. The abdominal uncini are similar to those of the thorax
and the abdominal setae (Fig. 6K) are slender and bordered. The posterior of
the worm has the usual blunt taper but a regenerating posterior tapers narrowly
(Fig. 6J).
Claparkde describes the tube as “rksistant mais form6 par une matiere semitransparente”. He does not describe the habitat but such tubes are sufficiently
substantial when lining burrows in calcareous substrata such as the
‘lithothamnion’ encrustations in Cherbourg pools.
Demonax medius (Bush, 1905 as Parasabella media)
The Alaskan type material has not been examined in these studies, Bush
describes her largest specimen as having a body 35 mm long and 55 mm wide,
narrowing at the base of the collar to 4 mm. The body has 8 thoracic and 100
abdominal segments. The radioles of the crown have long tapered tips. All the
thoracic segments are shallow on this comparatively short fat species, the ventral
shields being about 5 times as wide as long. The distal margins of the collar
cover the base of the crown. Dorsally the margins arise midway between the first
fascicles of setae and the midline. Ventrally the margins end as two small
triangular lappets. Superficially this species resembles D. leucaspis. Perkins (in
press) gives a detailed account of this species and figures dorsal lips with a taper
at least twice as long as the ventral ones, and thus unlike D.leucaspis.
Demonax Jecatus (Hoagland, 1919) new combination
‘Holotype’ AMNH 1 183 (as Parasabella) comprising two specimens collected
by A. L. Treadwell from Porto Rico.
The body of the largest specimen is about 18 mm long (nearly 2 mm wide)
with about 75 segments of which 8 are thoracic, the crown being a further 7 mm
TAXONOMY O F SOME SABELLIDAE
Figure 6 . Demonax brachychona: A, whole animal ventral view; B, distal part of radiole; C & D
anterior thorax and basal crown; C, dorsal view, with radioles splayed to show dorsal lips; D,
ventral view showing curved ventral lips and triangular collar lappets; E, thorax and anterior
abdomen, lateral view; F, thoracic hooded seta; G, thoracic uncinus side view with adjacent
companion setae in side view; H, companion seta three-quarter view; J, abdomen showing recent
regeneration posteriorly; K, abdominal seta. Scales: E & D as C; G, H & K as F.
26 1
262
P. KNIGHT-JONES
long (Fig. 7A). It has, however, lost its thoracic setae, so it is the smaller
specimen which is here described. The abdomen (in two parts) is 16 mm long
and 1.5 mm wide, the crown being a further 5 mm long. There are 16 radioles
each side with the usual basal fusion but neither specimen shows the ‘delicate’
web described by Hoagland. Below the angle of fusion are h e a t e d pigment
flecks whilst on the sides of each rachis are tiny clusters of light brown pigment
spots arranged in rows, with irregular spacing, along two-thirds of the rachis in
the position where the longitudinal ridges are usually found. However,
transverse sections of the base of a dorsal radiole showed no distinct outer
corners (i.e. ridges) but revealed that the cartilaginous cells are about 7,
arranged in a reniform group (Fig. 7B). The terminal filament is tapered, fairly
short and about 0.5 mm long, similar in length to the longest pinnules which
occur subdistally. The basal pinnule on each of the two dorsalmost radioles is
slightly enlarged and webbed to the adjacent dorsal lip. This tapers and is
supported by a radioiar appendage about 3 times as long as the average
thoracic segment.
The collar is set high on the first segment and dorsolaterally the distal margin
is also high, covering the fused part of the crown. Latero-ventrally, however, the
distal margin is low (Fig. 7D), forming shallow lappets each side of the ventral
midline, and the small separating cleft does not indent the first thoracic shield.
The other 7 ventral shields are 2-3 times as wide as long, indented laterally by
the ventral ends of the long thoracic tori. The uncini have short shafts no longer
than the distance between breast and crest (Fig. 7H). The companion setae,
many of which are broken, have the usual (Demonax) bulbous head and
filamentous blade (Fig. 75, K ) . These came from the 7th parapodium on the
right (the only one showing setal blades) and most of these were broken. The
setal fascicle forms two groups of shafts (4+7). The only intact setae were of the
inferior type. These have slender hoods 7 times as long as wide. They are similar
to those figured by Hoagland (1919) but she was mistaken in regarding them as
superior setae. It is common for the latter to be broken first as they are narrow
and project further. The abdominal setae are of two lengths in the same fascicle
(22nd), both types bordered and very slender (Fig. 7L, M), more so than that
figured by Hoagland (1919). No details of tube or habitat were given.
Of the species described above only D . saxicola is close but even that species
differs in being more slender in body with a relatively small crown, larger
ventral collar lappets, less slender hoods on the inferior setae, subcircular
skeletal axis and fewer and less distinct pigment clusters on each rachis.
Demonax brevithoracicus (Pillai, 1961 as Potamilla) new combination
Holotype BMNH 1960.3.13.25 from a somewhat sandy area at Nachikuda,
Ceylon.
The body is about 10 mm long (in two parts) comprising 60segments of
which just 5 are thoracic (Fig. 8A). The crown is just over 3 mm long with 9
radioles on one side and 8 on the other, each group fused basally with
longitudinal pigment flecks at the angle of fusion and bands of pigment across
the radioles. The terminal filament is long and gently tapered (Fig. 8C) and
often spiralled (Fig. 8D). A cross-section at the base of a dorsal radiole shows a
narrow even epithelium (no thickened ‘corners’), a substantial skeletal sheath,
and a reniform group of cartilaginous cells (Fig. 8B). The pinnules are fine and
TAXONOMY OF SOME SABELLIDAE
n
8
i[
F
Figure 7. DemonaxJlecatus syntypes showing the largest animal (A) and the smallest ( B M ) , two
specimens in all: A, whole animal; B, cross-section of base of a dorsal radiole; C, anterior thorax
lateral view; D, thorax and anterior abdomen, ventral view; E, anterior thorax, dorsal view, with
the lower part of the radioles splayed to shoy one of the dorsal lips; F, hooded thoracic seta
alongside the broken shaft of a superior seta; G, another hooded seta three-quarters view; H,
thoracic uncinus, side view, with broken shaft of companion seta; J & K, distal ends of companion
setae the latter with a broken filament; L, inferior abdominal seta; M, adjacent superior abdominal
seta. Scales: D & E as C; H-M as G; J as K.
263
264
P. KNIGHT-JONES
Figurr 8. Demonax breuithoracicus holotype: A, anterior thorax and crown, dorsal view; B, crosssection of base of dorsal radiole; C & D, tips of radioles; E, anterior thorax ventral view; F, thoracic
uncinus, side view; G & H, companion setae, shaft damaged posteriorly; J & K, inferior thoracic
seta; J , face view with slender hood; K, side view; L & M, abdominal seta; L, inferior; M, superior
seta from same fascicle; N, pygidium of the detached abdomen. Scales: G & H as B; J-M as F; N as
C.
fairly dense. The basal ones on the most dorsal radioles are fused to the adjacent
dorsal lip, which is supported by a long radiolar midrib about 4 times the length
of an average thoracic segment.
The collar is fairly deep, the dorsal margins arising midway between the first
fascicles of setae and the midline of the body (Fig. 8A). I n side view the base
and distal margins of the collar are oblique, ending distoventrally in 2
subtriangular lappets each with a rounded apex (Fig. 8E). A narrow cleft
separates the two midline margins of the lappets but it does not indent the
anterior margin of the thoracic shield below. Most thoracic shields are about
twice as wide as long. These are indented laterally by the ventral ends of the
long tori. Part of the fourth thoracic parapodium on the right was mounted.
The uncini have a short shaft, no longer than the distance between breast and
TAXONOMY OF SOME SABELLIDAE
265
crest (Fig. 8F). Pillai (1961) was unable to see the fairly short but very fine
‘blades’ on the companion setae. These arise from the usual (Demonax) bulbous
head (Fig. 8G, H ) . The superior setae (about 5 ) are narrow and bordered and
the inferior setae (about 9) have finely tapered winged hoods 4 times as long as
broad (Fig. 85) but narrow in side view (Fig. 8K). The abdominal uncini have a
shorter ‘shaft’ than those of the thorax. The abdominal setae (third
parapodium) are of two lengths (6 inferior and 5 superior) but similar in shape,
being geniculate with a narrow knee and a distal bordered blade (Fig. 8L, M ) .
Pillai (1961) recorded paddle-shaped abdominal setae. The setae of the separate
piece of abdomen were not examined. If its setae are paddle-shaped that portion
must belong to a different species. Its pygidium bears several dark eyespots
(Fig. 8N) as in Pillai’s description.
The tube is said to be horny and coated with sand particles and small
fragments of shell and algae. It was found in a somewhat sandy area.
This species seems to closest to Demonax brachychona, but that species differs in
that the first ventral shield is indented by the cleft separating the ventral lappets
(which are also longer), and the outer epithelium of the radioles is thick and
subquadrangular in cross-section (denoting longitudinal ridges) and probably
with more axial cells.
Demonax oculeus (Pillai, 1965, Fig. 21 as Potamilla) new combination
Holotype not examined in these studies.
Pillai (1965) describes a single small specimen with a body 12.5 mm long and
1.2 mm wide. The crown (12 radioles each side) is short and to judge from the
stubby radiolar tips may have been preserved whilst regenerating. The collar is
short, not unlike that of D.krusensterni, but has light orange eyes (spots) below
the small ventral lappets and above the first ventral shield which is not indented
distally and quite short. All thoracic shields are 2-3 times as broad as long and
all, except the first, are slightly indented by long tori. The figured inferior seta is
different from that of the foregoing species of Demonax in that the hood is very
slender, the length being over 7 times the width. The other setae and uncini are
similar and the pygidium has a cluster of eyespots on each side. The tube is
membranous and coated with sand. The species was found at an oyster farm in
Manila Bay, Philippines.
Demonax langerhansi nom. nov. pro Potamilla incerta Langerhans, 1884, non
Demonax incertus Kinberg 1867
The types of Langerhans’s Madeiran material are apparently lost (Banse,
1970). A single specimen was found (EWKJ) in Madeira in a matrix of
Spirobranchus tubes and calcareous algae between cobbles at a depth of 8 m and
100 m offshore of the Sheraton Hotel, west of Funchal. T h e material agrees well
with Langerhans’s description and with specimens found in Devon, Cornwall,
SW Wales and S Ireland. The following description is of the Madeiran specimen
now deposited as lectotype BMNH ZB 1982.46 with a mount of the left thoracic
torus, another of its left thoracic fascicle and a mount of the last abdominal
segments of the anterior portion (segments 30-33).
The body is in four parts having suffered damage extracting it from the
calcareous matrix. The measurements in brackets below are the maximum sizes
or numbers found in material other than the lectotype (e.g. L = those described
P. KNIGHT-JONES
266
by Langerhans; B=those found in Britain and K=Irish material found and
loaned by Professor J. A. Kitching).
The crown 6 mm long was shed whilst studying the worm in laboratory
conditions and the two halves were separated for observation of the interior lips.
Each half has 6 radioles fused basally and (L, 10; B, 7; K, 6) arising from a
cartilaginous foundation. The outer surface of the rachis bears distinct but blunt
longitudinal ridges which show as corners of a subquadrangular cross-section
(taken from the base of a dorsal radiole, Fig. 9B). The skeletal axis within the
epithelium consists ofjust 4 irregularly shaped cells in a reniform group. More
distally there are only 2 cells which in side view appear as a single row.
E
@!$)
.::
F
L
8
G
J
Figure 9. Demonax Zangerhansi nom. nov. Madeiran material: A, inside of the lower left half of
the crown, showing tapered dorsal lip on the left; B, cross-section of base of dorsal radiole; C, distal
part of same radiole; D & E, thorax and anterior abdomen; D, lateral view; E, ventral; F,
peristome and collar from above; G; superior thoracic seta; H & J, inferior thoracic seta; H, threequarter view of hood; J, ‘face’ view, same fascicles; K & L, companion seta; K, side view; L, plan
view of ‘head’; M, thoracic uncinus, side view. Scales: D, E & F as C; H & J as G; K & J as M.
TAXONOMY OF SOME SABELLIDAE
267
Subterminally there is just one cell. Although small numbers of cells can be a
juvenile character, the number in the larger specimens of the British and Irish
material is fairly constant in showing (T. S.) a group of 4 cells with just one
showing 5. The rachis tends to spiral when narcotized or placed in fixative but
the terminal portion characteristically remains straight, as this is thickened in
side view (Fig. 9C). In life this portion is pigmented interiorly with yellowish
white blotches. In British materials these blotches are quite white and very
granular. When the crown is viewed in situ (SW Wales) the distal part of the
rachis curves inwards towards the mouth, rather than outwards (like a fountain)
as seen in the co-existing species Pseudopotamilla renformis.
The body overall is 7 mm long (complete specimens: L, 15; B, 10; K 14) and
1 mm wide (B, 0.5; K, 1) with 5 thoracic setigers (L, 8; B, 5-7; K, 5-7) and 68
abdominal ones (L, 60; B, 58; K, 56). Langerhans, like other early authors, may
well have included the peristomium as a segment. The collar has a wide gap
separating the dorsal margins (Fig. 9F). It is oblique in side view (Fig. 9D), the
dorsolateral margins covering the base of the crown and extending to the
pointed ventral lappets (Fig. 9 E, F). These are separated by a V-shaped cleft
which indents the distal margin of the first thoracic shield. The other shields are
indented laterally by the ends of the long thoracic tori, and are nearly twice as
broad as long.
The 4th thoracic parapodium on the left side is now mounted. There are 4
slender superior setae (Fig. 9G) and about 7 inferior setae, with short winged
hoods only 23 times as long as wide (Fig. 9H, J). The torus is large (for the size
of the animal) containing 22 uncini with relatively short shafts (Fig. 9M) and
companion setae with the usual (Demonax) well developed head and a rather
long filamentous blade (Fig. 9K, L). The setae of the abdomen are slender and
similar to those of the thoracic superior setae but more geniculate. They are.
ranged in two sizes in a single fascicle, the smaller inferior one being a little
shorter above the ‘knee’. The Madeiran specimen lacks a pygidium but
Langerhans refers to 6 anal eyespots. Six Irish specimens out of 19 had clusters
of eyespots on each side of the pygidium and 8 of those without eyes showed
recent posterior regeneration (shallow and narrow abdominal segments). One
specimen 1.5 mm long, 0.25 mm wide with 20 segments has just three partially
developed segments anteriorly and a vestigial crown.
The Madeiran specimen had (in life) a straw coloured body with an orange
gut and whitish ventral shields, those of the abdomen divided longitudinally by
a distinct brown pigmented faecal groove. The crown is banded with pale
brown and the intermediate, somewhat transparent, bands have whitish yellow
blotches like those of each terminal radiolar process. Material from Britain often
lacked these bands but the distal white blotches seem to be characteristic.
Most of the tube is semitransparent hardened mucus with the last 2 mm
covered by sandy particles. In Britain the species was found in low water crevices
at Porthpean, Looe and Trevone (Cornwall); in scrapings off the piles on Mill
Bay Dock at Plymouth, and also at Castle Beach and West Angle on both sides
of the entrance to Milford Haven, SW Wales. The SEM micrographs of inferior
and companion setae (Fig. 10) are from West Angle material. In Ireland the
species is found in Lithophyllum incrustans in a tidal pool outside Lough Ine, Co.
Cork (Kitching & Thain, 1983).
This species differs from the species of Demonax described above and from
268
P. KNIGHT-JONES
Figure 10. Demonax Zangerhansi nom. nov. British material: A & B, second thoracic fascicle; A,
looking towards the dorsal midline, showing slender superior setae behind the tapered hooded
inferior setae; B, looking away from the dorsal midline, most superior setae having been removed;
C , rompanion seta (left) and adjacent thoracic uncini; P, detail of the head of a companion seta
viewed from above. Scale bars 0.02 mm.
TAXONOMY OF SOME SABELLIDAE
269
those described by Perkins (in press), in that the terminal apinnate parts of the
radioles are not tapered but even thicker than the rachis in side view, and the
cartilage cells at the base of the rachis are so few. It also differs from most
species of Demonax in that the hoods of the inferior thoracic setae are rather short
and therefore appear relatively wide. Of the two species which have a reduced
thorax; D. brachychona and D. brevithoracicus, the former (which overlaps
geographically) is much larger with a relatively short crown and longer ventral
lappets, whilst D. brevithoracicus lacks ridges on the radioles and has no anterior
indentation on the first thoracic ventral shield.
Genus Potamethus Chamberlin 1919 (nom. nov. pro Potamis Ehlers, 1887)
As Potamilla except that ( 5 ) the radioles are not webbed, but fused
basally above a cartilaginous foundation; (7) cross-section of radiolar skeleton
not noted (except in P. singularis, see below) but in each species the rachis is very
thin and rounded on the outer surface (no longitudinal ridges); (9) the paired
dorsal lips are small, triangular and usually obscured by the depth of the
cartilaginous base of the crown; (1 1) thejrst collar segment is 2 4 times the length of
the next thoracic segment; (12) ventral sacs very prominent fused distally to the ventral
lips and proximally to the base of the ventral cleft of the collar (Fig. 1 lC, M,
QJ; (14) the collar can usually be divided into four parts, the two main ones
extending obliquely backwards from the ventral cleft to, and usually beyond,
the collar setae; (16) the other two parts comprise a pair of dorsal lamellae
which flank the long dorsal midline of the first segment (e.g. Fig. 11A, L) but
are absent in two species (e.g. Fig. 11 P); (17) the dorsal margins of the main
collar are usually fused to the base of each lamella (Fig. 11A, L) except in
Potamethus malmgreni where the two parts are not fused; (22) inferior thoracic
setae have somewhat spoon-shaped hoods (Fig. 1 lE, R) and long shafts; (23)
thoracic uncini have very long shafts (Fig. 1 lF, U) between 4 and 12 times the length of the
distance between breast and crest; (24) companion setae have shafts as long as those
of the thoracic uncini, with a head bulbous and hooked in side view and with a
fine tapered blade or filament (Fig. 1 lG, T ) ; (26) abdominal setae of two
distinct lengths, the inferior setae being much shorter (0.6 as long overall)
including short bordered blades (Fig. 1 lJ, K); (28) abdominal uncini avicular but
virtually without a swelling at the curvature (breast) distal to the short shaft (Fig. 1 l H ,
V); (29) ventral glandular surface in the abdomen with narrow extensions
around the parapodia (Fig. 1 IN) which can extend onto the dorsal surface of
the posterior segments as narrow bands, but in some species the whole body
surface is glandular so that no shields or other special glandular areas can be
distinguished; (30) a narrow girdle (glandular?) can be seen behind the second
fascicles in the thorax (Fig. 1 lA, L), but this is obscured if the whole surface is
glandular (Fig. 1 1P).
DIAGNOSIS:
Type species Potamethus spathiferus (Ehlers, 1887 as Potamis)
Florida off Sambos at a depth of 130 m. Type not examined. The
longitudinal dorsal lamellae extend about halfway up the long first segment,
their lower parts fused to the dorsal margins of the collar, forming pockets at the
angie of fusion similar to those in Fig. 11A. Ehlers (1887) states that the collar is
undivided ventrally. Although this would be unique, the character cannot be
confirmed as the material is in poor condition (Perkins, in press). Ehlers also
P
Figure 1 1. A-K Potamethusjliformis holotype: A-C, anterior thorax; A, dorsal view; B, lateral view;
C, ventral view; D, superior thoracic seta; E, inferior spoon-shaped thoracic seta, same fascicle; F,
thoracic uncinus side view G, adjacent companion seta (not side view); H, abdominal uncinus side
view; J, abdominal inferior seta; K, as J, superior seta same fascicle. L-N Potamelhus scoltae holotype:
L & M, anterior thorax; L, dorsal view; M, ventral view; N, part of ventral abdomen. P-W
Potamethus stngularzs: P & Q, anterior thorax; P, dorsal view; Q, ventral view; R, inferior spoonshaped seta; S, superior seta same fascicle; T, companion seta side view; U, adjacent thoracic
uncinus, side view; V, abdominal uncinus, side view; W, optical (longitudinal) view of base of
dorsal radiole. Scales: B & C as A; E, J & K as D; M & N as L; Q a s P R-V as S.
TAXONOMY OF SOME SABELLIDAE
271
notes that “strongly marked ventral cushions” (glandular shields) “are not
present”. It is therefore likely that all or most of the body is glandular as in some
other Potamethus species (e.g. Fig. 1 lC, Q. The sabellid is very slender with a
body 28 mm long and 1 mm wide. The crown extends a further 20 mm with 9
radioles each side.
Potamethus malmgreni (Hansen, 1882 as Potamilla)
Holotype ZMUB 1947. North Atlantic a t a depth of 2127-2222 m.
The longitudinal dorsal lamellae extend two-thirds of the way up the collar
segment, but they are not fused to the dorsal margins of the collar which
terminate a t the first fascicles. This seems to be the only species with this
character. The whole epithelium is glandular (without shields or patches). The
companion setae are notable in having a filament 2 or 3 times as long as those in
Fig. 1l G , T. The body (damaged posteriorly) measures 15 mm by 1 mm, the
crown being 10 mm longer.
Annenkova (1952) suggested a new genus Gorbunovia with Malmgreni Hansen
as type, but she did not distinguish this from Ehlers’s genus Potamis, soon
renamed Potamethus. Furthermore her figures differ markedly from material as
described above and from the figures of the type by Eliason (1951).
Annenkova’s figures show short dorsal lamellae fused to the dorsal collar
margins, and distinct shields on the thorax. In size and form her material is
closest to Potamethus scotiae Pixell (e.g. Fig. 1lL, M), but the shafts of the thoracic
uncini are much shorter (less than one-quarter of the length). It would seem to
be a new species.
Potamethus elongatus (Treadwell, 1906 as Potamilla)
Holotype USNM 522 1. Molokai, Hawaii a t a depth of 130 m.
The longitudinal lamellae are short, occupying about one-quarter the length
of the dorsal surface of the collar segment. The dorsal margins of the collar are
united with the lower portions of these, forming pockets as in P. spathiferus and
P.jlzformis (e.g. Fig. 1lA, see below). T h e body lacks ventral shields or patches,
being glandular all over. A slender worm with a body 29 mm long, 0.6 mm
wide, and a crown extending a further 16 mm.
Potamethus scotiae (Pixell, 1913 as Potamis)
Holotype RSM 1921.143.1486. East of the Weddell Sea, Antarctica
at a depth of 650 m.
The dorsal lamellae extend two-thirds of the long broad collar segment,
united basally with the dorsal part of the main collar forming pockets
(Fig. 11L). T h e glandular areas are defined by ventral shields in the thorax
(Fig. 11M) and subtriangular patches in the ventral abdomen, the apices
extending around the parapodia (Fig. 11N). The hood of each inferior seta is
unusually small, the wings being narrow and short on either side of a relatively
wide core (the continuation of the shaft). Pixell ( 1913) wrongly figures the base
of the core as being much narrower than the shaft. Her figure of the distal part
of the thoracic uncinus is also inaccurate in appearing ‘hairy’. The uncini have
the usual dense short teeth at the crest. The sabellid is one of the largest in the
genus, with a body 75 mm long and 4 mm wide, the crown extending a further
48 mm.
15
272
P. KNIGHT-JONES
Potamethus murrayi (McIntosh, 1916 as Sabella) new combination
Holotype BMNH 1921.5.14202. North of the Hebrides at a depth of 1015 m.
Collar segment as in P. scotiae (Fig. 1 lL, M) except that the longitudinal
lamellae extend distally as far as the base of the crown. It also differs in that the
whole surface is glandular (no shields or patches). Like P. scotiae it is a large
species with the thorax 4.5 mm wide and a crown 13 mm long. The type
material is in two halves but McIntosh (1916) records the body as being 1.5 in
long (38 mm).
Potamethus mucronatus (Moore, 1923 as Notaulax)
Holotype USNM 17351. Found off Catalina Island, California at a depth of
1000 m.
Collar and lamellae are similar to those of P. scotiae (e.g. Fig. 1 l L , M j and the
abdomen has similar subtriangular glandular patches (e.g. Fig. 11N) but the
thorax lacks ventral shields, its whole surface being glandular. The type is small
with a body length of 22 mm, width 1.8 mm and crown 20 mm long, but larger
specimens have been described by Moore (1923) (body 57 mm long, 5.5 mm
wide).
Potamethus dubius (Eliason, 1951 as Potamis)
Type ZMUB not examined. Nares Deep W Atlantic at a depth of
5850-5860 m.
The dorsal margins of the collar meet the midline groove. There are no
dorsal lamellae and the collar is thus like that of P. singularis (Fig. 11P and see
below). The glandular areas form very indistinct shields in the ventral thorax
but there are more conspicuous patches in the first 5-6 setigers of the abdomen,
where “they branch around the bristles”, but more posteriorly the patches are
also found around the “bristles” (parapodia) but extending in narrow bands
dorsally. A very slender worm with a body length of 29 mm, width 0.6 mm and
the crown a further 20 mm.
Potamethus singularis Hartman ( 1965)
Holotype HFLA 11036. Off New England at a depth of 200-2000 m.
The collar (Fig. 1lP, C) is similar to P . dubius in lacking dorsal lamellae, but
this species differs in that all of the body is glandular with no shields or patches.
The cartilaginous axis of each radiole is just a single row of large disc-shaped
cells (Fig. 11W). It is not known whether this is a specific or generic character as
this character was not looked for in other Potamethus species. The species is
extremely small and the character could be specific. The holotype is damaged
posteriorly but it is 0.5 mm wide with a crown 5.5 mm long. Hartman (1965)
however describes a slightly larger specimen 8 mm long and 1.4 mm wide.
The legends of plate 52 in Hartman’s (1965) paper are partially reversed. Her
figures referring to P. singularis should read: c = distal part of thoracic uncinus,
d = thoracic inferior setae and e = (?) thoracic superior seta.
Potamethusjliformis Hartmann-Schroder (1977j
Holotype Z M U H P 13685 found off the Portuguese coast at a depth of
1370-1430 m.
TAXONOMY OF SOME SABELLIDAE
273
The dorsal collar (Fig. 11A) is similar to that of P. sputhifeerus. T h e ventral
margins of the collar are widely separated and form a shallow obtuse
indentation centred on the midline (Fig. 1 IC). Like P. sputhifeerus the ventral
shields are not distinct. The ventral glandular areas of the thorax extend
without differentiation to, and around the base of, the parapodia. Dorsally the
surface appears to be less glandular as the girdle below the second fascicles is not
obscured (Fig. 11A) like it is ventrally (Fig. 1 lB, C). I n the anterior half of the
abdomen the glandular areas are discrete subtriangular patches, the apices
encompassing the base of each parapodium as in P. scotiue (Fig. 11N). The
sabellid is a slender worm with a body length 22 mm, width 0.7 and the crown a
further 16 mm.
Species wrongly associated with Potumethus
Potumethus littoralis Hartman (1967): Holotype USNM 55570 from the South
Shetlands Islands at a depth of 79 m does not belong to this genus but is typical
of a new one described below.
Poturnethus breuiuncinutus Hartmann-Schroder ( 1977): Paratypes ZMUH 13687
and 13688. Found off the Moroccan coast a t a depth of 72 m.
Dr Hartmann-Schroder has already pointed out that this species may need to
be placed in a different genus but the characters of some of the more obscure
genera must first be explored. It can not be a species of Potumethus on account of
the shortness of the first segment and the shortness of the thoracic uncini. The
inferior abdominal setae are certainly much smaller than the superior setae
(former 0.7 of the latter) but there is only one of each in an abdominal
parapodium and the inferior seta has the compact spoonshaped hood commonly
seen in Hypsicomus sensu luto. As Hartmann-Schroder (1977) pointed out, it can
not be placed in ‘Hypsicornus’ as the first thoracic seta are not in a longitudinal
row nor are there eyes on the radioles. The material is in poor condition, owing
to the fact that much of the anterior epithelium has sloughed off, and eyes and
fine webbing would be the first to suffer.
Genus Perkinsiana gen. nov.
Radioles (1) without eyes but (2) with indistinct ridges along the
outer surface of each rachis, (3) without appendages (other than pinnules), (4)
without flanges or (5) webbing, though the radioles are (6) fused for a short
distance basally; the axial skeleton of each radiole (7) appears in a cross-section
near the base as a rounded group of cells; these axial supports arise from the
cartilaginous base of the crown which forms two semicircles, one each side
separated ventrally and connected mediodorsally, (8) without flanges or notches
along the dorsal margins; (9) the dorsal lips are paired, tapered and grooved,
each being supported by a radiolar midrib, (10) with its outer lamella fused to
the adjacent pinnule which is often enlarged; (11) the first segment is up to
twice the length of the adjacent one; (12) the ventral sacs are small and adjoin
(13) a ventral cleft between the lobes of the collar; (14) the collar is in two parts
each extending from the cleft to the dorsal side, (15) without lateral notches or
(16) dorsal lappets; ( 17) dorsally there is a wide gap between collar margins;
(18) the collar setae are arranged in oblique oval tufts and (19) the superior
setae of the thorax in sublongitudinal arcs; (20) both superior and collar setae
DIAGNOSIS:
274
P. KNIGHT-JONES
are slender with narrow borders; (21) the inferior setae are arranged in
transverse rows close to (and within) the arc of the superior setae; (22) each
inferior seta is slightly curved distally, with tapered wings forming a hood and
with a shaft about 5 times the length of the hood: (23) each avicular thoracic
uncinus (see subfamily diagnosis) with a shaft similar to, or up to 3 times as long
as the distance between its breast and its crest; (24) each companion seta with a
flat tapered blade at right-angles to, and covering the oval distal part of the
shaft that is slightly hooked in side view (Fig. 12H, K & L); (25) abdominal
setigers more numerous than those of the thorax; (26) superior and inferior
abdominal setae similar in shape, and either slender or with a bulbous knee
(27) and arranged in two close short transverse rows; (28) the abdominal uncini
avicular with a shaft similar in length to the distance between breast and crest;
(29) the glandular areas are obvious as ventral shields; (30) there is no
glandular girdle around the second setiger.
Type species Perkinsiana rubra (Langerhans, 1880 as Sabella (Potamilla))
Reg No. BMNH ZB 1882 47-48, mounted parapodia 48a, b & c.
Specimens 3-8 cm long (including crown) and only 1.2 mm wide, found
boring into littoral and sublittoral limestone in S Wales, agree well with
Langerhans’s description ( 1880). The delicate crown (Fig. 12A) is easily
overlooked amongst the encrusting fauna but it is characterized by its pink
colouring due to the red blood in the vessels of the radioles (5-10 each side).
The basal cells of the radiolar skeleton appear in cross-section as a circular
mosiac (Fig. 12C). I n a lateral view this cartilaginous core appears to be about
3 cells wide but it diminishes towards the terminal filament, which is as slender
as the neighbouring pinnules. The dorsal lip is fairly short, about twice the
depth of the ventral lip (Fig. 12B) or 3 times the length of the average thoracic
segment. Each is webbed to an adjacent dorsal pinnule that is not particularly
enlarged.
The first segment is fairly short, its thoracic shield not much longer than the
others which are about 23 times as broad as long. The collar is slightly oblique
in side view (Fig. 12A), the higher ventral part separated by a V-shaped cleft
forming two subtriangular lappets which do not overlap at the midline. The
dorsolateral margins slope down to the dorsal side of the first fascicles leaving a
wide dorsal gap. These do not reach the base of the crown dorsolaterally. The
numbers of thoracic segments vary between 6 and 11.
There are about 4 or 5 slender superior setae in each thoracic fascicle, and 6
to 10 inferior setae with subcircular spoon-shaped hoods (Fig. 12F, G; Fig. 13A,
B) in each fascicle except the first. The thoracic tori are fairly short with a large
gap between the ends of the tori and the adjacent ventral shields (Fig. 12D).
Each torus has 8-1% uncini with fairly long shafts (about 23 times the length
between breast and crest, Fig. 12J), and a similar number of companion setae
with a tapered distal blade (Fig. 12K, L) at right-angles to the shaft (Fig. 12H,
Fig. 13C, D). The abdominal tori have uncini similar to, but smaller than, those
in the thorax (Fig. 12M, N). The superior and inferior abdominal setae are
geniculate, the latter with a more bulbous ‘knee’ and shorter between ‘knee’ and
tip (Fig. 12P, Q, & R). The pygidium lacks eyespots and has four lobes, two
lateral ones being larger than the two dorsal ones, the anus as usual being
ventral.
TAXONOMY OF SOME SABELLIDAE
R
C
Q
275
\
\ i\
!
E
E
0~
E
F
H
Figure 12. Perkinsiana rubra genotype: A, anterior of worm left side; B, inside of right half of crown
showing tapered dorsal lip on the right; C, cross-section of base of dorsal radiole; D, anterior thorax
ventral view; E, superior thoracic seta; F & G, spoon-shaped inferior seta with sharp distal taper; F,
three-quarter view; G, ‘face’ view; H, companion seta side view; J, adjacent thoracic uncinus side
view; K & L, blade of companion seta; K, viewed from above; L, viewed from the back; M & N
abdominal uncinus; M, partial ‘front’ view, shaft distorted; N side view; P & Q abdominal seta
with a bulbous heel (inferior); P, ‘face’ view; Q side view; R, superior seta (same fascicle) with less
bulbous heel. Scales: B as D; F, G, K & L as E; M-R as J.
276
P. KNIGHT-JONES
Figure 13. Perkinsiunn firbra gen. nov. British material: A, eighth thoracic fascicle, looking
towards the dorsal midline showing the spoon-shaped hooded inferior setae with the shafts of the
superior setae beyond; B, showing the hollowed side of the spoon-shaped setae beyond the superior
setae distorted by foreshortening (foreground); C, thoracic uncini (top) and adjacent companion
setae (bottom); D, details of the blades of companion setae viewed from above. Scale bars 0.02 mm.
TAXONOMY OF SOME SABELLIDAE
277
The body is reddish orange with prominent and rather white shields
throughout the ventral surface. In rocky areas bordering sand, the tube is
extended above the surface of the limestone and the outer surface of the
hardened mucous wall is decorated with sand and shell particles, the latter
obliquely attached by one edge. Whether adorned or not the mouth of the tube
can be closed by lateral flattening when the worm is withdrawn, but unlike the
tube of Pseudopotamilla reniformis it does not roll up. These two species occur in
mixed populations near Swansea but can be readily distinguished by the form of
the tube mouth.
‘Potamilla’ rubra Langerhans was recorded by Rioja (1918) at Gijon in Spain,
but in a later paper (1923) he doubted his identification, perhaps being
influenced by McIntosh (1922, as Potamilla) who thought that this was variety
of P. reniformis. McIntosh regarded the latter as being with or without
compound eyes on the radioles and did not seem to notice the other characters
which call for separation not only at specific but at generic level. Watson’s
(1966) red-blooded species of Potamilla sp. from Tenby is clearly that found
along the S Wales coast in low water limestone from West Angle (Dyfed) to
Bracelet Bay (Swansea) the latter being the source of SEM material in Fig. 13.
Langerhans found his Madeiran material in calcareous encrustations on rocky
shores. I n the present studies no fossil limestone was found on Madeira, but
there is plenty of calcareous material formed there by serpulids and particularly
by coralline algae. Pseudopotamilla reniformis was fairly common there, boring into
such material, but in several days’ collecting near Funchal only one abdomen
was found which may perhaps be P. rubra. T h e setae agree with material from
S Wales but the identification is inconclusive without the anterior part. The
abdominal setae are remarkably like those of P. reniformis, which no doubt
accounts for McIntosh’s confusion. Pending more successful collections on
Madeira it is by no means certain that the Welsh species is rubra. If there should
be a specific difference between them, it is the Welsh form that should be
regarded as the type species of Perkinsiana.
Perkinsiana antarctica (Kinberg, 1867, as Laonome)
This material seems to have been lost but Ehlers (1897) examined Kinberg’s
specimens from the shore of the Magellan Straits and found that his material
from the Beagle channel agreed well. One sample from Ushuaia ( Z M U H V
4954) is in good condition and is here described. I t should not be included in
Laonome on account of the presence of companion setae. Kinberg (1867) wrongly
amended the genus to include his species.
The body of the largest specimen is 21 mm long, 2.2 mm wide with 7-8
thoracic segments, the crown being a further 9 mm with about 18 radioles on
each side. Each dorsal lip with its tapered midrib is about equal in length to 4
thoracic segments, the outer lamella being fused to the adjacent but enlarged
pinnule at the base of the neighbouring dorsal radiole (Fig. 14H). The axial
skeleton in this area is about 4 cells wide in side view. I n transverse section it has
13-16 cells in a circular group, the surrounding epithelium being thickened to
form corners (Fig. 14B) which are in fact longitudinal ridges along the outer
surface of the rachis.
The margins of the collar are oblique in side view as the collar is shallow
dorsolaterally (with a wide gap separating the dorsal margins) and extended
278
P. KNIGHT-JONES
Figure 14. Perkinsiana nnkzrctica: A, anterior thorax and basal crown showing the left dorsal lip
within the radioles; B, cross-section of base of a dorsal radiole; C & D, thorax and anterior
abdomen; C , right side; D, ventral view; E, hooded thoracic seta (inferior) three-quarter view; F,
thoracic uncinus, side view; G, adjacent companion seta showing ‘back’ view with the blade
straightened; H, superior abdominal seta (inferior one not shown is scarcely wider a t the knee).
Scales: A & D as C; F, G & H as E.
ventrally into two large rounded lappets that extend outwards when the worm
is out of its tube (Fig. 14C) and are overlapping and pointed distally when it is
withdrawn (Fig. 14D) and thus different from P. rubra. One of the main
differences between it and the latter, is that there is no gap between the thoracic
tori and the ventral shields (Fig. 14D) even in small specimens, a character more
common to Demonax. The lateral margins of the shields are indented by these
tori and they are 3 times as broad as long. The 4th thoracic segment on the left
side has about 40 uncini (Fig. 14F) and companion setae (Fig. 14G). The
numbers of setae in the 4th fascicle are similar to those of P. rubra but the hood
TAXONOMY OF SOME SABELLIDAE
219
of each inferior seta is more oval with a gently tapered distal point, in all 23
times as long as broad (Fig. 14E). The abdominal setae also differ in being
scarcely geniculate with very little swelling at the ‘knee’ (Fig. 14H) even on the
inferior setae. The pygidium normally has clusters of eyespots on each side, but
the posterior is subject to damage and regeneration. Of 28 specimens (Ushuaia,
intertidal Z M U H 4950) 9 had undamaged posteriors and of these only 3 (about
8 mm long) had clusters of eyespots. Some without eyes had small closely
segmented posteriors with pygidium, typical of regeneration.
Fauvel’s material (1916, as Potamilla) from the Falkland Islands agrees well
with that of Ehlers. One of his specimens (MNHNP A412) has embryos
attached to the crown as noted by Kinberg. They are incubated in a mucoid
pod stuck to the base of a ventral radiole (Knight-Jones & Bowden, in press as
Potamilla) .
Perkinsiana antarctica (Kinberg) differs from P. rubra in that the longer collar
lappets overlap ventrally, the abdominal setae are more slender, the thoracic
uncini have shorter shafts and the species is much larger.
Gravier (1907) described what he regarded as a new species, Potamilla
antarctica from the Antarctic Peninsula. The whereabouts of the type is unknown
but to judge from his description it is indeed different from Kinberg’s, Ehler’s
and Fauvel’s species. I t may well be a species of Perkinsiana as it has several
characters in common with Perkinsiana littoralis (Hartman, 1967) see below.
Perkinsiana socialis (Langerhans, 1884 as Sabella (Potamilla))
The type material seems to have been lost with Langerhans’s other Madeiran
collections (Banse, 1970) but 4 specimens agreeing well with Langerhans’s
(1884) description were found at depths of 8 m both west and east of Funchal
(E.W.K-J). It is probably the smallest species in the genus with a body only
3 mm long and 0.3 mm wide (Langerhans’s specimens were even smaller) with
7-10 thoracic and about 25 abdominal segments. The crown is a further 1 mm
long with 3-5 radioles each side, each with few pinnules (one with just 8)
alternatively placed along the rachis (Fig. 15B). This has an axial skeleton ofjust
two rows of cells (only one row in side view). T h e dorsal lip is attached to the
dorsal radiole but is too small to ascertain the method of support. The margin of
the collar is extremely oblique in side view (Fig. 15A), the collar segment being
twice the length of the others in the thorax. The dorsal margins of the collar are
widely separated (Fig. 15B) and distally the ventral lappets are pointed and
very long, about the same length as the average somewhat square ventral shields
(Fig. 15C). The first shield is longer and rounded distally below the V-shaped
collar cleft and small ventral sacs. There is a wide gap between the shields and
the tori which are unusually small, with thoracic uncini (Fig. 15H) and
companion setae (Fig. 15G) in groups of two or three of each. The thoracic setae
are equally sparse per fascicle. The superior setae, usually just one per fascicle,
are slender and the inferior setae, just two per fascicle, have a small oval hood
(Fig. 15D) and a sharp distal point at about 45” to the shaft which in side view
(Fig. 15E) is about half the length of the hood. The abdominal setae, just two
per fascicle, are geniculate with a small bulbous knee (Fig. 155, K ) and the
abdominal uncini, just two per torus, have a shorter shaft (Fig. 15L) than those
of the thorax. This species is very different from the ones above particularly in
its size and the shape of the collar.
280
P. KNIGHT-JONES
Figure 15. Perkinsiana soczuhs: A, whole animal left side; B, anterior thorax and crown dorsal view;
C, anterior thorax, ventral view; D & E, inferior hooded thoracic seta; D, ‘face’ view; E, threequarter view; F, blade of companion seta viewed from above; G , whole companion seta shaft
unnaturally bent showing the underside of the blade in part profile; H , adjacent thoracic uncinus,
Fide view; J , inferior abdominal seta ‘face’ view; K, superior abdominal uncinus showing bulbous
knee in side view. Scales: C as B; E-K as D.
Perkinsiana assirnilis (McIntosh, 1885 as Sabella)
Holotype BMNH ZK 1885.12.1.386 dredged a t 1100 m off the River Plate,
Argentina.
This species resembles P. antarctica in having no gap between the thoracic tori
and the adjacent thoracic shields but it is a more slender species (body 26 mm
long, 1.25 mm wide, crown 12 mm long) the shields, other than the first, being
only twice as broad as long (Fig. 16A, B). One of its main characters was
mentioned by McIntosh (1885), namely that each dorsal lip (tentacle) appears
to be ‘bifid’ on account of the very elongated basal pinnule to which it is
webbed. The tapered midrib is also very elongated (Fig. 16D) equal in length to
about 6 thoracic segments. It is however not particularly long compared with
the length of the radioles. There are 11 of these each side, each with blunt ridges
on the outer surface. I n cross-section there are about 6 axial cells arranged in a
pear-shaped core (Fig. 16K) with three rows of cells showing in side view. Each
terminal filament is long and tapered (Fig. 16C).
The collar also differs in that it is less extensive ventrally, with smaller b u t
rather angular lappets overlapping at the midline (Fig. 16A). The margins are
oblique in side view, low dorsolaterally (not reaching the base of the crown), and
with a wide gap separating the dorsal margins (Fig. 16D). The thorax has 7
segments on the left and 8 on the right, showing evidence of incomplete
TAXONOMY OF SOME SABELLIDAE
28 1
Figure 16. A-K Perkinsiana assirnilis holotype: A, anterior thorax ventral view; B, whole animal
showing left side anteriorly; D, anterior dorsal thorax and basal crown with radioles splayed to
show the dorsal lips supported by very long radiolar midribs and enlarged pinnules; E, thoracic
uncinus, side view; F, adjacent companion seta partial side view; G & H inferior thoracic seta; G,
‘face’ view; H, side view; J, inferior and superior abdominal setae in same fascicle; K, cross-section
of base of dorsal radiole showing the regular arrangement of axial cells in a pear-shaped outline and
epithelial corners (bottom) representing longitudinal ridges on rachis. L & M Perkinsiana acuminata:
L, companion seta with long tapered blade that appears filamentous in side view; M, cross-section
of base of dona1 radiole showing even epithelium around the periphery. Scales: D as A; F-H & L as
E.
reorganization after regeneration. Each torus has about 40 uncini (Fig. 16E),
and companion setae (Fig. 16F). McIntosh failed to record the latter. Each has
a flat tapered blade which is about 4 times as long as the width of the head of
the shaft. The blade is very thin and in true side view appears as a thin filament
at a sharp angle to the shaft. The other thoracic setae are also similar in shape
to those of P. antarctica (particularly the shape of the hoods of the inferior setae
Fig. 16G, H) and their number per fascicle. Abdominal setae are very damaged
and were not examined.
Perkinsiana acuminata (Moore, in Moore & Bush, 1904 as Patamilla)
Type: USNM 5500 found at Sagami Bay, Japan at a depth of 280 m.
Moore suggested that this species closely resembles P. assirnilis. Having studied
the types of both, they are indeed very similar in the angle and shallowness of
the collar, the depth of the base of the crown, the proportions of the ventral
282
P. KNIGHT-JONES
shields and the shape of the uncini and setae, particularly the hoods of the
inferior setae which are in fact more eggshaped (e.g. Fig. 16G) than those
depicted by Moore. The abdominal setae are slender as depicted by Moore and
geniculate in side view (e.g. Fig. 16J), the inferior setae having more girth at the
knee and a shorter ‘blade’ between knee and tip, than the superior ones. The
companion setae agree well with Moore’s figure, the blade being 5-6 times the
width of the head of the shaft in side view (Fig. 16L). Other views show a
tapered blade.
One of the two main differences between this species and P. assimilis is that
the dorsal lips are much shorter, only about twice the length of the second
setiger (it too has an adjacent enlarged pinnule) but the worm is larger with a
body 30 mm long and 2 mm wide, crown 19 mm long with 19 radioles each side,
and therefore the difference in length of the dorsal lips may well be specific. The
other difference is that a section across the base of a dorsal radiole shows a thin
epithelium without thickened corners (Fig. 16M) and it would seem that the
outer surface lacks the ridges found in P. assimilis.
Moore also noted that this species agrees fairly well with that identified as
Potamilla torelli from Japan by Marenzeller (1885) and perhaps that of
Langerhans (1880) from Madeira. He seems to have been the first author to
realise that many previous records of ‘torelli’ did not agree with Malmgren’s
(1865) description.
Perkinsiana fonticula (Hoagland, 1919 as Parasabella)
Two specimens described here, from flat reefs of dead coral at Galeta Island,
Panama, Caribbean (Fauchald, 1977b, USNM 066405) show complete
agreement with the holotype (AMNH 1275).
I n side view the collar obliquely crosses a large first segment (Fig. 17A)
rather similar to that of P. socialis (Fig. 15). The difference between the two
species could well be put down to ontogeny, but the two populations, Madeiran
and Caribbean, appear to differ in maximum size. Those of P. fonticula are much
larger, with a body length of 30 mm (width 1 mm) with up to 20 thoracic
setigers. The crown is a further 5 mm, with 6-8 radioles each side. These are
fused at the base, presumably accounting for the ‘web’ mentioned by Hoagland
(1919), but there is no fine web like that found in Potamilla (Figs. 1, 2), nor did
Hoagland figure such a web. The lower part of the crown is supported by a
deep cartilaginous base that unites the two halves dorsally. The dorsal lips are
tapered (not sharply so) with the outer lamella fused to a slightly enlarged
pinnule (Fig. 17B). The ventral lips lead to small ventral sacs behind the
median cleft between very elongated collar lappets (Fig. 17A, C). These are
about 4 times the length of the average ventral shield, the latter being about 3
times as broad as long. A cross-section of the base of one of the dorsal radioles
shows a small core of irregular cells (4-7) and blunt epithelial corners
(Fig. 17D) representing the outer ridges on the rachis.
The thoracic fascicles are very small with only one or two slender superior
setae (Fig. 17G) and about 5 inferior setae. The hood of each of the latter is
small (oval or subcircular) with a short distal filament continuing the curve of
the outer circumference of the spoon-shape (Fig. 17E, F). The adjacent tori
have 5-8 (sometimes 10) uncini with long shafts more than twice the distance
between breast and crest (Fig. 175) and companion setae with long tapered
TAXONOMY O F SOME SABELLIDAE
283
5[
0
0
L
E
F
Figure 17. Perkinsianafonticula: A, basal crown, thorax and anterior abdomen, left side; B, anterior
thorax and basal crown showing dorsal lips within; C, as B, ventral view; D, cross-section of base of
dorsal radiole; E & F, spoon-shaped inferior seta; E, ‘face’ view; F, three-quarter view; G, superior
seta from same fascicle (side view); H, companion seta, three-quarter view with J, adjacent thoracic
uncinus, side view; K, abdominal uncinus, three-quarters view; L, inferior abdominal seta with a
more bulbous knee than that of the adjacent superior seta (not shown). Scales: B & C as A; F-L as
E.
blades (Fig. 17H). The abdomen has uncini with much smaller shafts
(Fig. 17K) and geniculate setae each with a very bulbous knee (Fig. 17L) and
little difference between superior and inferior setae which together total 9 in
each fascicle.
Perkinsiana ctylonica (Augener, 1926)
Type: UZMC. Collected a t Trincomalee, Ceylon at 12 m.
This species resembles P. rubra in that (1) the thoracic tori are fairly short
leaving a gap between their ventral ends and the adjacent thoracic shields
(Fig. 18E); (2) the thoracic uncini have very long shafts, 2-4 times the distance
between breast and crest, (Fig. 18H); and the abdominal setae ( 1 1 in segment
29) have a similar bulbous knee (Fig. 18K, L, M ) . The thoracic setae are also
similar, superior ones (3-4) slender (Fig. 18G) and inferior ones with oval hoods
(Fig. 18F). There are 11 of the latter, but the thoracic fascicles are very
damaged.
Perkinsiana ceylonica, however, is much larger, particularly in breadth, with a
body nearly 2 mm wide and with closely packed segments (Fig. 18A) of which
45 (46 on the other side) are thoracic. Fauvel (1953) noted specimens with
15-23+ thoracic setigers and it would therefore seem that high numbers of
284
P. KNIGHT-JONES
E
Figure 18. Perkinsiana ceylonica holotype: A, anterior part ofworm showing unusually long thorax; B,
anterior thorax and basal crown showing dorsal lips, the one on the left being complete with
enlarged pinnule (crosshatching shows damaged area); C, cross-section of base of dorsal radiole
showing an irregular arrangement of axial cells, but these may have a regular arrangement as in an
adjacent section, e.g. D; E, anterior thorax ventral view; F, inferior thoracic seta; G, superior
thoracic seta; H , thoracic uncinus side view; J, companion seta three-quarter view: K, abdominal
seta with a very bulbous knee, side view; L & M, as K, superior and inferior setae respectively,
‘face’ view; N, abdominal uncinus, side view. Scales: G, H, J, K, N as F; M as L.
TAXONOMY OF SOME SABELLIDAE
285
thoracic segments are normal for this species. The crown, 12 mm long, has 13
radioles each side with very fine terminal filaments. A transverse section from
the base of one from the dorsal side shows a core of about 8 cartilaginous cells
with a thin sheath, and thickened epithelium particularly at the outer corners,
representing blunt ridges along the rachis. The axial cells are very thin. The
arrangement may be like a mosaic (Fig. 18D) in one plane of focus whilst an
adjacent layer can show a more regular pattern (Fig. 18C). In side view these
cells are about four deep. The radioles are fused for a short distance at the base
where they arise from a deep cartilaginous foundation, the left and right halves
of which are joined by a massive bridge. The tapered dorsal lips are extensive,
equal in length to about 6 segments, each outer lamella being fused to a very
elongated pinnule at the base of its dorsal radiole (Fig. 18B). This arrangement
is very similar to that in P. assimilis.
The collar is gently oblique in side view but higher dorsolaterally than in
P. rubra. The ventral lappets are similar in being separated by a wide V-shaped
cleft. This cleft indents the distal margin of the first thoracic shield, which is
quite large (Fig. 18E). The other thoracic shields are shallower, being 4 times as
broad as long.
Perkinsiana minuta (Treadwell, 1941)
Holotype AMNH 2894, collected at the island of S5o SebastiBo, Brazil.
The body is 13 mm long (0.75 mm wide) with 42 segments on one side and 43
on the other, the imperfect setiger occurring halfway along the abdomen. Only
4 setigers are thoracic (Fig. 19A). Treadwell counted 5 and it is possible that he
counted thoracic shields rather than parapodia. The crown is a further 3 mm
long and has 7 radioles on each side which are fused basally. Each has pinnules
about 4 times as long as the rachis is wide and a gently tapered but blunt ended
terminal filament. The axial skeleton at the base of a dorsal radiole has 4 cells in
cross-section surrounded by a thin sheath and thick epithelium (Fig. 19B). The
outline is subquadrangular with rounded outer corners so that there are blunt
ridges along the rachis. Between one-third and two-thirds of the length, the
radiole bears faded transverse bands of pigment as well as minute irregularly
placed brown spots mostly on the sides of the rachis. The basal pinnule of each
of the most dorsal radioles is webbed to a dorsal lip in which the radiolar
support (tapered) is equal to the length of about 2 thoracic segments (Fig. 19C).
In side view, the margin of the collar is oblique and covers the base of the
crown (Fig. 19D) the most distal part forming tapered lappets which do not overlap
and are separated by a cleft that is as deep as the average thoracic segment is
long (Fig. 19E). The dorsal margins of the collar end low on the first segment
not far from the first fascicles of setae (Fig. 19C). The tori of the second, third
and fourth segments are small, but their ventral ends nevertheless indent the
sides of the ventral shields. These are not much more than twice as wide as long.
The thoracic setae form small fascicles but they are mostly represented by
broken shafts. Just one of the inferior seta could be seen by dissecting microscope
and that parapodium (3rd) was chosen for mounting. This seta has a tapered
hood 3.3 times as long as broad. In outline it is not unlike some Demonax inferior
setae, but it is nevertheless slightly spoon-shaped, to judge from the abrupt taper
of the shaft ‘within’ the hood and the thickened appearance of the lateral
margins of the ‘wings’ (Fig. 19F). Each uncinus in the adjacent torus (about 10)
286
P. KNIGHT-JONES
\\
I
I
I
0
0
F
L
Figure 19. Perkinsiana minula holotype: A, whole animal right side; B, cross-section of base of dorsal
radiole; C, anterior thorax and basal crown with a radiole splayed to show the left dorsal lip; D &
E, thorax and first abdominal segment; D, right side; E, ventral view; F, thoracic inferior seta; G,
thoracic uncinus, side view; H & J, adjacent companion seta; H, nearly side view; J, viewed almost
from behind; K, inferior abdominal seta, side view; L & M, abdominal setae from same fascicle,
inferior and superior respectively, face view. Scales: D & E as C; G-M as F.
TAXONOMY OF SOME SABELLIDAE
287
has a fairly short shaft, only a little longer than the distance between breast and
crest (Fig. 19G). The companion setae have tapered blades (Fig. 19H, J). The
superior abdominal setae are only slightly longer than the inferior setae, both
being slightly geniculate (Fig. 19K) and narrow at the heel (Fig. 19L, M). Each
abdominal uncinus has a short shaft no longer than the distance between breast
and crest.
Treadwell admitted that a specimen with so few thoracic setigers could be a
juvenile, but material misidentified variously as occelata Jlecata and microphthalma,
(USNM) agrees well with the holotype, especially the reduced numbers of
thoracic segments. USNM 44906 from Key West has 5 thoracic segments and
USNM 42793 from Puerto Rico has 4. I t would therefore seem that the reduced
thorax is characteristic. The species also differs from others in the genus in that
the hooded setae are slim distally.
Perkinsiana linguicollaris (Day, 1961 )
Holotype Cape Town. Dredged off Cape Agulhas, rocky bottom at 18m.
Material not examined as Day’s figures (1961, 1967) seem to be sufficiently
explicit.
This species is closest to P. socialis and P. fonticula with regard to the depth of
the first thoracic segment and the oblique (in side view) collar with long ventral
lappets. The collar differs however in having a small dorsolateral notch in the
already low margin near the first dorsal fascicles. The most characteristic
feature may well be the very pointed spear-like hoods of the inferior thoracic
setae (about 6 per fascicle). This is more reminiscent of hoods in some Demonax
species but the companion setae figured are certainly not Demonax in character.
The species is smaller than P. fonticula (but larger than P. socialis) with a body
length of 11 mm and width 0.3 mm. There are 12 thoracic setigers. The crown,
a further 2 mm, has 6 radioles each side with fairly long tips and a pair of
slender dorsal lips (‘palps’) hardly larger than the pinnules on the base of the
dorsal radioles. The thoracic tori are very short, with only 8-9 uncini and
companion setae, and the abdominal setae too, resemble P. fonticula in having a
bulbous knee to the presumably geniculate setae (3 per fascicle).
Perkinsiana corcovadensis (Hartmann-Schroder, 1965, Figs 27 1 & 272 as Potamilla)
Type: ZMUH p. 15224. SW Chile at a depth of 190 m.
This species seems to be closest to P. acuminata and P. assimilis in having long
tori indenting the ventral shields (such shields being about twice as broad as
long) and rounded ventral collar lappets that overlap on the midline. It may be
closer to P. acuminata in that a transverse section of the radiole shows that the
rachis is without outer ridges, but it nevertheless differs in having more cells in
the axial core, about 16 in a subcircular group, appearing in side view ‘like
bricks in a wall’ with 4-5 cells across.
It further differs from these and other species in that the collar is not oblique
in side view. The distal margin dorsolaterally is sufficiently high to cover the
fused bases of the radioles and dorsally the margin curves sharply downward
ending close to the first fascicles (not fully outlined in Hartmann-Schroder,
1965, fig. 271). Most of the setae are similar to these of P. acuminata and
P. assimilis, but the hoods of the inferior setae of P. corcovadensis (about 14
per fascicle) seem to be more spoon-shaped, similar to those of P. rubra (Fig. 12G)
16
P. KNIGHT-JONES
288
but a little longer, nearly twice as long as broad. The shafts of the thoracic
uncini are about 14 times the length between breast and crest, and the shafts of
the companion setae are similarly long but the length of their blade is modest
and more like that of P. assimilis. Hartmann-Schroder’s figure of an abdominal
seta is one of the inferior series. The superior setae, also geniculate, are more
slender at the knee and a little longer between knee and tip.
Perkinsiana littoralis (Hartman, 1967 as Potamethus)
Holotype: USNM 55570, South Shetlands at 79 m.
This species resembles P . rubra and P. ceylonica in that there is a gap between
the thoracic tori and thoracic shields, whilst the collar is slightly oblique in side
view (Fig. 20C). It differs from both in that the small ventral lappets of the
collar overlap medially, (Fig. 20A). The thoracic uncini have a shaft little more
than the length between breast and crest (Fig. 205) and both superior and
inferior abdominal setae have slender blades (cf. Fig. 20M).
Perkinsiana littoralis is a large species with a body over 40 mm long and 3 mm
wide, with 14 thoracic segments on the left and 15 on the right, showing signs of
reorganization after damage. The thoracic fascicles are large with about 10
slender superior (Fig. 20F) and about 16 inferior setae (Fig. 20G, H) which
have in face view suboval hoods and gently tapered distal points. There are
A
Figure 20. Perktnsiann Lhrulis: A-C, anterior thorax; A, ventral view; B, dorsal view; C, lateral
view; D, whole detached crown; E, showing inner view of left of crown with tapered dorsal lip and
curved ventral lip; F, superior thoracic seta, side view; G & H, inferior thoracic seta; G, side view;
H, facr view; J , thoracic uncinus, side view; K & L, adjacent companion seta; K , ‘back’ view; L,
details of another with damaged blade, side view; M, abdominal seta. Scales: B-E as A; G-M as F.
TAXONOMY OF SOME SABELLIDAE
289
about 30-45 uncini in each torus. In the 10th thoracic segment (right) all uncini
have a shaft scarcely longer than the distance between breast and crest
(Fig. 205) and the companion setae have stout shafts with moderately long
tapered blades (Fig. 20K, L).
The crown, about 13 mm long, has a deep cartilaginous base and about 18
radioles fused basally on each side. A radiole was not sectioned but no ridges
could be seen on the outside of the rachis. The tapered dorsal lip is long, equal
to the length of about 4 segments, the average thoracic shield being about twice
as broad as it is long. Fusion of the dorsal lip to a basal pinnule was not noticed.
As mentioned earlier ‘Potamilla’ antarctica Gravier, 1907 (non antarctica
Kinberg, 1867; Ehlers, 1897; Fauvel, 1916) may be this species. It is also large
(60 mm long) with a high number of thoracic setigers (up to 13) and large
numbers of thoracic setae, 10 superior and 20 inferior in one fascicle.
Unfortunately Gravier figures only setae, the most noticeable difference there
being that the thin tip of the inferior seta is longer. It would be difficult to prove
synonymy without Gravier’s type material and anyway Hartman’s name is to
be preferred because a different species has been called ‘Potamilla’ antarctica (e.g.
by Fauvel, 1916).
SUBFAMILY FABRICIINAE
Two or more sinuous filaments (possibly enlarged pinnules) adjacent to the
two most ventral radioles (Fig. 21A, B); Thoracic uncini with crest, anterior
peg and long straight shafts (Fig. 21F); companion setae absent.
Genus Oriopsis Caullery and Mesnil 1896
Radioles (1) without eyes, (2) without ridges and (3) without
appendages other than pinnules but (4) thin lateral Janges Junking each rachis
but not Junking two or more ventral filaments (without pinnules); (5) these flanges may
or may not form an obvious web basally; (6) radioles arise direct from the first
segment with no basal fusion or obvious or cartilaginous foundation; (7)
radiolar skeleton thin, only 1 or 2 cells wide; (9 & 10) dorsal lips triangular but
so small that they can only be seen easily when the two halves of the crown are
separated; (1 1) length of the first segment up to 13 times the length of the next
thoracic segment; (12) ventral sacs vestigial on the apex of the peristomium
which may or may not be fused to the ventral margin of the collar; (13) the
collar if present can be with or without a ventral cleft and therefore (14) forms
just one or two sections extending to the dorsal side (15) without lateral notches
or (16) dorsal lappets, (1 7) the gap separating the dorsal collar margins ranging
from narrow to wide; (18-22) the collar fascicles and other thoracic fascicles
all similar, each composed of a few bordered setae and a few capillaries; (23)
thoracic uncini very few, their long shafts arranged in a fan-like group; (24)
companion setae absent (here a subfamily character); (25) abdominal setigers fewer
than those of the thorax but not less than 4; (26) abdominal setae a few long
capillaries (27) in clusters of two or three; (28) abdominal uncini
subquadrangular, each with a truncated flattened shaft, the distal toothed
margin more or less parallel to the wide base and usually with a well-defined
notch at the anterior margin; (29) glandular areas indistinct, just visible
DIAGNOSIS:
16*
290
P. KNIGHT-JONES
c
b
A
H
E
Figure 21. Oriopsis hynensis: A, B & C, holotype, D-H, paratype: A, whole animal ventral view; B &
C , anterior thorax and crown; B, left side; C; dorsal view; D, inferior capillary seta; E, superior
thoracic bordered seta, ‘face’ view; F, thoracic uncinus side view; G & H, abdominal uncinus; G ,
viewed from above; H, side view. Scales: C as B, E-H as D.
ventrally as shields, each of which is divided by a transverse groove; (30) no
glandular girdle around the second setiger.
Type species Oriapsis armandi (Claparcde, 1864) senior synonym of
0. metchnikovi Caullery and Mesnil 1896.
Oriopsis h p e n s i s sp. nov.
Holotype BMNH ZB 1982.49. Paratypes BMNH ZB 1982 50-51 and NMI
1982, mounted paratype without crown ZB 1982.52.
A very small species with a body about 1.2 mm long with 8 thoracic and 5
abdominal segments (Fig. 21A). Crown 0.4 mm long with 3 radioles on each
side, each with fine membranous lateral flanges that taper distally, ending at the
base of the terminal filament. This is slender, similar in thickness to the few
pinnules below. There are just two bare filaments at the base of the two most
ventral radioles. The apex of the peristomium is not obviously bilobed and is
TAXONOMY OF SOME SABELLIDAE
29 1
distinctly separated from the collar which lacks a midventral notch. T h e collar
is unusual in being set low on the first segment, its oblique margin (side view
Fig. 21B) lower than the oblique peristome. The dorsal margins are separated
by a wide gap (Fig. 2 1C ) . The glandular shields, confined to the ventral surface,
are not very distinct, particularly posteriorly. Each except the first has a
transverse groove midway between the intersegmental ‘division’ and all have
indented lateral margins at that groove.
The thoracic fascicles including those of the collar are alike in having 3 very
slender bordered setae (Fig. 21E) and 3 shorter capillary setae (Fig. 21D). The
long thoracic uncini (3-6 per torus) have coarse teeth of which about 3 show in
profile (Fig. 21F). The abdominal setae are long paired capillaries, whilst the
abdominal uncini (6-9 per torus) are small and subquadrangular, the distal
surface covered with teeth (Fig. 21G, H). Anteriorly below the teeth is a notch
and a prow which has a thick edge and appears to be gouged. The pygidium is
simply rounded and eyespots cannot be seen in the preserved material. Tube
unknown, but all small fabriciines have fine mucous tubes that are readily
abandoned and easily replaced (Day, 1967; Lewis, 1968; Knight-Jones, 1981;
Knight-Jones & Bowden, in press).
The species was found amongst Laurencia platycephala, Codium fragile ssp.
tomentosoides and weedy rocks at about 0.25 m below LWS on the south and
east shores of Lough Hyne (Ine), County Cork, Ireland (Kitching & Thain,
1983).
Comparison with other species of Oriopsis
Oriopsis hynensis clearly differs from 0. alata (Ehlers, 1897), 0. rivularis
(Annenkova, 1929), 0. minuta (Berkeley & Berkeley, 1932), 0. neglecta, Banse
(1957)’ 0. gracilis Hartman (1969) and 0. paciJica (Rullier, 1977) which have no
thoracic collar, and from 0. armandi (Claparkde, 1864) and 0. alatoides
Hartmann-Schroder ( 1962) which have a vestigial collar. Oriopsis hynensis also
differs from the remaining species of Oriopsis in that the collar is distinct and set
low on the first segment showing the ventral apex (and the sides) of the
peristomium above the collar. Oriopsis eimeri (Langerhans, 1880), 0. crenicollis
(Annenkova, 1934) and 0. tristanensis (Day, 1954) differ further in having
scalloped margins on the collar; 0. coalescens Banse ( 1959), 0. bansei Day ( 1961) ,
0. magellanica and 0. tatalensis both Hartmann-Schroder (1962) have a narrow
gap separating the dorsal collar margins; 0. magna Banse (1957) and 0. ehlersi
Day (1961) have collars each with a ventral notch; 0. limbata (Ehlers, 1897) and
0. michaelseni Banse (1957) appear to have the triangular collar lappet free from
the apex of the peristome, but study of live material of 0. limbata from southern
Chile shows that the tip is fused to the apex of the peristomium. Oriopsis
michaelseni nevertheless differs in that the lateral part of the collar is set high
obscuring the distal margin of the first segment. Another species 0. parvula
Ehlers (1913) was given a somewhat vague description. He seemed to regard the
first setiger as being different from the first body segment like many of the early
authors. The latter is presumably the distal margin of the first segment and the
fused ‘Kopflappen’ may represent a simple apex (not bilobed) similar to that in
Fig. 21A. He describes a ventral three-cornered part of the collar but it is not
clear whether this is free from the apex of the peristomium. T h e lateral part of
the collar is described as ‘niedrig’ but it is difficult to determine whether this
292
P. KNIGHT-JONES
relates to being set low or being just shallow as in 0. michaelseni. Day (1961)
comments that 0. paruula (Ehlers, non Augener, 1918) is reminiscent of
0. michaelseni, but draws attention to Ehler’s description of capillary setae in the
thorax, compared with Banse’s bordered setae. Both types of setae are present in
0. hynensis but the superior ones are so narrow that they could almost be
mistaken for capillaries (Fig. 21E). I n view of the inadequate description of
0. parvula, the apparent absence of type material and the misidentifications of
material attributed to C. paruula (Banse, 1957; Day, 1961) it seems best to regard
0. hynensis as distinct, a t least until such time as material can be found in the
type locality Simonstown, South Africa.
SUMMARY
Potamilla neglecta, the type species of the genus and the confusingly similar
Potamilla torelli are fully described for the first time. These are arctic-boreal forms
and, unlike most species commonly put in the genus, have each dorsal lip
without the support of a radiolar midrib and the two halves of the collar
separated only by the narrow dorsal groove. Pseudopotamilla is characterized by
compound radiolar eyes, thin flanges bordering the lower dorsal margins of the
base of the crown, and a collar in four parts. I n Demonax the dorsal margins of
the collar are widely separate and the companion setae have bulbous heads and
very narrow blades. Several well known species are now included in that genus
and redescribed. Demonax langerhunsi nom. nov. is proposed for a species
which is known from Madeira, SW Wales and S Ireland under the name Sabella
(Potamilla) incerta Langerhans. This is different from the South American form
Demonax incertus Kinberg.
Potamethus, which is more remote from the foregoing genera, is redefined and
reviewed, for there are mistaken concepts of the genus in the literature. Species
typically have an elongated first segment with enlarged ventral sacs, long shafts
to the thoracic uncini, and inferior bordered setae of the abdomen only 0.6 as
long as the adjacent superior capillary setae.
Several species that used to be placed in Potamilla (and one from Potamethus)
lack the characters typifying the above genera. One of these, an abundant
sabellid of S Wales, bores into limestone and appears to be the same as
Potamilla rubra Langerhans from Madeira. This is redescribed (together with
other related species) and made the type of a new genus Perkinsiana. Oriopsis
hynensis sp. nov. from Lough Ine (Hyne) Ireland, is also described. It differs
from other species in the genus by having a collar set low on a fairly long first
segment, its ventral margin entire and free from the apex of the peristome.
ACKNOWLEDGEMENTS
I am indebted for facilities, material and information to the Directors and
staffs of the MBA Laboratory Pymouth; Dale Fort Field Centre; the Museu
Municipal do Funchal, Madeira; the Centro Investigaciones Submarinas,
Coquimbo, Chile, and other South American Institutions; The British Museum
(Natural History), London (BMNH); the Museum National d’Histoire
Naturelle, Paris (MNHN); the American Museum of Natural History, New
York (AMNH); the Hancock Foundation, University of Southern California,
TAXONOMY O F SOME SABELLIDAE
293
Los Angelos (HFUSC); the National Museum of Ireland Dublin (NMI); the
Naturhistorika Riksmuseet, Stockholm (NRS); the Royal Scottish Museum,
Edinburgh (RSM); the United States National Museum Washington (USNM);
the Universitetets Zoologiske Museum, Copenhagen (UZMC); the Zoologisches
Museum Humboldt-Universitat zu Berlin (ZMHUB); the Zoologisk Museum
Universitetet, Bergen (ZMUB); the Zoologisches Museum Universitat
Hamburg (ZMUH).
I am also much indebted to Professor J. S. Ryland for laboratory facilities at
Swansea; to Mr Thomas H. Perkins of the Florida Department of Natural
Resources, for invaluable advice on sabellid taxonomy; to Miss Iffat Chughtai
(UC Swansea), Drs K. Hiscock, J. P. Hartley (Oil Pollution Research Unit,
Dyfed) and B. O’Connor (UC Galway) for providing material from their
sabellid collections; to Professor J. A. Kitching and Dr Vivian Thain for
providing the Lough Ine material in which they had recognized the Oriopsis
species as being different from the common European form; to Dr M. Fordy and
Mr Carl Stockton for help with the scanning electron micrographs; to my
husband Wyn for advice and help in making collections; and to the Royal
Society for travel grants.
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