Download Spirorbinae (Polychaeta : Serpulidae) from Chios

Zoo1.J. Linn. Soc., 4 8 , p p . 363-385. With 13figures
August 1969
Spirorbinae (Polychaeta : Serpulidae) from Chios (Aegean Sea)
JULIE H. BAILEY
Zoology Department, University College of Swansea
Accepted for publication October 1968
Twelve species are described, of which two are new, viz. Spirorbis (Janua)gnomontcus and
Spirorbis (9.) parvulus, and three others have been recognized only very recently, viz. S.
(Pileolaria) clavus, S. ( P . ) endoumenris, S . (P.) heteropoma. Some such as S. (J.) corngatus and
S. ( P . )koehleri were widely distributed between depths of M O m, but others showed a tendency
towards depth zonation, with S.(J.)pagenstecheriabundant in shallow and S. (Spirorbis)cuneatus
in somewhat less shallow water. S. ( P . ) pseudomilitaris and S. (Marsipospira) striatus were
confined to greater depths except where the former occurred amongst ships fouling and the latter
in a dark cave. S. (P.)
militaris seems rather uncommon.
A remarkably high percentage of the species (over 80%) are opercular incubators. Outside the
Mediterranean, the only areas known to have similar percentages are Ceylon and the West
Indies, suggesting that the habit may have originated in the Sea of Tethys.
CONTENTS
Introduction and methods .
Key to the species observed.
.
Descriptions of species
.
Spirorbis (Spirorbis) Cuneatus .
Spirorbis (Marsipospira) striatus
.
Spirorbis (Pileolaria) militaris .
Spirorbis (Pileolaria)pseudomilitaris .
Spirorbis (Pileolaria) heteropoma
.
Spirorbis (Pileolaria) clavus
.
Spirorbis (Pileolaria) koehleri .
Spirorbis (Pileolaria) endoumensis
.
Spirorbis (Janua)pagenstecheri .
Spirorbis (Janua) corrugatus
.
Spirorbis (Janua) gnomonicus sp. nov.
Spirorbis (Janua) paruulus sp. nov.
Distribution and relative abundance
.
Zoogeographical considerations .
Acknowledgements .
References
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PAGE
363
365
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383
INTRODUCTION AND METHODS
Accounts of Spirorbinae from the Mediterranean include those of Claparkde (1868),
Marion & Bobretsky (1875), Marion (1879), Caullery & Mesnil (1897), Harris &
Knight-Jones (1964), Thiriot-Quibvreux (1965), Zibrowius (1967, 1968) and Harris
(1968a), but all these deal with species found in the western part of the Mediterranean,
mainly from France and Naples. The nearest records to the Aegean are those of
Sterzinger (1909) and Pixel1 (1914), from Suez, Fauvel(l955) from Israel and Sterzinger
(1910) from the northern Adriatic. Nothing seems to have been published about the
Spirorbinae of the Aegean.
JULIE H.BAILEY
364
Chios is one of the larger Greek islands and lies at 38'20" and 26'00'E, about five
miles west of the nearest point on the Turkish coast. Collections of Spirorbinae were
made at several locations off the southern and south-western coasts and from one more
northern area, during August 1967, by an expedition from the University College of
Swansea, sent at the invitation of the Chios Committee. The main collecting localities
are indicated in Fig. 1. The expedition's camp site was by Station 1, at Emborios.
10 miles
lOkm
FIGURE
1. Map of Chios showing collecting stations. 1, Emborios bay, cave and cobble beach;
2, submarine peak; 3, Cape Mastikho;4, Venetico; 5, MoulaPetra;6, Nisos Aspro;7,Foradhas;
8, Nisos Margaritis.
T o collect Spirorbinae, rocks, stones, shells, erect plants and incrustations of coralline
algae were picked up or prised out of crevices by divers, examined briefly and taken
ashore. They were then sorted over, usually with the help of magnlfying lenses, and
most of the Spirorbis were scraped with a knife into specimen tubes, to reduce the volume
of preserved material. The recorded notes did not usually include details of the substrata sampled, but gave depths and locations. Some of the collections were from a
succession of shallow depths near Emborios, one was from a shore-line cave, another
from a local fishing vessel and the rest from various greater depths down to about 60 m.
Most collections were preserved in sea-water with 5% formaldehyde and these
showed, even after five or six months, colours which have been noted in descriptions
of species. Some other material was preserved in 50% alcohol, however, to ensure that
calcareous opercular structures were not dissolved. Polyvinyl-lactophenol was used as
a mountant, for laboratory examination of setae and uncini, the detailed structure of
which was studied by phase-contrast microscopy, using oil-immersion and iodinequartz illumination.
SPIRORBINAE FROM THE AEGEAN SEA
365
Twelve species were recorded, eight of which were sinistral and four dextral. Five
of the twelve had not previously been described, but correspondencewith other specialists revealed that two of the five were known from Marseille (Zibrowius, 1968) and
. remaining two are described here as new.
another from Naples (Harris, 1 9 6 8 ~ )The
The Spirorbinae are here regarded as comprising the single genus Spirorbis, which
may be split into several subgenera according to the scheme proposed by Bailey (1969 u).
Compared with Spirorbinae from the Atlantic and most other areas, a remarkably
high proportion of the Chios species incubate their embryos in opercular brood chambers, and the physiological and zoogeographical implications of this peculiarity are
discussed later.
KEY T O THE SPECIES OBSERVED
Mouth of tube faces clockwise
.
Mouth of tube faces anticlockwise
1.
2. (1) Operculum has a massive conical talon
.
.
Operculum lacks a massive conical talon
3. (2) Tube opaque (p. 367) .
Tube translucent (p. 368)
.
.
.
.
13
.
.
3
4
. S . cuneatus Gee
S . striutus (QuiCvreux)
4. (3) Flat to concave opercular plate, peg-talon and no embryos
Embryos in a calcareous brood chamber.
2
.
.
.
-
5
8
.
.
6
Narrow talon, with a row of spines projecting from one side (p. 370)
juvenile S. militaris Clapartde
5. (4) Simple peg-like talon
6. (5) Peg-talon reaches floor of ampulla and passes through it into the stalk
(p. 372) .
. juvenile S. clavus Harris
Peg-talon does not reach floor of ampulla
.
.
7
7. (6) Tube with three longitudinal ridges, peg-talon small and eccentric
(p. 371) .
.
juv. S. heteropomu Zibrowius
Tube smooth or with indistinct longitudinal ridges ; peg-talon small
and eccentric (p. 370) .
juv. S. pseudomilitaris Thiriot-QuiCvreux
.
8.
(4) Brood chamber associated with a stack of two or three concave opercular
plates
.
.
Brood chamber a simple calcareous chamber .
.
9
10
9. (8) Two or three opercular plates, stacked and interlocking; eggs incubated
below them in a calcareous sac (p. 373)
S. koehleri Caullery & Mesnil
Two or three opercular plates stacked, but not interlocking; eggs
incubated between the plates (p. 374) .
. S. endournensis Zibrowius
.
23
JULIE H. BAILEY
366
10. (8) Brood chamber with one or more spiny processes
Brood chamber lacking spines (p. 372) .
.
11. (10)Brood chamber with many distal spiny processes
Brood chamber with a single distal process (p. 371)
.
.
11
S. clavus Harris
.
12
S . heteropoma Zibrowius
12. (1 1) Brood chamber flat or slightly domed, witha peripheral spiny rim (p. 370)
S . pseudomilitaris Thiriot-Quikvrewr
Brood chamber helmet-shaped, with some spines forming a peripheral
rim and others scattered over the surface (p. 370) . S . militaris Clapartde
13. (12) Tube with three longitudinal ridges
.
Tubesmpoth
.
.
.
14
17
14. (13) Opercular plate flat, with a round peg-like talon
Opercular plate concave or convex
.
S . pagenstecheri Quatrefages
. 15
15. (14)Opercular plate with a bilobed eccentric talon (p. 375)
juvenile S. corrugatus Caullery & Mesnil
.
. 16
Operculum lacks a talon
16. (15) Opercular plate with no distal process, except for a thin peripheral
. older S. corrugatus Caullery & Mesnil
collar (p. 375) ,
Opercular plate with a distal median crest (p. 376) S. gnomonicus sp. nov.
17. (16) Concave plate forming the top of a brood chamber containing only about
two embryos; mouth of tube coiled into an almost apical position
(p. 377) .
. S.parvulus sp. nov.
DESCRIPTIONS OF SPECIES
Genus Spirorbis Daudin (1800)
Hermaphrodite serpulids living in a dextrally or sinistrally coiled tube. The thorax
has no more than four segments, and the opercular stalk lacks pinnules.
Subgenus Spirorbis sensu stricto
Sinistral tube; thorax of three segments; each collar seta with a proximal fin and a
distal toothed blade ; egg-string incubation inside the tube.
'
Spirorbis (Spirorbis) cuneatus Gee (Fig. 2)
Sinistral, white tube, 1-1-5 mm in greatest diameter across the coil, with three
longitudinal ridges forming teeth over the mouth. The last whorl may partially overlap
previous whorls and forms a flange against the substratum.
SPIRORBINAE FROM THE AEGEAN SEA
367
The operculum has a flat or slightly concave plate and a massive conical talon with
three pointed lobes, one of them on each side and the other terminal.
Three thoracic segments. Collar setae each with a cross-striated distal blade and a
proximal fin. Second and third segments have simple setae, in addition to which the
third has recurved, toothed setae.
Ten to fourteen abdominal segments with finely toothed geniculate setae. Egg
strings incubated in the tube.
A
B
"i
a
E
P
in
N
C
H
FIGURE
2. Spirorbis (Spirorbis)cuneatus. A, Tube; B, face view and C, lateral view of operculum;
D, collar seta; E, capillary seta of first segment; F, seta of second or third segment; G, seta of
third segment; H, thoracic uncinus; J, abdominal uncinus; K, abdominal seta.
The specimens agreed closely with Atlantic coast material studied by Gee (1964) and
with the description of Zibrowius (1968) from Marseille. The lobes of the talon vary
somewhat in their arrangement and it seems possible that this species provided the
material on which Caullery & Mesnil (1897) founded their description of Spirorbis
mediterraneus. That older name might thus be adopted here (Nelson-Smith & Gee,
1966 ;Nelson-Smith, 1967), but Zibrowius (1968) reported from Marseille two specimens which were distinct from S . cuneatus and agreed perfectly with the original
description of S . mediterraneus. They seemed to represent either a distinct species or
immature and somewhat aberrant S . militaris. Pending further studies the older name
should perhaps lapse, as having been based upon an inadequate description. Distribution elsewhere: Naples (Harris, 1968a) ; Marseille (Zibrowius, 1968) ; Costa Brava
(Harris & Knight- Jones, 1964) ;west coasts of Britain.
JULIE H. BAILEY
368
Subgenus Mursipospiru Bailey (1969a)
Sinistral coiling ; tube usually translucent, with transverse ridges ; thoracic brood
pouch with a stalk; traces of a fourth thoracic segment ;collar setae each with a fin and
cross-striated blade ;abdominal setae brush-like.
Spirorbis (Marsipospiru) striutus (Quitkreux) (Fig. 3 )
Sinistral translucent tube, with a round mouth and prominent transverse ridges.
The last whorl partially covers the preceding whorls. Diameter across coil 1-2.5 mm.
Slightly concave opercular plate, with a massive conical talon bearing a ventral spur,
which is more developed in large specimens than in smaller ones, and two lateral lobes,
each with a row of knobs.
_I
D
F
0
N
0
E
ni
Ln
0
C
FIGURE
3. Spirorbis (Mursipospiru) striutus. A, Tube; B, face view and C, lateral view of operculum; D, collar seta; E, capillary seta of first segment; F, seta of second and third segments;
G, seta of third or (when present) of fourth segment; H, thoracic uncinus; J, abdominal
uncinus; 11, abdominal seta.
I n the original material from Roscoff, the type locality, the thorax has ‘three and three
quarters segments’, the fourth segment being represented by a complete parapodium
on the convex side and a neuropodium on the concave side. Most specimensfrom Chios,
however, only have ‘three and a quarter segments’ (i.e. a neuropodium on the concave
side only) ;but one of the larger specimens may be said to have three and a half, the
extra half segment being represented by the neuropodia on both concave and convex
SPIRORBINAE FROM THE AEGEAN SEA
369
sides. The rudiments of the extra thoracic segment probably develop during growth,
as is known to occur in Spirorbis (Paralaeospira) malardi and Spirwbis (Paradexiospira)
witreus (Crisp, Bailey & Knight-Jones, 1967). It is probable that the notopodium on
the convex side of S. striatus at Roscoff appears during further growth, for the largest
Chios specimens were smaller than those recorded from Roscoff. The absence of these
parapodial rudiments from the Chios specimens seems better regarded as a neotenous
trend, than as a basis for distinguishing them specifically from the Roscoff forms (see
also Bailey, 1969a).
Collar setae each have a proximal fin separated by a gap from a distal, cross-striated
blade. The fin has three large and three smaller teeth. Setae on the second segment are
simple blades, those on the third and fourth when present bear recurved hooks. The
abdominal setae are brush-like.
Eggs are redlorange and incubated in a thoracic brood pouch in the tube.
This species differs from S. ambilateralis Pixell (1912), which has four complete
thoracic segments, and S. racemoms Pixell (1912) which has three and a quarter and
lacks lateral knobs on the opercular talon (see also Rioja, 1962). S. translucem Bailey
(1968) also lacks rounded knobs laterally and has angular ridges instead.
Distribution elsewhere : Roscoff and Cap Ferrat (France).
Subgenus PiZeolaria Claparkde (1 868)
Sinistral coiling ;incubation in an opercular brood chamber; three thoracic segments ;
fin and blade collar setae (which may bear cross striations).
Spirorbis (PiZeolaria)miZitaris Claparkde (Fauvel, 1927)
Sinistral, smooth tube with a wide, round mouth, transverse growth lines and the
whole spiral loosely coiled. Diameter across coil 1-3 mm.
This species shows opercular dimorphism (Fig. 4). The opercula of two immature
stages have previously been described as separate species (Zibrowius, 1967). The firstformed operculum has a three-pronged talon and was called s.cornuarietis (Caullery
& Mesnil, 1897). The second stage has a long talon, with a row of projecting spines,
and was called S. beneti Marion (1878). The brooding operculum develops below the
'beneti' type operculum as a calcareous dome. A frill of spines develops around the dome
and the distal surface becomes covered with individual spines.
There are three thoracic segments. The collar setae each have a cross-striated distal
blade separated by a gap from a proximal toothed fin. The second segment has setae
with simple blades, the third bears some simple setae and others with recurved teeth.
Abdomen of 8-10 segments with geniculate toothed setae.
Distribution elsewhere : English Channel, Madeira and Naples (Caullery & Mesnil,
1897) ;north-west Spain (Rioja, 1923) ; Mexico (Rioja, 1942).
Spirorbis (Pileolaria) pseudomilitaris Thiriot-QuiCvreux, 1965
Tube sinistral, white and smooth, diameter of coil 1-2 mm.
There is opercular dimorphism (Fig. 9, similar to that in S. militaris but with only
one type of primary operculum. In immature specimens the opercular plate is concave,
with an eccentric peg talon. The incubatory chamber can be seen developing under-
JULIE H. BAILEY
370
FIGURE
4. Spirorbis (Pileolaria) miiitaris. A, Face view and B, side view of primary operculum
(‘S.cornuarietis’ type); C, face view and D, side view of primary operculum (‘S.beneti’ type);
E,‘S.bmeti’ type operculum which lay in the distal groove of F,which is a developing opercular
brood chamber; G, side view and H,face view of a brood chamber with eggs.
neath the plate, as a thin calcareous dome. The adult loses the primary plate and the
flat-topped incubatory chamber has a peripheral frill of spines. There may also be a
few spines placed more centrally on the distal surface.
Three thoracic segments. Collar setae each with a cross-striated distal blade, separated
by a gap from a proximal toothed fin.Second and third segments both have simple
blades. Abdomen of 15-20 segments with geniculate toothed setae.
C
1:
FIGURE5 . Spirorbis (Pileo1aria)pseudotnilitaris.A,Face view and side view of primary operculum;
B, primary plate with incubatory chamber developing underneath; C, views of incubatory
chamber; D, views of chamber full of eggs.
SPIRORBINAE FROM THE AEGEAN SEA
371
Distribution elsewhere:Villefranche, Toulon, Tamaris-sur-mer (Thiriot-Quikvrew,
1965); Marseille (Zibrowius, 1968); Naples (Harris, 1968a, as S. berkeleyanus). I n
the French material the tube bears longitudinal ridges.
Spirorbis (Pileolaria)heteropoma Zibrowius, 1968 (Fig. 6)
Sinistral tube 1-2 mm in diameter, with two or three longitudinal ridges. The mouth
is round and often coils up on previous whorls.
There is opercular dimorphism. The primary plate is very similar to that of S.
pseudomilitaris, with a concave plate and eccentric peg-talon.
0
FIGURE
6. Spirorbis (Pileolaria) heteropoma. A, Tube; B, face view of primary operculum; C &
C1, two views of incubatory chamber full of eggs, with primary operculum still attached; D &
D1, two views of another incubatorychamber full of eggs; E, collar seta; F, capillary seta of first
segment ; G, seta of second or third segment;H, seta of third segment;J, thoracic uncinus; K,
abdominal uncinus; L, abdominal seta.
A domed chamber develops underneath this, forming a helmet-like calcareous brood
chamber, with a distal spine. The peg of the talon remains attached to the spine until
it is sloughed off.
There are three thoracic segments. Collar setae each have a cross-striated distal
blade separated by a gap from a proximal toothed fin.The second segment has simple
setae, the third both simple and recurved toothed setae.
Abdomen with 8-12 segments bearing geniculate setae.
Tentacles tinged yellow, thorax red, hepatic region brown, abdomen pale yellow.
Eggs in the domed opercular chamber pale yellow.
372
JULIE H. BAILEY
Distribution elsewhere : Marseille (Zibrowius, 1968); Naples (Harris, 1968a as
S . moerchi).
Spirorbis (Pileoluria) cluvus Harris, 1968a (Fig. 7)
Sinistral, white tube with two faint longitudinal ridges towards the inner curve of
the whorl. Round mouth with a thickened rim, diameter of coil 1-2 mm.
There is opercular dimorphism. In small individuals the operculum has an ampulla
topped by an uneven calcareous plate, with a peg-like talon extending to the floor of the
chamber and into the stalk.
Prior to brooding the ampulla increases in length considerably, and the opercular
plate and talon appear relatively smaller, so that the peg no longer reaches far into the
ampulla. Calcareous thickening of the roof and walls takes place under the opercular
plate. This plate eventually drops off, leaving the elongated ampulla borne eccentrically
on the stalk and calcified except for one side. When incubating, the eggs can be seen
clearly in this area and do not appear to be covered over.
FIGURE
7. Spirmbis (Pileolaria) clawus. A, Tube; B. primary operculum; C, developing incubatory chamber;D, incubatory chamber with eggs; E, collar seta; F, capillary seta of first segment;
G, seta of second and third segments; H, seta of third segment; J, thoracic uncinus; K,
abdominal uncinus; L, abdominal seta.
There are three thoracic segments. The collar setae each have a cross-striated, distal
blade separated by a gap from a proximal toothed fin. The second segment has simple
blades, the third a mixture of simple blades and recurved, toothed setae.
Abdomen of 10-12 segments with simple, toothed setae.
Tentacles colourless, thorax orange, hepatic region orange/yellow, abdomen
colourless. Eggs in operculum yellow.
Distribution elsewhere: at a depth of 55 m off Mergellina, near Naples (Harris,
1968~).
SPIRORBINAE FROM THE AEGEAN SEA
373
Spirorbis (Pileoluriu) koehleri Caullery & Mesnil (Fig. 8)
Sinistral, white tube with three obvious longitudinal ridges, which are divided into
short spines along the outer whorl and form teeth over the round mouth.
Operculum consists of two or three concave, calcareous plates, which interlock by
means of a peg and socket arrangement at the periphery. Eggs are incubated below
the last plate and the peg of this plate is elongated to form a keel, which contributes to
the calcareous wall of the chamber. A fragile calcareous sac forms under this, interlockingwith the keel of the plate above, which thus helps to keep the eggs in the chamber.
FIGURE
8. Spirorbis (Pileolaria) koehleri. A, Tube; B, operculum face view; C, incubatory
chamber;D, incubatory chamber showing eggs ;E, collar seta ;F, capillary seta of first segment;
G, seta of second or third segment;H, seta of third segment ;J, thoracic uncinus ;K, abdominal
uncinus; L, abdominal seta.
There are three thoracic segments. The collar setae each have a simple, distal blade,
separated by a gap from a proximal fin of three large and six smaller teeth. The second
segment has setae with simple blades, the third some with simple and others with
recurved, toothed blades.
The abdomen consists of 10-15 segments, bearing geniculate setae with large,
rounded teeth.
Tentacles pale yellow, thorax yellow/buff, hepatic region light brown, ovary yellow,
rest of abdomen colourless, eggs in operculum pale yellow.
Distribution elsewhere : La Ciotat ; Marseille (Zibrowius, 1968) ; West Indies
(Bailey, 1969b).
374
JULIE H. BAILEY
Spirorbis (Pileolaria) endournensis Zibrowius, 1968 (Fig. 9)
Sinistral, white tube with three smooth l o n g i t u d d ridges, which extend over the
mouth as teeth. The outer whorl lies flat against the substratum and forms a closely
adherent peripheral flange. Diameter across coil 1-2 mm.
Operculum bears one or more plates, each with a deep eccentric concavity, from which
a peripheral projection extends proximally and rests in the concave part of the plate
below. Eggs are incubated between the plates and calcareous walls are formed round
the brood chamber by thin peripheral and proximal extensions of the plate which
lies immediately above it.
D
FIGURE
9. Spirorbis (Pileolaria) endoumpnsis. A, Tube; B, side and rear views of opercular plate;
C, operculum with eggs; C1,operculum after liberatinglarvae; D, collar seta; E, capillary seta
of first segment; F, seta of second or third segment; G, seta of third segment; H, thoracic
uncinus; J, abdominal uncinus; K, abdominal seta.
There are three thoracic segments. Collar setae each have a distal blade, separated
by a gap from a proximal fin, which comprises three large and five smaller teeth. Setae
of the second segment have simple blades ; some on the third have simple blades and
others recurved hooks.
The abdomen has 12 segments, bearing geniculate setae with large rounded teeth.
Distribution elsewhere :Marseille (Zibrowius, 1968).
Subgenus Junuu Saint-Joseph
Dextral coiling ;incubation in an opercular brood chamber, three thoracic segments ;
collar setae without a gap between the fin and blade, or lacking a fin altogether.
SPIRORBINAE FROM THE AEGEAN SEA
375
Spiror6is (Janua)pagemtechmi Quatrefages (Fauvel, 1927)
Dextral, white tube with three longitudinal ridges, the last whorl not overlapping
previous whorls. Diameter across coil 1-1.5 mm.
Opercular plate flat, with a peg-like eccentric talon. Opercular walls form the sides
of an incubatory chamber, but do not appear to be strengthened by calcareous deposits.
There are three thoracic segments. The collar setae are simple geniculate blades,
each with a fin not separated from the blade. The blade is not cross-striated. Setae of
the second and third segments have simple blades without fins.
Abdomen has 6-10 segments bearing geniculate setae with finely toothed blades.
Distribution elsewhere : west Mediterranean, north-west coasts of Europe, Israel
(Fauvel, 1955), Ceylon (de Silva, 1965), Galapagos (Bailey & Harris, 1968).
Spirmbis (Janua)corrugutus Caullery & Mesnil (Fig. 10)
Dextral, white tube with three longitudinal ridges and a round mouth. The last
whorl may partially overlap the preceding whorl.
1
E
F
G
H
FIGURE
10.Spirorbis (~anua)
corrugatus. A-D, Views of opercula with developing embryos at
different stages; E-H, views of juvenile opercula.
Opercular plate slightly concave, bearing in young individuals a peripheral talon
which is often bilobed. When the plate forms the top of a brood chamber, however, it
often lacks any talon, but has a distinct rim protruding all round the periphery.
The side walls of the brood chamber are strengthened distally by a thin calcareous
wall, which has numerous minute perforations. A basal plate is formed under a brood
chamber containing well developed embryos.
There are three thoracic segments. The collar setae are simple geniculate blades,
those on the convex side being larger, with cross-striations, whilst those on the concave
side lack cross-striations. The second and third thoracic setae are simple blades.
The abdomen has 8-14 segments, bearing geniculate setae with fine teeth.
JULIE H. BAILEY
376
These abundant forms agree quite well with the descriptions of S. corrugatus from
north-west Spain by Rioja (1923) and from the Adriatic by Steninger (1910), except
that the tubes of those forms were smooth. Caullery& Mesnil(1897), however, who gave
the first adequate description of this species, found that their material (from near
Cherbourg) had ridged tubes. The species is fairly close to S. ceylonicus Pillai (1960)
and to other forms with which, indeed, it may be synonymous (Zibrowius, 1968).
Distribution elsewhere : Adriatic, west Mediterranean, north-west Spain ; Roscoff
(L’Hardy & Quikreux, 1964) ; Cherbourg ; Caribbean (Augener, 1936).
Spirmbis (Janua) gnomonicus sp. nov. (Fig. 11)
Dextral, white tube with three longitudinal ridges and obvious transverse growth
lines. The mouth is round and the last whorl coils up on the previous ones. Diameter
across coil 0.75-1 mm.
Opercular plate flat or slightly convex with a very distinctive median crest projecting
distally, like the gnomon of a sun-dial. S. unicornis Bailey (1968) and S.epichysis Bailey
(19693) seem to be the only other known species of theJunua complex, which have the
operculum bearing a distal process. Eggs are incubated below the opercular plate and
there is then some calcareous thickening of the ampulla walls. A basal calcareous plate
develops below the embryos.
C
FIGURE
11. Spirorbis (Janua)gnomonicussp. nov. A, Tube; B, operculum with developing brood chamber; C, operculum full of eggs; D, collar seta; E, seta of second or third segment; F, seta of third segment; G ,abdominal uncinus; H, thoracic uncinus;J, abdominal seta.
SPIRORBINAE FROM THE AEGEAN SEA
377
Three thoracic segments. Collar setae are simple striated blades, with no trace of
fins and without cross striations. On the second segment the setae have simple blades
and on the third there is a mixture of some simple and other recurved, hooked blades.
There are five abdominal segments, bearing geniculate setae with fine teeth.
Found commonly on stones in samples from stations 1 (four samples), 2(3), 3(2),
4(2), 7(1). Depths from 5 to 40 m.
Holotype and paratypes deposited in the British Museum (Nat. Hist.), Register
Nos. 1968.91 and 92.
Spirorbis (Janua) parvulus sp. nov. (Fig. 12)
Dextral, white smooth tube. Last whorl coils up, so that the round mouth occupies
an almost apical position over the umbilicus. Diameter across coil 0-75 mm.
Slightly concave opercular plate with a thickened edge which forms the top of the
brood chamber. There is a basal plate and no more than two embryos have ever been
seen through the transparent walls of the chamber.
0
D
FIGURE
12. Spirorbis (Janua)parvulussp. nov. A, Tube; B, operculum with two embryos;
C, collar seta from convex side of the thorax; D, collar setafrom concaveside ;E, seta of second
or third segment; F, thoracic uncinus; G, abdominal uncinus; H, abdominal seta.
There are three thoracic segments. The collar setae are simple blades, cross-striated
on the convex side of the body, simple on the concave. The setae of the second and third
segments are simple blades.
There are only three abdominal segments, each with uncini and a single geniculate,
fine-toothed seta on each side.
378
JULIE H. BAILEY
Found occasionally on stones in samples from Stations 1 (4 samples), 2(1), 4(1),
7( l),with no more than ten specimens in each collection. Depth 0-25 m. The shape
ofthe tube is characteristic. It is only rarely that the tubes of S. corrugatus approach this
shape. Some individuals of S. cormgatus, measuring less than 1mm in diameter across
the coil, also bore embryos. These usually contained three or more embryos, however,
and the bilobed eccentric talon, which is characteristic of juvenile S. corrugatus, could
be seen clearly through the transparent walls of the brood chamber. Lack of a talon in
S.parvuzus is highly characteristic.
Holotype and paratypes deposited in the British Museum (Nat. Hist.), Register
Nos 1968.89 and 90.
DISTRIBUTION AND RELATIVE ABUNDANCE
The distribution of species between collections is indicated in Table 1, in which the
locations are arranged in order of depth, with the shallowest on the left. The main
collecting site was Emborios bay (Station 1in Fig. l),which is very enclosed and shallow.
At the mouth of the bay, on the south side, is a large cave, with a small wave-washed
entrance, an average depth of 10 m and very reduced illumination. Outside the cave
the bottom shelves to about 15 m depth. All this area is littered with stones and small
boulders, very suitable for Spiror6is. Just south of the bay is a storm beach, of cobble
stones.
Station 2 was a rocky submarine peak, about 2.5 km south of Emborios and 100-200
m off shore. It rose from a sandy bottom, about 25 m deep, and came to within about
10 m of the surface. Station 3 was just south of Cape Mastikho and Station 4 south of
the island of Venetiko. At both those sites the shore cliffs continue down almost vertically
to about 30 m and then go on quite steeply to beyond diving range. Station 5, just
south-east of an isolated rock, Mikro or Moula Petra, was also fairly steep and deep.
Station 6 was off the small island of Aspro or Kokkina. Collections off the west point
of Aspro went down to about 30 m, others from the south side were from a more shallow
rocky bottom. Station 7 was around some rocks (Foradhas) which just break the surface
about 600 m off shore. Station 8 was near Marmaron, off the north-west point of Nisos
Margaritis. There were few Spirorbis from there, perhaps because those collections
were rather general ones, which did not include many suitable substrata. A few dives
were made off the north-west part of the main island, from Volissos to Point St. Nikolaos,
but that coast is shallow and wave-washed, with a high sand table. The rocks and stones
off shore are therefore too abraded to be suitable for Spirorbinae, which were not seen
there, whilst the shore rocks were inaccessible, because of heavy surf. All those collections remote from Emborios involved overland journeys and coastal voyages in an
inflatable speed boat.
Table 1 indicates the distribution of species between various depths. Spirorbis
corrugatus was the dominant species in shallow water, along with S. endournensis and
S. cuneatus. Of the Janua complex, only S. gnomonicus seemed to favour the deeper
locations. S.heteropoma was perhaps the most generally distributed, at all depths except
the water’s edge. Marsipospira and most of the species of Pileolaria occurred predominantly at greater depths. They also occurred very commonly, however, in collections
JULIE H. BAILEY
380
made from near the back of the cave (shown on right of Table 1).Their absence or
scarcity in shallow water would therefore appear to be due to their shunning strong
light.
A small series of samples were scraped from cobble stones at measured depths off
the beach just south of Emborios. The percentage contribution of each species to the
Spirorbis population at each depth is indicated in Fig. 13. As in the bay, the species of
the Janua complex were dominant in the shallow water samples, but Spirorbis
endoumertSiS was not found off the beach, perhaps because it favours more sheltered
environments.
Scale
A
I
B
C
;j
I
D
E
F
0-5
G
FIGURE
13. The distribution of Spirorbis on cobble stones at various depths off the beach south
of Emborios. A, S. cormgatus; B,S. pagmtecheri; C, S. gnomonicus;D, S.parvulus; E,S.
cuneatus F, S. koehleri; G, S. heteropoma. Scale indicates percentage of each species in the total
population of Spirorbis at each depth sampled ( 0 , 2 , S , 10,12 m).
The caique, a local fishing vessel from which scrapings were made by Mr A. R. D.
Stebbing, was often tied alongside the Emborios jetty and was quite heavily fouled by
Spirorbis and by Balanus amphitrite (the latter not being otherwise seen in Emborios
Bay). Evidently Spirorbis may be readily transported by coastal vessels.
ZOOGEOGRAPHICAL CONSIDERATIONS
Of the species found on Chios two (S. cuneatus and S.pugenstechmi) extend to
British waters (Gee, 1964) whilst another three (S.strktus, S.militaris and S. corrugatus)
reach the Channel Islands and Roscoff. The pattern of distribution of the Chios species
between subgenera, however, is very different from that on the Atlantic coast of Europe.
Remarkably enough, the Atlantic coast pattern resembles that on the Galapagos
(Bailey & Harris, 1968) rather than that on Chios, the latter being characterized by a
very high proportion of species with opercular incubation. It is interesting to consider
the nature of this pattern in some other of those few areas of the world from which the
Spirorbinae have been studied.
Table 2 shows the distribution of species between 15 such areas, including Chios.
The Spirorbinae of the rest of the Mediterranean are dealt with by the authors named
382
JULIE H. BAILEY
in the introduction. The record of S. borealis from Banyuls (Laubier & Paris, 1962) is
perhaps questionable in the light of a later survey (Harris & Knight-Jones, 1964) which
included the Banyuls area. The species of the western basin include most of those found
at Chios, with the addition of S . (Spirorbis) infundibulum Harris & Knight-Jones,
S. (Leudora)laevis (Quatrefages) and S . (Junua) pusilluides (Saint-Joseph). The Spirorbinae at Suez need further study.
Outside the Mediterranean, the only other areas knownto have such a high proportion
of opercular incubators are Ceylon (Pillai, 1960; de Silva, 1961, 1965) and the West
Indies (Bailey, 1969 b). With the single exception of S. (Junua) pagenstecheri the
Ceylon species are different from the Mediterranean ones, but some are obviously quite
closely related and the pattern of representation in subgenera is very similar. In the
Pacific only the eastern part is well known and the pattern there, revealed for Mexico
by Rioja (1942,1962) and for the Vancouver area by Pixell (1912) and Potswald (1969,
includes only between 40% and 50% of species incubating in the operculum.
In western Europe there are records for north-west Spain (Rioja, 1923), Roscoff in
north-west France (L’Hardy & QuiGvreux, 1964) ; south-west Britain (Gee, 1964;
Harris, 1968b); Norway (Bergan, 1953) and the Arctic (Fauvel, 1909; Augener, 1913).
Off Spain about half the species are opercular incubators, but off Roscoff this fraction
falls to about a third. On analysis it seems that the Bay of Biscay acts as a barrier more
for northern, littoral, tube-incubating forms t h n for the opercular incubators from the
Mediterranean.
In the South Atlantic two areas have been studied, Patagonia by Caullery & Mesnil
(1897) and the tip of South Africa by Day (1961, 1963). S . adeonella Day (1963) was
originally placed in Paralaeospira, but in Table 2 it is placed provisionally in Marsipospira, because of the form of the tube. This point needs further study, but it has no
important bearing on the present discussion.
Table 2 shows a clear tendency for forms with traces of a fourth thoracic segment
(subgenera Paralaeospiru, Paradexiospira and Marsipuspira) to be favoured by low
temperatures. It is only to be expected that tropical conditions would favour neotenous
loss of this character, which develops gradually in older individuals (p. 369). High
temperatures appear to favour opercular incubators, and might be expected to present
special problems for the embryonic respiration of tube incubators, particularly where
there is not much water movement. In barnacles embryonic development may be
retarded or terminated by anoxia, at high temperatures (Crisp, 1959). Opercular
incubation would presumably help respiration, but it seems likely that calm conditions
would be necessary for this habit to be evolved.
That conclusion may be drawn tentatively from two considerations. First, opercular
incubators are less common off Galapagos and the west coast of Mexico, where the
temperatures are high but the waters are presumably more disturbed by wave action
than are those of the Mediterranean. Secondly, the broods of ill-adapted opercular
incubators would be lost very readily in turbulent conditions, for in several species the
opercular brood chambers are open proximally. It seems likely that the embryos are
pushed into them by that route, the parents probably withdrawing into their tubes
whilst spawning takes place (Bailey, 1969~).This problem, of how the eggs reach the
brood chamber, is still not properly resolved, but the balance of opinion seems to favour
SPIRORBINAE FROM THE AEGEAN SEA
383
their being pushed in basally, from the outside. The opercula of several Chios species
look like inverted cups, which are more or less open basally or laterally (Figs 6 and 7),
and the embryos of S . cluvus are readily displaced through the lateral opening, when
material is being manipulated during laboratory study. This species would seem to be
ill-adapted to retain its embryos, in turbulent conditions, and it may be significant
that so far it has been found only in fairly deep water. One British opercular incubator,
S. gsanulatus, is notably restricted to calm environments (Bailey, Nelson-Smith &
Knight-Jones, 1967), although it seals its brood chamber effectively. Admittedly some
others, like S. pugemtechmi, can live on wave-washed coasts, but it seems likely that
turbulence would be an important hazard during the early evolution of opercular
incubation, if the embryos are pushed into the brood chambers basally from the outside.
One can thus imagine that the partially enclosed, semitropical Sea of Tethys may
well have been the cradle of evolution for opercular incubators. Time has allowed more
speciation of such forms in the area centred on the Mediterranean than in other regions.
Fully adapted opercular incubators have subsequently reached latitudes where winds
and tides are stronger, but have not yet been recorded from Patagonia. It should perhaps
be noted that none of the three methods of brood retention found in modern Spirorbinae
can be regarded credibly as ancestral to the others. It is more likely that each method
(1, egg strings fixed to tube wall as in the subgenus Spirorbis; 2, thoracic pouch fured
to a tentacular stalk as in the subgenus Mursipospiru; 3, opercular brooding as in the
subgenus Pileoluriu) has been derived independently, from ancestral serpulids spawning
into their tubes (Bailey, 1 9 6 9 ~ ) .
The links between the Mediterranean, Ceylon and the West Indies, suggested by
Table 2, involve relatively few species, which agrees with the remark of PCr&s(1967)
that the zoogeographical links between the Mediterranean and the Indian Ocean
involve genera and not species. The subgenera of Spirorbis could well be regarded as
separate genera (see Bailey, 1 9 6 9 ~ )The
. suggestionthat opercular incubators may have
originated in the Tethys region does not, of course, require them to have lived in the
Mediterranean area ever since. Ruggieri (1967) has suggested that the Tethys fauna
was probably removed from the Mediterranean area by adverse conditions in the
Miocene, after the connection with the Indian Ocean had been broken (though he
admits the possibilities of survival in some basins).
ACKNOWLEDGEMENTS
I am indebted to the S.R.C. for a research studentship and to my academic supervisor
Professor E. W. Knight- Jones for collecting the material, with the help of other members
of the expedition. The Chios Committee, particularly Dr Philip Argenti and Mr D. J.
Chandris, initiated and supported the project, in collaboration with the University
College of Swansea. Dr Helmut Zibrowius, of Marseilles, gave me helpful information
and Mr Tegwyn Harris, of Exeter, allowed me to use a prepublication copy of his
invaluable key to the Spirorbinae of the world.
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