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THE

EAY SOCIETY

INSTITUTED MDCCCXLIV.

V&iiyy^

This Volume is issued to the Subscribers to the Ray Society for the Year 1873.

LONDON

MDCCCLXXIY.

BRITISH MARINE ANNELIDS.

It is intended, on the completion of this Monograph, to issue Title Pages, Tables of Contents,
and Lists of the Plates for each volume, and an Index to the whole work. An Index to the species,
etc., described in Vol. I is inserted at the end of the present part.

THE

EAY SOCIETY

INSTITUTED MDCCCXLIV.

This Volume is issued to the Subscribers to the Ray Society for the Year 191.0.

LONDON

PRINTED BY ADLARD AND SON, LONDON AND DORKING.

MONOGRAPH

/

OF THE

BRITISH ANNELID^.

YOL. II.-PART II.

POLYCH^ETA,

SYLLITLE TO ARICIHLE.

Pages 233—524; Plates LI— LVI, Coloured, and LXXI— LXXXVII, Uncoloured.

BY

WILLIAM CARMICHAEL McINTOSH, M.D.Edin.,

L.E.C.S.E., LL.D, P.E.S., F.R.S.E, F.L.S., O.M.Z.S., V.P. LIT. & PHIL. SOC. St. AND, V.D., etc.

PROFESSOR OF NATURAL HISTORY IN THE UNIVERSITY OF ST. ANDREWS ; DIRECTOR OF THE UNIVERSITY
MUSEUM AND OF THE GATTY MARINE LABORATORY.

LONDON :

PEINTED FOE THE EAT SOCIETY.

1910.

MAY 3 1977

TEMPORABY PREFACE.

About 70 species are described in the present part (Part II, Vol. II), a smaller
number than in the previous part, but the coloration in some is not less beautiful than in
those previously depicted.

For the adequate rendering of the coloured plates the Carnegie Trust gave a grant
of £200, and the Royal Society £100 for the previous part. I cannot too gratefully
acknowledge the liberality and courtesy of both of these important bodies.

To Dr. R. Dohrn of Naples, Dr. Allen of the Plymouth Marine Laboratory,
Dr. Goodrich of Oxford, Canon Norman, Dr. Haddon, and Dr. Scharff, I am under
obligations for valued aid in connection with specimens ; whilst to Mr. Kappel of the
Linnean Society, Mr. White of the Royal Society, Mr. Waterhouse of the Zoological
Society, Mr. Maitland Anderson and Mr. Smith of the University Library, St. Andrews,
and to many fellow- workers in the department, I am equally indebted for aid in
connection with the literature of the subject.

I have also to thank Miss Walker for her continued and valued labours with pencil
and brush.

Mr. Hopkinson, the Secretary of the Ray Society, has been indefatigable in the
correction of proofs, in kindly verifying certain references, and in various ways giving
advice and help.

W. C. M.

GtAtty Marine Laboratory,

St. Andrews;

September 1st, 1910.

SYLLIDJE. 23:

Family IX. — Syllim-, continued.

By the careful search of the tidal region at Torquay, Major E. V. Elwes has made a
notable addition to the Syllidge of Britain since the issue of the last part. He followed
the plan of immersing roots of tangles and similar growths from the rocks in sea-water,
when the Syllids crawl to the water-line and are thus easily removed for examination.
Major Elwes kindly forwarded such preparations as he had made, and has thus
rendered it possible to include them in the present part. Should living forms be in
future obtained they may yet be represented in the work. The success of this observer
indicates what may yet be accomplished in the group by skilful and persevering
search in suitable localities. Further knowledge of the development of the Syllids has
been gained by Mr. Gravely, 1 who found a pelagic stage probably of Odontosyllis gibba,oS
Port Erin, Isle of Man. He also gives an account of three larvse which he connects
with Syllids, 2 his A, B, and C respectively, from the same locality, but further
examination would appear to be necessary.

The notice of these also affords an opportunity for reference to the remarkable
aberrant form Iclithyotoinus sanguinarius, found by Lo Bianco on the Neapolitan eel,
Myrus vulgaris, and ably described and figured by Eisig 3 and referred by him to the
Syllids. It is apparently a more modified condition of parasitism than that detailed
on p. 140, vol. ii, where a parasitic Autolytus, found by C. Crossland at Wasin,
East Africa, fixes itself by its permanently extruded proboscis to Nemerteans and Poly-
chaets. Ichthyotomus is attached by two stylets to the median fins of the eel, and
resembles a broad Syllid, measuring from 5 to 7 mm. in length, or in the case of a
ripe female to 10 mm., and having from seventy to eighty segments. The head is round
and truncated, with indications of median and lateral tentacles, a minute pair of eyes
posteriorly, rudimentary palpi, a ciliated (nuchal P) groove on each side in front, and a
pair of tentacular cirri on the first segment. The body ends posteriorly in a tapered
tail with a pygidium, at the tip of which are the anus and two cirri of moderate
length. The remarkable boring apparatus recalls the armature in the mastax of
certain rotifers, and consists of two twisted blades with serrated edges, sharp points,
a "locking" apparatus of the stem, and a posterior bifid process fixed to the complex
muscular proboscis in the centre of a cup-like sucker. The proboscis is globular or
ovoid, and is followed by a second globular region which Eisig considers to be the

1 ' Quart. Journ. Micros. Sci./ N.s. ; vol. liii, p. 600, 1909.

2 f L. M. B. C. Memoirs/ xix, Polycli. Larva?, pp. 4—10, pi. i, figs. 1—5, 1909.

3 ' Fauna u. Fl. Neapel/ Monographie xxviii, 10 pis., 1906.

87

234 GRUBEA.

homologue of the larval pharynx (dipharyngeal stage), then the stomach and posteriorly
the intestine which sends several lobes into the feet. In connection with the pharynx
is a pair of " hamophilinen " glands (dorsal and ventral). The nerve-cords are separate,
the ganglia being united by commissures. The typical foot has a dorsal cirrus
supported by a bristle, and one of the diverticula of the gut enters it, while at the tip
is a stylode. Beneath is the setigerous lobe with a spine and compound bristles of
two kinds, which vary from front to rear as regards the condition of the tip.
The first kind has long, slender, sharp tips which have a homogomph articulation
with the shaft; the second kind appears to have in some cases a homogomph
and in others a heterogomph articulation with the shaft, the terminal piece being
short with a spinous edge and occasionally a joint at the base. A process of the
gut also enters this region of the foot. Inferiorly is the ventral cirrus, into the base
of which a diverticulum of the gut passes. The segmental organ opens ventrally at
its base. Eisig mentions Schmarda's 1 remarkable Gnathosyllis dijplodonta, which has two
bifid stylets in its proboscis, but in many respects it agrees with a typical Syllid
like S. armillaris. It comes from the Atlantic. The intermediate form is rather to
be sought in the parasitic Autolytus found by Mr. Crossland, though this does not
in any way detract from the interesting and important treatise of Dr. Eisig.

Lately Akira Izuka 3 has found Try pa.no syllis misalciensis with similar caudal buds
to those described by Parlin Johonson. The group, indeed, is remarkable for its
budding, phosphorescence, and viviparity, though it is not clearly demonstrated whether
the last named is due to parthenogenetic development, or impregnation from without or
within. Still more recently Michel, who had made remarks on the diverse types of
stolons in Syllids, 3 found a more remarkable condition in Syllis arnica which presented
no less than twelve supplementary heads in successive rings anteriorly, each as a rule
having eyes, articulated tentacles, and palps. 4

Gravier, 5 in his account of the regeneration of the anterior region in the Polychastes,
gives various instances of this in the Syllids ; e. g. JEhlersia rosea, Langerhans, Syllis
gracilis, Grube, Typosyllis prolifera, Krohn, Syllis alternosetosa, De St. Joseph, and others.

Genus XLV A. — Giutbea, De Quatrefages, 1865. Ohar. emend.

Syllids with palpi coalesced almost to the tip, whilst a furrow separates them inferiorly.
Proboscis with a single tooth. Preventriculus with follicular walls; short stomach with
a pair of glandular casca; three smooth tentacles; two tentacular cirri. Dorsal cirri
smooth, fusiform at the base, attenuate at the apex. Ventral cirri short and tongue-
shaped. Bristles compound ; tips often bifid. Carry ova on the dorsum or elsewhere.

i ' Neue Wirb. Tliiere/ ii, p. 69, pi. xxviii, fig. 220.

2 < Annot. Zool. Japon./ vol. v, 1906.

3 ' Compt. Rend. Acad. Sci./ cxlviii, p. 365, 1909.

4 Ibid., p. 438, 1909.

5 f Ann. Sc. Nat./ Zool., 9 e ser., t. ix, p. 131, 1909.

GPUBEA CLAVATA. 235

Grubea clavata, Claparede, 1864. Plate LXXXVII, fig. 10— foot; Plate LXXXVI,

fig. 11 — bristle.

Specific Characters. — Head with three fusiform tentacles, four eyes arranged in a
trapezoid and with lenses, a pair of palpi soldered at the base, and two pairs of tentacular
cirri, similar in shape to, but longer and larger than the tentacles. Body slightly tapered
in front and more so posteriorly, 2—4 mm. in length and of thirty-four bristled segments.
Proboscis with a single tooth shaped like the tip of a lance ; proventriculus with twenty-
two rows of points, and the stomach with two lateral cseca. Tail with two fusiform cirri 1
longer than the ordinary forms. Foot presenting a fusiform dorsal cirrus, a somewhat
conical setigerous process with two short clavate papillae clistally, and a short ventral
cirrus. Bristles translucent, terminal pieces of moderate length with minutely bifid tips.
Ova (eighty to one hundred) borne on the dorsum by a pedicle of the capsule.

Synonyms.

1863. Syllis clavata, Claparede. Beobacht., p. 41, pi. xii, figs. 28 and 29.

1864. Sphserosyllis clavata, idem. Glanures Zoot., p. 90.

1865. Grubea fvsifera, De Quatrefages. Arm el., ii, p. 35, pi. vii, figs. 16 — 21.
1865. ,, clavata, idem. Ibid., ii, p. 40.

1868. „ „ Claparede. Annel. Nap., p. 207.

1874. ,, dolichopoda, Marenzeller. Sitzb. K. Akad. der Wiss. Wien, p. 26, pi. iv, fig. 1.

1879. ,, clavata, Langerlians. Zeitsclir. f. wiss. ZooL, Bd. xxxii, p. 564, Taf. xxxii, fig. 21.

1881. „ „ idem. Nova Acta Leop. -Carol. Deutsch. Akad., Bd. xlii, p. 105.

1886. „ „ De St. Joseph. Ann. Sc. Nat,, 7 e ser., t, i, p. 200, pi. x, figs. 75, 76.

1908. „ „ Elwes. Journ. M. B. A., n.s., vol. viii, p. 197.

Habitat: — A single example from a laminarian root at a low spring-tide at Oddicombe
Beach, Torquay (Major E. V. Elwes).

Mediterranean (Claparede) ; Atlantic, Channel (De Quatrefages, De St. Joseph).

The head is furnished with four eyes, the two anterior being wider apart and with
lenses, a median and two lateral tentacles, fusiform at the base and with a tapered tip,
and a pair of palpi, which are coalescent and ciliated. The buccal segment bears a pair
of tentacular cirri on each side larger and longer than the tentacles, but of similar shape.

The body is almost o — 4 mm. in length, and has thirty-four bristled segments. It is
a little tapered at either end, the greater part being of nearly equal breadth. Posteriorly
it diminishes to the tail, which has two caudal cirri of similar fusiform shape to, but longer
than the dorsal cirri. The proboscis occupies the first five segments, and has a single
tooth shaped like the tip of a lance. The proventriculus has twenty-two rows of " points
gris," followed by the stomach with two small lateral ca?ca. The colourless dorsal vessel
passes to the left of the proboscis and forms a ring as in PasdojjhyJax verruger of Claparede
(De St. Joseph).

The foot (Plate LXXXVII, fig. 10) has dorsally the fusiform cirrus with a rather
long tip, a conical setigerous region with two clavate papillge distally, and bearing a series

1 Langerlians states that there are four short anal cirri.

236 PIONOSYLLIS LAMBLLIGBRA.

of translucent bristles with cylindrical shafts and a distinct backward bend at the tip
(Plate LXXXVI, fig. 11) which is dilated and bevelled for articulation with the terminal
piece, which is of moderate length and has a hook at the tip. A minute secondary
process appears beneath it, and the front edge of the terminal piece is probably minutely
spinous, though this was not actually seen. Besides these both in the anterior and
posterior regions a single simple bristle is present, and the tip of the solitary spine is
slightly bent. The ventral cirrus is small and somewhat conical.

In the examples described by De St. Joseph (1886) the body was 2 mm. long, and
the number of segments ranged from twenty-seven to thirty-four. Both pairs of eyes
had lenses, the anterior being directed to the front, the posterior backward.

Reprorhiction. — De St. Joseph found the ripe males of a pale orange hue from the
sperms, which extended from the tenth to the ante-penultimate segment. The females
bore the eggs on the dorsum ; and in an example with thirty-one segments there were
eighty-six eggs from segments eight to twenty-six, and in another with thirty-four bristled
segments four eggs occurred on each segment from the ninth to the twenty-eighth — in all
eighty. The eggs are attached by the transparent capsule, and he described the appear-
ance of the advanced embryos. Swimming bristles are absent in all the females.

This species was first found by Claparede (1863) on the coast of Normandy, his largest
example having twenty -two segments. He correctly interpreted the structure of the foot,
but he did not allude to that of the bristles. The same form he obtained at Port-Vendres
next year. De Quatrefages (1865) then described it from the coast of Brittany under
the title of Grubea fusigera, having constituted the genus Grubea for its reception, and
he gave a detailed account of its general structure with figures; but he made several
misinterpretations which were corrected by Claparede in his ' Annelides de Napoli.'

Langerhans procured the same species at Madeira, and described it as having four
anal cirri and bifid tips to the compound bristles.

Pionosyllis 1 LAMELLIGER.A, De St. Joseph, 1886. Plate LXXXVI, fig. 12 — bristle; Plate

LXXXVII, fig. 11— foot.

Specific Characters. — Head rounded in front; four eyes with lenses in a trapezoid,
and two smaller specks in front ; a longer median and two shorter lateral tentacles. Palp
large, oblong or ovoid ; two tentacular cirri on each side. These and the tentacles in the
preparations are moniliform distally. Body slender, little tapered anteriorly, but
distinctly so posteriorly, where there are two anal cirri. Length 6 — 7 mm., and segments
forty-five to fifty-five. Translucent, marked by three violet bars on the dorsum of each
segment. The proboscis has ten soft papillae and a large tooth anteriorly. Proventriculus
with twenty-two to twenty- five rows of points. Foot has dorsally a moderately long smooth
cirrus ; setigerous region bluntly conical, with translucent compound bristles, the upper
series of which has long tips, the lower considerably shorter, and all are bifid, and thus
somewhat approaching Syllis cornuta, H. R. Ventral cirrus tongue-shaped. Repro-
duction direct ; male of rose-orange hue when ripe ; female has grey or violet eggs (De
St. Joseph).

1 See p. 161.

PIONOSYLLIS LAMELLIGERA. 237

Synonyms.
1886. Plonosyllis lamelligera, Dj St. Joseph. Ann. Sc. Nat., T ser., t. i, p. 163, pi. viii, figs. 30—38.
1908. „ „ Blwes. Journ. M. B. A., n.s., vol. viii, p. 198.

Habitat. — Common amongst Laninarian roots between tide-marks at Torquay (Blwes).

Between tide-marks at Dinard, France (De St. Joseph).

Head rounded in front with four eyes in a trapezoid, the anterior pair the larger
and wider apart, and all furnished with lenses. Besides, a pair of smaller black specks
occur near the anterior border. Three tentacles, a longer median, and two shorter
lateral, are attached to the head, and whilst the basal region of each is smooth, the distal
is moniliform, a result to some extent of the mode of preparation. In life De St. Joseph
found them hirsute with palpocils. The palpi are large, oblong in life (De St. Joseph),
but ovoid in the preparations, soldered at the base (De St. Joseph), though in the
preparations this could not be made out. The buccal segment is separated from the
head by a brown bar, and on each side are two tentacular cirri, a longer dorsal and
shorter ventral.

The slender body is very little tapered anteriorly, but posteriorly it gradually
diminishes to a slender tail which has two anal cirri. The length is 6 — 7 mm., and the
number of bristled segments varies from forty-five to fifty-five, the anterior being less
distinctly separated than the posterior. It is more or less translucent, marked by three
violet bars on the dorsum of each segment (De St. Joseph).

The proboscis has ten soft papillae anteriorly, and a large tooth. The proventriculus
has from twenty-two to twenty-five rows of points, and the stomach follows with two
lateral pouches. Abutting on the anterior circle of papillae in the proboscis is on each side a
glandular tube, enlarged into a sac on reaching the level of the proventriculus, and filled
with little "pits " like those in Eusyllis (De St. Joseph).

The foot (Plate LXXXVII, fig. 11) presents dorsally a moderately long cirrus,
which, though slightly crenate distally, does not present in the preparations the moniliform
condition of the cephalic and buccal appendages. The setigerous process is bluntly
conical, and bears a series of rather small translucent compound bristles, the tips of
which vary from the inferior series with short bifid tips to those at the dorsal edge
of the fascicle with much longer and apparently minutely bifid tips (Plate LXXXVI,
fig. 12). The condition is thus parallel w r ith that in the Syllis cornuta of H. Ratlike,
though none of the shorter series in the latter corresponds with the structure in
Pionosyllis lamelligera. Moreover the spines are much stouter in 8. cornuta, and have
not the modification of the tip indicated by De St. Joseph, though it cannot be said that
this condition has actually been seen in the mounted examples. The ventral cirrus is
a somewhat lanceolate structure with large hypodermic reticulations.

Reproduction. — De St. Joseph mentions that reproduction is direct, and that when the
sexual elements are ripening the four posterior eyes increase sensibly in size. In the
female grey or violet eggs appear from the fifth or sixth to the ninth segment from the
posterior end. All the segments bear swimming bristles, and the intestine atrophies.
Similar changes occur in the male ; only those segments filled with sperms have a trape-
zoidal form and a rose-orange hue. There are eighteen pairs of brownish testes (De St.
Joseph).

238 PIONOSYLLIS ALTEKNOSETOSA

Pionosyllis alteenosetosa, Be St. Joseph, 1886. Plate LXXXVI, fig. 15 — bristle;

Plate LXXXVII, fig. 12— foot,

Specific Characters. — Head generally as in P. prolifera,, but sometimes with an
additional pair of eye-specks in front. Median tentacle with twenty, the lateral with fifteen,
and the longest tentacular cirrus with twenty articulations. Body somewhat roseate with
madder-brown or roseate transverse bands in front; 15 — 28 mm. in length and with
110 — 144 segments, the latter having a stolon of thirty-two segments (De St. Joseph). Anal
cirri with twenty articulations, and a median pygiclial papilla. Foot with a moniliform dorsal
cirrus of twelve segments. Setigerous process carries compound bristles, the bifid tips of
which are of moderate length anteriorly and posteriorly, but are shorter in the central
region and may have neither spinous edge nor secondary process, which is minute and
slants forward, as if a stouter spike at the end of the spinous row. Ventral cirrus some-
what fusiform in outline, the tip less tapered than in P. prolifera, with which the ten soft
papillae in front of the proboscis and the large single tooth agree.

Synonyms.

1886. Syllis (Typiosyllis, Lang.) ulternosetosa, De St. Joseph. Arm. Sc. Nat., 7 e ser., t. i, p. 150.
1904. Typosyllt's alternosetosa, Allen. Journ. M. B. A., n.s., vol. vii, p. 219.
1908. Syllis alternosetosa, Elwes. Ibid., vol. viii, p. 199.

Habitat. — Eddystone Grounds, Plymouth (Allen) ; Torquay, between tide-marks
(Elwes).

Dinard, coast of France (De St. Joseph).

In 1886 De St. Joseph described, under the name of Syllis (Typosyllis, Lang.)
alternosetosa, a new species which he had found extremely common both in dredgings and
between tide-marks at Dinard. Major Elwes met with the same form in considerable
abundance at Torquay, and to his kind courtesy in forwarding mounted specimens the
present remarks are due.

This Syllid bears in size, external appearance, and coloration a near resemblance to
Pionosyllis prolifera, Krohn ; and as this widely distributed form shows very considerable
variation, both in coloration and the length of the cirri, there is room for re-consideration,
especially as Malaquin considers P. alternosetosa identical with S. hyalina, Grube, whilst
Giard thinks it is a variety of Syllis cornuta, H. Pathke. The head in these examples
closely resembles that of Pionosyllis prolifera, though differing in having an extra pigment-
spot in front and shorter tentacles. The median tentacle has about twenty articulations,
the lateral fifteen, and the longest tentacular cirrus twenty. The dorsal cirri have about
twelve articulations, the anal cirri twenty, and between them is an unpaired median
papilla, as in P. prolifera, the papilla in the latter occasionally being bifid, as an
abnormality. The cirri and tentacles certainly differ considerably in character, those of
P. alternosetosa being shorter and even more distinctly articulated. They are pro-
portionally thicker, and taper more abruptly. In the structure of the foot (Plate
LXXXVII, fig. 12) the ventral cirri of the two forms diverge, for whilst that of
P. prolifera has a tendency to be subulate, that of P. alternosetosa has a tip less tapered,

PIONOSYLLIS ALTBRNOSETOSA. 239

and a tendency to enlargement in the middle (according to the preparations), so that it is
somewhat fusiform, for it is narrowed at the base. It more nearly resembles the ventral
cirrus of Pionosyllis hyalina than that of P. prolifera. The proboscis, again, agrees in
structure with that in the latter form, haying ten soft papillae in front, and a single large
tooth.

De St. Joseph distinguishes the form thus : " Caractere distinctif . — Soies composees
a serpe bidentee dans les segments anterieurs, faisant place a des soies composees a
serpe unidentee clans les segments posterieurs nietees a celles-ci." The examples from
Torquay show a general resemblance externally to Pionosyllis prolifera, whilst a scrutiny
of the bristles from front to rear demonstrates that the tips of the compound bristles
(Plate LXXXVI, fig. 13) become somewhat shorter at the eighteenth or nineteenth, seg-
ment and for a considerable distance thereafter, but no abrupt change is noticeable.
The secondary process, which is minute, has the aspect of a stouter spike terminating
the spinigerous edge of the terminal piece, and therefore is subject to the same influences
which affect the latter. The spines from the edge of the terminal piece are generally
absent in the middle region of the body, but in almost every foot a secondary process or
a trace of it can be detected in one or more bristles from the anterior to the posterior
region of the body, where this process is again evident. Thus in all probability the
normal condition is to have a secondary process or stouter spike terminating the row
along the edge, but with the removal of the latter, by friction or otherwise, the secondary
process likewise often disappears. Moreover, in a male stolon, the secondary process
of the compound bristles could be observed in almost every bristle in each foot from
front to rear, and it has to be borne in mind that the secondary process is minute and by
no means easily seen in any case. It would also be exceptional in the Syllicls for a
compound bristle with a bifid tip to lose, without external influences, the secondary
process after eighteen or twenty segments so as to become simply hooked, and again to
become bifid posteriorly. The view that friction in the intermediate region removes the
spines and the secondary process, therefore requires consideration, especially as the
bifid tips are more numerous in the middle region of the stolon.

In comparing allied forms it is observed that in Pionosyllis hyalina the secondary
process of the tip of the compound bristle more resembles that in P. prolifera than in
P. alternosetosa. The same process is also nearer the tip in Sijllis cornuta, H. Rathke,
in which the tips of the bristles on the dorsal edge of the fascicle are much longer. The
tip in P. alternosetosa has a different character from that of P. prolifera, where the
secondary process is stronger, and the spinous edge seems to be less liable to injury. It
may be a question whether this secondary process represents the last of the spines on
the edge or a special development, but it has not the individuality seen in most forms.
The imperfection in figures renders it difficult to be precise as to the relationship stated
by De St. Joseph to exist between it and Syllis solida, Grrube, 1 but the latter has longer
dorsal cirri.

1 ' Annul. Semper./ p. 120, Taf. vi ; fig. 7.

240 TRYPANOSYLLIS (LELTACA

Teypanosyllis 1 c^liaca, Claparede, 1868. Plate LXXXVI, fig. 14 — bristle; Plate

LXXXVII, fig. 3— foot.

Specific Characters.' — Head somewhat small, transversely elongated in the prepara-
tion, with four distinct eyes arranged in a trapezoid, the anterior pair being wider apart.
The median and two lateral tentacles are short, and in the preserved specimen have only
six articulations (nine to eleven, De St. Joseph). Palpi somewhat ovoid. Tentacular cirri
short ; dorsal with ten and the ventral with six articulations. Body in preservation rather
thick, slightly tapered in front, more distinctly so posteriorly, and with two caudal cirri of
seven articulations, the distal being the largest. Colour a deep yellow. Length 8 — 12 mm.,
and segments seventy to ninety (De St. Joseph), the British forms being smaller. Proboscis
has ten denticulations in front; proventriculus has twenty to twenty-five rows of points.
Stomach with a pair of large yellow caeca. Foot has dorsally a cirrus of seven to eight
segments, the thickest part in the preparations being a little above the base, so that the
organs are fusiform. Setigerous region a blunt cone with a single spine and a fascicle of
compound bristles with rather short and boldly bifid tips. Ventral cirrus comparatively
large and lanceolate. Reproduction by stolons.

Synonyms.

1868. Trypanosyllis cseliaca, Claparede. Annel. Nap., p. 203, pi. xiii, fig. 3.

1875. ,, „ Marion and Bobretzky. Ann. Sc. Nat., 6 e sei\, t. ii, p. 37, pi. xiii, fig. 3.

1879. „ „ Langerhans. Zeitschr. f. wiss. Zool., Bd. xxxii, p. 557.

1886. „ „ De St. Joseph. Ann. Sc, Nat., 7 e sei\, t. i, p. 181, pi. ix, figs. 57—63.

1908. „ „ Elwes. Jonrn. M. B. A., n.s., vol. viii, p. 201.

Habitat. — Oddicombe rocks, Torquay, between tide-marks (Major Elwes).

Port Vend res (Claparede); Marseilles (Marion and Bobretzky); Madeira (Langerhans);
Dinard, France (De St. Joseph).

Head somewhat small, transversely elongated, and with four distinct eyes arranged
in a trapezoid, the anterior pair being wider apart. The median and two lateral tentacles
are short, and in the preserved specimen have only six articulations, the terminal being
the largest, so that each is somewhat clavate. The palpi are ovoid. The tentacular cirri
are likewise short, the dorsal having nine and the ventral six divisions.

The body, when preserved, is rather thick, slightly tapered in front, but more
distinctly so posteriorly, and with two caudal cirri of about seven articulations, the distal,
as in the case of the tentacles, being the largest. The colour is of a deep yellow, and
thus it is in contrast with the French examples of Trypanosyllis Krohnii. De St. Joseph
gives the number of segments as seventy to ninety. The proboscis has ten soft papillae
and ten teeth in front. It occupies segments three to eleven. The proventriculus
has twenty to twenty-five rows of points. The stomach which follows has a pair of
large caeca of a yellow colour. De St. Joseph states that he found air in these caeca as
in other forms. Claparede considered that the intestine was prolonged in front of the
latter in the form of caeca, but De St. Joseph shows that this is only apparent.

1 See p. 169.

EURYSYLLIS PAPADOXA. 241

The typical foot (Plate LXXXVII, fig. 13) Las dorsally a short cirrus of about
seven or eight segments, the thickest part in the preparations being a little above the
base, so that the organ is somewhat fusiform. The setigerous process forms a blunt cone
supported by a single strong spine and a fascicle of bristles having the distal ends of the^
shafts enlarged, curved backward, and bevelled for the terminal piece (Plate LXXXVI,
fig. 14) which is rather short and boldly bifid, the front edge, moreover, being minutely
spinous. The ventral cirrus is comparatively large and lanceolate.

Claparede first procured this species at Port-Vendres, and distinguished it from
8. Krohnii by the difference in coloration, though it resembles it in size, and in the
shortness of its tentacles and cirri. He noted especially the prolongation of the intestine
forward in front of the proventriculus, and its remarkably moniliform character.

De St. Joseph found the species propagating in July by stolons which have the form
of Tetraglene. The nurse-stock is of a deep yellow and the stolon of a fine rose-colour,
the former consisting of thirty-five to forty-four segments, and the latter (female) of
thirty-five to forty-one with natatory bristles. As exceptions he has seen a nurse-stock
of only fifteen setigerous segments without proboscis, proventriculus, or stomach, and a
stolon of thirty-seven segments without swumming bristles. The male stolon with
natatory bristles may have thirty-five to thirty -nine segments with testes from the second
or third to the third or fifth from the posterior end. Its head is less deeply cleft, the
feet less elongated, the body less massive, and the coloration of a deeper rose-colour
than in the male of T. Krohnii.

Genus LIV A. — Eurysyllis, 1 Elders, 1864.

Head large, fused dorsally with the first segment, without palpi, bearing anteriorly
three flattened processes ; four eyes. First segment undifferentiated dorsally, without a
foot, but with two stumpy tentacular cirri, the basal process being cylindrical and the
terminal ovoid or almost globular, and the cirri which follow have a similar shape. The
body is comparatively broad.

Eurysyllis paradoxa, Claparede, 1864. Plate LXXXVI, fig. 15 — bristle; Plate

LXXXVII, fig. 3— head, and fig. 14— foot,

Specific Characters. — Head small, transversely elongated, with two lateral processes
anteriorly. Eyes four, reddish, furnished with lenses, and occasionally an additional pair
of specks in front. The median tentacle is a broad, flattened truncate process, and the
lateral are rounded flattened processes. The palpi are flattened and ovoid. Two short
tentacular cirri, the cirrus being globular and attached to a basal ceratophore. The
ventral is considerably shorter than the dorsal. Body comparatively short and broad,
resembling Sphserosyttis, slightly tapered anteriorly and more so posteriorly, and ending in
two globular anal cirri with an intermediate point. Bristled segments about fifty-seven,
and the length 3 — 4 mm. Proboscis preceded by six large papillse and with ten denticles
on its anterior edge ; proventriculus short and ovoid, narrower in front ; stomach without

1 Evpvg, broad, and Syllis.

88

242 EURYSYLLIS PARADOXA.

lateral cseca. The foot has a more or less globular dorsal cirrus with a basal cirrophore, a
short conical setigerous region with translucent bristles, with an enlarged distal end to the
shaft and a bevelled region with a very acute point. The terminal piece of the bristle
has a simple hook at the tip. Ventral cirrus lanceolate, soldered at the base to the foot.
Reproduction by stolons.

Synonyms.

1864. Polymastia paradoxus, Claparede. Glanures Zoot., p. 109, pi. viii, fig. 3.

1864. Eurysyllis tuberculata, Ehlers. Borstenw., i, p. 264, Taf. xi, figs. 4—7.

1870. „ paradoxa, Langerhans. Zeitschr. f. wiss. Zool., Bd. xxxii, p. 574, pi. xxxii, fig. 27.

1886. „ (Polymastus) paradoxa, De St. Joseph. Ann. Sc. Nat., 7 e ser., t. i, p. 191, pi. ix,

figs. 68—74.
1908. „ „ „ Elwes. Journ. M. B. A., n.s., vol. viii, p. 201.

Habitat.— Between tide-marks at Oddicombe Rocks, near Torquay (Elwes).

Mediterranean (Claparede) ; Atlantic (Langerhans) ; Dinard, France, on dredged
shells at a depth of 10 — 25 metres, and on Rytiphloea pinastroides from a depth of 4 — 7
metres (De St. Joseph).

Head (Plate LXXXVII, fig. 3) comparatively small, transversely elongated, and
having two lateral processes anteriorly, the anterior eyes being at the base of these
or even on them, as Ehlers shows, with four reddish eyes furnished with lenses, and De
St. Joseph describes an additional pair of pigment-specks in front. The median tentacle
is a broad truncate process, and the two rounded lateral flattened processes represent
the lateral tentacles. The palpi form two flattened and somewhat ovoid processes on the
ventral surface, their tips only projecting forward in a dorsal view. The tentacular cirri
are two; the dorsal have a globular tip articulated to a basal ceratophore. A shorter
adnate process of similar shape ventrally represents the ventral cirrus.

The body is comparatively short and broad as in Sphserosyllis, a little tapered
anteriorly before reaching the truncate frontal border, and also slightly tapered pos-
teriorly to the two globular anal cirri with a point attached. The bristled segments are
about fifty-seven, so that the British examples approach those from the French coast,
which have in all sixty-five segments. The colour is reddish or greyish-red. The feet
form a somewhat even and regular series from end to end, the dorsal cirri, however, being
globular in front and somewhat more elongated posteriorly so as to make a greater
proportional difference between their breadth and that of the cirrophore in this region.
Moreover, in a form with a reproduced tail these organs and the anal cirri have a
tendency to greater elongation than in the ordinary form. Each segment has four
tubercles, the buccal alone having two, and these agree in structure with those on
the feet.

The proboscis has ten minute points on the edge anteriorly, besides a large dorsal
tooth, preceded by six large papillge (De St. Joseph). The proventriculus is short
and almost ovoid in outline and slightly narrower in front, and the stomach has no lateral
caeca. The reddish intestine has lateral processes which De St. Joseph thinks contain air
as in Trypanosyllis.

The foot (Plate LXXXVII, fig. 14) has dorsally the cirrus with its basal cirrophore

AUTOLYTUS EHBIEXSIS. 243

and its rounded or ovoid distal region containing, as De St. Joseph points out, yellowish
corpuscles and rod-like bodies or bacilli (" batonnets," De St. Joseph). The setigerous
region forms a short cone with the lanceolate ventral cirrus soldered to it, the tip alone
being free. The bristles are translucent, with shafts slightly curved at the tip, the
bevelled region having an acute tip, and the terminal piece is comparatively short (Plate
LXXXVI, fig. 14) and has a simple hooked tip. The single rather strong spine
supporting the setigerous region has a mucro at the tip. In the posterior segments De
St. Joseph found a simple bristle with a curved point.

Reproduction. — This was unknown to the earlier observers, but De St. Joseph found
that it showed the so-called alternation of generations. Thus in one form of sixty-five
segments, thirty-eight of the anterior are devoid of eggs, but the twenty-four following
have them, whilst the last three are free from them. The violet, green, brown, or dark
red female buds have the same dorsal tubercles as the nurse-stock, four eyes, of which
two are inferior, are devoid of proboscis, proventriculus, and stomach, but have a
rudiment of intestine. The feet possess bristles and a spine'like the adult, but in addition
the annelid has long swimming bristles. The male buds are pale rose- orange, with brown
testes from the third to the fourteenth. De St. Joseph further notes that the two
segments of the nurse-stock in front of the head of the bud are often full of ova or sperms.

Eiirysyllis paradoxa was introduced to science (1864) by the indefatigable Claparede,
who, in spite of delicate health, did so much for the marine Annelids. He found it at
Port-Vendres, and described it under the name of Polymastia paradoxus, giving fairly
good figures of its general structure. He regarded the dorsal processes as homologous
with those of Ephesia, and he figured the compound bristles with simple tips.

Simultaneously (1864) with Claparede, Ehlers gave an account of a new genus and
species — Eiirysyllis tubercidata — which he procured at Quarnero in the Adriatic. He
also alluded to the pinnate intestine, and described the compound bristles as having
simple tips, and the tips of the spines as furnished with a knob. So far as can be
observed this form would appear to be the same as Claparede's.

Langerhans next (1879) met with it between tide-marks at Madeira, and he agrees
with the preceding authors in the structure of the tip of the terminal piece of the
compound bristles. To the spine he gave a clavate tip with a central process.

Autolytus 1 ehbiensis, De St. Joseph, 1886. Plate LXXXVI, fig. 16— bristle; Plate

LXXXVII, fig. 15— foot.

Specific Characters. — Head rounded in front with four eyes having lenses
posteriorly and arranged in a trapezoid, the anterior pair being wider apart. Tentacles
normal, the median being much longer than the lateral, and the dorsal tentacular cirrus
fully as long. The dorsal tentacular cirrus is the longest appendage. Body of the
nurse-stock greyish, slightly flattened clorsally and ventrally, generally has a series (one
to seven) of buds, and much resembles Autolytus prolifer. Proboscis firm, sinuous and
narrow, with thirty minute teeth on its front edge, though a form with twenty is identical

1 See p. 209.

244 AUTOLYTUS EIIBIENSIS.

(De St. Joseph). Proventriculus with thirty-two to forty-two rows of points, barrel-
shaped. Foot has clorsally a longer or shorter cirrus (for they appear to be more or
less alternate). Setigerous lobe is fused with a large globular mass behind it, perhaps
representing a modified ventral cirrus. It bears a group of translucent compound
bristles of the usual shape, the terminal piece being boldly bifid. Reproduction by
stolons, and it is rarely without one or more.

Synonyms.

1886. Autolytus ehbiensis. De St. Joseph. Ann. Sc. Nat., 7 e ser., t. i, p. 228, pi. xi, figs. 106, 107;

pi. xii, figs. 114, 115.
1908. „ „ Elwes. Journ. M. B. A., n.s., vol. viii, p. 201.

Habitat. — On Fucus covered with Sertidaria pumila growing on the breakwater at
Babbacombe (Elwes).

On the shores of Dinard, France, it occurs at some depth off the coast, as well as
between tide-marks, amidst the forests of Sertalaria ojperculata, accompanied by swarms
of Gaprella linearis (De St. Joseph).

The head is rounded in front, with four eyes arranged in a trapezoid posteriorly,
the anterior pair wider apart, and all with lenses. The median is much longer than the
lateral tentacles, and the dorsal of the tentacular cirri is fully as long. The palpi are
not visible, and De St. Joseph states that they are small.

The body of the nurse-stock, which is slightly flattened dorsally and ventrally,
generally has a series of buds attached to it ; indeed De St. Joseph never found it without
a bud, whilst Major Elwes met with only two or three out of fifty or sixty examples
without one. It has the form of Autolytus prolifer, and has quite as remarkable a furor
for budding. According to the last-mentioned author, chains of five or six buds are
common. In the nurse-stock the dorsal cirrus of the first bristled segment is the longest,
the next longer than the succeeding, whilst the rest appear to have an alternate arrange-
ment of long and short, the two caudal cirri being considerably longer.

The proboscis is firm, convoluted and narrow, with about thirty minute sharp points
on its anterior end. As, however, De St. Joseph records an example not distinguishable
from this species, with only twenty minute teeth, there would seem to be a considerable
margin for variation. The proventriculus has from thirty-two to forty-two rows of
points, and is somewhat barrel-shaped, slightly narrowed at each end.

The foot (Plate LXXXVII, fig. 15 in the nurse-stock) has dorsally the longer or the
shorter cirrus; the former, to judge from the preparations, may reach the diameter of the
body. The setigerous region is fused with a large globular mass behind it, and perhaps
represents a modified ventral cirrus, and bears a series of translucent bristles (Plate
LXXXVI, fig. 16) with the ends of the shaft enlarged and bevelled for the terminal
piece, which is very boldly bifid as in the typical Autolytus.

De St. Joseph found Ophryodendron annulatorum parasitic on the tentacular cirri.

Reproduction.— This species reproduces by a male or female stolon, or by a chain of
stolons. According to De St. Joseph if a single stolon is found, the bristle with " article
en alnee " appears on the sixth segment, whereas it occurs on the fifteenth segment
where there is a chain of stolons. Moreover, in the case of a single bud, which is

AUTOLYTUS LONGIFERIENS. 245

produced between the last and the penultimate segment, these segments show no trace of
sexual elements. After it separates the nurse-stock regenerates the posterior end and
the caudal cirri. In the best-developed bud a single simple bristle accompanies the
compound.

The ripe buds agree with those of Aiitolytus prolifer, having a head with a frontal
notch, three tentacles, and four eyes furnished with lenses, the ventral pair being larger than
the dorsal. The first segment of the body has two tentacular cirri, and in a female bud
(De St. Joseph) eighteen segments follow, sixteen of which have large grey eggs. All
the segments except the first two and the last two have swimming bristles. In the first
a simple bristle, "a l'article en alene," is amongst the falcigerous forms. After they are
detached they acquire twenty-five segments, the swimming bristles are shed, and an
ovigerous pouch containing about three hundred embryos of the stage described by Max
Miiller with small eyes and cilia is found.

The description of the male bud as given by De St. Joseph does not seem to differ
from that of A. prolifer. The same author gives an account of nurse-stocks of thirty-
eight and twenty-seven segments, with buds varying in degree of development, in the
latter case a region without bristles following the nurse-stock ; then a bud of twelve
segments without a head ; a second of seventeen segments with the head indicated ; a third
of twenty-one segments with four eyes, three tentacles, and traces of eggs; a fourth of
twenty-five segments with the head well developed and advanced eggs; a fifth bud of
nineteen segments with large head and eggs ; and a sixth and last bud of twenty
segments with large eggs and natatory bristles, twisting its body with energy in order to
detach itself. Bach bud had a rudimentary intestine and compound bristles, but the last
only had the natatory bristles. Only a single fully formed male bud (Pohjbostriclms) of
twenty-one segments was seen attached to a nurse-stock of thirty-six segments. He
found a nurse-stock with seven male buds without swimming bristles, and only the last
showed sexual elements.

One example forwarded by Major Elwes had seven buds, the first having only a
pair of lateral processes posteriorly. The second had no bristles, and the third a few
in front.

Autolytus longifeluens, De St. Joseph, 1886. Plate LXXXVI, fig. 17 — bristle; Plate

LXXXVII, fig. 16— foot.

Specific Characters. — Head fairly large ; four considerable eyes with lenses. Central
much longer than the lateral tentacles. A median line indicates the palpi. Tentacular
and first dorsal cirri as in A. ehbiensis. Body rather massive, from 10 — 20 mm. in
length, and having from fifty-one to eighty-eight segments. Dorsal cirri of moderate
length and alternately long and short. Posteriorly are two long and large caudal cirri.
General colour orange, with three longitudinal reddish bands, the two lateral forming
" epaulettes " in the first two segments. Proboscis remarkable for its great length, coiled
condition, and slenderness, as well as for its crown of ten large, obtuse teeth, and
its intermediate series of two or three smaller pointed teeth, which have two spines

246 AUTOLYTUS LONG1FEKIENS.

posteriorly (De St. Joseph). Proven tri cuius reddish-orange, rather long, barrel-shaped,
and narrower anteriorly. It has forty to fifty-four rows of points (De St. Joseph). Foot
with a longer or shorter, slightly tapered, dorsal cirrus, a rather massive setigerous
process soldered to the rounded posterior mass, and bearing a series of somewhat short
bristles, which have short, deeply bifid terminal pieces, the base in front being so narrow
as to resemble a third tooth. Spine with a small mucro, and a single bristle with a short
delicately tapered tip is present.

Reproduction by stolons (De St. Joseph).

Synonyms.

1886. Autolytus longiferiens, De St. Joseph. Ann. Sc. Nat., 7 e ser., t. i, p. 217, pi. x, figs. 95—97.
1908. „ ,, Elwes. Journ. M. B. A., n.s., vol. viii, p. 202.

Habitat. — Between tide-marks, Oddicombe, Torquay (Major Elwes).

Common in dredged material at Dinarcl, coast of France (De St. Joseph).

Head fairly large ; four distinct eyes with lenses. The median and lateral
tentacles have the usual proportions. A central line indicates the palpi anteriorly. The
tentacular cirri are also normal, whilst the dorsal cirrus of the first bristled segment
agrees with that of A. ehbiensis in being the longest appendage of the kind.

Body rather massive, from 10 — 20 mm. in length, and having from fifty-one to
eighty-eight segments (De St. Joseph), the British examples so far as observed being inter-
mediate in size. The dorsal cirri are of moderate length, and long and short alternately,
a condition met with in other forms; but, as De St. Joseph points out, the relatively
greater length of the anterior cirri is noteworthy, and the dorsal cirrus of the fifth
segment is longer than the third, whereas in Proceroea it is small (De St. Joseph).
Unfortunately the examples clo not afford information on these points. Posteriorly the
body terminates in two long and large caudal cirri. The general colour is orange. In
front of the proventriculus are three longitudinal reddish bands, the two lateral forming
" epaulettes " in the first two segments.

The proboscis is remarkable for its great length, coiled condition, and slenderness, as
well as for the fact that its anterior edge has a crown of ten large obtuse teeth separated
by three smaller pointed teeth which terminate posteriorly in two spines (De St. Joseph).
Major Elwes, however, notes that the British examples have only two of the intermediate
smaller teeth instead of three, and this appears to be right ; but the French examples are
much larger, for the proboscis, instead of being only 2 mm. (Elwes), is from 18 — 20 mm.
(De St. Joseph) in length. The latter author agrees with Claparede 1 in defining three
regions of the organ, the anterior the most muscular. The reddish-orange proventriculus
is rather long and barrel-shaped, narrower anteriorly than posteriorly, and has from
forty to fifty-four rows of points (De St. Joseph).

The typical foot (Plate LXXXVII, fig. 16) has dorsally the long and slightly tapered
smooth cirrus (long or short), a rather massive setigerous region soldered to the
rounded posterior mass, and bearing a group of somewhat short compound bristles which
have very short bifid terminal pieces (Plate LXXXVI, fig. 17), the curve from the second

1 < Anuel. Nap./ p. 218.

AUTOLYTUS INERMIS. 247

process running to the base, which is thus pointed, giving a pseudo-trifid aspect to the
terminal piece. The distal end of the shaft of the compound bristles is minutely spinous
along both edges. The spine has a terminal mucro, and a single bristle with a pointed
terminal piece is present.

Reproduction. — De St. Joseph observed specimens without swimming bristles
carrying greyish ova behind the proventriculus, but he has rarely seen buds, though two
short examples with reproduced posterior ends may indicate this condition, and indeed
in one specimen of fifty-one segments a bud began at the forty-first.

Autolytus inermis, De St. Joseph, 1886. Plate LXXXVI, fig. 18 — bristle; Plate

LXXXVII, fig. 17— foot.

Specific Characters. — Head with large lateral connate eyes, and traces of a dimple
in the centre anteriorly. Colour reddish. Proboscis devoid of teeth and with a
distinct rim. Bristles rather long, with somewhat long terminal pieces minutely bifid
at the tip.

Synonyms.

1886. Autolytus iiiermis, De St. Joseph. Ann. Sc. Nat., 7 e ser., t. i, p. 237, pi. xii ; fig. 117.
1908. „ „ Elwes. Journ. M. B. A., n.s., vol. viii, p. 202,

Habitat. — Between tide-marks, Liver mead, Torquay (Elwes).

Shores of Dinard, France, rarely dredged on Bytlphlea pinastroides (De St. Joseph).

Head of the typical form with two very large anterior eyes laterally situated,
furnished with lenses and confluent with the posterior eyes, also provided with lenses.
De St. Joseph states that the lateral tentacles are a little longer than the median, but
this is not the case in the example procured by Major Elwes. A slight notch occurs in
the median border anteriorly with traces of a fissure behind.

Body of a reddish colour, from 3 — 5 mm. in length, slightly tapered in front and more
distinctly so posteriorly, and composed of forty-nine bristled segments, the pygiclium
having two cirri of moderate length. The characteristic feature, according to De St.
Joseph, is the absence of teeth in the proboscis, which is remarkably long and coiled like
that of Autolytus longiferiens. It terminates anteriorly in a smooth rim, The pro-
ventriculus is ovoid with twenty- five to thirty rows of points, and occupies segments nine
to eleven. The hypoderm throughout is remarkably granular and areolar, and the tissues
of the annelid are comparatively soft.

The foot (Plate LXXXVII, fig. 17) is typical, with a group of rather long bristles,
having a proportionally long terminal piece and a minutely bifid tip (Plate LXXXVI,
fig. 18). The first dorsal cirri are the longest, and the rest are comparatively short.

De St. Joseph found two examples with male stolons of two regions. In the first
nurse-stock of twenty-seven segments the stolon had thirty-one segments ; in the second
the nurse-stock had thirty-one segments and the stolon thirty segments, the first three in
the stolon devoid of natatory bristles. In the example from Torquay sperm-cells appear
to be present posteriorly.

248 AUTOLYTUS INERMIS.

This species may be accepted at present, but it is probable that an exhaustive study
of the British Autolytidse may reveal relationships at present unknown.

Another Autolytus, viz. A. rnacrophthalma, Marenzeller, was procured by Major
Elwes at Babbacombe, 1 but no example was preserved, and hence only a brief notice can
at present be given.

The bead has four large eyes and often a few additional specks. Palpi fused
dorsally. The median tentacle longer than the lateral. Tentacular cirri normal. The
proboscis has a rim of twenty teeth anteriorly. Second dorsal cirrus the longest, the third
considerably shorter (about one quarter the length), but longer than those which follow.
Two rather thick anal cirri. The general aspect is like that of Autolytus aurantiaca, with
forty to sixty segments. The colour is rose-orange, the tips of the tentacles and cirri
being deepest. Various examples had stolons.

Marenzeller 2 found it in the Adriatic, and both Langerhans and De St. Joseph think
that it closely resembles that author's A. luxurians s from the same region. It is common
between tide-marks at Madeira (Langerhans), as well as in dredged material at Dinard,
on the shores of France (De St. Joseph). 4

1 < Journ. M. B. A./ n.s., vol. viii, p. 201, 1908.
1 c Wiener Akad. Ber./ 1875, p. 37.

2 ( Wiener Akad. Ber./ 1875, p. 37.

3 Ibid., 1874 ; p. 50, Taf. vi, fig. 1, and Taf. vii, fig. 1,

4 ' Ann. Sc. Nat./ 7 e ser., t. i, p. 226.

NEEEIDJE

249

Family X. — Nebeihe:.

This family is characterized by the activity and muscularity of its members. The
head is well formed and bears eyes and prostomial tentacles. The cephalic ganglia and
nervous system generally are highly developed. The palps are massive and two-jointed.
The chitinous buccal membrane is continuous with that covering the proboscis, which in
extrusion is readily divided into two regions, and armed with a pair of horny jaws and a
series of horny teeth (paragnathi), which after Kinberg may be arranged in eight groups,
as in the accompanying outlines from Nereis pelagica (Figs. 58 and 58').

Figs. 58 and 58'.— Outlines of the extruded proboscis of Nereis pelagica, the dorsal surface being on the
left, the ventral on the right, a. Maxillary region, b. Basal region. The areas of the parognathi are
in Roman numerals, viz. — I, unpaired median dorsal maxillary ; II, lateral dorsal maxillary ; III, unpaired
median ventral maxillary; IV, lateral ventral maxillary; V, dorsal unpaired median basal ; VI, dorso-
lateral basal; VII, ventral unpaired basal; VIII, ventro-lateral basal.

The alimentary canal behind the proboscis is cylindrical and is furnished with a pair
of glands. This is followed by a more or less simple straight intestine.

The body is elongated, rounded or somewhat flattened, segments similar, the first
devoid of feet, but with two tentacular cirri on each side; the feet are usually biramous
and nearly alike throughout, though the first two are simple and the caudal somewhat
modified. It terminates in two cirri beneath the vent.

The body-wall (Fig. 59) in this family is surrounded by a well-developed cuticle with
a considerable thickness of hypoderm beneath. The latter coat is bounded internally
by the basement-membrane, within which is the circular layer of muscular fibres. The
dorsal longitudinal muscles in transverse section are lappet-shaped, that is, massive
externally and inferiorly, and thinning off towards the raphe in the mid-dorsal line. The
ventral longitudinal muscles, on the other hand, have a characteristic dorsal and external
inflection with a somewhat pennate arrangement of the fibres, the ventral portion of the
muscles being spindle-shaped. The inner ventral band of muscular fibres from the bases
of the spines passes to the exterior of the curvature of the ventral longitudinal muscles.

1 Lycoridea, Savigny, Griibe, Elilers, etc.

89

250

NEREID M.

At the outer border of the dorsal muscles a strong series of fibres stretches obliquely
outward to interdigitate with the vertical. The oblique muscles slope downwards and
almost meet in the middle line, only a central pedicle connected with the nerve-area
above separating them. On each side of the median raphe dorsally is a strong fascicle

vm

Fig. 59. — Transverse section (diagrammatic) of the body-wall of Nereis diversicolor. bd., bv. Eespectively
dorsal and ventral divisions of the foot. d. Alimentary canal, d. m. Dorsal longitudinal muscles.
d. v. Dorsal blood-vessel. n. c. Nerve-cord. o. m. Oblique muscles. s. o. Segmental organ, v. m.
Ventral longitudinal muscles, v. v. Ventral blood-vessel; the main branches of this and of the dorsal
vessel are indicated. (After Goodrich and nature.)

of fibres ensheathing the blood-vessel and decussating on the wall of the alimentary
canal beneath. The nerve-cords lie rather above the attachment of the oblique muscles
to the basement-tissue — a central pedicle occurring beneath. As a rule two large infero-
lateral neural canals (giant fibres) occur, with a median superiorly and occasionally two
smaller below the latter. The circulatory fluid in some contains hemoglobin. 1 Feet

Pig. 60.— Diagrammatic sketch of the nephridium of Nereis diversicolor, O. F. M., with its canal, which is
divided into four regions, viz. 1—4. n. f. Nephridial funnel, n. p. Nephridiopore, (Goodrich.)

somewhat complex with bifid dorsal lobe. Rarely is the dorsal lobe represented only bv
bristles. Bristles homo- and heterogomph.

The nephridium of N. diversicolor (Fig. 60) consists of a compact mass, perforated by
a convoluted canal, which can be divided into several regions, and communicates with the

1 Ray Lankester, ' Proc. Roy. Soc./ No. 140, p. 2, 1873.

NEKEHLE,

251

exterior by a short duct leading to a sac on the ventral surface. A long canal springs
from it, and ends in front by a nephrostome provided with long ciliated processes, which
opens into the next segment.

Goodrich describes the segmental organ as forming a gland-like mass of coiled and
ciliated tube opening externally on the ventral surface at the base of the foot by a non-
ciliated portion, and internally by a funnel — studded with cells having long cilia (Fig. 61) —
into the preceding segment. He considers that the dorsal ciliated organ is a genital
duct, not fully developed. The recent description by Fage 1 does not materially add to
this information, but he advances the view that the separate dorsal ciliated organ of
Goodrich is to be included in the segmental series, viz., as the rudimentary " pavilion

Fig. 61. — Optical section through the nephridial funnel of Nereilepas fucata, Sav. The long' processes on
the summit are external cilia, whilst the processes on the left are septa, not cilia. (After Goodrich.)

genital" He is not inclined to think it a phagocyte organ. It is noteworthy in this
respect that long ago Dr. Thos. Williams credited the segmental organ in Nereis with
both a dorsal and a ventral opening. It may be remarked that Fage insists throughout
his memoir on the essential difference between the " pavilion " (which may be freely
termed genital duct) and the nephridium, the former being connected with reproduction,
the latter with excretion.

The sexes are separate; reproductive elements shed through the body-wall. Some
undergo no change, whilst many species towards the period of sexual maturity pass through
a metamorphosis into the Heteronereis condition. In this the eyes increase in size, the
palps are reduced (Gravier), the prostomium is altered, and the body forms two regions,
viz. , an anterior in which the feet are unchanged, and a posterior bearing the genital

1 ' Ann. Sc. Nat./ 9 e ser. ; pp. 331—347, figs. 29- 35 ; and pi. vi ; figs. 19—23.

252 NEREIDS.

products, and in which the feet acquire foliaceous lobes and swimming bristles. Again
in some the males at maturity present a varicose or serrated condition of the dorsal
cirri, whilst the females show this in a slighter degree or not at all. Dignet mentions that
in Mav, 1901, in the Gulf of California he found male Nereids in which the body was
composed of three regions, the anterior characterized by the enormous development of
the dorsal cirri, which formed powerful natatory organs; a middle region with ordinary feet
and cirri, and a posterior division in which the feet were metamorphosed for the epitokous
condition. The eyes were very large and encroached on the ventral border, and the
palps much reduced 1 (Gravier).

The comparatively great size of the ova of Lycastis quadraticeps, Gay, from Chili
and Punta Arenas, in the Strait of Magellan, as described by H. P. Johnson, 2 is a feature
of interest in this small species, for they are about half the diameter of the body, and,
moreover, the species is hermaphrodite. It frequents not only the sea, but brackish and
even fresh water.

In Nereis Dumerilli young forms may become ripe whilst still Nereids, or they may
assume the Heteronereid condition during the ripening of the sexual elements; and as
the larger Heteronereids frequent the bottom whilst the smaller are pelagic, there are thus
three different kinds of males and females in this species (Benham).

Mr. A. Treadwell 3 is of opinion that the muscles of the anterior (unmodified) region
of such as Nereis hobiensis, Mcintosh, are greatly reduced in the Heteronereis condition,
whilst those of the posterior region are not. He also thought that the septa had
disappeared. His preparations, however, were imperfectly preserved, so that he speaks
with reserve.

The family of the Nereids is, as Claparede observes, a homogeneous one, and there is
little difficulty in assigning the representatives to it. Yet the foreign species are still in
want of elucidation, and those from widely divergent regions may yet be brought into
closer relationship than is at present possible.

It is interesting that, as a general rule, the two heterogomph bristles of the upper
group in the inferior setigerous process are stouter and often more deeply tinted yellow,
just as in certain Syllids.

The older authors, such as Linnasus, 0. Fabricius, 0. F. Midler, and others, included
the Nereids under the Vermes along with parasitic worms, Gordii, Cestodes, Planarians,
leeches, Ascidians, parasitic Crustaceans, Echinoderms, and Mollusca. Savigny placed
them under his second family of Annelids along with Nephthys, Artcia, Glycera, Oplielia,
Hesione, Myriana, Phyllodoce, and Syllis, under the Nereides, whilst the genus Lycoris
comprehended the Nereids proper. He was followed by Lamarck, who made the genus
Spio an appendix to the former groups. Some of the early authors again confounded
the young of the Nereids with Syllids.

Audouin and Milne Edwards (1834) made them the first genus of their Nereidiens,
which included all those treated in this Part. The Nereids were characterized by having
the tentacles (the author's antennas) well developed; tentacular cirri on the first segment ;

1 l Bullet. duMus. d'hisfc. nat.' (C. Gravier), No. 4, 1901.

3 ' Biological Bulletin/ vol. xix, No. 6, May, 1908, pp. 371—385, with text-figs.

3 Ibid., vol. ix, No. 4, Sept., 1905, p. 226.

NEREIDS. 253

filiform dorsal cirri ; feet of. two divisions, branchial lobes at the extremity of the feet;
tentacles dissimilar; two jaws. In this description the lobes of the feet were termed the
branchiae, and the great vascularity of these in certain forms renders the interpretation
physiologically correct, though it is not necessary to place them on the same footing as
those with special branchial organs attached to the dorsal cirri.

Grube 1 (1850) grouped the Nereids under his fourth family, Lycoridea, Savigny
s. str., two genera only being given, viz., Nereis and Lycastis, but he subdivided the first-
mentioned into three divisions according to the structure of the feet. The family was
placed between the Eunicea and the Nephthydea, and next year 2 he gave a synopsis of
the genera and species, basing his distinctions on the head and tentacles, the structure of
the feet, and the proboscis.

Dr. Johnston did much in his earlier papers to elucidate the British forms, and in
the Catalogue of Worms in the British Museum (1865) he gave an excellent summary of
the history and general characters of the group ; but at that elate the relationships of the
heteronereid phase were obscure, and they were regarded as forming a distinct genus. No
less than fourteen species are entered in the Catalogue, which does not include either
Mieronereis or Leptonereis ; but of the list five are Heteronereids, though one of these refers
to a species not entered elsewhere, viz. Nereis (Eunereis) longissima, giving in all six Nereids
which stand. An examination of his collection in the British Museum 3 showed that JV".
fimbviata included various forms such as N. d< 'versicolor, N. pelagica, and N. Marionii, but
in all probability they had been mixed after his death. Nereis imbecillis was found in
1872 to be a Nereilepas with the paragnathi for the most part absent. It is uncertain
what the Nereis pidsatoria of Montagu and mentioned by Dr. Johnston is. The foot
as figured by Audouin and M. Edwards somewhat resembles that of Nereilepas, but the
arrangement of the paragnathi diverges.

Dr. T. Williams i considered that the Nereids had a large amount of " milky
perivisceral fluid," which fluctuates from one end of the body to the other. The
corpuscles, he says, are superseded by true ova in autumn. The same author 5
describes the segmental organs in Nereis margaritacea as consisting of a tube, richly
ciliated, both ends of which open externally, the ingoing dorsally and the longer outgoing
ventrally at each foot, and there is no communication with the perigastric chamber.
The longer limb conveys the generative elements externally. The areolated tissue of
the foot receives the ova or sperms, and it is a development of the segmental organs.
The female in this group is much larger, he says, than the male.

Kinberg (1865) made a step in advance in the classification of the Nereids by
placing the structure of the proboscis (his pharynx) on a proper footing. He termed the
distal region in extrusion the maxillary ring, and the proximal the basal ring. Moreover,

1 'Arch. f. Naturges./ Bd. xvi, p. 294, 1850.
3 'Familien der Anneliden/ p. 125, 1851.

3 And in this connection I have to remember the kindness and courtesy of Dr. Gray, Dr. Baird,
Dr. Grunther, Prof. Jeffrey Bell, and others.

4 ' Philos. Tran?./ 1852, pp. 627—628.

5 Ibid., 1858, p. 124.

6 < Of vers. k. Yet.-Akad. Fork./ 1865, No. 2, p. 167.

254 NEREIDS.

be numbered the eight groups of paragnathi and utilized their presence or absence as
an aid in classification. Thus, to arrange them according as they occur on the
maxillary or the basilar ring, as Gravier 1 has done, they are as follows :

Maxillary ring. Basal ring.

Unpaired median dorsal group I. V.

Lateral dorsal pair . . . . II. . VI.

Unpaired median ventral group . . . III. . VII.

Lateral ventral pair .... IV. . VIII.

He made five families, or rather, as they might be termed, sub-families of the Nereids,

according to the presence of these papillse and their nature, or their absence, using the

structure of the feet, the bristles, and other features in separating the subordinate groups.

De Quatrefages (1865) classified the Nereides according to the condition of the

body, viz., those having one region or two, the uniramous or biramous nature of the feet,

and the foliaceous condition of the latter. Since he included as separate genera the

Heteronereids, the classification is now obsolete. In his characters of the genus

Nereis he terms the palpi the external lateral antennas, and the tentacular cirri

tentacles. The lobes of the feet are considered as branchiae.

Malmgren (1867) arranged the genera according to the uniform or dissimilar feet
along the sides of the body, the characters of the bristles, and the structure of the
paragnathi. He made two great groups, according to the foliaceous or non-foliaceous
condition of the superior lamella of the dorsal lobe of the foot. Then again he subdivided
them into those having the feet uniformly constructed throughout, and those in which
the feet in the posterior region were transformed. His other characters rest on the
form of the dorsal lobe and the condition of the paragnathi. As the epitokous forms
are included in this arrangement, considerable changes are necessary.

This author 2 has the credit of having subsequently discovered the relationships
between the Heteronereis-condition and the ordinary one in Nereis pelagica, in which
the male is Heteronereis grandifolia, and between Heteronereis fvcicola and Nereilepas
variabilis, (Erst., which he deemed equivalent to Nereis Dumerilii. He also showed that
examples of Nereis Dumerilii were either male or female, and that others with ova
assumed the Ijphinereis form. He was followed at no long interval by Ehlers,
Claparede, and others on this important question.

Cosmorici (1880) described the segmental organs in N. bilineata as in pairs on
the posterior faces of the dissepiments in the middle region of the body. 3

Following Malmgren, Ehlers 4 pointed out the relationship of Heteronereis to Nereis
and Nereilepas, taking especially Nereis Dumerilii with its conspicuous characters as an
example, and alluding to the analogous condition in the Syltids with their swimming
bristles.

The same author (1868) characterizes the family Lycoridea, Savigny, Grube, as
having an elongated and distinctly segmented body. Head clearly differentiated with

1 ( Nouv. Arch. Mus. Paris/ 4 e ser., t. iii, p. 151.

3 'Zeitschr. f. wiss. Zool./ Bd. xix, p. 466; and in ' Arch. f. Naturges./ Bd. xxxv, p. 58, 1869.

3 ' Thesis/ p. 113, pi. xxvi, fig. 14.

4 ' Nachricht. d. k. Gesellsch. zu Gottingen/ pp. 209—217, 1867.

NEREIDS. 255

two tentacles, two palpi, and four eyes. First segment devoid of a foot, with two
pairs of tentacular cirri on each side, and two cirri below the anus. Proboscis with
two regions in extrusion, furnished with paragnathi, rarely naked, and with two jaws,
which in retraction fall behind the two regions.
He divides the family Lycoridea as follows :

1. Feet single — Lycastis, 1 Aud. & Edw.

2. Feet bifid :

(a) With simple dorsal cirri.

1. With dorsal and ventral lamellae — Nereis, Cuvier.

2. Devoid of dorsal lamellae — Cevatocephale, Malmgren.

3. Devoid of central lamellae — Tylorhynchus, Grrube.

(b) With plumose dorsal cirri — TJendronereis, Peters.

Ehlers did good service in showing that many of the generic terms used by
Kinberg, Malmgren, and others, were due either to sexual variations or were other-
wise unnecessary.

He grouped the species as follows :

A. The dorsal lobe of all the feet simple, that is, without an enlarged lamella.
I. The feet on all the segments uniform or only little altered.

1. Paragnathi complete, including the British Nereis cultrLfera.

2. Paragnathi incomplete, comprising Nereis jpelagica, N. longissima.
II. The feet unequal.

1. Four or five of the anterior segments have the lobes of the feet little

developed, the following feet with prominent lobes.
(a) Paragnathi conical.

Nereis irrorata, N. Dumerilii.

2. The posterior feet elongated, with long lobes and processes.

3. The posterior feet elongated, the dorsal border, from which springs the

cirri, very convex.
Nereis fucata.

4. The dorsal lobe of the posterior feet broadly triangular.

Nereis diversicolor.

B. The upper lobe with the cirrus bearing a large lamella.
I. The upper lobe in the first feet not leaf-like.

Nereis virens, Sars.
II. The upper lobe only in the middle, and posterior feet foliaceous.

1. The dorsal cirrus terminal.

(a) Paragnathi complete, conical.

Nereis Marionii, Aud. & Edw.

2. The dorsal cirrus prominent in the posterior feet (no British form).
Claparede, in 1868, placed the Nereidae under the sub-genera Leontis, Malmgren,

Lipephile, Malmgren, Oeratonereis, Kinberg, Nereilepas, Blainville, Johnston, Malmgren,
making a special genus of Beteronereis. The same author, in an elaborate investiga-

1 Vide G-rube's view, p. 256.

256

NEREIDES.

tion into the Heteronereicl condition of the Neapolitan Nereids, in 1870, groups the
bristles of the Nereids into two divisions :

(1) Heterogomph bristles in which the distal end of the shaft is prolonged in a point,
so that an oblique articulation takes place.

Fig. 62 a and b. — Homo- and heterogomph bristles of Nereis pelagica.

(2) Homogomph, in which the distal end of the shaft forms a kind of sheath
for the terminal piece ; but the fact that in the ventral division of such forms as
Nereis pelagica, L., and Nereis cultrifera, Grube, a gradual transition from the former to
the latter occurs, shows that the distinction is artificial. He places some weight on the
distribution of the various bristles (four kinds) in the foot. He further describes two
canals in the maxillae of the Nereids in communication with the base, and opening
between the first two (distal) teeth, that is, the tip and the first tooth. In the heteronereid
of Nereis Dumerilii he found the cavity of each maxilla occupied by a peculiar proto-
plasmic tissue with nuclei, but whether the reticulated tissue secreted poison he
could not say. On the whole the beautiful drawings and the numerous observations by
Claparede on the Nereids in the two works just noticed make an epoch in their history.

Grube, 1 in 1873, entered into an elaborate review of the characters upon which he
based the classification of the Nereidae, these being chiefly the armature of the proboscis
and the structure of the feet. His main divisions are as follows :

A. Foot with two bristle-bundles, but not fissured accordingly.

Lycastis, Sav. The foot is not split into tongue-like processes. It has small

lobes with the bristle-bundles. Proboscis without paragnathi.
Foot divided into a dorsal and a ventral division.
The dorsal cirrus simple.

Nereis, L. s. str., Aud. & Edw. The border of the foot split into three tongues,
the upper and middle pertaining to the dorsal, the ventral to the ventra
division.
Proboscis with paragnathi, seldom with soft papillae.

1 ' Scliles. Gesell. f. v. Cultur/ Dec. 19th, 1873, p. 13

B.
B 1 .

NEREIDS. 257

Geratocephale, Malmgren. Without the upper tongue or lobe, but with lip-lobes

and middle tongue, the inferior division as in Nereis.
Tylorlnjnchus, Gr. Only the upper and middle tongues of the feet present.
The proboscis beset with hard papilla.

B 2 . The dorsal cirrus in certain segments furnished with branches (an arbuscle).
Ex. Dendronereis .

In his 'Philippine' Annelids (1878) he groups the Nereids under the family Lycoridea
— with Nereis and Dendronereis as sub-families. He relies on the condition of the proboscis
as the basis of his subdivisions, e.g., Leptonereis, Geratonereis, Platynereis, Lycoris, and
Perinereis. He had previously reviewed the group on similar lines, his main divisions
agreeing with those of Ehlers ; but he clearly indicates that Lycastis has two bristle-
bundles in the foot, though it appears to be simple.

Levinsen (1883) made the Nereids (Lycoridaa) the first family of his Syllidiformia
vera, whereas Benham places them as the sixth family of his Nereidiformia. There is less
objection to the associations of the former than to those of the latter. He follows the
previous authors, Kinberg, Grube, Malmgren, and Ehlers, in his generic and specific
characters.

R. Horst 1 (1889), in dealing with the Nereids belonging to the sub-genus Perinereis,
follows Grube in showing that the lateral dorsal paragnathi of the basal ring of the
proboscis (Group VI) are all, or some of them, transversely ridge-shaped, the remaining
paragnathi being conical or pin-shaped (pectiniformes, Kinberg) or compound. The feet
are all of the same structure, or those of the posterior region enlarged. The group comes
under Kinberg's Aretidea, but not his genus Perinereis. Horst includes Nereis cultrifera,
Grube, under this series.

De St. Joseph, 2 in 1898, adopted the classification of Kinberg, founded mainly on the
condition of the paragnathi, though the primary divisions are based on the presence or
absence of dendritic branchiae. In addition he uses the condition of the buccal segment
and the uni- or biramous nature of the feet. It is a question how much importance can
be attached to this method of classifying, though it may be useful. The British species
would chiefly fall under the fifth group of the second great division, viz., Nereis,
characterized by separate, horny, conical paragnathi, Alitta and Hediste, p.p., having all
the groups complete, whilst Group V or Groups V and VI are wanting in Nereilepas,
Hediste, p.p., and Praxithea. In Eunereis, again, all the groups except VI are absent.
In the sixth division are those in which there are separate, horny, conical, and transverse
paragnathi, all the groups being complete in Lipephile, Mgrn., Hedyle, Mgrn., Hediste,
Mgrn. p.p., Stratonice, Mgrn., and Nereilepas, Blv. sensu Johnst. In the eighth section
the horny paragnathi are very small, numerous, and ranged in rows, e.g. the genus
Platynereis including Leontis of Malmgren. As genera occur under more than one head,
confusion is apt to arise by too strict adherence to such a classification.

Gravier (1901), in his < Annelids of the Red Sea,' followed in the lines of Grube and
Kinberg, that is, made the arrangement of the paragnathi when present, and, in their
absence, the structure of the feet, the foundations of his genera, except in the case of

1 ' Notes from the Leyden Museum/ vol. xi, pis. vii and viii, 1889.

2 ( Ann. Sc. Nat./ 8 e ser., v, p. 283 et seq.

90

258

NEREKLE.

Dendronereis from Mozambique and the Philippines, in which the ramifications of the
dorsal cirrus are alone diagnostic. In his specific characters he makes use of the number
of the different bristles in each division of the foot, a method that has not been followed
here. Of the genus Nereis he makes four sub-genera, viz., (1) those in which all the eight
groups of paragnathi are present (Neanihes, Kinberg) ; (2) those in which one or more
groups are absent (Nereis s. st. Kinberg), whilst the third and fourth sub-genera have the
paragnathi only on one ring, — Geratonereis, Kinberg, having them on the maxillary, and
Eunereis, Malmgren, having them only on the basal ring.

For the British species of Nereis it is unnecessary to make a special classification.
The main lines of previous efforts have been indicated in the foregoing summary, and
that of Ehlers, for instance (p. 255), will suffice for reference. The structure of the
body-wall throughout the series presents no basis for differentiation, so that characters
derived from the armature of the proboscis, the structure of the feet, and other points, are
alone available. Moreover it is doubtful how far the presence of a continuous bar of
chitin in Group VI should be relied on as a satisfactory distinctive character, the two
British forms which fall under this head being Nereis (Perinereis) Marionii and Nereis
cuUrifera. Little is gained by its adoption, since otherwise there are no bonds of note.
Then, again, the British forms with a complete series of paragnathi from I to VIII are
N. cuUrifera and A T . virens, yet in other respects there is no special connection. Neither
is the absence or scanty presence of paragnathi, as in Nereis longissima, followed by other
special characters, though the condition is interesting (e.g. in the foreign Lycastis). All
the British forms have distinctly biramous feet, that is, a dorsal and a ventral division,
no example of Lycastis having yet been procured on our shores, though a persevering
search on the southern coasts might be successful.

In his researches on the eyes of Annelids, Andrews 1 (1892) regards the whole eye as
epidermal, and he gives figures of the minute structure in Nereis, showing that behind
the lens is a central axis with its sheath followed by blue pigment, then yellow pigment
in the central nerve-axis with its nucleus, and nerve-processes posteriorly.

Various allusions are made to the development of Nereids, and amongst others the
youngest larva3 mentioned by Milne Edwards 2 (1845) had three complete segments and a
fourth developing, two anterior cirri, two tentacular cirri, and two anal cirri.

Busch 3 (1851) describes a free-swimming larva of Nereis with six bristled segments,
two sub-tentacles, two anal cirri, and four eye-specks.

Wilson 4 (1892) describes a monotrochous larva of Nereis limbata, which exhibited
marked heliotropism. In the following stage on the twelfth clay it is still pelagic, but is
less active.

Goette 5 gives an account of a small prototroch of Nereis, and Salensky 6 alludes to
the pelagic stages of Nereis Dwnerilii with three pairs of feet and a pair of eye-specks.
He also describes the ninth day larva of Nereis cuUrifera as having three pairs of feet
with bristles, two frontal cirri, two tentacular cirri, and two anal cirri.

1 ' Journ. Morphol./ 1892.

2 ' Recherches/ p. 167, Taf. x and xi.

3 'Beobacht./ p. 69, Taf. ix, fig. 11.

4 ' Cell Lin./ 1892, p. 120, figs. 82 and 91

5 'Unters./ 1882, p. 84, Taf. vi, fig. 19.

6 < Stud/ ii, 1882, Taf. xxiv, fig. 10 n.

NEPEIDJE. 259

Hacker (1896) describes and figures the metatrochophore stage of Nereis with three
pairs of feet and three pairs of eye-specks, and later 1 (1898) a Nectochaste stage with four
bristled segments and two eyes. In Nereis limhata the prototroch appears in ten hours,
in another twenty hours it swims as a monotroch, and in two and a half days it assumes
what he calls a three-segmented Nauplius-stage. It ceases to be pelagic about the
twelfth stage. This author groups the Nereids with the Nephthydicla?, Phyllodocidse,
Eunicidse, and Aphroditidaa in having a secondary nectochsete stage.

The Nereids are generally found in tubes between tide-marks or near low-water
mark — sometimes under the nest of Amphithoe, at others in an independent dwelling in
fissures of rocks, under tangle-roots, in decayed stalks of the same sea-weed, or in
sponges. They also frequent the laminarian and coralline regions, and range to consider-
able depths, as in certain examples from the ' Challenger,' and those frequenting holes in
telegraph cables. Some occur in masses of peat, either having made the tubes or having
driven out those which made them, and others occupy tubes composed of fragments of
shells, corallines, algse, sand, gravel, and secretion ; or the tubes are more or less
transparent. Many are found in the stomachs of cod, haddock, and other fishes at St.
Andrews. Mr. Cyril Crossland communicates a note on a brilliant orange-red Nereid,
common at Dongonab in the Red Sea, in a thin collapsible tube attached throughout its
length. If the annelid is removed and placed in a vessel of sea-water, the tube is rapidly
re-formed. They are muscular annelids that wriggle most dexterously out of the forceps,
but unlike the common eel do not always make good use of their opportunities on the
ground, though they swim with vigour and agility. They soon die when kept with
other collections in a bottle, and rapidly putrefy. Their skins are extremely sensitive to
vapour of alcohol.

A few Nereids are commensalistic, like Nereile/pas fucata with the hermit crab in the
shell of Buccinum, and the orange species procured by Prof. Giarcl 2 in the folds of
Balanoglossus 3 on the rich sandy beaches of l'lles Glenans. The unarmed proboscis and
the reduced condition of the dorsal lobe of the foot with its simple bristles would indicate
its relationship with Lycastis, species of which have been found at Rochelle and elsewhere
on the shores of France.

Some devour other annelids, Crustacea, and sponges, the spicules of which occur in
their intestines; whilst others, like Nereis Dumerilii and N. cultrifera,Sire partial to Algaa.

All the British forms are marine, though N. pelagica moves far up the inlets in
North Uist, and N. diversicolor also is found in brackish regions; Cuvier, indeed,
suspected that true Nereides may be found in the lakes of North America, though the
reasons for this view are not stated. A considerable number of foreign species, however,
occur in fresh water and many in brackish water. 4 Such forms chiefly pertain to the genus

1 ' Plankton Exped. Polyclnet./ p. 9, Taf. 1, figs. 3 and 3 a.

' 2 Since this was written the distinguished French naturalist Las died. He was one of the
foremost marine zoologists of his day in Prance, and equally esteemed in other countries.

3 ' Compt. Rend./ Aug. 21, 1882, p. 389; < Ann. Nat. Hist./ 5th ser., vol. x, p. 230.

4 Interesting accounts are given by C. Gravier, ' Bullet. Soc. Philomathique de Paris/ 1901, and
1905, pp. 1—12, sep. copy. See also H. P. Johnson, ' Mark. Anniv. Vol./ art. x, pp. 208—223, pis.
xvi and xvii, 1903.

260 MICRONEREIS.

Lycastis, in which the dorsal division of the foot is rudimentary, being represented as a
rule by a spine and a few bristles. A recent contribution by Dr. R. Horst 1 on one of
Johnson's species (Lycastis hawaierisis) shows that the dorsal cirrus becomes greatly
developed. This form seemed to have been conveyed to the fresh-water pond of the
Botanical Gardens at Buitenzorg in transplanting mangrove trees.

No example of Ceratocephale (a form with soft papilla in the proboscis) has yet been
found in British waters, yet it occurs on the opposite shores of the Atlantic (Canada), in
Sweden, and in Japan. The Japanese species, Ceratocephale osawai, Izuka, 2 is the
"Palolo" of their seas, occurring in swarms in October and November close to the new
and the full moon, and thus in four different periods each lasting a few days. It, like the
swarming Nereis Dumerilii of Britain, is much used as bait, and is captured at night with
nets, each boat having a lantern to attract the worms. It differs from the Atlantic and
Pacific "Palolos" not only in family, but in the fact that its posterior end is shed as a
useless product, the ova and sperms being discharged from the open posterior end of the
anterior region, which swims freely. Dr. H. C. Sorby 3 has found in the summer months
Heteronereids swimming very actively off the estuary of the Thames and elsewhere, and
then suddenly disappearing.

SECTION I.

NEREIDS WITHOUT PARAGNATHI IN THE PROBOSCIS, AND WITH

BIRAMOUS FEET.

1. Micronereis variegata, Claparecle. 4 Head without tentacles or palpi; peristomial
segment with feet and bristles. Bristles all homogomph with long slender tips. Length
4 — 6 mm.

2. Leptonereis vaillanti, De St. Joseph. Head rounded, buccal segment passing
below, so that the tentacular cirri appear at its sides ; conspicuous glands in feet. Bristles
of more than one kind. Length 30 mm.

Genus LVIII. — Micronereis, Claparede, 1863.

Buccal segment bears feet and bristles. Proboscis muscular, only with maxillae.

Foot deeply cleft, with a dorsal and a ventral cirrus. Bristles uniform throughout ;
homogomph.

This genus, therefore, would naturally fall into the group devoid of paragnathi, and
is specially distinguished by having bristled feet on the buccal segment.

1 ' Bullet. Dep. Agric. aux Indes Neerlandaises/ No. xxv, p. 1 (sep. copy).

2 ' Journ. Coll. Sci. Imper. Univ. Tokyo/ vol. xvii, art. ii, pp. 1—37, 2 pis.

3 ' Journ. Linn. Soc./ vol. xxix, p. 434, 1906.

4 See a useful key to the British species by Major Elwes, ' Journ. M. B. A./ n.s., vol. viii, p. 357,
1909.

MICRONEREIS VARIEG-ATA. 261

1. Microneeeis variegata, Claparede. Plate LXXXVI, fig. 8 — bristle.

Specific Characters. — Head rounded in front, with traces of palpi, and four large eyes,
the anterior pair wider apart and with lenses. Tentacular cirri four, enlarged at the base,
which is coloured greenish or yellow.

Body 4 mm. long, very little tapered in front, but diminished from the third
segment to the tail, which has two caudal cirri of the same tint as the tentacular cirri
(Claparede). Bristled segments nineteen to twenty. Proboscis muscular; two horny
maxillae with five teeth.

With the exception of the first two, the feet are deeply biramous, the setigerous
lobes dilated distaJly and furnished with a subulate and subterminal dorsal and ventral
cirrus. Each lobe has a group of translucent homogomph bristles, with long, finely
tapered terminal pieces, the dorsal being somewhat longer than the ventral.

Synonyms.

1863. Micronereis variegata, Claparede. Beobacht. u. Anat., p. 57, Taf. xi, f. 5 — 7.

1864. ,, „ idem. Grlanures Zoot., p. 122, pi. viii, f. 4.

1865. „ „ De Quatrefages. Annel., i, p. 578.

1893. „ „ Eacovitza. Compt. Rend., 12 June, 1893.

1894. „ „ idem. Ibid., 15 Jan., 1894.

1900. ,, ., Fauvel. Mem. Soc. Nation, etc., Cherbourg, T. xxxi, p. 315.

1909. „ „ Elwes. Journ. M. B. A., n.s., vol. viii, p. 350.

Habitat. — Under a stone between tide-marks, St. Peter Port, Guernsey, July, 1868;
amongst littoral red algae and Lithothamnion between Oddicombe and Babbicombe
(Elwes).

St. Vaast-la-Hougue, Normandy (Claparede).

Head (Fig. 63, p. 262) rounded in front, and devoid of tentacles or palpi, though in
a preparation kindly forwarded by Major Elwes a bluntly conical process projected
ventrally on each side, but such may have been connected with the proboscis. Pour
large black eyes arranged in a trapezoid occur on the head, the anterior pair with
lenses and wider apart. The posterior pair show minute, glistening points as if
from crystalline cones. Tentacular cirri, four on each side, enlarged at the base and
banded with green or yellow and tapered distally, one of each pair being dorsal, the
other ventral.

Body about 4 mm. long, little narrowed in front, and tapering from the third foot
gradually to the tail, which ends in two long, tapering cirri. Bristled segments nineteen
or twenty. The general colour is brown, as Claparede notes.

The proboscis is a proportionally large muscular organ with two powerful maxillae
of a golden colour, short and broad in form and with five teeth on the edge in front
(Fig. 63), the largest tooth being anterior. In his first figure of the maxilla Claparede
indicated only three teeth on the edge, but his next representation shows six on one
side and seven on the other.

The first foot is small and uniramous, and is borne apparently by the buccal
segment. It has a somewhat clavate setigerous region, that is, the process enlarges

262

MICRONEREIS VARIEGATA.

toward the distal end, a pale spine, and a tuft of translucent bristles like those
following.

The second foot has a much larger setigerous region also enlarged distally and
curved, the convexity being in front in the preparation. A spine and a group of bristles
are also present. Claparede found a cirrus in both this and the first foot. The pre-
parations had lost them.

The third bristled segment bears a bifid foot, and this condition continues to the
posterior region. The typical foot (Fig. 63) consists of two widely separated divisions,
the split proceeding almost to the base, the spine in each division arising close to its
neighbour, and then gently diverging to the tip of its respective lobe. The superior
division in the specimen from the Channel Islands is somewhat clavate — that is, distinctly
enlarges till a well-marked knob for the cirrus on the upper surface is reached. The tip
then narrows to an acute angle (in spirit) occupied by the spine, which slightly projects.

Fig. 63. — Head and anterior region of Micro7iereis variegata, Claparede, somewhat compressed. (After Claparede.)

The inferior division of the foot has a similar shape reversed, and the ventral cirrus
is about the same size. The dorsal surface of the upper division is ciliated and may
assist in respiration (Claparede).

The bristles (Plate LXXXVI, fig. 8) are slender, translucent structures with a
slight curvature of the shaft, and an enlarged tip which is homogomph, the long, slender,
tapering terminal piece being sunk in the pit of the bifid region. In the example from
Guernsey the upper bristles (four to five) lay beneath the spine, whereas the inferior
bristles appear under pressure on both sides of the spine, and they are somewhat shorter
than the upper. As Major Elwes remarks, the tips of the bristles represented by
Claparede differ in being shorter, wider, and with a more distinct tendency to curvature.

Reproduction. — Claparede's first example was a female, and he found eggs 0*08 mm.
m the coelome in summer (July). As mentioned below, Racovitza more recently has
given an excellent account of the reproduction of this form.

This is one of the numerous discoveries of the accomplished and indefatigable
Claparede on the southern shores of Europe. He found a somewhat larger form (6 mm.)
at Port Vendres, 1 which had a third small, horny point between the tips of the

1 ' Grlanures Zoot./ p. 122, pi. viii, fig. 4 ; 1864.

LEPTONEREIS. 263

maxillae, but gave it no distinctive name. It probably is an older example of the same
species.

De Quatrefages (1865) thought that in regard to the fusion of the head and buccal
segment and in the structure of the feet it diverged from the Nereids ; indeed, he would
have considered it a larval form if Claparede had not found ripe ova. Ehlers, on the
other hand, is doubtful concerning its position, and would even connect it with the
Aphroditidae.

De St. Joseph 1 places Micronereis under the group devoid of paragnathi and having
a biramous foot, by which it is distinguished from Lycastis ; whilst it is separated from
Leptonereis, which he places next it, by having bristled feet on its buccal segment.
Gravier 3 follows a similar arrangement.

Racovitza (1893) gave an interesting account of the sexual distinctions of this form,
only the females of which, with twenty-one pairs of feet, had been seen by Claparede and
De St. Joseph. He found the male very minute, only 2 mm. long, or half the length of
the female, and with fewer feet — sixteen to seventeen pairs. Moreover, it has copula-
tory ventral hooks on the third pair of feet which do not exist in the other sex. The
maxillae likewise differ, for they have in addition to the smaller teeth three larger
posterior points. On the other hand the female has only five teeth. The reddish eggs,
which are about sixty in number, are deposited in glairy mucus, and the larvae escape
from this when they have five pairs of feet. The mature female undergoes considerable
changes. Thus tlie cirri enlarge, become ovato-lanceolate, and the anal cirri are elongated.
The posterior region of the body seems to be paralysed, and the animal drags it through
the glairy mucus by aid of the anterior region. This author a little later 3 gave a
description of the amoebocytes, the ovogenesis, and the deposition of the eggs in the same
form.

When observed in life in Guernsey it was at first sight, from its activity, associated
with the Hesionidae, but a more careful examination, independently of any reference to
literature, shoAvecl that it most nearly approached the Nereids, a conclusion which
substantiates the original view of Claparede and of others subsequently.

Genus LIX. — Lkptonereis, Kinberg, 1865.

Head rounded in front, eyes large, buccal segment and tentacular cirri shifted
forward ; anterior dorsal cirri peculiarly modified ; tail with two bluntly conical lamellae
and two slender subulate cirri above them.

Foot without spine or bristles dorsally in the first two segments, the rest boldly split
into two divisions.

Leptonereis was constituted by Kinberg 4 one of the genera of his family Niconidea in
which the paragnathi are absent. He characterizes the new genus as devoid of paragnathi

1 ' Ann. Sci. Nat./ 8 e ser., t. v, p. 285, 1898.

2 ' Nouv. Arch. Mus./ 4 e ser., fc. in, p. 155, 1901.

3 'Compt. Rend. Acad. Sci./ January, 1894.

4 ' Ofvers. k. Vet.-Akad. Forh./ 1865, p. 179.

264 LEPTONEREIS VAILLANTI.

(or soft papillae ?) ; form of foot altering by degrees, the superior branchiae (lamellae)
becoming large posteriorly— elongate and compressed; the dorsal cirri terminal; jointed
bristles both long and short. His single example L. Isevis came from a gulf beyond
Guayaquil.

Claparede 1 thought that the changes in the feet of the three genera, Mcon,
L&ptonereis, and Nicomedes proposed by Kinberg under his family Niconidea, were of
secondary importance, and he united them all under Leptonereis.

Grube 2 likewise minimised Kinberg's group Niconidea, and placed his example under
Nereis (Leptonereis).

1. Leptonereis Vaillanti, Be St. Joseph, 1888. Plate LXXXVI, figs. 9 and 9 a — bristles.

Specific Characters. — Head large, rounded in front, typical in regard to appendages;
four large eyes with lenses ; buccal segment twice the breadth of the others ; four pairs
of tentacular cirri, short, and appearing from above to be cephalic in position. Body
15 — 20 mm. long, segments forty-eight to sixty, tapering posteriorly, and ending in two
bluntly conical processes closely applied to each other, and two slender subulate cirri
above them. Anus in front and dorsal to these processes. Colour pale rose with darker
touches.

Proboscis has only a pair of well-developed maxillae with twelve to fourteen teeth.
First two feet have dorsal cirri, a pointed lamella beneath devoid of spine and bristles,
whereas all the other feet are complete, viz., dorsal cirrus, upper lamella, rounded
setigerous lobe with a black spine and homogomph bristles, whilst the inferior division
has a double setigerous region of a triangular shape, with two groups of bristles and a
powerful black spine. Upper group of bristles homogomph with some falcate hetero-
gomph, lower group with homogomph and a large number of heterogomph. Beneath
is an inferior lamella and a small ventral cirrus.

Synonyms.

1888. Leptonereis Vaillanti, De St. Joseph. Ami. Sci. Nat., 7 e ser., t. v, p. 246, pi. x, f. 113—123

pi. xi, f. 124.
1909. „ „ Elwes. Journ. M. B. A., n.s., vol. viii, p. 351.

Habitat. — A male heteronereid was found between tide-marks in February at
Oddicombe, Devon, by Major Elwes.

Shores of Dinard, France (De St. Joseph).

Head (Fig. 64) in the ordinary form (female nereid), according to De St. Joseph,
is large, rounded in front, with four large violet eyes with lenses, the anterior pair
elliptical, the posterior rounded. The buccal segment is twice the breadth of the others,
and the four pairs of tentacular cirri are short and appear from above to be almost
cephalic in position.

Body is from 15 to 30 mm. long, and has from forty-eight to sixty segments.

1 l Annel. Chretop. Nap./ SuppL, p. 90, 1870.

2 l Annulat. Semper./ p. 61, 1878.

LEPTONEREIS VAILLANTI.

265

It tapers posteriorly, and ends in two bluntly conical processes closely applied to each
other, and two slender subulate cirri above them. The aims is in front and dorsal to
these processes. Mounted as a transparent object, rows of glands cross the dorsum
of each segment between the feet. Colour, pale rose with darker touches.

The proboscis (Fig. 64) has well-developed maxillae with about twelve or fourteen
teeth, and two canals pass from the hollow base forward to the tip. These De St. Joseph
considers to be poison-canals. The roseate stomach has two lateral pouches which, after
Eisig, De St. Joseph considers to be air-chambers.

The first two feet have only the dorsal cirrus which is of a slightly bent fusiform
shape, a pointed lamella beneath in the superior division which is devoid of spine and
bristles, whereas all the other feet have this division complete, viz., dorsal cirrus, upper
lamella, rounded setigerous lobe with a black spine, and homogomph bristles (Plate
LXXXVI, fig. 9) and a second lamella.

Fig. 64. — Head and anterior region of Leptonereis Vaillanti, from a preparation forwarded by Major Elwes.

The inferior division has a double setigerous region of a triangular shape, the one
covering the other, with two groups of bristles and a powerful black spine. In the upper
group are homogomph with some falcate heterogomph ; the lower group is composed of
homogomph and a large number of heterogomph falcate bristles (Plate LXXXVI, fig. 9 a).
De St. Joseph states that the bristles are identical in structure with those figured by
Claparede 1 in Leptonereis glauca. Beneath the foregoing is an inferior lamella and a
small ventral cirrus. The feet vary in the several regions ; thus the first four are small
and have a large dorsal cirrus which projects beyond the upper lamella, whereas at the
fifth the cirrus is of the same length. Then the lamellae become prominent, and continue
so to the posterior end, where, for ten segments, the cirrus again extends beyond it.
After the tenth or twelfth three greyish glands occur on the dorsum of each foot, and they
are conspicuous in the posterior segments (De St. Joseph).

De St. Joseph found both male and female heteronereids between June and October,
yet he did not secure a male in the nereicl condition. The female heteronereid resembles

1 ' Annel. Nap./ Supply pi. vif, fig. 3 b.

91

266 NEREIDS.

the nereid female except that the large eyes are almost coalescent, the colour of the body
is dark orange, and from the sixteenth to the fourth in front of the tail swimming
bristles are present. The reddish-brown male heteronereid he procured in August, and it
measured about 20 mm. and had fifty-seven segments. The head has enormous eyes,
whilst the body has three regions. The first region is composed of fifteen segments
having bristles resembling those of the nereid form. The first four feet have the
peculiar hook-like dorsal cirrus, the first two, as in the nereid, having neither spine nor
bristles clorsally. Segments five to eight have the dorsal cirrus spatulate, whilst from
nine to sixteen it is short and subulate. The second region consists of thirty segments with
swimming bristles on the feet, and the heteronereid lobes are greatly developed, the
largest being the foliaceous lobe of the inferior division. The third region comprises
eleven segments and is translucent, or, if the sperms are developed, white. The feet
resemble those between the ninth and sixteenth, only their lobes are less developed.
Moreover, the inferior division has curious brown spines with a crotchet-like tip. The
tail resembles that of the nereid form, and the tissues are transparent (De St. Joseph).

Unfortunately the single preparation of the heteronereid is incomplete, so that only
an imperfect account of this form is at present possible. The foregoing remarks,
however, will aid in the discrimination of the form, the life-history of which is still in
need of investigation.

SECTION II.
NEREIDS WITH SEPARATE HORNY PARAGNATHI.

The British forms 1 in this group may for convenience be arranged according to the
condition of the paragnathi thus : —

1. All the eight groups of paragnathi present.

Nereis (Alitta) virens, Sars. With a leaf -like dorsal lamella. Large bluish-green
worm, varying in size and occasionally reaching three feet in length.

2. Group V or V and VI absent.

Nereis pelagica, L. Marked distinction between proximal and distal rows on

ventral surface of basal segment. Dorsal cirri longer than feet. Length

4—8 ins.
Nereis (Ilediste) dirersieolnr, 0. F. Midler. Distinguished from N. pelagica by

the much smaller paragnathi and the broader head ; dorsal cirri not longer

than feet.
Nereis (Praxitliea) Sclimardwi, De Quatrefages. Conspicuous glands in feet;

feet bluntly lobate in front, but upper lobe much elongated posteriorly.

Length 4 — 6 ins.
Nereis (Leontis) Dumerilii, Aud. & Echv. Paragnathi very minute, closely

arranged in pectinate rows. Two glands on each foot posteriorly. Very

long tentacular cirri. Length 2 — 3 ins.

Nereis (Nereilepas) fucata, Savigny. Dorsal lobe of foot large with the cirrus

1 Vide a useful key to the British species by Major Elwes, Mourn. M. B. A./ vol. viii, p. 357
1909.

NEREIS PELAGICA. 267

arising from the dorsal edge. Two white longitudinal dorsal bands. Com-
mensal of hermit crab. Length 4 — 9 ins.

3. Transversely elongated as well as conical paragnathi.

Nereis cultrifera, Grube. Paragnathi conspicuous, VI transversely elongated,
and the basal ventral in two regular rows. 6 — 12 inches in length.

Nereis (Stratonice) Marionii, And. & Edw. Paragnathi small, VI transversely
elongated at the base, pointed distally. Considerable dorsal elevation
commences at thirtieth foot and increases posteriorly to a long lobe with the
cirrus at its extremity. Length 3 — 4 ins.

4. Paragnathi only present on the basal region of the proboscis in extrusion, or

absent.
Nereis (Eunereis) longissima, Johnston. Paragnathi VI present in some.
Maxillary and basal regions of proboscis mapped into areas by grooves.
Cirri short. Length, a foot or more.

Genus LX. — Nereis, Linnseus, 1731. (Char, emend.)

Cephalic lobe somewhat rectangular at the base, on which the four eyes are
situated, whilst the anterior part is narrower and bluntly conical. Two tentacles ; palpi
two, lobate, with a terminal process. Nuchal organs primitive, consisting of a ciliated
surface in direct contact with the exterior (Racovitza) . Body sublinear, convex clorsally,
flattened ventrally, slightly diminished in front, more distinctly tapered posteriorly,
where it terminates in two caudal cirri below the vent. The first segment bears only
four tentacular cirri. Feet subequal throughout, sometimes foliaceous, with spines and
falcate bristles. Third and all succeeding feet biramous. Superior division of the foot
bifid, the spine not reaching the surface, and the bristles issuing from the base of the
fork. Proboscis armed with right and left maxilla3, strongly denticulated, and horny
dentiform processes (paragnathi) variously arranged, provided with powerful muscles, and
protrusible. (Esophagus with a pair of glands ; stomach simple and intestine straight.

1. Nereis pelagica, Linndeus, 1756. Plate LII, figs. 1 and 2; Plate LX, figs 6 and 6 a —
head; Plate LXXI, figs. 7-7 i— feet; Plate LXXX, figs. 25-25 b— bristles.

Specific Characters. — Head having a triangular anterior and a rectangular posterior
region with four black eyes of considerable size; the pairs wide apart and obliquely
situated. Tentacles and tentacular cirri of moderate length and typical. Body firm
and rounded, with a peristomial segment about twice the breadth of the succeeding;
segments sixty to eighty. Colour iridescent greenish-brown, fawn, reddish-brown, or
yellowish-brown, the central dorsal vessel forming a thin red line except at the moment
of contraction. Opposite each foot are yellowish-white grains, and often the sides and
feet generally present a greenish or olivaceous tint. Under surface paler, iridescent.
Anus often marked by a whitish opacity. The feet anteriorly are somewhat blunt and
bulbous at the tip. The first foot has three lobes, a dorsal, a ventral, and an intermediate
setigerous process which bears both homogomph and heterogomph bristles. The feet vary

268 NEREIS PELAGICA.

little after the three lobes and the two setigerous processes are complete, except that
behind the anterior third the tips of the lobes become more pointed, and posteriorly the
cirri are elongated. The posterior feet also diminish in size and become more transparent,
and the heterogomph bristles, both upper and lower, are proportionally stronger.

Proboscis of the typical two regions in extrusion (Fig. 58). From the maxillary
project the two dark brown, horny jaws, having five to seven teeth. Dorsally behind
them is on each side a crescent of dark brown paragnathi (II) formed of three rows
of points. Between these are two paragnathi (I), one anterior to the other. On
the eminence in front of each palpus is a group of four long teeth which probably
correspond with VI, whilst a narrow triangle with the apex in front probably represents
V, though devoid of paragnathi. Slightly behind the foregoing a line of paragnathi (VII)
trends to the ventral surface and joins VIII. Ventrally a belt of maxillary prickles (IV),
broader at the outer or ventral edge, occurs below the jaws in a similar position to the
dorsal crescents, but with a different arrangement of the prickles. In the median line is
a transverse belt of paragnathi (III), which in some forms a central group of about eight
and a short lateral crescent of three prickles. The basal division of the organ has a
conspicuous belt (VIII) extending more or less from the dorsal elevations across the
lozenges formed by the lines of the region. The larger prickles are generally in a single
row in front, whilst a belt of more numerous, minute prickles — diminishing from front to
rear — occurs behind. 1

Ejpitokous form. — In the males, which are found from January to June, the head is wide
and the eyes are larger; the dorsal cirri of the first seven feet are thickened throughout
the greater part of their length, the short tip being filiform. The ventral cirrus in the first
four or five feet is also enlarged. At the seventeenth foot the change from the nereid to the
heteronereid condition commences with increased vascularity; a dorsal lamella, a ventral, a
large inferior setigerous lamella, and a superior subsetigerous lamella all being developed,
with fan-like groups of swimming bristles. The dorsal cirrus has a row of prominent
papillae along its lower edge. Sperms extend far forward to the anterior or nereid region.
The colour is greenish anteriorly, posteriorly pale or cream-coloured from the sperms,
except in those first mentioned, Avhich are pale anteriorly. The tip of the tail is pinkish
from the vessels.

Female. — Deep russet-brown and iridescent in February from ova, which are of a
faint salmon-colour. The tips of the palpi are dull brownish-orange like the proboscis.
The same whitish patch occurs in front, between and behind the eyes. In September the
feet become very vascular, so as to give a deep red aspect to the sides, whilst their tips
are pale. At St. Andrews pelagic females have been found, but none have been observed
from other British localities.

Synonyms.
1746. Nereis pelagica, Linnasus. Fauna Suec, Ed. 2, p. 2096.
1758. „ „ idem. Syst. Nat,, Ed. 10, i, p. 654.

1762. ,, „ Baster. Opusc. subsec, Tom. ii, p. 133, Tab. 6, f. 6.

1 Grube characterizes the proboscis of the group containing N. pelagica and N. diver sieolor as
having VI of four or five in the form of a quadrangle or a cross, and Y absent.

NEREIS PELAGICA.

269

1767. Nereis pelagica, Linnaeus. Syst. Nat., ed. 12, i, pt. 2, p. 1086.

1770. „ ferruginea, Griinner. Skrift. Kjobenh. Selskab, x, p. 169, Tab. e, f. 10.
„ Soe-Skollorm, Strom. Ibid., x, p. 9, Tab. b, iv.

1771. Die warzige Nereicle, 0. F. Muller. Yon Wiirmern, p. 144.
1776. Nereis verrucosa., idem. Zool. Dan. Prodr., p. 217.

,. „ fimbriata, idem. Ibid., p. 217.

1780. „ verrucosa, Fabricius. Fauna Gronl., p. 292.

1791. „ pelagica, Linnaeus. Syst. Nat. (Gmelin), ed. 13, i, pt. 6, p. 3116.

1806. „ „ Turton's Linnaeus, iv, p. 86.
„ „ fimbriata. Ibid., iv, p. 86.

1807. „ pelagica. Ibid., Brit. Fauna, p. 135.
1817. ,, „ Stewart. Elements, i, p. 390.
1820. „ subulicola, Leach. Brit. Mus. Coll.

1825. „ margaritacea, De Blainville. Diet. So. Nat., xxxiv, p. 433.

„ „ pelagica, idem. Ibid., p. 433.

1827. ,, „ Encycl. Meth., 1 Vers, i, p. 134, Tab. Iv, f. 21—23.
„ „ fimbriata. Ibid., p. 134, Tab. Iv, £. 18—20.

1828. „ margaritacea and pelagica, De Blainville. Diet. Sc. Nat., lvii, p. 470.
„ Lycoris margaritacea, Johnston. Zool. Journ., iv, p. 419.

„ ,, viridis, idem. Ibid., p. 420.

1830. Nereis pelagica, idem. Ann. Nat. Hist., iii, p. 290, pi. vi, f. 1.

1833. „ margaritacea, Aud. & Edw. Ann. Sc. Nat., t. xxix, p. 217.

1834. Lycoris margaritacea, Johnston. Loud. Mag\ Nat. Hist., vii, p. 230, fig. 42.
1840. Nereis pelagica, idem. Ann. Nat. Hist., v, p. 172, figs. 3, 4.

1842-3. „ „ (Ersted. Kroyer's Nat. Tids., p. 115.

1843. „ ,, idem. Annul. Danic. Consp., p. 21, f. 72, 75, 76.

„ „ „ H. Rathke. Nova Acta Acad. Leop.-Cur., xx, 1, p. 158, Tab. viii, f. 1 — 3.

„ „ ,, Oersted. 3 Gronl. Annul. Dorsibr., p. 175, f. 52, 53.

1844. ., „ Thompson. Rep. Brit. Assoc, 1843, p. 273.
1844-5. ,, „ (Ersted. Kroyer's Nat. Tids. (Dr^bak), p. 408.
1847. ,, „ Frey and Leuckart. Beitr., p. 154, Tab. ii, f. vii.
1849. „ „ Leuckart. Arch. f. Naturges., xv, i, p. 207.

1851. „ „ Sars. Nyt Mag. f. Naturvid., vi, p. 209 (Reise Lofot. og Finmarken).

,, „ „ Grrube. Fam. Annel., pp. 48 and 125.

1853. ,, fulgens, Daly ell. Pow. Creat., ii, p. 153, pi. xxii, f. 6 — 8.

1858. „ pelagica, Grrube. Sibirien's Annel. Mem. Sav. etrang., St. Petersburg, viii, p. 4.

1860. ,, „ Packard. Canad. Nat. and Greol., vii, p. 418.

1863. „ „ Stimpson. Ann. Grreenl., in Proc. Acad. Nat. Sc, Philad., p. 140.

1865. „ „ Johnston. Cat. Worms Brit. Mus., pp. 148 and 341.

„ ,, diversicolor, idem. Ibid., p. 152 (excl. syn.).

„ ,, fimbriata, Johnston. Cat. Worms Brit. Mus.

,, ; , pelagica, De Quatrefages. Annel., i, p. 542.

„ ,, margaritacea, idem. Ibid., p. 510.

„ „ Reynaudi, idem. Ibid., p. 519.

pp. 155 and 341

1 Bruguiere, Lamarck, and Deshayes.

2 It has not been thought necessary to quote a resume in Kroyer's ' Naturh. Tidskrift/ Bd. iv,
2 Hefte.

270 NEREIS PELAGICA.

1865. Nereis Boiverbanckli, idem. Ibid., p. 541.

„ ,, pelagica, Carrington. Annel. Southport, p. 8.

1866-9. „ „ Packard. Mem. Bosfc. Soc. Nat. Hist., i, p. 294.

1867. „ „ Malmgren. Annul. Polych., p. 47, Tab. v, f. 35.

1868. „ „ Ehlers. Borstenw., ii, p. 511, Taf. xx, £. 11—20, and var. lunulata,

p. 511.
1870. „ „ Grube. Arch. f. Naturges., Bd. xxxvi, p. 308.

,, ,, margaritacea. idem. Ibid., p. 314.

1873. „ pelagica, Verrill. Invert. Yiny. Sound, i, pp. 319, 591, pi. xi, f. 52—55.
„ „ „ Kupffer. Exped. Osfcsee, ' Pommerania/ p. 150.

,, „ „ Sars. Bidrag Christ. Fauna, p. 22.

1874. „ ., and heteron&reis, Mobius. Die Zweite cleutsch. Nordpol., p. 254.
„ ,, „ Grube. Jahresb. Schles. Gesell. for 1873, p. 70.

„ ,, „ Smith and Harger. Trans. Conn. Acad., iii, pp. 12, 16, 20.

,, „ ,, Mcintosh. Ann. Nat. Hist., ser. 4, vol. xiv, p. 198.

„ „ „ Malm. Goteb. vet. o. vitt. Handl, xiv, p. 83.

1875. „ ,, Mcintosh. Invert, and Fishes St. Andrews, p. 122.
,, ,, ,, idem. Proc. Roy. Soc, vol. xxv, p. 217.

,, „ ,, Mobius. Jahresb. Comm. deutsch. Meer. ii, p. 168.

1878. ,, „ Lenz. Jahresb. Comm. deutsch. Meer. iv, Anhang., p. 13.
„ „ ,, Marenzeller. Denks. k. Akad. Wien., Bd. xxxv, p. 396.

1879. „ ,, idem. Ibid., t. xli, p. 122 (Sudjap. Ann.).

„ Verrill. Check. List, U.S. Comm. F. and F., p. 8.
„ „ ,, Tauber. Annul. Danic, p. 96.

„ „ ,, Mcintosh. Trans. Linn. Soc, 2 ser., Zool., vol. i, p. 503.

„ „ ,, Hansen. Nyt Mag. f. Naturvid., xxiv, p. 268.

1881. ,, „ Leslie and Herdman. Proc R. Phys. Soc Edin., vol. vi, p. 274.

„ „ „ Horst. Niederl. Arch. Zool., Suppl. Bd. i, p. 12. (Wm. Barent's Exped.,

1878.)
„ ,, Levinsen. Yidensk. Meddels., 1881, p. 234.

„ „ Webster and Benedict. Pep. U.S. Comm. F. and F., p. 718.

Mcintosh. Annel. < Challenger' (t. xii), p. 216.
„ Cams. Fauna Medit., p. 218.
,> „ Harvey Gibson. Proc Lit. Phil. Soc. Liverp., vol. xi, p. 154.

„ „ Jeffrey Bell. Proc Zool. Soc Loud., p. 3.

procera, De St. Joseph. Ann. So. Nat., 7 C ser., t. v, p. 266, pi. xi, f. 132.
„ pelagica, Marenzeller. Archiv f. Naturges., 1889, i, p. 128.
„ „ Giard. Bull. Soc Sc Fr. Belg., t. xxii, p. 77.

} , „ Malaquin. Annel. Boulon, p. 29.

„ „ Hornel. Trans. Biol. Soc Liverp.. vol. v, p. 241.

» » Biedermann. Jenaische Zeitschr. f. Nat., Bd. xxv, p. 450.

>, (eyes) Andrews. Journ. Morph., p. 178, pi. ix, f. 13^

„ „ Levinsen. Yidensk. Ud. " Hauchs/' p. 329.

De St. Joseph. Ann. Sc Nat., 7 e ser., t. xx, p. 221, pi. xiii, f. 40.
;; „ Michaelsen. Gronl. Annul. (Zool. Ergebn. IX), p. 126.

» » De St. Joseph. Ann. Sc Nat., 8 e ser., t. v, p. 293.

„ ■> Whiteaves. Geol. Surv. Canada, No. 722, p. 80.

Mcintosh. Ann. Nat. Hist., ser. 7, vol. x, pp. 256 and 258.
,, Allen. Journ. M. B. A., n.s., vol. vii, p. 224.

1883
1884

1885

1886
1887
1888
1889
1890

1891

;;

1892
1893

1895
1898

>>
1901
1902
1904

NEREIS PELAGICA. 271

1906. Nereis pelagic a, De St. Joseph. Ann. Sc. Nat"., 9 e ser., t. iii, p. 2] 5.

1908. „ „ Horst. Tijdschr. Nederl. Dierk. Yereen (2), xi, p. 143.

1909. ; , „ Blwes. Journ. M. B. A., n.s., vol. viii, p. 351.

„ „ „ Moore. Proc. U.S. Nation. Mus y vol. xxxvii, p. 137.

Forma Epitoca. Male.

1800. Diefaserige Nereicle, O. F. Muller. Wurm-Arten., p. 144, Tab. viii, f. 1—3.

1840. Nereis renalis, Johnston. Ann. Nat. Hist., y, p. 176.

1843. Heteronereis arctica, CErsted. Gronl. Annul, dors., p. 179 (27), f. 50*, 51, 60, 65, 68, 70*.

„ Nereis grandifolia, H. Kathke. Beitrage Fauna Norweg., p. 155, Tab. vii, f. 13, 14.
1849. „ „ R, Leuckart. Archiv f. Naturg., Jrg. 15, Bd. i, p. 207.

1851. ,, renalis, Williams. Rep. Brit. Assoc., p. 197.
1865. Heteronereis renalis, Johnston. Cat. Worms Brit. Mus., p. 163.

,, „ arctica, De Quatrefages. Annel., i, p. 566.

„ „ renalis, idem. Ibid., p. 580.

Female.

1842-3. Heteronereis arctica, CErsted. Kroner's Nat. Ticls., p. 117.

„ „ assimilis, idem. Ibid., p. 117.

1843. „ „ idem. Gronl. Annul. Dorsibr., p. 180, f. 54, 61, 72,

„ ? Nereilepas fusca, idem. Annul. Panic. Consp., p. 21, f. 49, 50.
1853. Nereis denticulata, Stimpson. Invert. Grand Manan, p. 33, f. 23.
1865. Nereilepas fimbriatus, Pe Quatrefages. Annel., i, p. 559 (excl. syn.).

„ Heteronereis assimilis, idem. Ibid., p. 566.

„ „ „ Malmgren. Nord. Hafs- Annul., p. 108, Tab. xi, f. 15 e, c.

1866-69. ; , arctica, Packard. Mem. Bost. Soc, Nat. Hist., i, p. 293.

1874. „ grandifolia, Malm. Coteb. vet. o. vitt. Handl., xiv, p. 84.

Male and Female.
1865. Heteronereis grandifolia, Malmgren. Nord. Hafs-Annul., p. 108, Tab. xi, f. 15 ? , £. 16, b,

1867. ,, „ idem. Annul. Polych., p. 60, Tab. iv, f. 31.

1873. „ „ Sars. Bidrag Christ, Fauna, p. 23 (sep. copy).

Habitat. — Abundant in tubes under stones between tide-marks on both shores of
Britain and Ireland ; under layers of slaty rocks, in fissures of sandstone, in sanely mud,
and under old limpets. It is less common in the Channel Islands, where medium or
small examples are found only in chinks of rocks and under Balani and Ascidians in the
Grouliot Caves, Sark, a single small example of Nereis cultrifera, and one or two of
N. DumeHlii accompanying them. Occasionally the tube formed by it in fissures of
sandstone is not half the length of its body.

Many young forms occur on the tangle-blades (Laminaria saccharina and L. digitata),
to the surface of which their tubes are attached, and also on Delesseria sanguined in the
laminarian region. It extends to the coralline ground off the East coast, and now and
then it takes possession of an empty tube of other forms, such as Thelepus.

It ranges from the shores of France, though De St. Joseph found it bat seldom at

272 NEREIS PELAGICA.

Dinard, to the south of Halifax, Nova Scotia ; Japan (Marenzeller) ; the eastern shores
of Siberia near Anion (Grube) ; Behring's Sea (Marenzeller) ; and var. lunulata, Ehlers,
to the Strait of Magellan. Large examples were procured by the 'Valorous' in
20 fathoms in the Arctic seas. Specimens were also found on floating seaweed off (rough's
Island, South Atlantic, long. 9° 49', lat. 40° |' (Prof. Dickie, Aberdeen), and a variety
from Kerguelen.

This cosmopolitan species thus extends from the British shores to those of northern
and southern Europe, the Mediterranean, the Atlantic shores of America, the Pacific, the
Arctic, and the Antarctic seas.

Head (Plate LII, fig. 1, and Plate LX, figs. 6 and 6 a) somewhat rectangular
posteriorly and bluntly triangular in front. Four black eyes of considerable size are
situated on the rectangular region. The pairs are wide apart and obliquely placed, —
that is, the anterior pair is wider apart than the posterior. A pair of short tentacles
occurs on the truncated anterior end of the head. From the lateral regions of the snout
the palpi project forward and outward as large tumid processes in contraction, and having
a terminal articulation which is retractile within the basal region. From the anterior part
of the peristomial segment on each side spring two pairs of tentacular cirri, which form
a kind of cluster, though the anterior pair arises somewhat lower than the posterior.
Each has a long dorsal cirrus and a shorter ventral cirrus, with a basal cirrophore, the
longest being the dorsal of the second pair. The peristomial segment is about double the
breadth of the average segment, and probably the tentacular cirri indicate that the
segment is double, the appendages having been shifted forward.

Body 4 — 8 ins. long, slightly diminished anteriorly, then remaining of a uniform
diameter for some distance, and again narrowing to the tail, where it terminates in a
prominent vent, beneath which are two long cirri. The segments are convex dorsally,
flattened ventrally, and of nearly uniform breadth and appearance. In colour it is of an
iridescent greenish-brown, reddish-brown, or yellowish-brown, the dorsal vessel forming
a thin, dull red line, except at the moment of transmission of the blood. Opposite each
foot in some are yellowish- white grains.

There is considerable variety in the coloration, though most show a brownish or
fawn-colour dorsally with a greenish or olivaceous shade at the sides and feet. In some
a dark grey stripe occurs along each side dorsally at the bases of the feet. Many of the
larger specimens have calcareous deposits in circular patches in the skin of the dorsum.
The under surface is of a paler, iridescent, or roseate hue, and the anus is often marked
by a whitish opacity.

Proboscis. — In extrusion the organ presents dorsally (Plate LX, fig. 6) two regions
— a distal or maxillary and a proximal or basal— mapped out into distinct and somewhat
lozenge-shaped areas by linear depressions. From the distal project the two curved
horny maxilla) or jaws, with five teeth. On the dorsal margin behind them (Area II) is
a crescent of dark brown horny teeth, formed of a posterior curved row, another which lies
within (in front of) the former, and a third short row composed of four small prickles at
the inner border in front. In the central line (Area I) between these are two teeth, one
in front of the other, on an elevation. In front of the palpi is a prominence on each side
(VI) with four long prickles or teeth, and slightly behind each a line of prickles (VII and

NEREIS PELAGICA. 273

VIII) trends to the ventral surface. Area V seems to be represented by an acutely
triangular space between and behind VI. Ventrally (Plate LX, fig. 6 a) a belt of prickles
(IV), broader at the outer or ventral edge, occurs below the jaw r s in a corresponding
position to the dorsal crescents, though with a different arrangement of the constituent
prickles. In the median line on an eminence is a central group of small prickles (III),
about eight in number, and on each side a short lateral crescent of three prickles.
The band of points (VII and VIII) from the dorsal elevation with the four horny teeth
extends across the ventral surface, the arrangement of the prickles being somewhat
modified by the mapping out of the area by the depressed lines. The general effect,
however, is to produce a belt, with the larger prickles, in some cases, forming an anterior
row, and a belt of more numerous small prickles behind, the smallest points being
posterior. In some of the areas marked off by the depressed lines two or three of the
large points occur in front and a group of the smaller behind, and an indication of
symmetry is seen especially in the lateral areas. Little variation is found in the
arrangement and proportional size of the paragnathi.

The food consists of delicately branched Algae, coarse fragments of Fuci and other
sea- weeds, sandy mud, sponge-spicules, small ova, bristles of other polychaetes — probably
swallowed with portions of the bodies. The faecal masses are enveloped in a coating of
mucus.

The first foot (Plate LXXI, fig. 7) has dorsally a prominent edge ending in the
cirrus, then a large lobe smoothly rounded distally. Beneath is a somewhat pointed
conical setigerous process bearing two tufts of bristles and a spine, the upper tuft having
the terminal pieces long and serrated, with the exception of one with a short serrated
terminal process. The lower group consists of bristles with short terminal pieces.
Inferiorly is a lobe of similar shape externally to the dorsal, but as the ventral cirrus
springs from the base close to the body, it is more conspicuous and has a slightly sinuous
ventral margin. The second foot is more or less like the first, the elevation at the base
of the dorsal cirrus being more prominent, but the third has two lobes above the conical
setigerous process, viz., a smaller upper, and a lobe resembling the dorsal of the first foot.
Between them, however, is a tuft of bristles with long spear-tips. There are now two
spines. The setigerous region remains conical, and the inferior lobe has increased in
bulk. The dorsal cirrus is shorter.

The tenth foot (Plate LXXI, fig. 7 a) presents in the dorsal division two nearly
equal lobes with the bristles between, the dorsal bearing the cirrus, whilst the ventral
division consists of the conical setigerous and a rounded ventral lobe with a cirrus at its
base. All the bristles in the upper division have long slender tips. Those of the ventral
division form two groups. In the upper group most have long tips, but three or four
have short tips. In the lower group a few of the upper have long terminal pieces, the
rest short tips (Plate LXXX, fig. 25).

The thirty-seventh foot (Plate LXXI, fig. 7 b) has the two lobes of the upper division
considerably more pointed, so that they form elongated cones. The shafts of the bristles
have greatly increased in strength, and one or two with short tips and strong shafts are
present. In the inferior division the two groups are equally differentiated, those with
slender shafts having long slender tips (Plate LXXX, fig. 25 a), whilst those with stout

92

274 NEREIS PELAGICA.

curved shafts have short tips. The conical setigerous region is less prominent than the
dorsal lobes. The ventral lobe is also pointed and more prominent than the setigerous
region. The cirrus is shorter than the latter, and the foot generally is less massive.

The fifty-seventh foot (Plate LXXI, fig. 7 c) has considerably diminished in general
bulk, and the lobes are more acute. This diminution and increased transparency make
both the spines and bristles very prominent in mounted preparations ; the deep black of
the former and the golden hue of the stout bristles with the short tips are characteristic.
Both dorsal and ventral cirri are proportionally longer and more slender, the former, as
in the thirty- seventh foot, far exceeding the latter. While this foot closely resembles in
general configuration the thirty-seventh, except that the dorsal cirrus is longer, and the
proportions of the two upper lobes are in contrast with the lower, the bristles with the
short tips both in the upper and lower groups are proportionally stronger.

An interesting account of the minute structure of the bristles of this species is given
by Schepotielf, 1 who shows that the rows of cross-bars are continuous with the upper
longitudinal streaks.

Just in front of the tail the foot presents a still further diminution in size, the hump
of the dorsal outline at the base of the cirrus is marked, the upper lobe is smaller, and it
and the others have less acute tips. The dorsal cirrus is much lengthened, and the
ventral projects beyond the adjoining lobe. The vascularity of the foot generally is
well shown in these translucent organs. The bristles retain the characters already
indicated.

In this species the ventral longitudinal muscles are more elongated transversely, and
have a longer and more pointed dorsal (external) fold than in Nereis diversicolor.

In a specimen from St. Andrews in which injury had occurred, the third and fourth
feet of the right side were in process of reproduction. In these the dorsal and ventral
cirri were proportionally large, appearing more conspicuous than the lobes, yet all were
new. Many examples show reproduction of the tail. Prof. Jeffrey Bell, again, has
recorded an example from Guernsey in which the tail was bifid.

Habits. — Though the majority of examples obtained from deep water off St. Andrews
are small, very few reaching medium size, large specimens are found in the stomach of
the cod and also in that of the flounder. In the whole series of British examples no finer
ones occur than those from St. Andrews. The other features in the habits of this form
are indicated under those of the family.

This species has been supposed to bore in wood and also in telegraph cables. Thus
Dr. Carpenter forwarded a specimen of medium size along with a portion of a telegraph
cable (1874). So far as could be observed, however, the perforations were due to other
forms.

Reproduction. — Specimens having the general form of Iphinereis fucicola, (Erst, are
not infrequently met with at low water early in February, or even as early as the 18th
January. The anterior region of the body resembles that of Nereis pelagica typica, only the
head and palpi are of a richer green with a slight tint of blue, and the tentacles are bluish-
green, thus contrasting with the dull orange proboscis, which is beautifully marbled or
cut into areas by pale streaks. The terminal processes of the palpi are paler. The
1 ' Zeitschr. f. wiss. Zool./ Bd. lxxiv, p. 21, Taf. xxxiv, figs. 5—15, 1903.

NEREIS PELAGTCA. 275

brownish-purple eyes are larger, have lenses, and a median patch of white occupies the
region behind the posterior pair and passes forward as a wedge in front. A brown
patch exists behind the posterior pair, mimicking a third pair on each side. A little
white also occurs to the exterior of the anterior eye. The prickles of the proboscis agree
with the typical form. The anterior feet, whilst also resembling those of Nereis pelagica
generally, have the dorsal cirri of the first seven enlarged in the middle. At the
seventeenth foot the changes in the lobes of the feet commence with increased vascularity,
and this increases in the posterior feet. The coloration above alluded to (greenish, with
a slight tinge of red) extends at least as far backward as the seventeenth foot, gradually
fading behind, all the feet being paler than the body. The dorsal blood-vessel enlivens
the region, and by-and-by fine, dark-greenish, granular transverse lines mark each
segment, but the posterior region is only granulated. A little later, as in the beginning
of March, those with the nuptial form as described have the sperms extending far
forward — in front of the region with the modified feet. The body behind the greenish
anterior region is pale — whitish or cream-coloured — from the sperms ; indeed, some
specimens are whitish up to the peristomial segment, laterally and ventrally. The tip of
the tail is slightly pinkish from the blood-vessels.

These forms are likewise found in June and July under stones near low water-mark,
or in runlets of water at half -tide. They swim freely about in the water assisted by the
large lamellas. In some many of the feet are only half the size of the others, having
apparently either been later in assuming the change, or in process of reproduction. The
dense whitish mass which exudes on rupture consists of sperm-cells not yet matured. The
cells are not entirely granular throughout.

In a male with typical paragnathi (Plate LII, fig. 2), procured on the 18th January,
the first fifteen feet are more or less normal, with the exception of the enlargement of the
dorsal cirrus in the first seven segments and the slight enlargement of the ventral cirrus in
five. The sixteenth foot has a minute flap close to the internal border of the dorsal cirrus,
and a like rudiment occurs in a similar position to the ventral cirrus. A rudiment of a
lamella also appears posteriorly at the tip of the inferior setigerous process. At the
seventeenth foot the three lamellae or flaps are larger, but the bristles (homogomph) are
unaltered. The heterogomph bristles are absent. The full development of the
heteronereid foot occurs in the thirty- seventh and neighbouring feet (Plate LXXI, fig.
7 f), in which the lamella on the ridge internal to the dorsal cirrus is fan-shaped. The
dorsal cirrus is rather long with nine or ten papillae on its lower border. Each papilla
has an opaque mass internally. The dorsal lobe is lanceolate and comparatively short,
the cirrus extending more than double its longer diameter beyond it. The dorsal setigerous
process bears only swimming bristles (Plate LXXX, fig. 25/;). The lobe beneath has a
short conical tip, and a heel projecting beyond its base internally. The inferior
setigerous process has a large fan-shaped lamella posteriorly. The bristles are of two
kinds in the specimen, viz., the dense fan of swimming bristles, and a few of the atokous
forms, both homo- and heterogomph. These in all probability would soon be shed. The
ventral lobe is tongue-shaped. The ventral cirrus has a fan-shaped lamella internally and
a pointed one externally.

The average number of nereid feet in front of the heteronereid region seems to be

276 NEREIS PELAGICA.

about sixteen, though sometimes seventeen or eighteen are present. Levinsen (1883)
likewise mentions that the first sixteen are unaltered.

The fifty-seventh foot (Plate LXXI, fig. 7 g) differs little from the preceding
except perhaps in the narrower dorsal and more attenuate dorsal and ventral lobes, and
the diminished number of crenations on the under surface of the dorsal cirrus. These
crenations disappear before the eightieth foot, which presents a somewhat fusiform
dorsal cirrus and a tapering ventral cirrus.

The eightieth foot (Plate LXXI, fig. 7 h) shows considerable modification of
the parts, though the plan is similar to the fifty-seventh. The dorsal cirrus is now
smooth.

In a large epitokous example procured by Mr. Gray in the stomach of a salmon at
Berwick-on-Tweed, a curious transverse mark, simulating a pit on one side, occurred
behind each posterior eye.

In certain examples from the deeper water of St. Andrews Bay the paragnathi of
the basal segment of the proboscis in extrusion present certain variations. Thus the
anterior row of large isolated points is very regular and distinct, each occupying its area,
whereas the smaller teeth behind are very minute and at first sight almost invisible. The
widening of the head and the increase in the size of the eyes are often marked.

In a pelagic male the dorsal cirri of the first seven segments are much enlarged, the
filiform tip only extending from the bulbous end of the cirrus. Moreover, the ventral
cirri have the same shape. These curious cirri thus approached those anteriorly in
N. Burner llii.

The first foot in an epitokous male more or less agrees with that in the atokous
form, consisting of a single lobe superiorly and another below the setigerous lobe with the
single spine. In this, a large specimen, the dorsal and ventral lobes are bluntly rounded
and short, but in the smaller forms (males) they are more elongated, though still with
a bluntly rounded tip. The dorsal cirrus in the large example is long and tapering
(Plate LXX, fig. 7) with a slight thickening of its base dorsally, extending about a third
of the length of the process. The dorsal and ventral lobes form short, bluntly rounded
cones. The intermediate setigerous lobe ends in a short bifid cone, bearing superiorly
bristles with long tapering tips and interiorly some with shorter tips, considerably
longer, however, than the ordinary types.

In another epitokous male the changes are only evident in the twenty-third foot,
though a small process occurs at the base of the ventral cirrus of the twenty-second foot.
In this foot (twenty-third) a process appears at the base of the dorsal cirrus, and the
cirrus itself is clavate. The dorsal lobe is not much altered. The superior setigerous
lobe bears a dense tuft of the sabre-like swimming bristles and is closely connected with
the lobe beneath it, which has a nearly normal outline, but in addition a large translucent
fan-shaped process projects downward behind the inferior setigerous lobe. The latter is
elongated, and a dense series of swimming bristles is developing, whilst the shafts of two
of the stronger bristles of the original series with short tips still remain. This lobe has
a large foliaceous and translucent lamella projecting from its outer and upper border.
The ventral lobe is tongue-shaped, and the ventral cirrus has a flattened leaf-like
expansion on each side of its base.

NEREIS PELAGIOA. 277

The Johns tonia prolifera of De Quatrefages 1 is a Heteronereis of this or other species
of Nereis. An epitokous example of this form is in all probability the Nereis bipinnata of
Montagu (MS. Drawings, Linn. Soc., Plate XXIX, fig. 3, 1808).

Female Heteronereid. — In September the feet of the female become very vascular,
so as to give a deep red appearance to the sides (Plate LII, fig. 1), and as the tips of
the lobes of the feet are pale, the aspect is the more striking. The perivisceral cavity
is loaded with eggs which have one large oil globule. They are opaque whitish, and the
majority are of equal size. There is as yet no indication of structural change in the feet
or other organs.

At the beginning of February large females show a somewhat deeper and beautifully
iridescent russet-brown of the body, probably from the tint of the eggs, which are faint
salmon or buff coloured in mass. The head and tentacles are of the same greenish hue as
the males, the palpi having the tips dull brownish-orange like the proboscis. The same
whitish patch (though less developed) occurs behind, between and in front of the eyes.
The latter are not enlarged. The armature of the proboscis is typical. The feet show
no evident change in front and apparently not much posteriorly, their extremities being
pale. The ova are of comparatively large size. The Nereis ferruginosa of Montagu
(MS. Drawings, Linn. Soc, Plate LII, fig. 2) may represent this stage, and it appears to
be the Nereis fulgens of Dalyell.

Before the heteronereid condition is developed, the female Nereis pelagica is thus
distinguished by the lustrous deep brownish or bronzed colour just mentioned, and the
size is frequently larger than that of the male. As the ova develop and distend the
posterior region the latter assumes a somewhat pale hue, and by-and-by also the body
generally becomes somewhat shorter and broader from contraction. The change of colour
does not at first much affect the general aspect, and no sign of alteration in the structure
of the feet is visible externally, but they are highly vascular, the dorsal edge of each
being connected with the body by a reticulated bar of blood-vessels. The eggs at this
stage are dull yellowish and minutely granular, and have none of the large globules of a
subsequent stage; moreover, in the lustrous brown condition of the adult they are more
minute. The eyes as yet show no perceptible change.

The next stage is characterized by the lustrous pale-greenish condition of the body
from the twentieth foot backward, whilst the brownish-olive hue remains in front. The
head has a pale trifid mark between the eyes, the median spur going forward. The eyes
do not show marked alteration, but the anterior dorsal cirri have a slight enlargement at
the base. The pale region is loaded with greenish -white ova measuring -^ mm. The
vascularity of the feet is pronounced.

Then the deep brownish or olive colour anteriorly gives way to a paler greenish
brown, the posterior region becoming still more pale as the eyes enlarge. Various isolated
and fixed black specks appear, but whether in the gut or body-wall is not clear. The
pallor makes the blood-vessels of the feet and of the dorsum conspicuous. The head and
anterior region retain in some the brownish hue mingled with green for some time, but
the tentacles and tentacular cirri are pale greenish.

At a further stage (24th February) the anterior region of nineteen segments is olive
1 f Ann. Sc. Nat./ 3 e ser., fc. xiv, p. 350, pi. viii, fig. 1, August, 1850.

278

NEREIS PELAGICA

green, and the palpi, tentacles, and tentacular cirri are pale green, the head itself being olive
with the white trifid mark posteriorly. The pelagic posterior region is developing, and
it is pale, while the feet are vascular. The ova are pale greenish with numerous large
globules amongst the smaller.

The epitokous females hitherto observed are about the size of the males, though one
or two are larger, and all possess the triradiate band between the eyes and a pale peacock-
green colour anteriorly. The first five dorsal cirri are enlarged at the base and the
ventral cirri of the same feet are slightly swollen. In the altered posterior region the
dorsal cirri are also somewhat dilated towards the base (cirrophore), and have inf eriorly
on the enlarged region low papillae or warts, to which groups of stalked Infusoria and
also slender filaments of Algae adhere. Thus, although the prominent row of papillae
which projects from the lower edge of the cirrus in the male is absent, there is a tendency
to such growths. The ventral cirri present even a more marked tendency to these
growths, about four prominent cones being found on the ventral edge, and they also

otn

vfn

Fig. 65. — Transverse section (made by Capt. F. H. Stewart, M.D.) of a nearly ripe Nereis, ov. Ova.

The other letters as before.

have Infusoria and Algae attached. The swimming bristles in the posterior region are
very large. Ventrally the same marked distinction between the anterior and posterior
regions of the body occurs as in the male, the anterior region being pale, a slightly
greenish shade only being visible in the head-region, whereas the posterior is of a
deep pinkish purple. This hue is apparently not due to the muscles, as during the
movements of the animal the intensity varies, as if from vascular changes.

The heteronereid feet of the female agree generally in the position and shape of
the lamellae and other parts with the male.

The changes which ensue in the body of the ripe female Nereis are indicated in
the accompanying section (Fig. 65).

Two examples, about 5 ins. in length and apparently answering to this species in the
heteronereid condition, were procured by Mr. Gray in October, 1894, in the stomach of
a salmon at Berwick-on-Tweed. Both were females, still with a few dark-coloured ova
in groups at the bases of the feet. The condition of the jaws and paragnathi, which
were unusually large, showed that these were old specimens.

NEREIS PELAGICA. 279

On the 28th March three Heteronereids were procured at the margin of the beach,
near high water-mark. They swam swiftly and gracefully through the water. No
sperms could be observed in the vessel, which contained a large quantity of water.

In contrast with the condition in this species it is interesting to find that in the
Japanese Geratocephale osawai of Izuka, 1 the epitokous form presents no marked change
in the structure of the feet though they increase in size, and develop swimming bristles.
Moreover, a considerable region posteriorly shrivels even before swarming, and is cast
off by the pelagic anterior moiety, which discharges its sexual elements by the ruptured
posterior end. The distension of the anterior region of the epitokous Nereis pelagica.
and the more slender and mobile posterior region with its modified feet thus foreshadows
the condition in the former.

Habits. — Like other Nereids this species in confinement forms tough tubes of
secretion on the sides of vessels, and the same habit occurs in the early stages of the
epitokous condition.

Development. — The development of Nereis pelagica is still in need of further elucida-
tion, though it probably closely corresponds with that of other forms, such as N
Dumerilii and N. diversicolor — indeed, the very early stages may have been confounded
with these. In the bottom -net at the beginning of November many small ova of a dull
yellowish hue, and larvae of the same tint rotating by aid of cilia, are captured, but
their relationships are doubtful. By transmitted light the larvse are yellowish green.
Externally are more minute cells, within are globules of yolk.

A young Nereis pelagica of forty bristled segments and measuring 8 mm. was
obtained at Lochmaddy in August. In this the tentacles are larger in proportion
than in the adult, whilst the tentacular cirri are shorter. The longest (in spirit)
extends to the fourth segment. The eyes are wider apart and have the long axis
longitudinal. The bristles, though small, and with indistinct camerse, appear to follow the
normal type. This is probably the young of the year, just as another procured at St.
Andrews in September probably is. The latter has forty-four bristled segments. The
eyes are more rounded. Attached to the twenty-eighth foot on the right is a parasitic
body of a clavate outline (like a Demo&ex) with traces of segmentation and apparently a
chitinous cuticle (Plate LXXI, fig. 7 i).

An older form with fifty-seven bristled segments was procured at St. Andrews in
May and it probably represents the young of the previous year, as also does a somewhat
larger form brought from deep water the same month. The lobes of the feet in these
are proportionally more elongate than in the adult.

Similar young in tubes are common in deep water and they show long tips to the
heterogomph bristles.

At Lochmaddy in August an example with sixty-three bristled segments was found,
measuring about 1 in. in spirit. Such may have been hatched very early the same year,
but this diagnosis is uncertain, for it is possible that it may have belonged to the
previous year.

1 ' Journ. Coll. Sci. Univ. Tokyo/ vol. xvii, art. ii, 1903; also ' Ann. Nat. Hist./ ser. 7, vol. xv,
p. 35. I am indebted to the author for excellent examples of this form.

280 NEREIS CULTRIFERA.

Buscli 1 (1851) figures a larval Nereid with four eyes and six bristled segments, the
posterior end having two short caudal cirri, but its identity with this species is
problematical.

In the British Museum this widely distributed species has various titles, probably in
some cases from confusing preparations and their bottles.

Baster 2 (1759) gives a fairly good figure of this species, with an enlarged drawing
of a foot. He considered the marine Nereids as the analogues of the Centipedes
on land.

So far as observed the Nereis fimbriata of 0. F. Miiller, Dr. Johnston, and others is
only a deep-water variety, very common on the eastern shores, with somewhat longer
cirri and more pointed lobes to the feet. 0. F. Miiller noticed that this form, his N.
verrucosa, readily reproduced lost parts.

This is probably the species figured by Montagu (MS. Drawings, Linn. Soc, Plate
VIII, fig. 2, 1808) as Nereis pulsatoria.

Malmgren 3 showed that Heteronereis grandifolia, H. Rathke, was the male epitokous
condition of N. pelagica.

Ehlers (1868) points out that Lyeoris margaritacea and Nereis fimbriata of Dr.
Johnston belong to this species, and so with the Nereis fidg ens of Dalyell. It is doubtful
if this author's Nereis procera from the Gulf of Georgia is other than a variety of
N. pelagica, and so with the same form mentioned by Langerhans from Madeira.

Marenzeller 4 (1889) distinguishes Nereis zonata, Malmgren, from this species by the
colour, by the fact that only a single tooth occurs in Group I of the paragnathi, seldom
two, a single tooth in V, whilst in Group VI there are from five to ten, generally seven,
whereas only three or four are present in N. pelagica. The bristles likewise differ, and
whilst N. pelagica is rare in Spitzbergen, N. zonata is common.

Cunningham 5 (1888) states : " In N. pelagica I find (neural) canals placed in the
positions ascribed by Mcintosh to those of N. virens. There is one dorsal median in
the fibrous partition between the canals, a pair corresponding to the typical pair of
Sigalion, and another pair consisting of one on the external border of each cord."

Biedemann (1891) gives a minute account of the nervous system of this species, and
he shows large neural canals (giant fibres).

2. Nereis cultrieeha, Grube, 1840. Plate LII, fig. 3; Plate LX, figs. 7 and 7a — head;
Plate LXXI, figs. 8-8 cl— feet; Plate LXXII, figs. 1-1 c— epitokous feet;
Plate LXXXI, figs. 1 and 1 a — bristles.

Specific Characters. — A larger and less rounded species than the foregoing. Head
with a rectangular posterior part on which are the four large eyes often obscured by the
mottled pigment. The anterior part is furrowed and bears two short tapering tentacles.

1 ' Beobach. Anat. Entwickel. Wirb. Seeth./ Taf. ix, figs. 11 and 12, 1851.

2 ' Opuscula Subseciva/ Tom. ii, Lib. ii, Taf. vi, fig. 6.

3 < Nord. Hafs- Annul./ p. 106, 1865.

4 ' Arch. f. Naturges./ lv, p. 128.

5 Loc. cit v p. 270.

NEREIS CULTRIFERA. 281

Palpi massive, the distal process encircled by a band of brown and with a pale tip, fused
to the anterior region of the prostominm almost to the bases of the tentacles. The dorsal
tentacular cirri of the second pair are nearly twice as long as those of the anterior pair.
Peristomial segment narrower than in Nereis pelagica.

Body 6 — 12 ins., greenish anteriorly, pale brownish in the middle and again greenish
posteriorly, with the sinuous red or purplish blood-vessel branching in each segment, and
an orange or golden iridescent hue exists on each side of the vessel, whilst the feet are
greenish. Posterior part of each segment with a greenish-black bar, and another exists
anteriorly. Whitish grains on every segment after the anterior fourth, and a distinct
white patch at the base of the caudal cirri. Ventral surface pale and iridescent.

The proboscis in extrusion has dark brown horny jaws with about five teeth.
Dorsally behind these are a single median tooth (I) and two lateral crescents (II) of
paragnathi, which are less developed than in Nereis pelagica . Veritrally this maxillary
region has two lappet-shaped patches or crescents of similar small teeth (IV), the
convexity being inward. In the middle line between the foregoing is a group (III) of
about six small teeth. In the basal division of the organ ventrally is a belt formed of a
conspicuous double row of isolated paragnathi (VII and VIII), the size in each row being
nearly equal, though, if irregularity occurs, the smaller are in the posterior row. Dorsally
the basal region has a transversely elongated tooth of a bluntly conical shape on an
elevation on each side (VI), whilst between and rather behind them are a median and
tw T o lateral teeth (V).

First foot has a greater development of the dorsal and ventral lobes than in
N. pelagica, whilst the setigerous lobe is less. Feet generally with longer and more
pointed lobes than in N. pelagica, but the dorsal cirrus is rather shorter. Bristles of
two kinds, viz., those with long, slender, tapering tips (homogomph), and secondly
those with short tips (heterogomph), which have a more decided curve at the base,
and the spinous region is shorter. The strias are less regular than in N pelagica.
Toward the tip of the tail the dorsal region of the foot becomes much enlarged.
Claparecle 1 observes that this and the second foot have no spines, a feature formerly
mentioned by Rathke 2 in connection with Nereis piilsatoria. Whilst no spine occurs
in the first foot of the British examples examined, a spine is present in the second
foot.

Epitohous Form. — Eyes large and convex, almost confluent on each side, and with
lenses. Anterior (nereid) region has nineteen feet ; the succeeding region has about
seventy-two feet furnished with swimming bristles and lamellae. Dorsal and ventral
cirri of the first seven feet as in N. pelagica, only the enlargement is slight in the ventral,
and it ceases sooner. In the fully developed pelagic foot the dorsal cirrus in the male
has seven or eight papillae on its lower border. An accessory fan-shaped lamella at the
base of the lobe beneath the superior setigerous process. Ventral lobe with a spur at its
base. An accessory small lamella in addition to the two at the ventral cirrus.

1 ' Annel. Nap./ p. 163.

2 ' De Bopyro et Nereide/

93

282 NEREIS CULTRIFERA.

Synonyms.

1761. Nereis ceruloea, Linnaeus. Fauna Suec, ed. 2, p. 508.

1787. „ „ Pennant. Brit. Zool., iv, p. 47.

1791. „ coerulea, Linneeus. Syst. Nat. (G-melin), ed. 13, i, pt. 6, p. 3117.

1806. „ ceruloea, Turton's Linnaeus, iv, p. 88.

1807. „ coerulea, idem. Brit. Fauna, p. 135.

1812. „ ceruloea, Pennant. Brit. Zool., ed. 4, p. 93.

1817. „ coerulea, Stewart. Elements, i, p. 390.

1826. Lycoris lobulata, Risso. Hist. Nat. Europ., iv, p. 416.

1840. Nereis cultrifera, Grube. Actin. Ecliinod. u. Wiirm., p. 74, f. 6.

1849. „ margaritacea (Leacli). M. Edw., Cuvier's Regne Anim., pi. xii, f. 1.

1851. „ cultrifera, Grube. Fam. Annel., pp. 49 and 126.

„ „ lobata, idem. Ibid., pp. 50 and 127.

1862. „ Beaucoudrayi, Keferstein. Zeitschr. f. wiss. Zool., Bd. xii, p. 94, Taf. viii, f. 1 — 6, 12.

1864. „ cultrifera, Grube. Insel Lussin, p. 81.

1865. „ coerulea, Johnston. Cat. Worms Brit. Mus., pp. 154 (excl. syn.) and 341.
„ Heteronereis lobulata, idem. Ibid., p. 161 (excl. syn.).

fulva (Sav.), De Quatrefages. Annel., i, p. 507.

ventilabrum, idem. Ibid., p. 517.

cultrifera, idem. Ibid., p. 527.

bilineata, idem. Ibid., p. 535 (excl. syn. Johnst.).

incerta, idem. Ibid., pi. vii, f. 3—10. Explic. des PI., p. 12.

Beaucoudrayi, idem. Ibid., p. 533.

viridis, idem. Ibid., p. 539.

1867. Lipephile margaritacea (M. Edw.), Malmgren. Annul. Polych., p. 50.
,, Hedyle lobulata, idem. Ibid., p. 58, forma epitoca.

1868. Nereis cultrifera, Ehlers. Borstenw., p. 461, Taf. xviii, f. 31—37, Taf. xix, Taf. xx ; f. 1—3,

Taf. xxi, f. 31—36.
1870. „ [Lipephile) cultrifera, Claparede. Annel. Nap. Suppl., p. 75, pi. vii, f. 1.

1874. „ cultrifera, Mcintosh. Ann. Nat. Hist., ser. 4, vol. xiv, p. 199.
„ ,, coerulea, Malm. Goteb. vet. o. vitt. Handl., xiv, p. 83.

1875. „ cultrifera, Mcintosh.. Invert, and Fishes St. Andrews, p. 122.

„ „ „ Marion and Bobretzky. Ann. Sc. Nat., 6 e ser., t. ii, p. 16.
1879. Perinereis cultrifera, Langerhans. Zeitsclir. f. wiss. Zool., xxxiii, p. 289.

1881. „ ,, idem. Canar. Annel. in Nova Acta Leop., p. 110.

1883. Nereis „ Levinsen. Vidensk. Meddel. Foren. Kjobenh., p. 236.

1885. „ „ Cams. Fauna Medit., p. 218.

1886-7. „ „ Meyer. Mitth. Zool. Stat. Neap., vii, p. 740, pi. xxvii, f. 1—8 (young).

1887. „ „ (development), Salensky. Arch. Biol. Belg., vi, p. 589.

1889. ,, „ Horst. Contrib. from Leyden Mus., t. xi, p. 162.

1890. Lipephile „ Giard. Bull. Sc. Fr. et Belg., t. xxii, p. 77.
„ „ „ Malaquin. Annel. Boulon, p. 31.

1892. Nereis „ (eye), Andrews. Journ. Morph., p. 179.

1895. Lipephile „ De St. Joseph. Ann. Sc. Nat., 7 e ser., t. xx, p. 215.

1896. Nereis margaritacea, Roule. Camp. f Caudan/ p. 449.

1898. Perinereis cultrifera, De St. Joseph. Ann. Sc. Nat., 8 e ser., t. v, p. 317, pi. xvii, f. 113, 114,

and pi. xviii, f. 115, 116.

NEREIS CULTRIFERA. 283

1902. Nereis cultrifera, Mcintosh. Ann. Nat. Hist., ser. 7, vol. x, p. 256.

1904. „

1905. „

1906. Lipephile
„ Perinereis

,, Nereis
1909. „

Allen. Journ. M. B. A., N.s., vol. vii, p. 224.
Graeffe. Arbeit. Zool. Stat. Triest, xv ; p. 322.
(movements), Bohn. Ann. Sc. Nat., 9 e ser., t. iii, p. 80.
De St. Joseph. Ann. Sc. Nat., 9 e ser., t. iii, p. 221.
Eisig". Fauna u. Fl. Neap., xxviii, p. 207.
Elwes. Journ. M. B. A., n.s., vol. viii, p. 352.

Heteronereid Form.

1825.

Nereis

1828.

>)

1833.

))

>)

>j

1834.

)>

1837.

Lycoris

1851.

Nereis

1820. Lycoris lobulata, Savigny. Syst. de Annel., p. 30.

„ De Blainville. Diet. Sc. Nat., xxxiv, p. 430.

„ idem. Ibid., lvii, p. 468.

„ Audouin and Edwards. Ann. Sc. Nat., t. xxix, p. 123, pi. xiii, f. 7, 8.

„ idem. Ibid., p. 124.

„ idem. Annel. da lit. France, ii, p. 191, pi, ivA, f. 7 and 8.

„ Rathke. Beitr. z. Fauna der Krym. in Mem. fAcad. St. Petersb., t. iii, p. 415,

Taf. vii, f. 2, 9—15.
, Grube. Fam. Annel., pp. 50 and 127.

1865. Heteronereid lobulata, Johnston. Cat. Worms Brit. Mus., pp. 161 and 341 (partim).
„ Nereilepas lobidatus, De Quatrefages. Annel., i, p. 560.
„ Hedyle lobulata, Malmgren. Nord. Hafs-Annul., p. 182.

1867. „ „ idem. Annul. Polych., p. 58.

1868. „ „ Ehlers. Borstenw., ut antea.

1870. Nereid (Lipephile) cultrifera, Claparede. Suppl. Annel. Nap., p. 75, pi. vii, f. 1.
1898. Perinereis cultrifera, De St. Joseph. Ann. Sc. Nat., 8 e ser., tome v, p. 318.

Habitat. — Between tide-marks on both eastern and western shores of Britain (from
Shetland to the Channel Islands), under stones, and in tunnels in masses of peat.
Extends also to both shores of Ireland, especially the western, as at Dingle Bay and
Valentia Harbour (J. G. Jeffreys).

Ranges to the Mediterranean, shores of France (De St. Joseph), Canaries (Langerhans),
Black Sea (Bobretzky).

Head (Plate LX, fig. 7) with a rectangular posterior region on which are the four
large eyes, the posterior pair often rendered obscure by the dark mottled pigment.
The anterior eyes are wider apart. The front part of the head is bluntly conical, the
truncated anterior region bearing two short tapering tentacles, which are somewhat less
flattened than in N. pelagica. Moreover, a marked median furrow, often boldly indicated
by the arrangement of pigment, passes along the centre of this region, leaving two lateral
fillets. The palpi are massive, the distal papillae encircled by a band of brown at the
base and with a pale tip. The inner border of each palpus is fused to the conical anterior
region of the prostomium almost to the bases of the anterior tentacles, a diagnostic
feature in comparing with N. pelagica. The two pairs of tentacular cirri are similarly
situated to those of the latter species, the dorsal being the longer in each case. The
dorsal of the second pair is nearly twice as long as that of the anterior pair.

Body proportionally more elongate than in N. pelagica, some nearly reaching a foot
in length, very little tapered anteriorly, but much more so posteriorly, where it terminates
in two long slender cirri. The peristomial segment is broader than the succeeding,

284 NEREIS CITLTRIFERA.

but considerably narrower than the corresponding segment in N. pelagica. The body
is greenish anteriorly, pale brownish in the middle and again greenish posteriorly. A
deep purplish blue line extends along the dorsum, with the sinuous dark red or purplish
blood-vessel branching in each segment and with long twigs directed backward. An
orange or golden iridescent hue exists on each side of the vessel, whilst the bases of the
feet and these organs themselves are generally greenish. The posterior part of each
segment has a greenish-black bar extending quite across the junction, and in addition
another on each side anteriorly — formed, as it were, of a single bar separated by the
dorsal blood-vessel. These markings become indistinct when the brownish median hue
takes the place of the anterior greenish. Moreover, in the latter region an oblique
(longitudinal) whitish streak occurs on each side in every segment, and whitish grains
on the segments behind the anterior fourth. A distinct white patch is on the tail at
the base of the cirri. The ventral surface is pale and iridescent. The inferior edge of
the foot is the most vascular, whilst a series of bright red transverse vessels passes from
the posterior border of each towards the interspace. The ventral vessels are of a much
brighter red than the dorsal.

After preservation it is somewhat duskier than Nereis pelagica, and the head and
anterior segments have small whitish specks. A pale central patch also occurs at the
anterior border of several of these.

Proboscis. — Dorsally (Plate LX, fig. 7) the strongly curved jaws present about five
teeth, and they vary in position, in some the left and in others the right passing
above the other. In extrusion the basal or proximal division of the organ has an eleva-
tion on each side with a transversely elongated tooth of a bluntly conical shape, whilst
between and slightly behind are a median and two lateral teeth, the former being slightly
in front. In the maxillary (distal) division of the proboscis a single median tooth
occupies the central region dorsally, whilst two crescents occur laterally, one on each side.
These are more feebly developed than in N. iielagica. Ventrally (Plate LX, fig. 7 a) two
lappet-shaped patches or crescents of similar small teeth occur in a corresponding
position, the convexity being inward. In the middle line between the foregoing is a
group of about six small teeth. In the basal division of the organ ventrally is a belt
formed of a double row of well-marked isolated teeth, the size of these in the rows being
nearly equal, though if irregularity occurs in this respect, the smaller teeth are found in
the posterior row in extrusion. There is thus a great difference between the single row
of large teeth and the "dusting" of smaller prickles as in Nereis pelagica, Some of the
large examples have numerous calcareous deposits on the proboscis.

Specimens frequenting tunnels in peat had stalks of Algge in the hind-gut, and in
others from the Channel Islands the fascal masses were enveloped in mucus, and consisted
of delicately branched pale Algas with muddy debris. The central parts of the filaments
of the Alga3 had been acted on by the gut, leaving only the cellulose exterior.

The perivisceral or coelomic corpuscles are translucent bodies, spindle-shaped or
elliptical in outline, and varying in size (December). Free nuclei lay near or on them.
The larger elliptical corpuscles show minute granules, and free granules (minute nuclei ?)
floated in the fluid. About a week later during very cold weather a female which had
been isolated showed its perivisceral fluid so loaded with small rounded corpuscles as to

NEEEIS CTJLTBIFERA. 285

be slightly opaque to the naked eye. In the fluid were also many of the larger cells with
coarsely granular contents. A few of the elongated coelomic corpuscles were still visible.
There were also a few ova. Such a condition raises the question as to the changes in the
fluid after confinement. In a dying example the fluid abounded with disintegrating cells
and granules, but the perivisceral corpuscles proper were few. There were also groups
of circular pellucid cells, and ova with nuclei and nucleoli, surrounded by the smaller cells.
Some of the bodies showed a punctate zone, which may have been connected with
Gregarinee, having a thickened anterior end. It is a large broad form. The fatty globules
in the eggs of the annelid are remarkably large, whereas the granules in the parasitic (P)
bodies are small. De St. Joseph 1 found gregarines in other forms, such as N. Biimerilii.

The first foot (Plate LXXI, fig. 8) is distinguished from that of N. pelagica by the
greater development of the upper and lower lobes and the rudimentary condition of the
setigerous lobe. The dorsal edge of the foot is more prominent, but the cirrus is some-
what shorter. The dorsal lobe is larger and more pointed, and the same may be said of
the inferior lobe, which is, however, less acute. The setigerous or median lobe offers a
marked contrast to the corresponding part in N. pelagica, since it is much less prominent.
The ends of the upper long-tipped homogomph bristles are shorter, whilst those of the
lower series are somewhat longer. The ventral cirrus is shorter. Both species have
black spines.

At the tenth foot (Plate LXXI, fig. 8 a) the dorsal cirrus is shorter, the upper lobe
more triangular in lateral view, and the distance between the dorsal edge and the first
bristle-bundle greater than in N. pelagica. Moreover, a small but distinct superior
setigerous lobe is present — none being visible in N. pelagica. The inferior setigerous
lobe is less acute than in the latter, and the ventral cirrus is less in proportion to the
bulk of the foot, and it extends outward only to the middle of the great ventral lobe.
The bristles of the dorsal process have long, slender, serrated tips, as in N. pelagica, but
the transverse strias of their shafts are bolder and less regular (Plate LXXXI, fig. 1 a).
The bristles with the shorter tips in the inferior division (Plate LXXXI, fig. 1), while
somewhat similar to those of N. pelagica, have a more decided curve at the base of the
terminal piece, and the spinous region, like the whole structure, is shorter. The trans-
verse lines or cameras in the centre of the shaft are less regular than in N. pelagica. The
tip in both shows a slight differentiation, indicated by a line within the margin.

In the thirty- seventh foot (Plate LXXI, fig. 8 b) the two upper lobes are proportion-
ally longer than in N". pelagica, and they have a wider space between them, since a minute
spinigerous process separates them. The dorsal cirrus is considerably shorter and
smaller. The inferior setigerous lobe is somewhat less, while the ventral lobe is more
deeply cleft superiorly, and thus appears to be longer. The ventral cirrus, like the
dorsal, is both shorter and smaller than in N. pelagica.

At the fifty-seventh foot the enlargement of the dorsal region is even more marked,
and the cirrus is carried further outward. In shape, however, the dorsal lobe and all the
others conform to the condition just described.

Towards the tip of the tail (Plate LXXI, fig. 8 c) the enlargement of the dorsal
region has altered the shape of the foot, since the dorsal lobe in lateral view occupies

1 ' Ann. Sc. Nat./ 7 C sei\, t. xx, p. 215.

286 NEREIS CULTRIFERA.

half the depth. Externally it terminates in an acnte tip, and the slightly elongated
dorsal cirrus is carried so far outward on the ridge of the flattened lamella that it extends
considerably beyond the dorsal lobe. The upper setigerous lobe is now indistinguishable
in the sulcus between this and the next lobe, the reduction in the size of which makes the
disproportion between it and the dorsal lobe very conspicuous when contrasted with the
condition in the fifty-seventh foot. The inferior setigerous lobe is reduced in size, and
the ventral lobe is so small that it projects little beyond the tip of the former. The
ventral cirrus is considerably smaller, and its base less prominent, while its tip goes little
further than the base of the ventral lobe.

This species is often infested in Guernsey and Herm by the crustacean parasite
Nereicola ovata, Keferstein, 1 which generally occurs towards the posterior end of the
annelid, the largest example of which had about fifteen specimens, most of them, however,
being small. The usual number is four or five. They adhere firmly either to the groove
between two feet or to the sides of the feet, but they do not seem to incommode the
worms, and it may be supposed that the Nereis, if so inclined, can at any time remove
them. It is, however, by no means an active species.

The male Nereicola (Fig. 66) is small and has a somewhat ovate carapace, which,
however, diminishes posteriorly. It has in front minute chitinous processes like hairs,
and a pair of antennules composed of four segments uniformly diminished from base to
apex. The edges of the last three, under pressure, have minute projections from which
the setaa spring. The tip of the organ has several setae which are longer than itself.
Behind these is a more cylindrical pair of antennas of the same number of segments, the
tip having five long setae, one of which surpasses the others in length. The mouth has
two pairs of appendages, probably representing mandibles and maxillae, the posterior
having its terminal process furnished with a single strong claw. Three appendages occur
on the ventral surface of the carapace behind the foregoing, the first two being biramous
limbs attached to a basal process, the third a rudimentary one in the form of a simple
process. The testes form a pair of symmetrical organs posteriorly, the narrowed region
behind being terminated by two styles.

The young female (Fig. 68) has an ovoid outline, with a blunt anterior and a narrower
posterior region, and both antennae and caudal styles are visible from the dorsal surface.
In the fully developed condition (Fig. 67) the thoracic region is greatly enlarged, so that
dorsally the outline is rounded ovate, with a narrower region in front, and it is only in a
lateral (Fig. 69) or ventral view that the snout and tail are fully seen. The antennae are
similar to those of the male, as also are the appendages of the mouth and the carapace
posteriorly. The ovisacs are attached by slender pedicles to slight prominences on each
side, and form cylindrical pouches filled with a vast number of ova. The same form was
found by Grube at St. Vaast-la-Hougue, 2 and by the author 3 in the Channel Islands.

The segmental organ of the annelid has a trumpet with somewhat short fimbriae or
lobes and long cilia (Goodrich).

1 'Zeitschr. f. wiss. Zool./ Bd. xii, p. 461, Taf. xlii, f. 1—4, 1860.

3 Grube termed it Nereidicola ovata in his ' Notes on the Fauna of St. Vaast-la-Hougue/ and he
gave two good figures.

3 ' Quart. Journ. Micros. Sci./ vol. 10, n.s., p. 89, pi. v.

NEREIS CITLTRIFERA.

287

Reproduction. — An almost ripe male was observed on July 1st, and as no alteration
of the feet was visible the epitokons condition wonld appear to be suddenly developed.

In a male towards the end of November the reproductive elements form large
compound cells with many nuclei, and they vary much in size. Most are more or less

Fig. 66.

Fig. 67.

Fig. 68.

Fig. 69.

/

Fig. 66. — Nereicola ovata, male, as a transparent object. Fig. 67. — Female from the dorsum. Fig. 68. — Young
female from the dorsum. Fig. 69. — The adult female in lateral view. All masfnified.

rounded. Besides these are translucent spheres and groups of smaller cells with nuclei —
apparently having escaped by rupture from the larger corpuscles. They measure about
•00021 in., the smallest being one sixth of that size. There are also elongated transparent
bodies. When breaking up, the smaller corpuscles show coarse granules, and probably
these pertain to the wall of the alimentary canal.

288 NEREIS OULTEIPEEA.

On the 18th of January the males, which are smaller than the females, have the
coelom laden with sperms, which are not very active, though they exhibit slight move-
ments. The " head " has a knob followed by a somewhat ovoid region terminating in
the filament posteriorly. The under surface of the worm is pale anteriorly, reddish from
the blood-vessels posteriorly.

On 6th December some of the females are laden with ova, which when extruded
in mass have a dull or faintly greenish aspect. The ova are surrounded by a coating of
yellowish cells filled with refracting granules, a coarser series also filling the ova,
which present a large nucleus and nucleolus. On the 18th of January the ova are still
numerous in the perivisceral cavity, and the nucleus and nucleolus can be made out
under a lens.

The only Heteronereis pertaining to this species comes from Ireland, the exact
locality not being indicated. The specimen, a male, is between two and three inches
(two inches in spirit). The eyes are large and convex, the anterior almost touching the
posterior on each side. They are chiefly developed laterally, and have a small lens in the
centre of each. The pigment of the dorsum of the body seems to have been specially
developed in bands and touches across the middle of each segment. The anterior region
has nineteen feet — distinguished as usual by their greater thickness. The succeeding
region has seventy-two feet, furnished with the swimming bristles and lamella. The
last, however, are very small. The vent is everted and richly papillose, and two long
cirri project posteriorly. The segments of the anterior region are wider, those of the
posterior narrow, but the sulci between the segments are in both transverse. The
paragnathi of the proboscis do not appear to differ from the typical form. The first
foot has its dorsal cirrus enlarged, as in N. pelagica, and a slight increase has also
occurred in the ventral. This enlargement continues to the seventh foot as regards
the dorsal cirrus, but it ceases sooner in the ventral. The feet remain of the
normal structure till the seventeenth, in which the dorsal cirrus has a papilla at
its ventral border near the tip, this marking off the filiform termination, and the
ventral cirrus of the same foot presents a swelling near its tip. The dorsal cirri of
the eighteenth and nineteenth segments show one or two additional papilla?, and the
terminal region is often bent at an angle. The tips of the ventral cirri of the eighteenth
and nineteenth are as in the seventeenth, but the nineteenth (Plate LXXI, fig. 8 d)
has a small process (lamella) arising on the inner side of the base of the cirrus, and a
more minute one to its exterior. A small lamella also appears at the lower edge of the
tip of the inferior setigerous process. An opaque granular mass (gland ?) occurs at the
base of the dorsal cirrus, and the tips of the five lobes of the foot and the base of the
ventral cirrus are all similarly opaque. The bristles of this foot retain the normal outline.

The heteronereid condition is more distinctly pronounced in the next (twentieth)
foot (Plate LXXII, fig. 1), which presents a small vertical lamella internal to the base
of the dorsal cirrus. The latter has four papillae and traces of two others below the
filiform tip. The inferior setigerous process has a considerable fan-shaped lamella with
a pit in the middle at the tip, as if from the coalescence of two folds. The lamellas at the
base of the ventral cirrus have increased, and the cirrus comes from the edge. The
margins of the second lobe and of the ventral also show traces of increase. A few bristles
of the atokous type remain in the superior setigerous lobe, but the ends of the shafts and

NEREIS CULTRIFEEA. 289

the broad tips of the swimming bristles project in a dense mass from the foot, showing
that they are rapidly developing. None are yet seen in the inferior setigerous process.

The pelagic foot is fully developed behind the foregoing, and at the thirty- seventh
foot (Plate LXXII, fig. 1 a) the parts perhaps attain their maximum. A large fan-like
lamella is internal to the dorsal cirrus, which shows seven or eight papillas on its lower
border. A small lamella projects at its base externally. The dorsal lamella is long,
thin, and tongue-shaped. The superior setigerous process has a lamellar fan of a semi-
circular outline in front of the dense group of swimming bristles. The next lobe of the
foot forms a large and lanceolate lamella, with an accompanying fan-shaped flap depending
from its base. A great fan-like lamella extends above, below, and beyond the swimming
bristles of the inferior setigerous process. The ventral lobe has a marked spur at its
base superiorly, A considerable fan-shaped lamella occupies the ventral (internal) region
of the ventral cirrus, whilst a narrow lamella externally and an accessory smaller one are
near the base on the same side. The cirrus projects from the edge of the lamella, an
opaque granular band connecting it with the body.

The condition at the fifty-seventh foot of the same epitokous male is shown in Plate
LXXII, fig. 1 b, and near the tip of the tail in fig. 1 c, the former presenting in proportion
to its size very large foliaceous lamella?-, the latter showing considerable diminution in all
the processes.

c — =3aj^asG>

»©fc=^

Fig. 70. — Long spear-like instrument used by the fishermen of Guernsey in digging for Nereis cultrifera. 1

These annelids form a favourite bait at various parts of the coast, especially in the
south. Thus the fishermen of St. Peter Port, Guernsey, dig for them with pointed
instruments resembling spears (Fig. 70) and keep them in vessels amongst a little sand
and seaweed. They are often used for capturing whiting, which again are cut into
portions for the capture of conger. They form a very successful bait, and in these rich
waters various species of wrasse, besides bream, whiting-pout, various species of blenny,
and other fishes are readily captured. *■

The Nereis megodon of De Quatrefages (1865) has certain resemblances to this species,
but it is difficult, with the materials at hand, to ascertain its precise relationships.

Ehlers (1868) gives a detailed account of the anatomy of this species as the type of
the family, and rectifies the synonymy.

Claparede (1868) found this Nereid common at Naples, where it is known by the
fishermen as esca. He was sure it was known to Delle Chiaje, but it is difficult to make
certain either in regard to figures or description. Possibly it may be his Nereis Rauzani.
He is doubtful whether Ehlers' view that the N. margaritacea of M. Edwards is this form
is correct, basing his opinion on the fact that while the figures in the ' Pegne Animal ' closely
resemble it, these were subsequently referred by De Quatrefages to N incerta, which
differs in paragnathi, and whereas the Neapolitan examples had only sixty-five to seventy
segments those of De Quatrefages had 170. Subsequently (1870) he gave special attention
to the heteronereid form, showing that the change is accompanied by the absorption of the
1 For the use of tins block the author is indebted to Messrs. Taylor and Francis.

94

Ventral division

290 NEREIS CULTRIFERA.

peritoneal pigment in the middle and posterior regions, thus giving a pinkish hue to
the animal from its vascularity. In the female the greenish coloration exists in the first
twenty segments, the abdominal region being pale, variegated with black. He thinks the
male heteronereids are smaller. The spermatozoa are similar to those of JV. DumeriliL
The eggs are colourless, and 0-15 mm. in diameter. He found the metamorphosis in progress
in specimens about the end of March at Naples, but they were rare. The sexual elements
are developed in individuals of large size— apparently before metamorphosis.

Claparede (1870) states that the following arrangement of the bristles is constant:—
Superior division . Homogomph with long tips.

' Superior . • Ditto, falcate heterogomph.

Inferior . . • Homogomph with long tips,

falcate heterogomph.

Bhlers again avers that there are no falcate bristles in the inferior groups of the
ventral division, but there seems to be a difference of opinion as to what is a falcate
bristle. So far as general description goes that of Claparede holds.

Grube (1869) gave a description and figure of Nereidicola bipartia which he had
found parasitic on the feet of this species at St. Vaast-la-Hougue. This is probably the
same as Keferstein's Nereicola ovata.

Ed. Meyer (1886 — 7) refers to a young form of this species having six bristled segments,
and especially to the development of the segmental organs and other structural features.

In the collection of the British Museum this form is contained in bottles under the
names, e.g., Nereis ceruloea, Nereis brevimana, and Nereis pelagica.

De St. Joseph 1 found considerable variation in the groups of paragnathi. Those least
variable were the two cultriform paragnathi on the dorsum of the basal segment, and the
double row on the ventral surface of the same region.

Benham 2 mentions that the species is known by the name of the "Red Cat" in the south.

Schepotieff gives in this form a careful account of the bristle- sacs, with the developing
bristles. 3

Claparede 4 states that the foot-glands and the dorsal muciparous glands increase in
volume at the period of sexual maturity. Langerhans met with an epitokous female at
Madeira.

On the shores near Cherbourg, M. Fauvel found the heteronereid in April and May.

De St. Joseph gives a careful account of both sexes in the heteronereid condition. In
the female the large eyes are almost coalescent, the head and four anterior segments
being dull green. In the first six segments following the buccal, the cylindrical dorsal
cirrus is abruptly narrowed towards its extremity, and a similar condition occurs in the
shorter ventral cirrus. The heteronereid condition of the feet commences at the twenty-
second bristled segment.

Lo Bianco 5 found JV. cultrifera mature and depositing ova at Naples from May to
June, the yellowish ova being enveloped in mucus on Algaa dredged at some depth.

1 'Aim. Sc. Nat./ 7 e ser., xx, p. 215 ; 1895.

2 f Camb. Nat. Hist./ ii, p. 316.

3 ' Zeitschr. f. wiss. Zool./ Bd. lxxvi, p. 589, Taf. xxvi, and Taf. xxvii, figs. 1—4, 1904.

4 ' Annel. Nap./ p. 164, 1868.

5 ' Mitth. Zool. Stat. Neap./ xiii, p. 488, 1898.

NEREIS SCHMAED^EI. 291

3. Nereis Sohmard^i, Be Quatrefages, 1865. Plate L, fig. 17; Plate LX, fig. 8 — head;
Plate LXXII, figs. 2-2 b— feet; Plate LXXXI, figs. 2-2 c— bristles.

Specific Characters. — Head typical in shape. Eyes large and furnished with lenses,
anterior pair oval, posterior circular. Tentacles rather stout and short. Palpi with
somewhat slender terminal joints, a ring of pale madder-brown occurring within the tip.
Tentacular cirri rather long. A madder-brown patch like a crown occurs in front, with
a longitudinal median streak, and behind it a brownish patch with a pale area on each
side of the posterior eyes. Body 4 — 5 ins. long, more definitely tapered anteriorly than
usual. Of a pale flesh colour throughout, the anterior third toning off to pale yellow,
and dull whitish posteriorly. The palpi, peristomial segment, and a considerable number
of the anterior segments are flecked with pale madder-brown touches or specks irregularly
scattered over each segment, and, after they decline, whitish touches occur at the
segment-junctions in the line of the blood-vessel, and posteriorly a double yellowish-
white streak occupies the mid-dorsal region of each segment. About the tenth bristled
segment a differentiated region is found near the base of each foot in the form of two or
three pale brownish dots, and as the body becomes paler the region is recognized by a
pale, transversely elongated area on each side at the base of the foot, and a minute black
speck often lies at its inner side. These lateral white patches continue within a short
distance of the tip of the tail. Proboscis in extrusion has a pair of horny maxillae with
twelve to thirteen denticulations. Dorsally the maxillary segment has a row of
paragnathi (II) at the bases of the jaws. Ventrally at the base of each maxilla are a
crescent and a spur of larger paragnathi (IV), with a group of smaller at the posterior
angle between crescent and spur. Dorsally in front of the palpi is on each side a group
of small paragnathi (VI) in two rows. The basal division has ventrally a long row of
paragnathi in lozenges (VII and VIII). As a rule a larger one is in front near the angle
of the lozenge and a row of smaller behind. The first foot is short, a brief space inter-
vening between the body and the dorsal cirrus, the tip of which scarcely reaches that of
the dorsal lobe. The latter forms a conical blunt cone. Setigerous lobe beneath is
obscured by a conical lobe behind it. Ventral lobe large and tongue-shaped. The chief
feature of the anterior feet is the rounded, bulbous nature of the lobes and the short
cirri, so that at the tenth foot the parts present a bluntly papillose appearance. The
lobes become considerably longer after the thirtieth foot, and at the thirty-fifth the dorsal
cirrus reaches the end of the lanceolate dorsal lobe. The superior setigerous process has
an elongated anterior papilla soldered to it inferiorly. The lamella beneath is shorter
and broader than the dorsal. The inferior setigerous lobe is irregularly conical and bifid,
a flap being in front and the shorter lip behind. Ventral lamella short and lanceolate.
Ventral cirrus small, projecting from an elevated base. Bristles of the dorsal setigerous
lobe homogomph, as in the typical Nereids. Those of the inferior lobe at its upper edge
are similar, though the tips are shorter, but those toward the ventral border have short
tips (heterogomph).

Epitokous Form. — The nearly ripe Heteronereid distended with large eggs presents
indications of change in the thirty-second foot in the form of a rudimentary lamella

292 NEREIS SCHMARDSEI.

behind the inferior setigerons process. In full development (e.g., the sixty-fifth foot) a
dorsal fan-shaped lamella, a large lamella of similar shape behind the inferior setigerons
lobe, and a larger and a smaller lamella at the ventral cirrus characterize the foot. The
dorsal, middle, and ventral lobes of the foot are flattened and elongated, and swimming
bristles occur in both divisions.

Synonyms.

1865. Heteronereis Schmardsei, De Quatrefages. Aimel., i, p. 569, pi. vii, f. 11, and pi. ix, f. 7.

1867, Praxithea irrorata, Malmgren. Annul. Polych., p. 51 (167), Taf. iv, f. 24.

1868. Nereis rubicunda, Ehlers. Borstenw., ii, p. 529, Taf. xxi, f. 5—9.

Heteronereis Schmardsei, Grube, Mitth. iiber St. Yaast-la-Hougue (and Jahresb.), p. 126.
1870. „ „ idem. Jahresb. der Scliles. Gesells. fur 1869, p. 52 (fide Auct.).

idem. Mitth. iiber St. Malo u. Roscoff (ibid. Jahresb.), 1869—72
p. 91.

idem. Arch. f. Naturges., Bd. xxxvi, p. 312.
1874. Praxilla irrorata, Malm. Groteb. vet. o. vitt. Handl., xiv, p. 84.
1879. Heteronereis Schmardsei, Tauber. Annul. Danic., p. 98.
1883. Nereis irrorata, Leviusen. Yidensk. Meddel. Foren. Kjoben., p. 235.
1885. ,, rubicunda, Cams. Fauna Medit., p. 220.

1888. Praxithea irrorata, De St. Joseph. Ann. Sc. Nat., 7 e ser., t. v, p. 263, pi. xi, f. 131.
1895. „ „ idem. Ibid., 7 e ser., t. xx, p. 215, pi. xii, f. 33 -36, pi. xiii, £. 37—39.

1898. Nereis „ idem. Ibid., 8 e ser., t. v, p. 299, pi. xv, f. 82,

1902. „ „ Mcintosh. Ann. Nat. Hist., ser. 7, vol. x, p. 256.

1904. ,, „ Allen. Journ. M. B. A., n.s., vol. vii, p. 224.

1906. Praxithea „ Bohn. Ann. Sc. Nat., 9 e ser., t. hi, p. 83.
„ Nereis „ De St. Joseph. Ibid., 9 C ser., t. iii, p. 219.

1909. „ „ Elwes. Journ. M. B. A., n.s., vol. viii, p. 352,

Hpitokous Form.

1864. Heteronereis signata, Baird. Journ. Linn. Soc, vol. viii, p. 8, Taf. i, f. 1 (?).

1865. „ „ Johnston. Cat. Worms Brit. Mus., p. 341 (?).

Habitat. — In tubes between tide-marks, St. Peter Port, Guernsey, July, 1868. In
similar regions at Plymouth, and small specimens also dredged (Allen).

Apparently chiefly a southern form. It is not uncommon on the shores of France,
such as at St. Vaast, Roscoff, and Dinard, amidst the prairies of Zostera. It is also found
in Sweden.

Head (Plate LX, fig. 8) has the typical outline, though the palpi have somewhat
slender terminal joints, with a ring of madder-brown within the tip. Anterior eyes oval,
posterior circular. They are furnished with lenses, and in the fully developed epitokous
form De St. Joseph found them almost coalesced. Tentacles comparatively stout and
short. Tentacular cirri rather long. A curious madder-brown patch like a crown with a
median longitudinal streak occurs in front of the head, and behind it a brownish patch
with a pale area on each side of the posterior eyes.

Body 4—5 ins. long (15—18 cm., De St. Joseph), more definitely tapered anteriorly
than usual, and posteriorly terminating in two long anal cirri. The anterior third is of
a pale flesh-colour toning to pale yellow or dull whitish posteriorly. The peristomial

NEREIS SCHMABJLEL 293

segment and a considerable number of the anterior segments are flecked with pale madder-
brown touches or specks irregularly disposed, and, after they begin to decrease, whitish
touches occur on the segment- junctions in the line of the blood-vessel, and posteriorly a
double yellowish-white streak occupies the mid-dorsal line of each segment. About the
tenth bristled segment a differentiated region, indicated by two or three brownish dots,
occurs near the base of each foot, and as the body becomes paler the region is recognized
by a pale, transversely elongated area on each side at the base of the foot, often with a
minute black speck at its inner side. These lateral white patches continue almost to the
tip of the tail. De St. Joseph gives the body a brick-red colour in life.

The 'proboscis in extrusion has a pair of maxillas with twelve to thirteen denticulations,
the apex being slightly twisted. Dorsally the maxillary segment has a row (II) of para-
gnathi at the base of each maxilla, and in a corresponding position ventrally are a crescent
and a spur of longer paragnathi (IV) with a group of smaller ones at the posterior angle
betweeu crescent and spur. Dorsally in front of the palpi (basal division) is on each side
a group of small paragnathi (VI) in two rows ; whilst ventrally a long row (VII and VIII)
of paragnathi occurs in lozenges ; as a rule, a large one is in front in the angle of the
lozenge, and a row of smaller behind — somewhat similar to the condition in Nereis
pelagica, but with fewer small paragnathi.

The intestine contains debris of TJlva, diatoms, and Grammatophora (De St. Joseph).

The first foot (Plate LXXII, fig. 2) is short, with but a brief space between the
body and the dorsal cirrus, which is comparatively short and stout, its tip scarcely reaching
that of the dorsal lobe. The latter forms a massive cone blunt at the tip. The setigerous
lobe beneath is obscured by a conical lobe behind it, so that in lateral view only the tips
of the bristles are observed. The ventral lobe is large and tongue-shaped, the ventral
cirrus short and rather stout (in spirit). The bristles of this part all have tapering tips
with finely serrated edges. The third foot shows two setigerous processes as in allied
forms.

On glancing along the feet anteriorly one feature is conspicuous, viz., the rounded,
bulbous nature of the lobes ; indeed, this condition gives a distinctive character to the
species. Thus at the tenth foot (Plate LXXII, fig. 2 a) the lobes in lateral view resemble
bluntly rounded processes. Between the dorsal cirrus and the body is a prominent
rounded elevation, and the cirrus is subulate, whilst its tip does not reach the outer edge
of the lobe. The superior setigerous lobe beneath forms a low convex process, with a
dense fan of bristles, the shafts of which are deeply tinted. All have tapering serrated
tips (Plate LXXXI, fig. 2). The lobe beneath is a rounded boss, which does not project
so far outward as the dorsal. The inferior setigerous process is a blunt cone, with its
long bristles in two divisions— an upper and lower, and each with two kinds of bristles
as in the typical Nereids. The upper has those with long tapering tips superiorly, then
a series beneath with shorter tips (Plate LXXXI, Hg. 2 a). The inferior has similar
bristles at the upper edge, but those toward the ventral border have longer tips than the
corresponding series in the upper group (Plate LXXXI, fig. 2 b). The ventral lobe
recedes inward as a massive rounded boss, and the small ventral cirrus springs from an
elevation of its own.

The specimen consisted of an anterior fragment, so that the structure of the feet

294 NEREIS SCHMARDiEL

could only be examined as far as the thirty-fourth or thirty-fifth. In this region
considerable elongation of the lobes has occurred, whilst the depth of the foot from
above downward has increased. The dorsal cirrus reaches to the end of the dorsal
lobe, which is lanceolate. The superior setigerous lobe has an anterior elongated
papilla soldered to it inferiorly. The lamella beneath is shorter and broader than
the dorsal. The inferior setigerous process is irregularly conical and bifid, a flap
being in front, and the shorter tip behind. The ventral lamella is short and lanceolate.
The ventral cirrus is small and projects from an elevated boss. The bristles by-and-by,
as at the thirty-second foot, show only the series with the long tapering tips dorsally ;
whilst ventrally are (1) two upper powerful bristles and a group of those with long
tapering tips only a little shorter than the dorsal, followed (2) by a ventral series,
all of which, with the exception of the two upper, which have long tapering tips, have

falcate tips.

In an epitokous female from St. Vaast, kindly sent by the late Prof. Grube of
Breslau, and much distended with nearly ripe eggs, the changes in the feet begin about
the thirty-second and thirty-third in the form of a process or rudimentary lamella behind
the inferior setigerous lobe. At the thirty-fifth foot a small vertical lamella appears
inside the dorsal cirrus, a considerable flap projects behind the inferior setigerous process,
and the ventral cirrus has a small outer and a larger internal lamella. The parts increase
in complexity until about the sixty-fifth foot (Plate LXXII, fig. 2 b) ; the fan-shaped
lamella on the ridge of the foot forms a leaf-like process, the dorsal cirrus extending
from its outer border beyond the tip of the dorsal lamella beneath. The latter is a long
lanceolate process. A dense group of swimming bristles (Plate LXXXI, fig. 2 e) is borne
by the superior setigerous lobe with which the lamella below appears to be fused. The
inferior setigerous lobe also carries a dense group of swimming bristles and the large
foliaceous expansion behind. The spatulate ventral lamella projects from its lower
border, and the ventral cirrus has a narrow lamella at its outer and a large fan-shaped
lamella on its inner edge.

The whole foot is largely distended by the ripe eggs, which escape on rupture of the
thinned integuments. The imperfect preservation of the specimen still further adds to
the distortion of the soft tissues of the foot, so that the drawing, like that of De St. Joseph,
might be improved.

The segmental organ has its trumpet fringed with more irregular lobes than in
N. cultrlfera (to judge from the figure) and also furnished with long cilia (Goodrich).

In the form of the anterior feet this species shows certain resemblances to N.
Marionii, Aucl. & Edw., but it diverges in the structure of the posterior feet. The
specimen from the Channel Islands is an epitokous female, partially developed, the ova
being about one-third grown. It has a dark speck on the ridge near the dorsal cirrus
for twenty segments, viz., from the eighth or ninth foot backward.

In the British Museum, in a tube labelled " Tube of Venusia from the coast of
Cornwall," is an example of this Nereid. The tube is composed of coarse sand, stones,
and shell-gravel, and has little secretion, so that it is more lax than that of Thelepus.
Bohn, in his recent paper on the movements of Annelids, considers this tube-dwelling
species very sensible to its surroundings, when free rapidly passing from the reptant

NEREIS (PERINEREIS) MABIONIL 295

to the pelagic condition. From the description of Dr. Baird, moreover, the form he de-
scribed as Heteronereis signata is in all probability the epitokous condition of N. Schmard&i,
and accordingly it has been entered doubtfully under that species. The specimen could
not be found in the British Museum at the period of examination.

The Nereis Sarsii of Rathke 1 has certain resemblances to this form, but it is difficult
to determine the exact relationships of the northern form.

The grounds on which Malmgren (1867) founded the genus Praxithea, though
interesting in themselves, do not seem to be sufficient to warrant the separation.

De St. Joseph procured his examples in coriaceous tubes amongst the mud of the
Zostera-flats in company with Gerianthus membranaceus, Haime. Gravier, 2 again, met
with this form in muddy tubes in the Zostera-region at St. Vaast-la-Hougue in August
and September, until it became transformed into a Heteronereis.

The species is chiefly a southern one.

4. Nereis (Perinekeis) Marionk, Audouin and Edwards, 1833. Plate LX, fig. 9— head ;
Plate LXXII, figs. 3-3 d— feet; Plate LXXXI, figs. 3 and 3 a— bristles.

Specific Characters. — Head greenish mottled with dark pigment, but the posterior
border is pale. Eyes normal, brownish, with lenses anteriorly and occasionally
posteriorly. Dorsal of the first pair of tentacular cirri longest, cirrophore greenish,
ventral short and thick. All cirri somewhat short. Body 3 — 4 inches long ; peristomial
segment twice as broad as the next. Dull olive green anteriorly, paler posteriorly ;
whitish patch in front of each foot, and a little white pigment in the dorsal median line.
Lobes of the feet pale. Posteriorly are two short caudal cirri. Proboscis with strong
curved jaws having five teeth. Paragnathi feebly developed. Dorsally the basal region
in extrusion has two conspicuous papillae, each surmounted by a large brown tooth (VI),
and on the fold between them are two or three smaller teeth (V). The folds are
thirteen in number, and ventrally have minute paragnathi arranged on two sides of
each triangular area (VII and VIII), the apex being in front. In some each area is
tinted brownish. Dorsally in the maxillary region is a group of small teeth (II) on each
side, ventrally in the same region are rows of small paragnathi generally forming a
somewhat elongated area (IV) on each side, whilst between them is a considerable group
of small paragnathi (III).

The first foot is characterized by the greater proportional bulk of the cirri; seti-
gerous lobe massive. About the thirtieth foot a considerable dorsal elevation appears,
and continues to the posterior end. It is marked by opaque granular glands. The
characteristic foot occurs near the fifty-seventh, in which the dorsal region — comprising
the dorsal lobe and the dorsal cirrus — is greatly hypertrophied, extending by-ancl-by, e g.,
at the sixty-eighth, as a long lamella with the cirrus at the tip. The bristles are of two
kinds, the tapering tips of the homogomph being somewhat short, as are also those of the
heterogomph.

1 ' Beitr. Fauna Xorweg./ p. 161, Taf. viii, f. 6—8, 1843.

2 f Xouv. Arch. Mas./ p. 148, 1901.

206 NEEEIS (PERINEKEIS) MARIONII.

Synonyms.

1833. Nereis Marionii, 1 Aud. & Eclw. Ann. So. Nat., t. xxix, p. 207, pi. xiii, f. 1—6.

1834. „ „ idem. Annel., p. 185, pi. iv a, f. 1—6.

1847. „ succinea, Frey and Leuckart. Beitrage, p. 154, Taf. ii, f. 9 and 11.
1851. „ „ Grube. Fam. Annel., pp. 49 and 127.

1865. „ Marionii, De Quatrefages. Annel., p. 549.
„ „ crassipes, idem. Ibid., p. 550.

1867. Stratonice Marioni, Malmgren. Annul. Polych., p. 56.

1868. Nereis lamellosa, Ehlers. Borstenw., ii, p. 564, Taf. xxii, f. 10—17.

succinea, idem. Ibid., ii, p. 570, Taf. xxii, f. 18—22,

1870. „ (Lipephile) macropus, Claparede. Annel. Nap. Snppl., p. 80, pi. viii, f. 1.

,, ., Marionii, Grube. Arch. f. Naturges., Bd. xxxvi, p. 305 (?).

,, ,, crassipes, idem. Ibid., p. 304.

1885. ,, lamellosa, Cams. Fauna Meclit., p. 221.

1898. „ longipes, De St. Joseph. Ann. Sc. Nat., 8 e ser. ; t. v, p. 314, pi. xvii, f. 107—112.

1902. ., Marionii, Mcintosh. Ann. Nat. Hist., ser. 7, vol. x, p. 256.

1908. „ (Neanthes) succinea, Horst. Notes Ley den Museum, vol. xxx, pp. 215 — 218, Text-

figs. 1-4.

1909. ,, „ „ idem. Tijdschr. NederL Dierk. Vereen. (2), xi, p. 144.

Habitat. — In galleries in cracks of rocks, St. Peter Port, under stones at Perrelle
Bay, Guernsey, and in similar places in Herm. South of England, Plymouth (Allen).

Shores of France (Milne Edwards and De St. Joseph) ; Naples (Claparede).

Head (Plate LX, fig. 9) marked by dark pigment on the dorsum, but with a pale
posterior margin so as to be abruptly denned from the peristomial segment. Eyes not
larger than in other species of Nereis, and having the same position ; brownish, with a
black speck in the centre, the anterior pair with lenses, and occasionally a minute
lens in the posterior. Dorsal of the first or upper pair of tentacular cirri longest, the
cirrophore being greenish, the ventral much shorter in proportion than in the other
species, and in addition rather thick. The other pair is longer than the latter, but
shorter than the former, the dorsal slightly exceeding the ventral in length. All the
cirri are somewhat short. The peristomial segment is about twice as broad as the
succeeding.

Body 3 — 4 ins. long, slightly tapered anteriorly, but more distinctly posteriorly, where
it terminates in two short cirri. 3 The peristomial segment is about twice as broad as the
succeeding. In large examples the dorsum anteriorly is dull olive green, paler green
posteriorly, the tints being the more pronounced from the pallor of the feet and their
processes, as well as from the whitish patch in front of each foot. There is also a little
white pigment in the median line of the dorsum. Though supplied with blood-vessels,
the lobes of the posterior feet appear to be pale, and this, with the whitish pigment at
the base, aids in the discrimination of the species. About the thirtieth foot a consider-
able dorsal elevation at the base of the foot occurs, and continues to the posterior end of
the body. It is marked by the same opaque granular glands as are found in the dorsal

1 Named after Dr. Marion de Proce, a naturalist and physician of Nantes. The Scottish
University Commissioners of 1889 do not encourage such talent in the department in our country.

2 In some four short filaments on each side of the vent (De St. Joseph).

NEREIS (PERINEREIS) MARIONII. 297

lobe of the foot itself. In spirit-preparations these give a character to the region, as in
N. Dumerilii.

The proboscis (Plate LX, fig. 9) has a pair of strong, curved maxillse with about
five or six serrations on the edge, a considerable portion of the tip being smooth. The
paragnathi are somewhat feebly developed, and apparently lose their brownish hue after
long preservation in spirit. Dorsally in front of the palpi (i.e., on the basal segment) are
two conspicuous papillae, each surmounted by a brown tooth (VI), and it may be one or two
smaller externally, transversely elongated at the base but pointed distally, and on the folds
between them are two or three smaller (V), though occasionally the solitary anterior one
is large. When the proboscis is half extruded these two papillae are very prominent,
and the edge of the proboscis at this stage of extrusion has thirteen crenations, each
furnished with minute paragnathi (VII and VIII). Those on the ventral surface are
arranged along two sides of a triangle, the apex being in the centre and toward the outer
border of the papilla. In some preparations this triangular area is tinted brownish and
thus sharply marked off from the proximal continuation of the fold, and from the distal
part of the fold forming the papilla in semi-extrusion. In the latter condition an inner
ring of six papillae occupies the centre, viz., two large lateral on each side, and a smaller
papilla dorsally and ventrally.

The maxillary region presents dorsally a group of small teeth (II) on each side;
ventrally in the same region are rows of small paragnathi forming a somewhat
elongated area (IV) on each side, whilst between them is a considerable group of small
paragnathi (III). The figure made by the artist from a bleached specimen is somewhat
abnormal, for it shows two large paragnathi with two or three smaller internally (VI),
and only a single small posterior tooth in V, instead of the foregoing typical arrange-
ment. The variations of the paragnathi in this species are sufficiently evident by referring
to the figures of Ehlers, and more recently those of De St. Joseph as indicated in the
list. The foot in such a case is, perhaps, individually more diagnostic than the
paragnathi.

The first foot (Plate LXXII, fig. 3) essentially differs from that of both Nereis
pelagica and N. cultrifera by the greater proportional bulk of the cirri. The dorsal edge
of the foot is short, and the short and massive dorsal cirrus extends considerably beyond
the dorsal lobe. The latter is large and bluntly ovate. The setigerous lobe beneath
(corresponding to the ventral setigerous lobe) is massive and projects much further than
in N. cultrifera, with which it agrees in being bifid. It bears dorsally a group of bristles
with slender tips of moderate length and finely spiked on the edge. The shafts have less
distinct transverse bars than in the former species. Beneath is a series of bristles with
short tips, the oblique part at the base being long, and curved near the prominent anterior
spur, whilst the spinous edge is curved and the tip scarcely hooked. The shafts show a
double camerated streak superiorly and a single one below the dilated region. The inferior
and larger group of bristles in this lobe corresponds in structure with the latter. The
spinous edges of all the bristles are directed upward in the preparation. Besides the
smaller upper and the larger inferior papilla of this lobe, the ventral edge presents a
distinct prominence.

The ventral lobe does not project as far as the setigerous, and is elongate ovate in

95

298 NEREIS (PERINEREIS) MAEIONII.

outline. The ventral cirrus arises at its base and extends somewhat beyond it. Its
proportions are large. A single spine passes to the furrow between the larger and
smaller terminal papillse.

The tenth foot (Plate LXXII, fig. 3 a) has a slightly tumid upper border with a faint
depression at the origin of the dorsal cirrus. The dorsal lobe forms a short, broad cone,
and is therefore less blunt than in the first foot. A dorsal setigerous lobe appears as a
small papilla below the former, and it bears a tuft of bristles with slender tapering tips
and a spinous edge (Plate LXXXI, fig. 3). These have rather shorter tips than in the
two foregoing species, viz., N. pelagica and N. cultrifera. Beneath is a somewhat bluntly
rounded lobe. The inferior setigerous lobe has the characters mentioned in the first foot,
viz., a small dorsal papilla, then a larger blunt process followed inferiorly by a projecting
rim under the bristles. The upper bristles of this lobe conform to those of the dorsal
division, whilst beneath is a dense group of stout bristles (Plate LXXXI, fig. 3 a) with
short tips, which may form a palisade of uniform length. The terminal piece is slightly
curved at the tip, and has a less projecting anterior inferior angle than in N. cultrifera,
and a spinous edge. The shaft shows two camerated streaks in the centre. The inferior
lobe is tongue-shaped with a blunt tip which does not extend so far outward as the
setigerous lobe above it. The ventral cirrus is considerably shorter than the dorsal and
does not reach the extremity of the ventral lobe.

The lobes of the foot undergo considerable changes, so that at the thirty-seventh the
dorsal becomes both diminished and pointed (Plate LXXII, fig. 3 b), the dorsal cirrus
is carried somewhat further outward, and is, compared with the lobe, proportionally large.
The upper setigerous lobe beneath is more distinctly seen than in the tenth foot and is
bifid. It bears similar bristles. The ovate lobe beneath is also considerably diminished,
and does not extend so far outward as the dorsal. The inferior setigerous process has
rather increased in lateral view, but projects less, and the distal end is more uniform, only
the small papilla at the upper bristles being prominent. The bristles are unaltered.
The tongue-shaped ventral lobe is much less, but it approaches the tip of the setigerous lobe
above more closely, since the latter is less prominent. The diminution of the foregoing
lobes brings the tip of the small ventral cirrus nearly in a line with that of the adjoining
lobe.

At the fifty-seventh foot (Plate LXXII, fig. 3 c) a great development of the region
between the base of the dorsal cirrus and the body has ensued, so that the diminished
dorsal cirrus with the minute lobe beneath is thrust far outward — as the terminal parts of
the great flattened vascular lamella thus formed. The dorsal border is occupied by
granular glands (Claparede's granular follicles), and similar opaque matter occurs at its
inferior edge. The spine and two bristles mark the dorsal setigerous lobe, but no papilla
is visible. The lobe beneath is about the same size as in the thirty-seventh foot. The
inferior setigerous lobe is rather less and has fewer bristles. Moreover the upper two
or three bristles with short tips have proportionally increased in size. The ventral lobe
and the cirrus are somewhat less.

The elongation of the upper region of the foot is fully developed at the sixty-eighth
foot (Plate LXXII, fig. 3 d) where it forms a greatly elongated vascular process, the base
of which occupies fully half the foot in lateral view. The region, indeed, forms a long

NEREIS (PERINEREIS) MARIONIL 299

cone with glands dorsally and ventrally and a great blood-vessel in the centre. The short
dorsal cirrus is now at the tip of the region, in a line with the dorsal border, a small
rounded papilla beneath indicating the dorsal lobe. The dorsal setigerous lobe is
represented by its bristles and the spine. The lobe beneath is considerably less. The
inferior setigerous lobe and the parts beneath have the same character as in the previous
foot.

Behind the sixty-eighth the characters of the foot present little alteration, even to the
tip of the tail. The greatly elongated and very slightly tapered dorsal region has the short
cirrus at its extremity, and the deeper angle at its base inferiorly is now the only trace of
the dorsal lobe. The upper setigerous lobe is represented by its spine and a few bristles.
The lobe beneath is ovate, but its base encroaches on that of the region above it. The
inferior setigerous lobe is conical, the longer slope being ventral. The ventral lobe is
small, but of similar shape to that in the sixty-eighth foot. The ventral cirrus projects as
far as the tip of the setigerous lobe above it. The vascularity of the entire foot is great,
so that the posterior region of the animal is thus provided with a series of leaf -like organs
for aiding, along with the other parts of the foot, in respiration. The bristles preserve
the same characters as in the sixty-eighth foot ; two of these in the inferior series with
short tips arise above the spine, and are stronger than the others.

Habits. — This species is less active than Nereis pelagica, N. cultrifera, or N Dumerilii,
and is considerably smaller than the two former species.

An example from Falmouth in the British Museum is labelled Nereis fimbriata.
Considerable confusion, indeed, has occurred in the synonymy of this species, the
characters of which, however, are, on the whole, distinctive. The variability and com-
parative feebleness of the paragnathi, and their tendency to become indistinct in spirit-
preparations, may also have tended to create doubt. These remarks are suggested by the
fact that Baron De St. Joseph, whose admirable work on the Annelids of Dinard and
generally of the shores of France has done so much to clear up the subject and increase
our knowledge of the group, does not include Nereis Marionii, And. & Edw., in his compre-
hensive series. Yet he constitutes two new species for forms having the closest resemblance
to that of Audouin and Edwards. The first of these is one which he places under the
genus Neanthes of Kinberg, viz., Nereis (Neanthes) Perrieri, 1 which he procured at
Villerville and Villers on the shores of France. It is true the longest tentacular cirrus is
longer than that usually found in N Marionii, and the paragnathi show certain differences,
but, as Horst 2 has already pointed out, a considerable margin should be allowed for
variation, and he is of opinion that this form is only the Nereis succinea of Leuckart, a
form here considered synonymous with Nereis Marionii, Aud. & Edw.

The second species, viz., Perinereis longipes, he doubtfully associates with the Nereis
crassipes of De Quatrefages 3 and of GrubeV critique on the Annelids in the Parisian
Museums; but it appears to be this form (Nereis Marionii), so well known in the
Mediterranean, on the southern shores of England, and in the Channel Islands. De St.

i ' Ann. Sc. Nat./ 8 e ser., t. v, p. 288, pi. xv, figs. 69—77, 1898.

2 Vide p. 301.

3 ' Anneles/ i, p. 550.

4 < Arch. f. Naturges./ 1870, p. 305.

300 NEEEIS- (PERINEREIS) MARIONII.

Joseph procured his specimens at Guethary and St. Jean-de-Luz under stones, and at St.
Anne near d'Hendaye in Mdobesia agariciformis.

The original description with its excellent figures by Audouin and Edwards, 1 drawn
up from a specimen from La Vendee, sent by Dr. Marion de Proce, a physician and
naturalist of Nantes, places the identification of this species beyond doubt, though the
paragnathi are not alluded to. That so competent a successor, therefore, should have
been unable to satisfy himself of the relationship is not easily explained, even though the
absence of the original example should have made some obscurity. The description
De St. Joseph gives of the body, the head, and its organs, the changes in the structure of
the feet — especially the remarkable increase of the dorsal process with the cirrus at the
tip (which Audouin and Edwards regarded as branchial in function), as well as
the structure of the bristles— all apply to Nereis Marionii The great variability in the
details of the paragnathi of N. Marionii probably has given rise to ambiguity in the
absence of a description or reference to the original specimen. Moreover, the changes
after long preservation are considerable. In every British example Group VI is repre-
sented by a prominent conical tooth, or by a transverse bar, akin to that indicated as
typical of Perinereis longipes by De St. Joseph. This, from its somewhat conical outline,
is prominent during the partial protrusion of the proboscis, as it is elevated on a papilla.
In the centre between there is (V) a single conical tooth, with a somewhat irregular series of
smaller points " dusted " between the larger pair (VI). The large number of very minute
specks shown in De St. Joseph's figure (Plate XVII, fig. 110) is apparently a variation.
In some of the British forms Group V consists of a central in front and two or three
similar ones behind, with a few smaller dotted around. Groups I and II on this aspect,
viz., those near the base of the maxillae in extrusion, are after the plan shown by De St.
Joseph. From the outer border of each of the larger posterior paragnathi (VI) a dotted
series runs to the ventral surface, and occasionally a line of paragnathi leaves Group V on
each side in the same direction. These lead to the basal ventral series VII and VIII,
arranged in lozenges after the manner shown by De St. Joseph (fig. 112), though in the
British specimens the minute points are less numerous. The teeth in the maxillary
(distal) region, near the base of the maxillse on the ventral surface (Groups III and IV),
are fewer in number, but have a similar arrangement to that figured by De St. Joseph.
The author, indeed, observes that the proboscis approaches that of Perinereis Marionii, but
that species has three dorsal lobes to the foot (as figured by Audouin and Edwards?).
So far as the British examples go, however, the structure does not differ from that given
by De St. Joseph, and it may be that Grube was right in considering Nereis crassipes,
De Quatrefages, as the young of N. Marionii, which, according to his descriptions, seems
altogether to have eluded the careful search of the French shores by De St. Joseph.
Periuereis longipes, De St. Joseph, therefore, as well as Nereis Perrieri, is regarded as a
synonym of Nereis Marionii, And. & Edw.

The Nereis Ockenii 2 (the N. oltoana of the ' Descrizione' 3 ) of Delle Chiaji may have
some connection with this species, if it is not identical. His artist may have made

1 c Annelides/ p. 185, pi. iv a, figs. 1—6, 1834.

2 'Memorie/ hi, pp. 166 and 175, Tav. xlii, figs. 2, 12, and 20, 1828.

3 Tomo iii, pp. 96 and 103, v, p. 102, Tav. 96, figs. 7, 12, 17, 20, 21.

NEREIS (PERINEREIS) MARIONJL 301

long lamellae anteriorly as well as posteriorly, for he was by no means scientifically
accurate.

Frey and Leuckart (1847) describe and give figures of a new species, Nereis
suednea, from Heligoland, which appears to correspond most closely with the present
form.

Grrube's Nereis vexillosa 1 from the shores of Eastern Siberia (near the Amoor) is an
allied form, and his Nereis arctica may be its epitokous stage.

The characteristic condition of the dorsal lamellae of the feet posteriorly induced
Malmgren (1867) to institute the genus Stratoniee for this species.

Ehlers (1S68) apparently refers to this form under two titles, viz., Nereis lamellosa
from the Adriatic, and Nereis suednea of Leuckart from Cuxhaven and Norderney. The
differences in the paragnathi are due to mere variation, as formerly explained. Both
have short tentacles and tentacular cirri, and the form of the foot is characteristic.

Claparede (1870), in his account of this form, his Nereis macropus, omits reference
t'o Nereis Marionii (And. & Edw.), the resemblance having apparently escaped him for the
moment. The shortness of the tentacles and the general aspect of the head, the armature
of the proboscis, the nature of the feet, bristles, and other parts are more or less identical
with A. Marionii. He truly considered the greatly elongated dorsal lamella as an aid to
respiration, his figures bearing out his interpretation, and he alludes to other forms in
which the superior lamella is largely developed, e.g., Mastigonereis, Schmarda, P sen doner eis,
Paranereis, Perinereis, and Mastigonereis, Kinberg, though he does not approve of the
latter author's classification.

It is remarkable how closely Nereis melarwcephala of the ' Challenger ' approaches
the British species, which probably has a much wider distribution than is at present
supposed. Slight variation may explain the differences in paragnathi and other respects
which are held to be of specific importance.

Gravier's 2 (1901) Pseudonereis anomala, from the Red Sea, is another form which
closely approaches this species.

An interesting criticism, with four good figures, on this species, as the Nereis
(Neanthes) suednea of Leuckart, and the N. Perrieri of De St. Joseph, has recently been
made by R. Horst, 3 who had the advantage of examining the original specimen of
Leuckart, as well as those of Metzger and others, in the collection of Ehlers. He comes
to the conclusion that these are one and the same species, and that the limited geographical
distribution no longer holds. If, however, he had included other synonyms in his review,
this distribution would have been still further extended.

The form termed Nereis glandulosa by Ehlers 4 approaches this form in the structure
of the feet and other features, but it does not appear to be identical, though too much
reliance should not be placed on the paragnathi.

1 ' Mem. des Sav. etrang./ St. Petersb., viii ; p. 4, pi. ii, figs. 1, 5, and 6.

2 <Nouv. Arch. Mus./ 4 e ser., t. iii, p. 191, t. ii, pi. xii, figs. 50—52.

3 ' Notes from the Ley den Museum/ vol. xxx, p. 215, 1908.

4 'Deutsch. Tiefsee Exped./ p. 74, Taf. viii, figs. 1—6, 1908.

302 NEREIS DTTMERILIL

5. Nekeis dum'eeilii, Audonin fy Edwards, 1833. Plate LII, fig. 5 ; Plate LX, figs. 10-10 c
—head, etc.; Plate LXX1I, figs. 4-4/— feet; Plate LXXXI, figs. 4-4 c— bristles.

Specific Characters. — Head of the normal shape with four large ejes. Occasionally
the elevated central region has a wedge-shaped patch of white from the posterior pair of
eyes to the tip. Tentacles of moderate length. Tentacular cirri long, ranging from one
third the length in small examples to one fifth the total length in larger ; the longest is
the dorsal of the second pair. Body 2\ ins. long, and of seventy to eighty segments.
Colour, various shades of yellowish-brown, enlivened by the red dorsal vessel. Anterior
fifth of the dorsum minutely dotted with brown, and more brightly iridescent than the rest
of the dorsum. Minute white grains occur over the central vessel. Behind the anterior
fifth white grains are distributed over the whole dorsum. Occasionally small specimens
are greenish-yellow, and have rather large yellow grains. Ventral surface greenish
anteriorly, the rest of the same hue as the dorsum. Proboscis has two slightly translucent
amber-like maxillge with six teeth. Paragnathi comparatively feebly developed. Dorsally
the maxillary segment in extrusion has in the ordinary examples no paragnathi, but a
short double row of small paragnathi (II) occurs on each side. Ventrally a patch of
oblique but parallel pectinate rows of minute denticles, the whole having a somewhat
triangular outline and a rasp-like surface, occurs near the base of each maxilla (IV),
and between are two rows of minute denticles (III). The proximal segment has dorsally
two patches of paragnathi (VI) in front of the palpi, each composed of about two rows
of minute points. Ventrally each lozenge-like area has two rows (VII and VIII) of
minute paragnathi.

The first foot has a proportionally large and long dorsal cirrus and three rather
long lobes, besides the setigerous process. A tendency to the increase of the dorsal
region is apparent at the thirty-seventh foot and becomes conspicuous at the fifty-seventh,
with the presence of two opaque glandular masses in each foot. The ventral cirrus is
somewhat short throughout. Bristles homo- and heterogomph, the former with tapering,
boldly spinous terminal pieces, the latter with short tips, and a very short spinous edge.

Epitokous Form. — The head in the epitokous male is short and rounded, the large,
sometimes connate, eyes occupying the lateral regions. Body soft and delicate. Dorsal
cirri of the first seven segments dilated and peculiarly modified. The ventral cirri of the
corresponding feet are also dilated. At the compressed sixteenth foot the lamellse appear,
and the pelagic condition is fully developed at the thirty-seventh foot, the great lobes
being very thin, and finely and somewhat regularly veined. The dorsal cirrus has a
series of prominent papillas on its lower edge. The long swimming bristles give a glassy
sheen to the sides.

Synonyms.

1833. Nereis Dumerilii, Aud. & Edw. Ann. Sc, Nat,, t. xxix, p. 218, pi. xiii, f. 9—12.

1834. „ „ idem. Annel., p. 196, pi. iv a, f. 10—12,
1840. ., ., Johnston. Ann. Nat. Hist., v, p. 174.

a a » G-rube. Actin. Echin. u. Wiirm., p. 73.

1843 « » » H- Ratlike. Beitrage Fauna Norweg., p. 163, Taf. vii, f. 4, 5.

NEREIS DUMEBILII.

303

1843.
1844.
1851.

>>
1865.

1867.
1868.

1869.

1870.

1873.

>■>

1874.

>>
1875.

1909.

Nereis zostericola, CErsted. Annul. Danic. Consp., p. 22, f. 20, 29, 67, 70, 71, 74.
„ Dumerilii, Thompson. Rep. Brit. Assoc, 1843, p. 273.
„ „ Grube. Fam. Annel., pp. 48 and 125.

„ zostericola, idem. Ibid., pp. 49 and 125.

„ Dumerilii, Johnston. Cat. Worms Brit. Mas., pp. 156 and 341.
„ ,, De Quatrefages. Annel., p. 502.

,i „ Carrington. Annel. Southport, p. 8.

Leontis Dumerilii, Malmgren. Annul. Polych., p. 52. Taf. iv, f. 25.
„ peritonealis , Claparede. Annel. Nap., p. 157.

Ehlers. Borstenw., ii, p. 535, Taf. xx, f. 21—37.
„ Massiliensis, Moquin-Tandon. Ann. Sc. Nat., 5 e ser., t. xi, p. 134.
,, peritonealis, Claparede. Annel. Nap. Suppl., p. 44, pis. iii— vi.
Nereis Dumerilii, Grube. Arch. f. Naturges., Bd. xxxvi, p. 308.

„ „ Kupffer. Exped. Ostsee, ' Pommerania,' p. 150.

Leontis „ Sars. Bidrag Christ. Fauna, p. 22.

„ „ Malm. Goteb. vet. o. vitt. Handl., xiv, p. 84.

Nereis „ Mcintosh. Ann. Nat. Hist., ser. 4, vol. xiv, p. 199.

„ „ Marion and Bobretzky. Ann. Sc. Nat., 6 e ser., t. ii, p. 16.

„ „ Mobius. Jahresb. Comm. deutsch., ii, p. 168.

„ „ Mcintosh. Invert, and Fishes St. Andrews, p. 122.

„ Lenz. Jahr. Comm. deutsch., iv, Anhang., p. 12.

„ Webster. Trans. Albany Inst., ix, p. 34.

„ Marenzeller. Sudjap. Annel., p. 123, Taf. ii, f. 4.

„ Langerhans. Zeitschr. f. wiss. Zool., xxxiii, p. 281.

„ Tauber. Annul. Danic, p. 98.

,, Langerhans. Canar. Annel. in Nova Act. Leop., p. 110.

„ Levinsen. Yidensk. Meddel. Foren. Kjoben., p. 235.

„ Carus. Fauna Medit., p. 220.

Mcintosh. Annel. ' Challenger/ p. 221.
„ Malaquin. Annel. Boulon, p. 27.

Nereis (Leonti*) Dumerilii, Hornel. Trans. Biol. Soc Liverp., vol. v, p. 241.
Dumerilii, Levinsen. Yidensk. Ud. ' Hauchs,' p. 329.

„ De St. Joseph. Ann. Sc. Nat., 7 e ser., t. xx, p. 214.

„ Racovitza, Arch. Zool. exper., 3 e ser., iv, p. 217, pi. iv, f. 35—39.

,, Wistinghausen. Mitth. Zool. Stat. Neap., t. x, p. 47, pis. vi and vii.

Mcintosh. Ann. Nat. Hist., ser. 7, vol. x, pp. 256 and 258.
„ Allen. Journ. M. B. A., n.s., vol. vii, p. 224.

G-raeffe. Arbeit. Zool. Stat. Triest, xv, p. 322.
Platynereis Dumerilii, De St. Joseph. Ann. Sc Nat., 9 e ser., t. iii, p. 222.
Nereis ,, Eisig. Fauna u. Fl. Neap., xxviii, p. 208.

„ „ Elwes. Journ. M. B. A., n.s., vol. viii, p. 351.

Forma epitoca.

1878.

y>

1879.

))

>>

)>

>>

Leontis

>>

Nereis

1881.

Leontis

1883.

Nereis

1885.

>>

j}

Leontis

1890.

))

1891.

Nereis (

1893.

„ i

1895.

Leontis

1896.

Nereis

1901

iy

1902.

>>

1904.

»

1905.

>>

1906.

Platynei

1843. Heteronereis fueicola, CErsted. Annul. Danic. Consp., p. 19, f. 17, 55—58, 61, 62.

,, „ viridis, idem. Ibid., p. 20 (?).

„ Nereilepas variabilis, idem. Ibid., p. 20, f. 18, 26, 51, 52, 54, 59, 60.
1851. Nereis fueicola, Grrnbe. Fam. Annel., pp. 50 and 127.
1865. Heteronereis fueicola, De Quatrefages. Annel., p. 574.

,, Iphinereis „ Malmgren. Nord. Hafs-Annul., p. 182.

304 NEREIS DUMERILII.

1867. Iphinereis fucicola, idem. Annul. Polych., p. 58, Taf. v, f. 29, 30.

1868. Heteronereis Malmgreni, Claparede. Annel. Nap., p. 173, pi. xi, f. 1.
1870. „ fucicola, idem. Annel. Nap. Suppl., pi. hi, f. 4, 5.
1874. Iphinereis fucicola, Malm. Goteb. vet. o. vitt. HandL, xiv, p. 84.

„ Hetercmereis viridis, idem. Ibid., p. 85.

Habitat. — Abundant on both the eastern and western coasts of the British
Islands, though its headquarters are on the western and the southern shores. On the
east coast, as at St. Andrews, it generally occurs in deep water or is tossed on
shore by storms, its tubes being attached to the empty valves of Gardium echinatum and
other mollusks. Off the western shores it frequents both deep and shallow (laminarian)
zones, being especially common in the latter in four to six fathoms. Its tough tubes are
often met with in the interior of decayed laminarian stalks, or in peat, whilst one took
advantage of a partially open egg-capsule of a skate. In Loch Portan, near Lochmaddy,
where there is a considerable admixture of fresh water, the tubes (with the annelids in
the interior) are common. In deep water they are often fixed to the large red Ascidians
of the Minch and other western seas.

In the south, as in the Channel Islands, it seems to take the place of Nereis pelagica
of the north, and is found in crevices and fissures of the rocks between tide-marks, as well
as in the adjoining sea. It is not uncommon in the stomachs of cod and haddock in St.
Andrews Bay, especially the epitokous Iphinereis fucicola (E. M.). It occurs on the
eastern, western, and southern shores of Ireland (e. g. Loch Slyne, Co. Cork, Roy. Irish
Acad. Expect., 1866) ; Bay of Galway (Prof. E. P. Wright). 1

It is very plentiful in the Mediterranean, and since it ranges to Norway and Sweden
its distribution is wide : Black Sea (Bobretzky) ; St. Vincent, Cape Verde Islands ; Japan
(Marenzeller) ; Norway (Malmgren) ; Canaries (Langerhans) ; Virginian coast (Webster).

Head longer than broad, with four large and sometimes reddish eyes, the anterior
pair wider apart. In a few preserved examples a little pigment occurs on each side in
front of them. In some from deep water the elevated central region has a wedge-shaped
patch of white, with a median furrow running from the posterior pair of eyes to the tip
of the snout. The tentacles are of moderate length and subulate. The long tentacular
cirri, for which the species is remarkable, range from rather more than one fifth the
entire length of the annelid to one third in small examples, and are pale greenish. These
tentacular cirri are obliquely situated, the longest being the most posterior and dorsal.

Body 2± ins. in length and of seventy to eighty segments. Colour, yellowish-brown
on the dorsum, some being more deeply tinted than others, and marked by the red dorsal
vessel. The surface of the anterior fifth is minutely clotted with brown, and more brightly
iridescent than the rest of the dorsum. At every segment-junction minute white dots
occur over the central vessel. Behind the anterior fifth minute white grains are
distributed over the whole dorsum, and very distinctly towards the tip of the tail, which is
terminated by two long cirri. The tips of the feet (not the cirri) are whitish in the
caudal region and for some distance forward. Some small specimens from deep water
are of a greenish yellow, the dorsum being speckled with rather large yellow grains,
which over the region tinted brownish by the intestine are arranged in two rows.
1 The recent death of this genial naturalist deprives both zoology and botany of an ardent worker.

NEREIS DUMERILIL 305

Anteriorly greyish-brown grains are mingled with the yellow. In preserved specimens
the two opaque, brownish, granular, glandular bodies on the dorsum of the foot are
visible about the tenth foot, and are very conspicuous posteriorly, where one or two dots
also occur on the side above the foot. Ventral surface greenish anteriorly, the rest being
similar to the dorsum, but without the white grains.

The proboscis (Plate LX, figs. 10, 10 a, and 10 6) has two slightly translucent
maxillae, like amber, with six teeth pointing or sloped distally. The paragnathi are com-
paratively feebly developed, and they become very indistinct in some after long preservation
in spirit. On the dorsum of the organ in extrusion are two patches (VI) in front of the
palpi, each composed of about two rows of minute points. No teeth occur on this surface
in the maxillary division at the base of the jaws in the ordinary examples, but in the
epitokous forms (Plate LX, fig. 10 b) a short double row of small paragnathi (II) are met
with on each side. On the ventral surface the proximal region of the organ presents two
rows of minute paragnathi (VII and VIII) at the outer end of each lozenge-shaped area,
whilst on the maxillary division a patch of oblique but parallel rows of similar minute
denticles (IV), the whole having a somewhat triangular outline, occurs near the base of
each jaw. In the middle line between these are two rows (III) of minute denticles.
Considerable variation is met with in the arrangement of the paragnathi on the various
areas.

The first foot (Plate LXXII, fig. 4) has a long and proportionally large dorsal cirrus,
which extends fully a third of its length beyond the dorsal lobe. The latter is of an
elongated tongue-shape, slightly diminished at the tip. Beneath it and partly over-
lapping the setigerous lobe is a smaller elongate lobe, also diminished at the tip. A small
papilla occurs opposite the elongated lobe just mentioned. The setigerous lobe bears a
series of bristles with long tapering tips. The shafts have a single camerated band in
lateral view, and in the comparatively small bristle these are faintly marked. The sabre-
shaped tip is not much tapered and has a finely spinous edge. The ventral lobe projects
considerably beyond the setigerous, but not so far as the two upper lobes. It also is
tongue-shaped, with a long inferior border. The ventral cirrus is smaller than the
dorsal, its tip reaching to that of the lobe above it, The foot thus differs from that of any
of the preceding species. A single spine, as usual, is present,

At the tenth foot (Plate LXXII, fig. 4 a) the lobes have become more obtuse, the
upper being the more prominent, as the outline slopes from above downward and inward.
The edge of the foot from the body to the dorsal cirrus is longer and the ridge more
convex. The dorsal cirrus is still large, though less in proportion to the bluntly conical
dorsal lobe, which has considerably increased in bulk. A dorsal setigerous lobe or
papilla is now present, but, as it lies in front, is not included in the outline shown in the
figure. The lobe beneath is a massive blunt cone, its base overlapping the inferior
setigerous lobe, which is somewhat posterior in position. The latter has one long papilla
and a projecting edge inferiorly. The upper group of bristles has tapering and boldly
spinous tips (Plate LXXXI, fig 4), besides others with short tips, the spinous edge being
very short (Plate LXXXI, fig. 4a). A double line occurs below the hooked tip as if a
differentiation were indicated. Ehlers' figure differs from the condition in our examples.
The camerated central region is less distinct than in Nereis pelagka or N. eultrifera.

96

306 NEREIS DUMERILII.

The ventral lobe is bluntly ovate and the small ventral cirrus does not extend so far
outward as its tip.

At the thirty-seventh foot (Plate LXXII, fig. 4 b) a considerable increase in length
has occurred, both by the elongation of the region between the body and the dorsal cirrus,
and the elongation of the lobes themselves. The dorsal cirrus is still long, and has two
opaque glandular masses near its base. The dorsal lobe forms a long triangle, and the
superior setigerous lobe is indicated by two small papillas in the sulcus. The next lobe is
nearly lanceolate, though there is an elevation on its ventral border near the base. The
inferior setigerous lobe forms a truncated cone (though in the larger specimens a longer,
pointed papilla and a shorter are present) bearing bristles with tapering tips
superiorly and those with short tips inferiorly. The ventral lobe is tongue-shaped, and
the ventral cirrus small, only reaching to the middle of the lobe above it.

The general size of the foot is diminished at the fifty-seventh (Plate LXXII, fig. 4c).
The dorsal border is convex and is entirely occupied by the opaque granular bodies now
in close apposition. The dorsal lobe does not differ much from the condition in the
thirty-seventh foot, forming a long, lanceolate process. The superior setigerous lobe has
at least one prominent papilla. The next lobe is elongate-lanceolate as in the thirty-
seventh foot, only the inferior elevation is less. The inferior setigerous process is conical
with a long and a short papilla, the latter for the spine. The ventral lobe is tongue-shaped,
and the cirrus is somewhat longer than in front. The upper bristles with short tips are
considerably stronger than those beneath them.

At the seventy-seventh foot the lobes, especially the two upper, are larger in propor-
tion. The opaque granular bodies have fused into one great mass which encroaches on
the base of the dorsal lobe. The other parts of the foot are similar to the fifty-seventh.
The upper bristles with short tips in the inferior setigerous lobe remain stronger than
those beneath, as if in this region of the body they had a special function— probably in
connection with the tubicular habits of the species. A slight indication of a wing is
observed beneath the hook in the form of a double line at the margin.

In the epitokous form (Iphinereis fucicola) the feet are furnished with long,
translucent swimming bristles (Plate LXXXI, fig. 4 b) the terminal piece of which is thin
and flat like the blade of a scalpel. The serrations on the edge (figured by Ehlers) were
not definitely made out in these examples (long preserved). The shaft is equally
translucent, and differs from that in the ordinary condition in having two stripes with the
transverse bars or cameras in the centre. Those bristles with short tips occurring in the
epitokous forms are exemplified in Plate LXXXI, fig. 4 c, from the tenth foot. The
head in these (Plate LX, fig. 10 6) has remarkably large eyes and long tentacular cirri,
which in spirit show slight crenations.

The epitokous forms occur most plentifully in the warmer months, e.g., in June.
Their rounded heads, and large, often connate, eyes, which severally have lenses, are
characteristic.

In a male heteronereid (Plate LX, fig. 10 c) the first fifteen feet (the average
number) are little modified, except the dorsal cirri anteriorly. The head is short and
rounded, the large eyes occupying the lateral regions and apparently having vision
dorsally, laterally, and anteriorly. The softness and delicacy of the tissues, as pointed

NEREIS DUMBEILTI. 307

out by Claparede, are diagnostic. In many the anterior region of the body has the
segment-junctions transverse, whilst those behind slant from the middle line outward and
backward, a feature observed in the heteronereid of iV. pelqgica. This, however, is not
seen in those from which the genital products have escaped, and the bodies of which are
semi-transparent.

The dorsal cirrus of the first foot is considerably dilated (Plate LXXII, fig. -id) so
that its dorsal outline forms a semicircle. The ventral cirrus is also similarly enlarged.
Three elongated lobes appear in lateral view. The second foot is similar. The third
has a tendency to an angular condition near the tip of the dorsal cirrus, from the abrupt
cessation of the enlargement. In this foot, as usual, the two setigerous processes occur.
The peculiar knobbed condition of the dorsal cirrus is more marked in the fourth foot
and attains its maximum in the fifth and sixth (Plate LXXII, fig. 4e). In the seventh
the cirrus, though longer, is more slender, and shows the angular process at the tip. The
eighth on the right has a slender cirrus with the tip bent at an angle, whilst its fellow of
the opposite side is normal, and so with the others which follow. The bristles are normal.

At the compressed sixteenth foot a decided change has taken place. An ovate
lamella arises internal to the base of the dorsal cirrus, which is long and furnished with
eight papillae on its ventral border, the tip tapering from the last, which is longest. The
dorsal lamella is long and lanceolate. The superior setigerous process ends in a blunt
point, and some of the old bristles are present, the rest being swimming bristles. The
lobe beneath it is lanceolate with a rounded spur at its base interiorly. The inferior
setigerous process is bifid, with an ovato-lanceolate lamella superiorly. Some of the old
bristles remain in the superior tuft (homo- and heterogomph) ; the rest are swimming
bristles. The ventral lobe is tongue-shaped. The ventral cirrus has an ovate lamella
internally, and a small elongated one externally.

The pelagic foot is seen in full development at the thirty-seventh in those
captured near the surface, as at Castlebay, Bona, on the 22nd May. The lobes of the
feet are very thin and the sides glisten with the sheen of the large and long swimming
bristles. Dorsally (Plate LXXII, fig. 4f) is the fan-shaped lamella close to the inner
edge of the dorsal cirrus. The dorsal cirrus itself is long, with the papillae (eight to ten)
on the lower border, and distally the filiform tip. The dorsal lamella is lanceolate. The
upper setigerous lobe is bifid and bluntly conical, the transparent bristles projecting from
it in two groups. The lamella soldered to it inferiorly is broadly lanceolate with a heel
at its inner and inferior border. The long inferior setigerous lobe has a huge lamella on
its posterior face, stretching almost to the tips of the bristles, which project like a great
fan. A long heel likewise extends inward from its base superiorly. The tongue-shaped
ventral lamella leaves the stem of the former lobe and turns outward as a process with
a spur on its upper border near the base. The ventral cirrus projects from a large fan-
like lamella internally and a tongue- shaped one, with an upper heel, externally.

All the foliaceous lamellae are finely and somewhat regularly veined from a midrib
or a basal region. The venation appears to be due to granular canals (?) of the hypoderm.
Many terminate between the pale end-organs at the margin of the lamella.

Little change occurs in the structure of the foot till near the tip of the tail, where
the diminution in the size of the foot is marked. All the lamellae become less ; the dorsal

308 NEREIS DUMERILII.

cirrus is shorter and devoid of the inferior papillae. The bristles become few and long,
but have the same structure.

Those captured in the pelagic condition at Barra were nearly uniform in size, viz.,
about 2 ins. in spirit. The vent is remarkably papillose.

A small example (about 1 in. in spirit) from the Mediterranean has very large eyes
which are completely connate. The anterior region of the body has twenty-one segments
of the ordinary structure. The rest are heteronereid. It probably is a male though the
sexual elements are absent.

The males of this form seem to be more frequently met with than the females at St.
Andrews — generally in March, and some reach fully 4 ins. If this is the male epitokous
form of N. Damer Mi the radical change in the structure of the first foot is interesting.

In what appeared to be a small male from Plymouth the swimming lamellae occur
on the seventeenth foot, the thirteen anterior being large, and the fourteenth, fifteenth,
and sixteenth smaller and more compressed, but having no foliaceous development or
swimming bristles. The change is thus sudden. The eyes are large and connate and the
anterior pair have lenses. The paragnathi are well developed.

The forms under which this species appears are : (1) The ordinary typical form
which produces ova and sperms in separate individuals without change in the feet.
(2) The same producing ova and sperms in the same individual. (3) The Iphinereis
transformation with swimming lamellae and bristles, and of separate sexes.

In a small specimen, probably about an inch when living or very little more, the
eyes were as large as in the foregoing, and connate. There were fourteen feet of the
ordinary form in front, the next were heteronereid. It came from Jersey (Mr. Horn ell).
Posteriorly the heteronereid condition abruptly ceased, and about a dozen of the terminal
segments were of the ordinary form.

Reproduction. — At Naples Lo Bianco l found this annelid mature in August, and the
larvae in the tube from December to February and April to August. The Heteronereid
condition occurred as a pelagic form from November to May.

A young example of Nereis Ihmerilii captured in the tow-net by Mrs. Collings of
Sark in July has thirty-seven feet. The jaws have eight or nine teeth. The anterior
pair of eyes have lenses and are considerably larger than the posterior. They are trans-
versely ovate, whereas the posterior pair are irregularly rounded. This may be an early
example hatched the same season.

In a small example of fifty-six segments, and measuring little over half an inch in
spirit, the eyes are considerably enlarged, and the dorsal cirri of the first seven segments
form large clavate processes with the filiform tip. It seems to be a very small male. No
change in the lobes of the feet or in the bristles have occurred. It was dredged in Tangiers
Bay in thirty fathoms by the ' Porcupine ' in 1870.

An interesting account of the earlier stages in the development of this species at
Madeira is given by Langerhans (1879). He only found one out of several hundred examples
in the Heteroneveis condition. The eggs are whitish with a greenish point. The youngest
larva he examined had three bristled segments, with a single dorsal and several ventral

1 The recent loss of this able observer at the Zoological Station, Naples, is to be deplored both
in the interests of science and of the workers there.

NEREIS DUMBRILII. 309

bristles, a head with four eyes, two frontal tentacles or papillae, and rudimentary palpi.
The proboscis is unarmed, and the rectum is differentiated. Two short anal cirri are
present. He noted for some days the development of the parts and the increase of the
segments to sixteen. He found that the armature of the proboscis appeared when there
were eight or nine bristled segments. He describes the bristles of the early stages as
embryonal, the tip being short with three spikes on the edge.

At St. Peter Port, Guernsey, in July, 1868, a female of the ordinary outline, and
measuring about 2 in. in length, was laden as far forward as the fifth foot with large
ova. These ova are larger than in the Heteronereid forms ; indeed, about six or seven
made up the diameter of the body. The eyes are of average size, and no change is
apparent in the structure of the feet or bristles. This is an example of that condition in
which the sexual elements are developed without external change.

A large female, again, of the ordinary type presents slightly enlarged eyes, but no
change in the feet. The body-cavity is filled with small ova. Another procured in
Birturbury Bay, Ireland, in June, is in the same condition, along with what appeared to
be an epitokous female, since no papillee occurred on the dorsal cirrus, but the specimen
was so badly preserved that doubt remains. A few stray ova are seen, but their nature
is doubtful.

Habits. — It is a less active species than Nereis pelagica, yet swims easily through the
water in an undulating manner, without coiling the long tentacles. It rapidly secretes
tubes even in a collecting bottle, and when deprived of them often enters those of
Terebellge or other forms. When in its own tubes in sea-water an undulating motion of
the body is kept up, apparently for purposes of respiration, as in Phijllodoce and other
annelids. The tubes in nature are tough, the mucus being strengthened by fragments of
seaweeds and debris, and they are generally attached to Fuci and Laminariaa, as CErsted
originally found, though occasionally to other structures. It is somewhat brittle : even
withdrawal from its tube suffices to cause fractures of the posterior region, and the same
happens when placed in spirit. In the separated fragments the dorsal blood-vessel
remains distended, and irregular contractions take place for some time.

At St. Peter Port, Guernsey, the tubes are sometimes stiffened by fragments of
shells, both univalve and bivalve, Foraminifera, minute fragments of gneiss, particles of
gravel, and fragments of Algse and mud. At Lochmaddy, in the Outer Hebrides, besides
the tough secretion and mud, the tubes are frequently strengthened and tufted with Algse,
and here and there a fragment of shell. Broad-leaved Algge are often bound together by
the secretions in Bressay Sound, besides tufts of filamentous Algas, and the walls have
many fragments of shells and occasionally fragments of Fuci. The tubes of some from
Valentia Harbour, again, are hairy all over with filamentous Alga3. At Herm the tube
of a small example with Algge and fragments of Trochus attached was neatly placed inside
the rim of a small Hallo tis between tide-marks. An example has the palps soldered in
the middle line, whilst the frontal tentacles are absent. Another from Norway shows
the reproduction of a tail.

The Lycoris cirrkosa of Risso * (1826) appears to be allied to Nereis Darner Mi, with
long tentacles and tentacular cirri, and may be that form.

1 < Hist. Nat. Europ./ t. iv, p. 417.

310 NEREIS DUMERILIL

So far as can be observed the Nereis Beaucondrayi of Audouin and Edwards 1 comes
nearest this species.

The Lycoris Dumerilii of H. Rathke 2 (1837) probably refers to this form, though the
figures are imperfect.

The Nereis agilis of Keferstein 3 (1862) appears to come nearest this species, though
the figures show certain differences, probably due to the artist.

For similar reasons to those already mentioned under Nereis Schmardaei it is
unnecessary to retain Malmgren's genus Leontis (1867) for this form.

The same author 4 pointed out the relationship of Ip/iiuereis fucicola, OErsted
(Heteronereis fucicola, Auct.), with N. Dumerilii, viz., a sexual stage in its life-history.

Ehlers (1868) gives a careful description of this form both in its atokous and
epitokous conditions, and in the different sexes. In the epitokous male he describes the
cirri of the first four feet as enlarged at the base, the change of the feet occurring on the
sixteenth, in the female on the twenty-first. The cirri in the female are simple ; those
of the male furnished with papillae or warts.

Claparecle (1868) describes Heteronereis Malmgreni, a pelagic Nereid from the surface
of those currents which occur between Capri and Sorrento. The tissues are most delicate
and the agility great in both males and females. The species is probably identical with
the present. He gives fifteen segments to the anterior region (his thoracic) in the
male, and twenty-two in the female. He draws attention to the ventral direction of the
palpi, which are thus invisible from the dorsum, and notes that the form differs from
II. CErstedi, De Quatrefages, from Palermo, in so far as the median tentacle in the latter
is biarticulate and the form of the foot different. Further investigation, however, is
needed.

Moquin-Tandon 5 found at Marseilles a hermaphrodite Nereid, which he termed Nereis
massilie?isis, with spermatozoa and ova floating freely in the body-cavity. This apparently
is the hermaphrodite form of N. Dumerilii, which occurs in a membranous tube amongst
Ulvas. He also found two under the same circumstances loaded with eggs only.

Claparede (1870) cites this form as one of the most puzzling of the group, for whilst
forms of 13 — 14 mm. may be ripe, others of 50 — 60 mm. in length give no trace of sexual
maturity. Again, at different epochs of their existence the males show sperms of
totally different form. In his elaborate study of the Nereid and Heteronereid stages of
this species he shows that there are five kinds of bristles in the Nereid, whereas in the
pelagic Heteronereid there are but three, viz., the awn-like homogomph, the falcigerous
heterogomph, and the sabre-like heterogomph. The falcigerous homogomph only appears
at the twentieth segment. He has met with individuals possessing all the characters
of Heteronereicls, yet the bristles were Nereid. The presence of peritoneal pigment
(purple, etc.), and the homogomph bristle in the inferior fascicle of the dorsal division of

1 ' Annel./ p. 192, pi. iv, figs. 1—7, 1834.

2 ' Fauna der Koym./ p. 419, Tab. vii, figs. 3, 16, and 17, 1837.

3 ( Zeitschr. f. wiss. Zool./ Bd. xii, p. 97, Tat*, viii, figs. 8—11, 1862.
i < Annul. Polycli./ p. 59, April, 1867.

5 'Compt. Rend./ April 12th, 1869, t. lxviii, p. 869; 'Ann. So. Nat./ 1869, p. 134; and < Ann.
Nat. Hist./ 4th ser., vol. iv, p. 73, 1869.

NEREIS DUMERILII. 311

about the twentieth segment, indicate the transformation. Careful examination of the
interior of the feet discloses in these the flabelliform pelagic bristles in process of forma-
tion. The spine of the superior division in the abdominal feet shows an enlarged base
like a spatula for the attachment of the muscles, which are much longer in the Heteronereid.
He points out the increase of the cutaneous glands, especially the verticillate glands of
the Heteronereid. The pigment at the moment of transformation undergoes marked
changes. In the Heteronereid the cutaneous pigment becomes diffuse and disappears,
except in the abdominal region, where brownish transverse lines occur. The peritoneal
pigment undergoes resorption, except in the head and the basal segments of the tentacles
and palpi. A brown line also occurs in the mid-abdominal region as a new development,
for it is absent in the Nereid. The blood-vascular system shows corpuscles (nucleus and
coat of protoplasm) in the Nereid, and the dorsal vessel has valves in pairs. These are
not seen in the Heteronereid. He also found contractile vascular casca in various
regions in this species, e.g., in the feet, and a dichotomous arrangement of vessels. These
vessels undergo rapid development during the change of the foot. The muscular fibres
become granular and indistinct in the Heteronereid. The proboscis presents remarkable
variations in its paragnathi, the epigamous condition having small parallel groups arranged
transversely. The maxillae also, he states, show an increase in the posterior or muscular
region in the Heteronereid, and generally seven or eight teeth are present.

Claparede shows a simple, long trumpet-like segmental organ. He says he did not
find large examples of N. Dumerilii with sexual elements. They become Heteronereids.
The sexual forms range from 15 — 35 mm., rarely the latter. The sexual elements in
these develop as in other Nereids, and the ova are 0*41 mm. in diameter. There is no
structural difference (externally) between male and female. In March he found a peculiar
form in the tubes, violet in front and delicate green posteriorly, the latter colour being
due to the development of the reproductive elements. Moreover, in the feet the developing
heteronereid bristles can be seen. The eyes by-and-by enlarge. Peritoneal pigment is
absorbed and other changes occur, as already noted. He truly calls this epigamy.
Some, moreover, are hermaphrodite in the Nereid form, developing ova and sperms,
as Mecznikow found at San Remo and Villefranche-sur-mer. The colour was violet or
"rosy vinous." Moreover, no transformation was apparent in December, January, and
February. This is probably the form found by Moquin-Tandon at Marseilles, 1 though
the foot, as Bgured by Mecznikow, shows certain differences. So far as known it is still
a question whether a specimen which has borne sexual elements as a Nereid can after-
wards become a Heteronereid.

Racovitza (1896) describes the structure of the cephalic lobe (brain) in this species
and compares it with that in certain other families. He finds three regions, viz., the
sincipital connected with the eyes and antenna?, the region of the palps, and the nucal
(posterior) connected with the ciliated nucal organs.

Mobius (1871), in the ' Pommerania ' Expedition, observed that Nereis zostericola,
Heteronereis fucicola, and Nereilepas variabilis were all forms of Nereis Dumerilii.

Wistinghausen 2 distinguishes three forms of this species : (1) In which reproduction

1 ' Ami. Sc. Nat./ 5 e ser., t. xi, p. 134.

2 f Mitth. Zool. Stat. Neap./ 1891.

312 NEREIS DIVERSICOLOR.

is carried on without metamorphosis. (2) A small pelagic Heteronereid which is found
at Naples in February and March. (3) A tnbicolous Heteronereid form of large size which
occurs in June and July. He gives a careful and well-illustrated account of the segmenta-
tion of the egg and the general development of this species up to the formation of the
embryo. The species is stated also to be viviparous. A hermaphrodite form Avas found
at La Hougue by Caullery and Mesnil ] amidst Lithoth amnion.

De St. Joseph 2 (1895) mentions that the homogomph bristles do not appear in the
dorsal division except in the anterior part of the body ; thus in a ripe female Nereid of
eighty segments they appeared at the forty-first segment ; at the thirtieth segment in a
small example of 30 mm. and sixty segments. The last had a vibratile fossa on each
side of the head between the posterior eyes and the base of the tentacles. He also notes
that the intestine of an example contained Doliocystis nereidis, a gregarine.

The same author 3 more recently describes a new species (0 era toner eis punctata) from
St. Raphael, the foot of which approaches that of N. Dumerilii somewhat closely.

6. Nereis diveksicolor, 0. F. Midler, 1771. Plate LII, figs. 4 and 4a; Plate LX, figs. 11
and 11a — head; Plate LXXII, figs. 5-5 J — feet; Plate LXXXI, figs. 5-5 & —
bristles.

Specific Character*. — Head somewhat triangular and mottled with brownish pigment.
Tentacles about a third the longitudinal diameter of the head. Palpi in life end in tapering
extremities. Eyes of moderate size situated far back. Tentacular cirri of moderate
length, the dorsal of the second pair being longest, and considerably exceeding the
diameter of the body. Body 3 — 4 ins. in length, and of 120 bristled segments. Peristomial
segment about twice the breadth of the succeeding. Tail terminating in two caudal cirri.
General colour yellowish-brown, dusky anteriorly and greenish at the sides — from the feet,
which are also vascular. Proboscis with two brownish maxillae translucent at the base ;
teeth five to seven. Paragnathi are small. Dorsally, in extrusion, the maxillary division has
a band (II) of small teeth on each side and one or two (I) in the middle line. Ventrally at
the base of each jaw is a slightly curved belt of minute teeth (IV), narrower in front,
broader behind, and directed nearly longitudinally. Between them is a broad belt (III)
of small teeth, formed in some of a series of longitudinal rows of about three teeth, but in
others they are less regular. The proximal segment has dorsally a group (VI) of small
paragnathi (five to eight in number) on the eminence in front of each palpus. Ventrally
a basal belt (VII and VIII) of minute teeth stretches from side to side, but with a gap
between it and the dorsal groups. The first foot is distinguished by the three long, pointed
conical lobes, the indistinct setigerous process, and the short cirri. The dorsal lobe
increases to the tenth foot, and is also of considerable size at the thirty-seventh. Both
setigerous lobes become prominent. The ventral lobe soon diminishes. Foot generally
presents an uneven outline when viewed from above, the second and third lobes being

1 c Ann. d. Univ. de Lyon/ Bd. x, fasc. xxxix, 1898.

2 ' Ann. Sc. Nat./ 7 e ser., t. xx, p. 214.

s Ibid., 9 e ser., t. iii, p. 219, pi. iv, figs. 90—93, pi. v, figs. 94, 95.

NEREIS DIVERSICOLOR, 318

anterior and the bristle-tuft posterior. In the inferior setigerous lobe the bristle-tuft
emerges between the smaller anterior and the larger posterior process. Posteriorly the
foot is less complex, and the ventral lobe diminishes. Bristles homogomph and hetero-
gomph, the latter with a small club and a slight differentiation at the tip.

Synonyms.

1767. Scolopendra marina, Linnaeus. Syst. Nat., ed. 12, t. i, pt. 2, p. 1064.

1776. Nereis diversicolor, 0. F. Miiller. Prod. Zool. Danica, p. 217, No. 2624.

1780. „ „ Fabricius. Fauna Greenland., p. 291.

1791. „ versicolor, Linnaeus. Syst. Nat., Gmelin, ed. 13, i, pt. 6, p. 3115.

1800. Die bunte Nereide, 0. F. Miiller. Naturges. einiger Wiirm-Arten., p. 104, Tab. vi, f. 1 — 6.

1806. Nereis versicolor, Turton's Linnaeus, iv, p. 86.

1825. „ „ De Blainville. Diet. Sc, Nat., xxxiv, p. 436.

1827. „ „ Bruguiere. Encycl. Meth., Yers. i, p. 134, Taf. 55, f. 1—6.

1828. „ „ De Blainville. Diet. So. Nat., lvii, p. 471.

1829. Lycoris viridis and margaritacea, Johnston. Zool. Journ., iv, pp. 419 and 420.
1834. „ margaritacea, idem. Loudon's Mag. Nat. Hist., vii, p. 230.

1839. Nereis pelagica, idem. Ann. Nat. Hist., iii, p. 290, pi. vi, f. 1.

1840. „ brevimanus, idem. Ibid., v, p. 170, f. 1.
„ „ viridis, idem. Ibid., v, p. 171, f. 2.

1843. „ diversicolor, CErsted. Grcenl. Annul. Dorsibr., p. 24 (176).
„ „ „ idem. Annul. Danic. Consp., p. 23, f. 66, Q8, 73.

„ „ ,, idem. Kroyer's Nat. Tids., p. 118.

„ „ Sarsii, H. Rathke. Beitrage z. Fauna Norweg., p. 161, Tab. viii, f. 6 — 8.

1847. „ d&pressa, Frey and Leuckart. Beitrage z. Kenntniss wirbell. Thiere, p. 156, Taf. ii,

f. 10, 12.
1851. „ diversicolor, G-rube. Fam. AnneL, pp. 49 and 126.

1853, „ versicolor (fulgens), Dalyell. Pow. Creat., ii, p. 153, pi. xxii, f. 6 — 8 (?).
1856. „ diversicolor, Max Schultze. Bntwickl. Arenicola u. Kiemenwur., p. 214 (Halle), 4to.

fi „ Sarsii, De Quatrefages. AnneL, i, p. 118.

1865. „ diversicolor, idem. Ibid., i, p. 508.
„ ,, brevimanus, idem. Ibid., p. 510.

99 „ brevimana, Johnston. Cat. Worms Brit. Mus., pp. 147 and 341.

w „ viridis, Carrington. Annel. Southport, p. 8 (183).

1867. Hediste diversicolor, Malmgren. Annul. Polych., p. 49, Tab. iv, f. 28.
„ Ehlers. Borstenw., ii, p. 554, Taf. xxii, f. 5 — 9.

„ Grube. Arch. f. Naturges., Bd. xxxvi, p. 311.

„ Mobius. Jahresb. Comm. deutsch. Meere, 1873, p. 112; and 1875, p. 168.

„ idem. Die Zweite deutsch. Norpol., p. 253.

„ Marenzeller. Sitz. d. k. Akad. Wiss., Wien, 1 Abth., p. 60.

Hediste „ Malm. Grofceb. vet. o. vitt. Handl., xiv, p. 83.

„ Mcintosh. Ann. Nat. Hist., ser. 4, vol. xiv, p. 199.

„ idem. Invert, and Fishes St. Andrews, p. 122.

Lenz. Jahresb. Comm. deutsch. Meere, iv, Anhang., p. 12.
„ Mai'enzeller. Op. cit. (Sudjap. AnneL), p. 122.

„ Tauber. Annul. Danic, p. 98.

Leslie and Herdman. Proc. B. Phys. Soc. Edin., vol. vi, p. 274.
Levinsen. Vidensk. Meddel. Foren. Kjobenh., p. 236.

97

1868.

Nereis

1870.

))

1873.

>>

1874.

>>

}>

1878.

y>

1879.

})

>>

>>

1881.

Hediste

1883.

Nereis

1886.

)}

1889.

>>

1890.

Hediste

i>

Nereis

a

y>

1891.

»

1893.

» '

1902.

>)

1904.

))

1905.

))

1906.

Hediste

1908.

Nereis

314 NEREIS DIVERSICOLOR.

1885. Nereis diversicolor, Cams. Fauna Medit., p. 221.

Schroder. Anat.-liistol. Untersnch., 1886, 8vo. (Ratlienow).
„ Menclthal. Untersuch. Mollnsk. u. Annel., iv, p. 8, pi. i, f. 1—9

(Konigsberg) .
„ Malaquin. Annel. Bonlon., p. 31.

Gh'ard. Bull. Sc. Fr. et Belg., t. xxi, p. 275.
„ idem. Ibid., t. xxii, p. 78.

(Hediste) diversicolor, Horn el. Trans. Biol. Soc. Liverp., vol. v, p. 241.
diversicolor, Levinsen. Yidensk. Ud. ' Hauchs/ p, 329.

„ Mcintosh. Ann. Nat. Hist., ser. 7, vol. x, p. 256.

„ Allen. Journ. M. B. A., n.s., vol. vii, p. 224.

Graeffe. Arbeit. Zool. Stat. Triest, xv, p. 322.
Bohn. Ann. Sc. Nat., 9 e ser., t. iii, p. 86.
Horst. Tijdschr. d. Nederl. Dierk. Yereen (2), Dl. xi, p. 139.

Habitat. — Abundant on various parts of the coast near high- water mark from the
northern to the southern shores of Britain and Ireland. Thus, at St. Andrews, it
frequents clayey ground near the shore-bridge in water that must frequently be brackish.
At Perrelle Bay in Guernsey it inhabits holes which it apparently bores in the peat not
far from high-water mark, the surface being studded with hundreds of holes of various
sizes. It is also tossed on shore at St. Andrews after storms, which dislodge it from its
tunnels. It occurs at Plymouth, where the density of the water is low (Allen), and
Benham found it in brackish water at Bembridge, Isle of Wight.

A species very generally distributed on the shores of the North Sea, the Baltic,
the Channel, and other European coasts, as well as in the Mediterranean. Japan
(Marenzeller).

Read (Plate LX, fig. 11) somewhat triangular and mottled with brownish pigment.
Anteriorly the apex of the triangle terminates in two short tentacles, which are about a
third the longitudinal diameter of the head. The palpi end anteriorly in tapering
extremities (in life). The eyes, which are of moderate size, are situated far back, the
posterior pair being near that border of the head, the anterior a little in front and wider
apart. The tentacular cirri are of moderate length, the longest, as usual, being the dorsal
of the second pair, which in life is considerably longer than the diameter of the body.
The basal segment (cirrophore) is tinted brownish, the cirri being translucent greenish.

Body of 120 bristled segments, and 3 to 4 ins. in length, thick and rounded anteriorly,
thinner posteriorly, where it tapers to a delicate tail with two cirri. It is only slightly
tapered anteriorly. The dorsal surface is rounded, the ventral flattened. The greatest
breadth is about the eighth or tenth segment. The peristomial segment is about twice
the breadth of the succeeding 1 .

o

The general colour is yellowish brown, duskier anteriorly and greenish at the sides,
from the feet. The dorsal blood-vessel makes a sinuous streak from the anterior to the
posterior extremity, and the vascularity of the feet also gives a tone to the greenish sides.
In some from Southport (Dr. Carrington) the colour ranged from dark green to orange.
De St. Joseph describes French examples as having a longitudinal brown line on each
side of the median dorsal vessel.

The proboscis (Plate LX, figs. 11 and 11 a) has two brownish maxillae, translucent

NEREIS DIVBRSICOLOE. 315

at the base, and with five to seven teeth. The paragnathi are small. Dorsally, in
extrusion, the proximal segment has a group of small paragnathi (VI), five to eight in
number, in front of each palpus. The maxillary (distal) segment bears a band (II) of
similar small teeth slanting obliquely outward on each side, and one or two in the middle
line (I). Ventrally (Plate LX, fig. 11 a) a belt of minute teeth (VII and VIII) stretches
from side to side of the basal division, but with a gap between it and the dorsal groups.
At the base of each jaw is a slightly curved belt (IV) of similar minute teeth, narrower in
front, broader behind, and directed nearly longitudinally. Between them is a broad belt
(III) of small teeth, formed in some of a series of longitudinal rows of about three teeth,
in others less regularly grouped.

Fragments of peat and Alga3 occurred in the alimentary canal of one from
Lochmacldy.

The first foot (Plate LXXII, fig. 5) is distinguished by the three long, acutely
conical lobes. Dorsally is a short cirrus, not reaching the tip of the dorsal lobe,
which forms in outline an isosceles triangle. There is no trace of a dorsal setigerous
lobe. The next lobe is fused at its base with the ventral lobe, pushing the inferior
setigerous lobe to the front. Both form acute cones. The ventral cirrus is short and
filiform. Dorsally the setigerous process bears bristles with long tapering tips, and
ventrally those with shorter tips, both furnished with well-marked spines. The shorter
bristles have a blunt hook at the tip.

At the tenth foot a view from above shows that the second and third lobes are
anterior and the superior bristle-tuft posterior. The inferior setigerous lobe has its
larger horn posterior, the bristle-tuft being between it and the smaller anterior horn.
The foot thus presents an uneven outline when viewed vertically. Laterally the foot
shows a greatly developed dorsal lamella of a triangular outline (Plate LXXII, fig. 5 a),
with the short cirrus at its base. The two lobes beneath seem to be associated with
the superior setigerous lobe. Both project on the anterior face of the foot, the bristles
being behind. The bifid condition of the inferior setigerous lobe diverges from the
foregoing in so far as the one (smaller) lobe is anterior and the larger posterior, the
bristle-tuft passing out close to the former. The ventral lobe is triangular or lanceolate
in outline, and the ventral cirrus reaches only a little beyond its middle. The bristles
of the dorsal lobe all have long, tapering, and finely spinous tips (Plate LXXXI, fig. 5).
The upper bristles of the inferior setigerous lobe are similar in structure, and so with
the upper bristles of the lower tuft, whilst the lower bristles have shorter tips, with
spinous edges and a hook at the tip, one of those with a longer tip being shown in
Plate LXXXI, fig. 5 a, and one with a shorter tip in the same plate, fig. 5 b.

The thirty-seventh foot (Plate LXXII, fig. 5 b) presents a greater elongation and a
more pointed condition of the superior lobe, an increase in size of the third lobe, a fusion
of the elements at the tip of the inferior setigerous lobe, so that the outline is simplified,
and a considerable diminution of the ventral lobe — formerly so conspicuous. The bristles
follow the same arrangement. The forms with the short tips (inferior edge of the ventral
series) show a differentiation of the club at the tip (Plate LXXXI, fig. 5 b).

The posterior feet (e.g., sixty-third) diminish much in size, and there is a fusion of
the superior setigerous lobe with that below it. The inferior setigerous lobe is very

316 NEREIS DIVERSICOLOR.

prominent as a single process. The ventral lobe is minute, and the ventral cirrus is a
small filiform organ behind the ventral edge of the foot.

Habits.— As the early authors observed, this Nereid burrows in the clayey mud often
in estuaries where the water must sometimes be brackish. Moreover, the entrance of
sewage into the water does not seem harmful to it, and the dark mud is often odori-
ferous, yet there it lives and breeds. Horst l gives some interesting notes and references
on this subject, quoting especially the experiments of Ferroniere and Metzger.

Mr. Goodrich 2 gives an account of a "new dorsal organ in every segment," except a
few after the first and before the last; and consisting of a pair of large ciliated patches
of coelomic epithelium present in most species of the Nereids. He considers it a genital
duct not fully developed. He also notes that the nephridium consists of a compact mass
perforated by a convoluted canal. The nephrostome has long ciliated processes com-
municating with the anterior by a short duct on the ventral surface (nephridiopore).

Reproduction. — Various statements have been made about the reproduction of this
species. Thus Max Schultze 3 refers to the occurrence of the young in the body-cavity,
having apparently considered Koch's statements in regard to Marphysa sanguined as
correct. He describes them as pea-shaped, reddish-yellow embryos, with mouth, rudi-
mentary digestive system, two eyes, and small pits or furrows (" Lochern ") under the
feet.

Cuvier and Grube, again, thought Arenicola was hermaphrodite, and Ratlike placed
Amphitrite in the same category, though each might be more or less male or female. An
interesting resume of the views of the period is given by Frey and Leuckart. 4 Claparede
and Mecznikow 5 give an account of Cirraiulus chrysoderma, Clpde., an adult of which
presented at each side in the median region of the body ovoid sacs, which were at first
thought to be parasitic Crustacea. They proved, however, to be young Cirratulids, one
figured by the authors showing an elongated body, two eyes, two pairs of filiform
branchiae, with several bristles and hook-bearing segments behind, and a fairly complete
alimentary apparatus. A. Krohn, 6 again, describes from Nice a viviparous Sijllis
(S. vivipara) allied to S. Armandi, only the new form has simple tips to the terminal
pieces of the bristles. The enclosed young form has twenty-three segments, and is like
the adult.

In F. M. Balfour's 'Embryology' 7 it is stated that "a few forms (e.g., Eunice
sanguinea, Si/llis vivipara, and Nereis diversicolor) are viviparous." Considerable reliance
in more recent times has been placed on the observations of Mendthal on the supposed
hermaphroditism of Nereis diversicolor. This author seems to have been attracted to the
subject by the previous work of Schroder, 8 but beyond the general and incorrect view

i l Tijdschr. d. Nederl. Dierk. Yereen ' (2), Dl. xi, p. 142.

3 ' Quart. Journ. Micr. Sci.,' April, 1893.

8 ' Entwickehmg Arenicola piscatorum, etc./ p. 214 (Halle, 1856).

4 ' Beitr. z. Kenntniss wirb. Thiere/ p. 82 (1847).

5 < Zeitschr. f. wiss. Zool./ Bd. xix, p. 192, sep. abdr., p. 30, Taf. xiv, f. 4.

6 l Arch. f. Naturges./ Bd. xxxv, p. 197, 1869.

7 Vol. i, p. 319.

8 ' Anat.-histol. Untersuch. von Nereis diversicolor ,' 0. F. M., Rathenow, 1886, p. 35.

NEREIS DIVEBSICOLOR. 317

that in most Polychasts the fertilization and development of the eggs occurs in the
interior of the mother, there seems to be very slender foundation in fact. It is true
Schroder states that in sections he has found the morula-stage of N. diversicolor in
the body-cavity, but the complex condition of things in this species before, at, and after
reproduction might well lead to misinterpretation. Schroder, at any rate, could not find
the later stages in April and May. Mendthal described the species from the Bay of
Pillau as carrying both eggs and testes in June, and he was the more inclined to believe
in its hermaphroditism since Schroder met with only three males out of forty-eight
examples. He places and figures the testes in the dorsal region as pear- or flask-shaped
masses to the exterior of the dorsal longitudinal muscles. The supposed testes, therefore,
occupy the same position as the outer limb of Dr. Goodrich's dorsal organ, and it may be
that the cilia were a source of misapprehension. On the other hand, he describes the ova
as developing at the bases of the feet, the figure representing them in the position of the
segmental organ.

De St. Joseph, however, could not corroborate the foregoing observations. Gravier,
on the other hand, considers that hermaphroditism and viviparity may exceptionally exist
in this form, and in the ' Cambridge Natural History ' it is stated that viviparity is
characteristic of it, whilst Goodrich 1 quotes it as a viviparous hermaphrodite along with
" Salmacina Dysteri and Pomatoceras triqueter."

The annelid is very common on many parts of the British shores, burrowing in clay,
sand, mud, peat, and similar media, and at St. Andrews it abounds at the upper end of
the harbour amidst muddy clay on the sides of the Kinness Burn which enters there.
Careful observations were carried out lately to test the condition as regards repro-
duction ; yet though both hermaphroditism and viviparity have been discountenanced,
it has not been possible to find the worm either discharging from its tunnel in the mud
its eggs and sperms or becoming pelagic at maturity and thus dispersing the sexual
elements. So far as can be observed, the former, perhaps, seems to be the more likely,
though no certainty exists on this head.

In the middle of October the majority of the examples — both large and small — are
females with fairly developed ova, which are almost visible to the naked eye and easily
under a lens. In these the vascularity of the feet has slightly increased, but no change
in the lobes of the feet or in the bristles has occurred. Amongst the ova in the ccelomic
space are numerous pale granular cells, apparently modified perivisceral corpuscles. Very
few males are obtained at this time, and these for the most part are undeveloped.

In November comparatively few ova are attached to the ventral plexuses, most
being free in the perivisceral space. On the other hand, the vascular plexuses in the
region of the ciliated organ are laden with dense groups of rather coarsely granular cells,
sometimes in lobular masses, and it is considered that these pale cells are associated
with the growth and maturation of the ova in the ccelomic space. The ova increased
considerably in size towards the end of the month.

In transverse section 2 the body of the annelid at this time differs little from the type

1 ' Journ. Linn. Soc./ vol. xxviii, p. 107.

3 I am indebted to Dr. Tosh, Assistant Professor, for valued aid in making these sections and in
other respects.

es

318 NEREIS DIVEES'ICOLOR.

except in the presence of ova, which appear most abundantly, in the sections, at the
bases of -the feet and extending into their lobes. The number in the cceloin is not lai
many, in all probability, having fallen out. The dorsal and ventral longitudinal muscl
show little change, the pennate fold of the latter being well developed. The oblique
muscles pass at each side slightly below the nerve-cord to be attached to the basement-
membrane ; and at least three neural canals are present, the larger lateral having a
coagulable fluid internally, and each runs along the outer border of the nerve-trunk
between the pairs of ganglia. The median or dorsal canal, again, appears to have a
separate strand in the interganglionic region, and when it reaches the ganglia it splits
into two trunks in certain sections. The muscles of the bases of the feet and of the
bristles are also strong. The wall of the alimentary canal appears to be of normal
thickness.

The annelids remained very much in the same condition during December, the great
majority of those examined being females, so much so that it was at first considered
probable that, as authors had stated, a complex sexual condition existed (e.g., alternate
development of the reproductive elements) ; but as a few males were still met with and
no trace of an intermediate stage occurred, such complexity could not be proved. This
month the only difference in the sections of the females is the increase in the size of the
ova, the abundance of the ccelomic corpuscles, and the distension of the body-cavity
and the bases of the feet, so that the muscles of the wall are stretched. The vascularity
of the outer surface of the gut also appears to be increased. The segmental organs
show no feature of note. In the males the perivisceral cavity and the bases of the feet
contain dense masses of translucent granular cells, the large nuclei of which stain deeply
with eosine.

The great increase of the large granular cells in the ccelomic cavity in January
is a feature of moment, especially in those females in which the ova are small. The
enormous masses of these cells distend the bodies of the females and they probably
increase by division, each being filled with spherules. In glancing at the living annelids
a pale, or greyish-green hue characterizes the posterior region of the body in the
females; whilst the anterior segments have their vascularity increased, the dorsal vessel
of the foot and its branches especially being distinct. When the posterior region is
punctured, the masses of ova have a pale greenish colour as in the previous months,
and are similarly unfertilized. As many have discharged ova, it would appear that
fertilization is external as in allied forms.

The sexes are not always distinguished by colour, both males and females being
greenish or dull yellowish, though the males are often paler. No change in eyes, feet, or
bristles is apparent. In many an opaque, dull whitish condition occurs in the anterior
feet, which are filled with the large granular perivisceral corpuscles. Moreover, almost
all have a touch of white at the base of the dorsal cirrus— from one end of the body to
the other, a similar touch occurring at the tip of the cirrophore of each tentacular
cirrus.

Mendthal x figures two examples, one of the general brownish-orange hue of the

1 ' Untersucli. u. d. Mollusken u. Amieliden des frischen Haffs/ p. 9, Taf. figs. 1 and 2 (Konigs-
berg, 1889).

NEREIS DIVERSICOLOR. 819

ordinary type, and the other of this colour in the anterior fourth, whilst the rest is of a
grass-green tint. He considers that the latter (green) colour is due to a diet of green
Algae, a condition which is not in accordance with the observations at St. Andrews, where
the greenish hue appeared to be the result of the presence of masses of more or less ripe
ova. There is no reason, however, why a diet of JJlva or other green Alga should not
tint the digestive canal green.

In the male the sperm-cells fill the various cavities even to the bases of the dorsal
cirri throughout January, but though the body- wall is frequently distended, no degenera-
tion of the muscular bands can be observed. In the females the size and abundance of
the ova in particular examples vary, but, as a rule, the granular ova with a slightly
stained nucleus and a more deeply stained nucleolus are more or less advanced towards
maturity. Occasionally an example with few and small ova appears. In some of these
the muscular walls of the body are contracted, and very few ova, and these of small size,
occur in the perivisceral space. More frequently they are found in the feet. Masses of
ovigerous tissue are attached to the vessels near the bases of the latter ; and cellular
masses (it may be parts of the dorsal organ) with boldly stained nuclei pass upward at
the bases of the feet within, as well as without, the lateral lobe of the dorsal longitudinal
muscles, and in specimens having a considerable number of large ova similar cells pass
into all the spaces of the feet. These loose coelomic masses probably represent the
cells alluded to in the living forms.

In February the increase in size and the division of the sperm-cells form the most
noteworthy feature, but no free sperms are seen. In the other sex the ova in many have
considerably increased in size, and from the appearance of the annelids a large number of
ova must have been discharged.

Signs of rapid division of the sperm-cells characterize many males in the middle of
March, and towards the end of the month the perivisceral cavity forms a vast reservoir
for the male elements. The body-walls are stretched, the muscular layers diminished, and
the alimentary canal collapsed. The tubes of the segmental organ have also enlarged, so
that sperms could readily find issue by the segmental papilla. Though the nuclei of the
walls of the organ and its tubes are everywhere distinct, no trace of sperms was at any
time found within it, so that they probably escape by rupture of the body-wall, as in
allied forms.

The ova in the various examples throughout March appear to attain fall size and
maturity, and they are remarkably large, viz., about '1524 mm., and they are probably
shed by rupture of the bod}- wall, the vast numbers set free, even from a limited area,
affording an indication of the almost illimitable resources of nature in the sea. Whether
the sexes discharge their elements in situ or by a terminal pelagic stage could not be
ascertained. None were captured in the tow-nets dragged, by day and by night, up
stream at ebb-tide, and no indication of any change in the structure of the feet, bristles,
or eyes, and no shrivelling of the posterior region of the body as in the very interesting
Ceratocephalus Osawai of Akira Izuka, 1 common in the estuary of the Sumicla River in
Tokyo, occurred. If one may hazard a conjecture, it is probable that the sperms and ova
are discharged on the sites inhabited by the annelids, and the larvae by-and-by carried
1 ' Journ. Coll. Sc. Univ. Tokyo/ vol. xvii, art. ii (1902).

320

NEREIS DIVERSICOLOB,

seawards. There is no doubt, however, that, without change in the condition of the feet
and bristles, the species is an apt swimmer, progressing through the water swiftly in
graceful screw-coils.

Towards the end of May (29th) and for some time previously signs of degeneration
appear in certain ova, as if they were in process of absorption. These ova are smaller,
minutely granular, and with larger oil-globules. The larger ova have lost the germinal
vesicle and spot, and transmit light more readily than formerly. Nothing is seen to
suggest the view that certain ova are undergoing development, for, when kept in vessels
both of sea- water and fresh water, they are rapidly disintegrated.

Some females at this elate have shed all their ova, and are of a pale brownish-yellow
colour, occasionally with a minute dusting of yellow grains along the dorsum.

Having failed to secure the early larvae by any of the methods alluded to, masses of
the clayey mud with the adults in situ were brought from the banks of the Kinness

Fig. 72.— Maxilla of
larval N. diversicolor.

Fig. 71. — Post-larval Nereis diversicolor
with three pairs of bristles.

Fig. 73. — Post-larval N. diversicolor
with five pairs of bristled feet.

Burn near the harbour towards the end of May, and a strict scrutiny was made of the tubes
and the mud lining them. Numerous post-larval forms were thus obtained, but no trace
of trochophores or other early stages. Whether these stages had been passed in such an
environment before the examination, or whether the later larvae to be subsequently
described had settled in the mud of the tubes after a pelagic stage, is yet undetermined.
The youngest stage observed has three bristled segments (Fig. 71), a head with two
short palpi, and two short frontal tentacles. The eyes are imperfectly differentiated,
consisting of an irregular group of black pigment-granules. A tentacular cirrus
occurs on each side opposite the eyes. The three feet are nearly alike, each with a
bristle-tuft and a small dorsal cirrus, or the first is rudimentary. A minute caudal
cirrus is at each side of the posterior end. Between the last foot and the pygidium is
a projection, indicating a segment. The proboscis has a pair of jaws each with three
long teeth, including the anterior fang (Fig. 72). The opaque part of the gnt extends
over the last two bristled segments. The minute bristles already present the typical

NEREIS DIVERSICOLOR. 821

structure of cameratecl shaft and homogomph articulation of the end of the shaft, and are
in two bundles in each foot.

No younger form has yet been found amongst the mud or the adults, so that it is
probable that they settle down at this stage. Moreover, no pelagic larvae appeared in the
vessels ; but as the post-larval forms with three bristled segments occurred in the mud of
the tubes of the adult, it would appear that all do not wander. Yet pelagic Nereids with
six bristled segments occur in the tow-nets of the period, but whether of this or other
species is yet uncertain.

When the post-larval form has four bristled segments the head has two eyes on each
side, placed close together and posteriorly, a pair of short frontal tentacles, short palpi
which present no distal articulation, a pair of tentacular cirri, and two short anal cirri.
Behind the head, which bears the tentacular cirri, is a region with only bristles on each
side; a foot with a large bristle-tuft and a minute dorsal cirrus follows, and then a second
foot of similar structure. A rudimentary foot comes next, with a bristle-tuft on one side.
A rudimentary ventral cirrus occurs on each foot. A pair of minute jaws having two
teeth behind the anterior fang is found in the proboscis. Vast swarms of Infusoria (like
monads) frequent the moist and odoriferous mud in which the young Nereids live.

The post-larval form of the 28th May (Fig. 73) has a head with two comparatively
large frontal tentacles and two stumpy palpi with very short terminal segments that only
occasionally are visible, two pairs of tentacular cirri, the last with a spine in the dorsal,
five pairs of distinctly bristled feet, with more evident dorsal and ventral cirri and slightly
brownish spines, besides two rudimentary ones behind and a large pygidium with two
caudal cirri. The cirrus of the first bristled segment is elongating, but the bristles are
shorter than in the succeeding foot. The mouth is a wide aperture, followed by the some-
what ovoid proboscis, which is armed with two translucent jaws having three prominent
teeth which are proportionately longer than in the adult. The proboscis is narrowed
posteriorly and joins a large opaque, yellowish, glandular region, broad and truncated in
front and diminishing posteriorly and terminating in the pale rectal part of the gut and
its dorsal anus. The opaque glandular region stretches from the space between the
second and third feet to the last bristled foot, and shows the groups of oily granules so
characteristic of the glands on the walls of the alimentary canal. Two slightly opaque
ovoid glandular bodies, the segmental organs, lie behind the first complete foot, just in
front of the opaque region of the gut, and active ciliary action is occasionally noticed
in them.

In the post-larval form with six bristled feet the eyes are better defined, and the
palpi, frontal tentacles, and tentacular cirri are larger. The dorsal cirrus of the first
bristled segment is the longest in the animal, but there is no spine. The spines are
darker, the posterior end remains bifid, and three teeth occur behind the anterior
fang in each jaw.

When seven bristled feet are present the caudal cirri are considerably longer, the
last foot (seventh) having a few short bristles, a short dorsal cirrus, two short spines, and
a minute rudiment of a ventral cirrus. Behind is an indication of the eighth foot as a
lateral projection, with a minute papilla representing the dorsal cirrus. Internally is the
tip of a minute spine, but no bristles. The ventral of the first pair of tentacular cirri

98

322 NEREIS DIVBRSICOLOR.

(opposite the eyes) is a short subulate process; that of the next segment is indistinct,
though, with the exception of the caudal cirri, the dorsal is the longest of the series.
The bristles of this segment are always short and in contrast with the next segment,
which shows the segmental organs immediately behind. Opacities in the corresponding
parts of the following feet indicate the early development of these organs throughout.
Traces of a fourth tooth behind the great anterior fang of the maxilla are visible. This
description might also apply to a post-larval form with eight pairs of bristled feet
(Tig. 74), the latest stage observed towards the end of May.

|

^ __qDaL.

iiv.

~pr.

..-:;.'■-
■Err so

q

7

i '"t

.'.■■: •

-T9

Fig. 74. —Young N. diver sicolor with eight pairs of bristled feet and four eyes. g. Glands of alimentary canal,
m. Maxillae, pal. Palpi, pg. Pygidium. pr. Proboscis, so. Segmental organ ?.

If the young Nereids of the foregoing stages are left in the vessels beside the adults
the larger examples rapidly disappear. They are probably devoured by the adults, only
the minute stages escaping capture, and in all likelihood they, too, would by-and-by be
found out and captured ; yet, as in the fishes, these checks have little influence on the
permanent abundance of the species.

On the 1 7th July the young forms are found on the same sites, but considerable progress
has been made in development. The head in outline nearly resembles that of the adult,
and is marked by whitish pigment in the younger, brownish in the older, in front of

NEREIS DIVEBSICOLOR. 323

the eyes, which are definitely formed on each side, the anterior pair being somewhat larger
and furnished with lenses. The frontal tentacles and the palpi are well formed and show
numerous fine palpocils, the mobility of the distal segment of the palpi being noteworthy.
The two pairs of tentacular cirri are now much longer, the dorsal stretching outward like
fine hairs as the animal pushes its snout forward. They retain the proportions to each
other as in the adult, though they have not yet reached full development in any case.
The ventral of each is considerably shorter than the dorsal, but projects clearly on each
side, the first rather longer and thicker at the base than the frontal tentacles. The body
is now elongated, having from sixteen to twenty bristled feet, and is marked along the
centre by a yellowish or brownish streak, often slightly moniliform or zigzag from the
intestinal contents, and a whitish dot is observed in some at the bases of the feet. The
massive and somewhat ovoid proboscis has six teeth behind the great anterior fang, and
the narrow part of the canal behind it is usually firmly contracted (and empty). The
first part of the gut behind the post-proboscidian narrow region is the largest, and for
some distance it is not marked by lateral constrictions, as in the succeeding portion.
Most of the feet anteriorly present a condition approaching that of the adult, though the
dorsal cirrus is shorter. Only the last two feet are devoid of bristles externally. The
caudal cirri are considerably longer. The circulation of the reel blood in the dorsal and
ventral trunks is now evident. The segmental organs extend backward to the posterior
feet, though not quite to the tail.

So far as observed at St. Andrews, therefore, there is no foundation for the statement
that the Scotch representatives are hermaphrodite, and still less that they are viviparous,
as mentioned by Max Schultze, by the i Cambridge Natural History,' and by Gravier.
Max Schultze 1 gives a circumstantial account of finding female examples in April at
Greifswald with ciliated eggs and pear-shaped larva3 in the body-cavity, and he figures
two of the latter, hundreds of which occurred in the cavities at the bases of the feet.
His figures, however, give rise to doubt as to the nature of the ciliated forms, which have
the narrow end of the pear anteriorly with the two eyes at some distance behind it. In
one, indeed, the mouth is indicated behind the eyes as a small radiated disk. The
posterior end of the larva is broad and rounded. In the light of the condition at St.
Andrews doubt arises as to correctness of the interpretation, more especially in view of
the structure of the larva; but it is right to give a margin for possible variations in regard
to internal fertilization under certain circumstances. Yet it seems unlikely, and De St.
Joseph 2 is of the same opinion.

The foregoing interpretation as to the mode of reproduction would seem to be most
in accordance with observation, since on the 28th May masses of clayey mud brought from
the habitat of the species and with the annelids in situ showed many free ova amongst
the mud coating the walls of the tube, and, further, various post-larval examples.

In connection with the reproduction of Nereis interesting experiments have been
carried out by Loeb and Fischer, in which they found that the unfertilized eggs of Nereis
limbata developed into swimming larvae by keeping them for some time in sea- water

1 ' Amer. Journ. Physiol./ vol. ix, p. 100, 1903.

2 ' Ann. Sc. Nat./ 8 C ser., t. v, p. 298, 1898.

324 NEREIS DIVERSICOLOR.

concentrated by the addition of potassium chloride, and then returning them to ordinary
sea-water.

In the British Museum specimens of this species from Bexhill and various parts
of the southern coast are labelled Nereis finibrlata, whilst others which bear the title
N divers loolor pertain to N. cidtrifera, N. pelagica, and AUtta virens.

The Nereis falgens {versicolor) of Dalyell has been placed under this head with some
doubt. The figure might pass for the reddish-brown female of N. pelagica.

This species is largely used for bait in the Channel Islands. Mr. Punnett also found
that at Gosport it was frequently employed for the capture of pollack.

In his original account of this Nereid, 0. F. Miiller (1800) enters into a disquisition
as to the history of this " sea scolopendra," as such forms were termed by Aristotle, and
its connection with the " Rodat."

The Nereis Beaacondrcuji of Aud. & Edw. and Keferstein x seems to come near this
form.

Max Schultze in 1856 refers to the occurrence of the young in the body-cavity of the
mother, but so far as observations in this country go this has not been corroborated. He
seems to have considered that Koch's statements in regard to Marphysa sanguined were
literally true.

Malmgren (1867) constituted the genus Hediste for this species, founding the distinc-
tion chiefly on the trifid nature of the superior division of the foot, but it seems unnecessary
to multiply genera on this basis.

Schroder 2 (1886) gave a general account of this species structurally and otherwise,
and added a note in regard to its reproduction. His observations were made at Kiel.

Mendthal (1889) thought the species was hermaphrodite, the ova being developed
at the bases of the feet, whilst the sperms occurred in the dorsal region external to the
longitudinal muscles. This has not been corroborated at St. Andrews. Like Schroder
he also found Gregarinae in the gut, and the early stages in the muscles. He also refers
to the respiratory function of the lobes of the feet.

An interesting account of the " swarming " and development of Nereis japonica, 3 n. sp.,
is given by Akira Izuka, who, while admitting close resemblances between it and Nereis
diversicolor, bases his distinction on the fact that the paragnathi in the Japanese form in
VII and VIII are arranged in a continuous row, whilst in N. diversicolor they are in the
form of three or four irregular rows, that the falcate bristles of the former are larger than
those of the latter, that the eyes have lenses in the Japanese, and that the breeding habits
of the latter differ from those of Nereis diversicolor. After a perusal of the description in
all its details, however, the very close resemblance of the two forms is apparent, and
though the " swarming" of N. diversicolor has not yet been observed in Scottish waters,
that and other slight divergences may be due to environment, and especially to tempera-
ture and food. It has to be remembered that some forms, e. g., Eunereis longissima, may
or may not have paragnathi, so that future investigations may show that changes of the
nature indicated by Izuka may not have the importance claimed for them.

* 'Zeitsclir. f. wiss. Zool./ Bd. xii, 1862, p. 94, Taf. viii, figs. 1-6, and 12, 1862.
' Anat. histol. Untersuch. von N. diversicolor, Rathenow/ 1886, 1 plate.
' Aimot. Zool. Japon./ vol. vi, part 4, 1908.

NEREIS LONGISSIMA. 325

7. Nereis longisslma, Johnston, 1840. Plate LXI, figs. 1-16 — head; Plate LXXIII,
figs. 1-1 d — feet; Plate LXXXI, figs. 6 and 6 a — bristles.

Specific Characters. — Head small, broader posteriorly, and tapering to a narrow snout,
bearing two tentacles of moderate length. Eyes obscure, in some preparations only
indicated by the duskiness of the thick cuticle in the ordinary position. Tentacular cirri
short. Body elongated, 10 ins. to a foot or more in length. The peristomium is considerably
wider than the next segment. Colour slate-blue in front and finely iridescent. Vent-rally
the surface is lilac. Proboscis dark bluish ; maxillae dark brown, seven teeth. Proximal
and distal regions in extrusion mottled, and marked by grooves into areas. In some no
paragnathi occur, but in an epitokous female the basal segment dorsally has on the usual
eminence on each side four small brownish-black paragnathi (VI). Moreover, on the
ventral aspect of the distal section an irregular dark brownish patch (IV) appears on each
side near the base of the jaws. The first foot has moderately developed dorsal and
ventral cirri, only a little shorter than the elongated and pointed dorsal and ventral lobes.
The intermediate setigerous lobe forms a small papilla. The feet generally are characterized
by three long lanceolate and pointed lobes, a trificl inferior setigerous lobe, and the cirri
are short. The bristles consist chiefly of those with long, slender, tapering tips, but beneath
the upper series (above the spine) in the inferior setigerous process is a group of those
with short falcate tips (heterogomph), longer, however, than those of the previous
species.

In the epitokous form great lobes are developed (e.g., sixty-seventh foot) dorsally, on
the posterior surface of the inferior setigerous lobe, and at the ventral cirrus, whilst all
the other lobes are expanded. Swimming bristles are present. This condition continues
till near the tip of the tail.

Synonyms.

Atohous Form.

J800. Die faserige Nereide, 0. F. Miiller. Naturges. einiger Wurm-Arten., p. 144, Tab. viii, f. 1 — 3.
1825. Nereis longisslma (Saw.), De Blainville. Diet. So. Nat., xxxiv, p. 443.
1850. „ regia, De Quatrefages. Ann. So. Nat., 3 e ser., Zool., t. xiv, p. 339.
1865. „ ,, idem. Annel., i, p. 511.

J} , } edenticidata, idem. Ibid., p. 538, pi. vii, f. 1 and 2.

„ longisslma, idem. Ibid., p. 581.
1869. „ „ Grube. Abh. S.chl. Gesell., 1868-9, p. 100.

1874. „ „ idem. Jahresb. Schl. Gesell., 1873, p. 69.

, Eunereis „ Malm. Goteh. vet. o. vitt. Handl., xiv, p. 84.

1876. Typhlonereis gracilis, Hansen (?). Nyt Mag. f. Naturvid., xxiv, 1 Hefte, p. 4, Tab. iii, f. 6 — 10.
1879. Nereis longisslma, Tauber. Annul. Danic, p. 100.

1883. ,, „ Levinsen. Vidensk. Meddel. Foren. Kjobenh., p. 233.

1890. Ceratonerels longisslma, Malaquiu. Annel. Boulon., p. 28.
1898. Eunereis ,, De St. Joseph. Ann. Sc. Nat., 8 e ser., t. v, p. 304, pi. xvi, f. 88—100,

and pi. xvii, f. 101.
1902. „ >j Mcintosh. Ann. Nat. Hist., ser. 7, vol. x, p. 256.

1904. „ ,, Allen. Jo urn. M. B. A., n.s., vol. vii, p. 224.

)}

Eunereis

1867.

Nereis

})

)>

1868.

j)

1874.

Eunereis

1875.

35

326 NEREIS LONGISSIMA.

Epitokous Form.

1840. Nereis longissima, Johnston. Ann. Nat. Hist., vol. v, p. 178, f. 9.

1843. Heteronereis paradoxa, GErsted. Gronl. Annul. Dorsib., p. 177, f. 50, 63, 64, 66.

1844. Nereis longissima, Thompson. Eep. Brit. Assoc, 1843, p. 273.
1851. „ „ Grube. Fam. Annel., p. 50.

1865. Heteronereis longissima, Johnston. Cat. Worms Brit. Mus., p. 164.

,, Malmgren. Nord. Hafs-Annul., p. 183.

,, idem. Annul. Polych., p. 57, Tab. v, f. 32.

., Parfitt. Trans. Devon Assoc., ii, p. 23 (sep. copy).

„ Ehlers. Borstenw., ii, p. 525.

., Mcintosh. Ann. Nat. Hist., ser. 4, vol. xiv, p. 199,

„ idem. Invert, and Fishes St. Andrews, p. 122.

Habitat. — Tossed on shore after storms in March and October, St. Andrews (E. M.) ;
Peterhead, advanced epitokous female (Mr. John Bain, Fishery Officer) ; Montrose Bay,
epitokous forms (Dr. Howden). A small example in the stomach of a flounder at
St. Andrews (E. M.). Epitokous forms with the lamellae but slightly developed occur
on the West Sands, St. Andrews, at the end of March (E. M.).

Extends to Ireland (Johnston).

St. Magnus Bay, 80 — 100 fathoms (J. G. Jeffreys) ; rather small example.

Ranges to the Mediterranean (' Porcupine,' and Claparede, if his Nereis (Ger atone reis)
guttata refers to the same form). Epitokous form, Greenland (CErsted). North Sea.

Head (atokous) (Plate LXI, fig. 1) somewhat small, broader posteriorly, and tapering
anteriorly to a narrow snout, bearing two tentacles of moderate length. The eyes are in
some preparations obscure, only indicated by a duskiness of the thick cuticle. They
occupy the lateral regions of the broad part of the head, the anterior pair being somewhat
wider apart than the posterior. They are best seen in the epitokous form as slightly
elevated pale areas with what appears to be a cuticular lens in the centre. The tentacular
cirri are short.

Body much elongated — 10 ins. to a foot or more — and having seventy to two hundred
and forty segments, gently narrowed anteriorly so as to give a character to the region,
and again diminishing to a slender tail. It is firm and resistant anteriorly, flatter
posteriorly, especially in the epitokous female. In the latter the dorsum is of a slate-blue
colour, finely iridescent, the corrugated regions at the bases of the feet having a greenish
sheen. The large lamellae of the feet are pinkish from the blood-vessels. Ventrally
the body is lilac in front, then greenish-blue. The first segment (peristomium)
is considerably wider than the succeeding. The body terminates posteriorly in two
long cirri.

The proboscis (Plate LXI, figs. 1 a and 1 b) is of a blackish-blue colour, and probably
increases the dusky appearance of the anterior region when retracted. The maxillae are
deep brown, with seven teeth (epitokous female). 1 The proximal and distal (maxillary)
regions, in extrusion, are mottled and marked by grooves into areas. In some no
paragnathi are present, but in an epitokous female the basal segment dorsally has in front
of each palpus, on the usual eminence, a group (VI) of four small brownish-black

1 Ehlers gives seven to ten teeth.

NEREIS LONGISSIMA. 327

paragnatlii. Moreover, on the ventral aspect of the maxillary section, an irregular dark
brownish patch (IV) occurs on each side near the base of the jaws. That on the left is
the larger, and is distinctly horny (Plate LXI, fig. 1 b).

The first foot (atokous form, Plate LXXIII, fig. 1) has dorsally a large cirrus which
extends considerably beyond the tip of the dorsal lobe. The latter forms a long and some-
what narrow cone. The setigerous lobe beneath appears as a short papilla in lateral view.
The ventral lobe has a similar outline to the dorsal, though its tip is less acute, and the
inferior line longer. The ventral cirrus is large and long, its tip reaching that of the
adjoining lobe. The bristles are apparently all furnished with tapering tips.

At the third foot another lobe appears, probably above that which was dorsal in the
first foot. At the tenth (Plate LXXIII, fig. 1 a) the diminution of the dorsal and ventral
cirri, the presence of a superior setigerous lobe, and the increase in the inferior setigerous
lobe are the chief features. The dorsal lobe is somewhat smaller than the next, the
ventral being considerably shorter than these. The bristles (Plate LXXXI, fig. 6) of the
superior setigerous lobe are of one kind with long, tapering, spinous tips, but, in the
typical foot, amongst those of the inferior setigerous process, are some falcate bristles
(Plate LXXXI, fig. 6 a). These occur as rather stout bristles near the spine, and also
more slender forms at the ventral edge of the series. The falcate forms are not readily
seen in a cursory glance, and had escaped the notice of Malmgren — unless the northern
form differs from the British in this respect.

Little change in the general form of the foot in the atokous condition takes place
posteriorly (to the one hundredth at least) except the increase of the gap between the
second lamella and the inferior setigerous lobe (Plate LXXIII, fig. 1 d), the diminution of
the dorsal and ventral cirri, and to some extent that of the ventral lobe. Toward the tip
of the tail, however, the cirri again become longer, and the inferior setigerous process
increases in size, whilst the ventral lobe is considerably less.

The bristles throughout conform to the same type, though in the dorsal tuft a
differentiation very early occurs, even in the tenth foot, two or three of these having
stronger shafts than the others with more evident cameraa. These bristles, which have
tips, become gradually stronger and more deeply tinted in the posterior feet, until toward
the tail only two very robust bristles and one or two more slender remain in this
division. The bristles of the inferior setigerous process are slender throughout.

In an example of the epitokous female, which had about one hundred and forty-seven
bristled segments, the feet remain of the normal pattern to the forty-seventh, on the ventral
border of which, close to the inner edge of the ventral cirrus, a small papilla occurs.
This in the forty-ninth foot projects as a narrow lamella. Simultaneously a tendency to
the development of a flap or lamella behind the superior setigerous process is observed,
and it increases in the succeeding feet until a large, fan-shaped lamella is formed. It
thus happens that, with the changes in the other parts of the foot, there is at the sixty-
seventh foot, for instance, the following arrangement (Plate LXXIII, fig. 1 c). Dorsally
is a large, thin, fan-shaped lamella, externally touching the comparatively short dorsal
cirrus. The dorsal lamella is a long, semi-translucent, and pointed triangle. The
superior setigerous lobe is opaque, blunt, and bifid, having a dense tuft of the swimmino*
bristles with the spine at the inferior edge. The next beneath is a long, pointed,

328 NEREIS LONGISSIMA.

translucent lobe with four separate frills or lobes at its base ventrally, and it generally
lies in front of the bristles pertaining to the former lobe. There the inferior setio-erous
process bears the large fan-shaped lamella, often with prominent crenations at its outer
border interiorly. In front is the great group of swimming bristles diverging in a fan-
like manner at the tip. The ventral lobe is considerably smaller than the dorsal lobe and
has a prominent rounded border inferiorly. The lamella to the inner edge of the ventral
cirrus is large and fan-shaped, and is joined to a smaller lamella on the outer side of the
lobe. From the connate lobes the ventral lamella extends outward but does not reach
the tip of the ventral lobe. The connate lamellae thus take the place of a cirrophore, a
change which also occurs subsequently in the dorsal cirrus.

At the hundredth foot the dorsal lamella is large, extending upward nearly as far as
the dorsal cirrus, which now projects from the translucent edge of a small process
(connate with the dorsal lamella) which bulges externally. The superior lobe of the
foot is somewhat less, but retains its elongated triangular outline. The superior
setigerous process, its bristles and the long lanceolate lamella beneath, are similar to the
sixty-seventh, and so are the parts situated ventrally. The surface of the foot has
various specks of black pigment, and in the interior are many ova.

A considerable change takes place at the two hundredth foot. The lamella internal
to the dorsal cirrus is now much less, the cirrus seems to be proportionally longer, and
it is less distant from the body and the centre of the foot (by the diminution of the entire
organ). The superior lobe beneath it is smaller and of a lanceolate outline. The superior
setigerous process has amalgamated with the lamella beneath it and the area of the latter
is circumscribed. The inferior setigerous process bears a smaller lamella, still with a
crenate border externally and inferiorly. The ventral lamella is tongue-shaped, with a
straight upper edge and a curved inferior border. The lamellge at the ventral cirrus are
much smaller, but the base of the cirrus is still connected with their border.

Towards the tip of the tail (Plate LXXIII, fig. 1 d) the lamella to the inner side of
the dorsal cirrus has disappeared, a rounded and somewhat thick eminence marking its
position. The dorsal cirrus is long. The superior lobe is more opaque, and the seti-
gerous lobe and its lamella beneath are narrow and long. The inferior setigerous lobe
has only a trace of a thin margin superiorly, and forms a long, narrow, pointed process.
The ventral lobe is tongue-shaped, and the ventral cirrus still retains a frilled lamella
inferiorly and a small process externally. The black pigment-touches continue to the
posterior feet, and so do the ova interiorly.

An elongated Nereid was dredged in the Bay of Tunis, on the 3rd September, 1870,
in the ' Porcupine ' Expedition of that year. In general form and in the absence of
paragnathi it resembles a small Eunereis longissima, though its coloration appears to
differ. The head agrees with that in Eunereis, but no eyes are visible. Slight elevations
devoid of pigment at the sides may indicate them. The tentacles are short, and the
tentacular cirri especially short and slender. The fragment of the body is about 4 ins. long
in spirit, and from the slight diminution at the posterior end it probably extended an inch
or two more. The surface is finely iridescent, and a dull pinkish band (from the intestine)
passes along the centre of the dorsum. The first segment (peristomium) is considerably
broader than the next. In the structure of the feet this form agrees very closely with

NEREIS LONGISSIMA. 329

E. longissima, the only feature being the occurrence of a somewhat regular series of four
heterogomph bristles below the upper group in the inferior setigerous lobe. These
bristles have the same structure as in the Scotch examples. This appears to be a
southern deep-water variety of N. longissima.

The segmental organ of Euneveis longissima has its trumpet furnished with rather
long, ciliated fimbriae, the latter thus contrasting with the lobes of N. cultrifera
(Goodrich).

This species was first described by Dr. Johnston, of Berwick-on-Tweed, and he
noted its great length, absence of horny processes on the proboscis, and the structure of
the feet. It is true De Blainville mentions Nereis longissima, Savigny, but no reference
to this form occurs in Savigny's ' Systeme.'

The Nereis pulsatoria? of H. Rathke 1 apparently comes very near this species, which
extends to the Mediterranean. This view is borne out by the structural features given
in his treatise, ' De Bopyro et Nereide.' 2

Levinsen (1883) observes that the epitokous male has the first thirty-five segments
and the female the first forty-six rings unchanged. It is used as bait for pollack and
other fishes (Malm).

Parfitt (1867) mentions that Mr. Hoarder observed millions of them swimming on
the surface of the water in May in Plymouth Sound.

Ehlers (1868), as in many other cases, simplified the synonymy of this form by an
investigation of the original types.

The Nereis (Cera toner eis) guttata of Claparede (1868) may be a southern variety of
this species which seems to vary considerably in regard to the development of the
paragnathi.

Claparede describes two species closely allied to N. longissima, from Naples, viz.,
Nereis (Cera toner eis) Ehlersiana, and N. (G.) Kinbevgiana, ?i both of which have paragnathi
on the maxillary division of the proboscis in extrusion, none in the basal, yet the first foot
of the latter is identical in outline with the British form, and that of the second is not very
divergent considering the variation in preparations. The variation in the paragnathi of
the British species and the similarity in the feet generally point to a very close relation-
ship between the three forms. Whether Claparede's Nereis {Lept oner eis) glauca, a small
form apparently approaching the heteronereid phase, has any connection with Eunereis
longissima, is an open question. The unarmed proboscis, the character of the feet, and
other features suggest it.

An allied species (N. irritabilis) from the Virginian coast is described by Webster 4
(1879). The mature males and females, after assuming their nuptial dress, are caught
swimming at the surface of the sea, the former in much greater numbers.

It is difficult to make out what the Typhlonereis gracilis of Hansen 5 (1882) can be
except this species.

i < Fauna der Krym/ p. 412, Tab. vii, figs. 1, 4 and 8, 1837.

2 ' Riga3 et Dorpati/ 1837.

3 ' Aimel. Nap./ Suppl., pp. 88 and 89, pi. viii, figs. 2 and 3, 1870.

4 < Trans. Albany Inst./ ix, p. 31, etc., 1879.

5 'Norsk. Nordhav. Expect./ p. 31, Tab. iv, figs. 14—19, 1882.

99

330 NEREIS (ALITTA) VIRENS.

The Leptonereis vasculosa of Griard 2 appears to approach this form closely, and is
also characterized by the feeble armature of the proboscis.

8. Nereis (Alitta) virens, Sars, 1835. Plate LIII; Plate L, figs. 12, 13, and 16— larval
and post-larval; Plate LXI, figs. 2 and 2 a — head, and figs. 3-5 — other parts;
Plate LXXIII, figs. 2-2 6— feet ; Plate LXXXI, fig. 7— bristles.

Specific Characters. — Head typical in outline, with two short tentacles anteriorly.
Eyes four, arranged in a trapezoid, toward the posterior part of the head, the anterior
pair wider apart, a line from each posterior eye sloping inward and backward to the
nuchal bridge. The longest tentacular cirrus is the dorsal of the second pair. Colour
bluish-purple, and in the preparations often presenting a longitudinal median groove, the
ridge on each side running forward to the frontal tentacle. Body from 1 — 3 feet in
length, and If inches in breadth across the feet in a large specimen, slightly tapered
anteriorly, diminishing posteriorly, and ending in an anal segment with two long cirri.
Peristomial segment twice the breadth of the succeeding. Dorsum of various shades of
iridescent bluish-purple, and the lamella of the feet are also finely tinted and iridescent ;
dorsal cirrus larger anteriorly than in those with foliaceous lamellae, and of a deep green.
Ventral surface pale iridescent pinkish, deeper posteriorly, but in some iridescent bluish,
with the belts at the anterior part of each segment of a deeper hue. Proboscis in
extrusion greyish-blue, somewhat lighter than the hue of the body. The maxillas have
six teeth. The maxillary segment dorsally has a triangular group of paragnathi with the
apex in front, on each side behind the jaw, sometimes forming a double row with the
limbs wider apart posteriorly, and two points — one in front of the other — between them.
Ventrally a long, rather irregular double row proceeds in the maxillary division backward
from each jaw, and in the middle is a transverse belt of points, or a larger group in the
central area, and a point or two in the lateral area on each side. The proximal segment
has dorsally on the eminence in front of each palpus one or two paragnathi, and between
them a single point. Ventrally is an irregular double row of paragnathi on the folds,
and a brief interval exists between them and the dorsal points. First foot has a dorsal,
a middle, and a ventral lobe, the setigerous process being attached to the anterior face of
the middle lobe. These lobes are subequal, and nearly lanceolate in outline. The dorsal
and ventral cirri are somewhat short, in neither case reaching the tip of the adjoining
lobe. The succeeding feet present a tendency to the increase of the dorsal lobe, so that
at the tenth foot it forms a large foliaceous lamella. The superior setigerous lobe has a
lanceolate lamella beneath it, whilst, after a gap, the inferior setigerous lobe has one above
it in lateral view. The dorsal lamella gradually increases until (as at the thirty-seventh
foot) it forms a great lanceolate leaf overlapping that behind. The middle lamella is also
large and foliaceous, and the cirri appear small beside it. The foot retains this structure
till near the tip of the tail. All the bristles have simple tapering tips, those at the
ventral edge of the inferior division having shorter tips of the same kind. The shafts are
finely camerated.

Ehlers describes the epitokous form as having the upper lamellse of the feet much
1 'Ball. Sc. Fr. et Belg./ T. xxii, p. 78, 1890.

NEREIS (ALITTA) VIRENS. 331

enlarged, and the hind lamellae of the inferior division also enlarged. The ventral cirrus,
further, has small leaf -like flaps on its thickened base.

Synonyms.

1835. Nereis virens, M. Sars. Beskrivelser, p. 58, Tab. x, f. 27.

1851. „ „ Grube. Earn. Annel., pp. 49 and 127.

1853. „ grandis, Stimpson. Synops. Grand Manan, p. 34, f. 24.

1865. Alitta virens, Kinberg. Of vers. Vet.-Akad. Forh., p. 172.

„ ,, „ Malmgren. Nord. Hafs-Annul., p. 183.

„ Nereis Yankiana, De Qnatrefages. Annel., i, p. 553, pi. xvii, f. 7, 8.

„ „ virens, idem. Ibid., p. 555.

1867. Alitta „ Malmgren. Annul. Polycli., p. 56, Tab. iii, f. 19.

1868. Nereis „ Ehlers. Borstenw., ii, p. 559, Taf. xxii, f. 29—32.
,, ,, grandis, Packard. Americ. Nat., ii, p. 275.

1874. Alitta virens, Mcintosh. Ann. Nat. Hist., ser. 4, vol. xiv, p. 199.

1875. „ „ idem. Invert, and Fishes St. Andrews, p. 123.

1876. Nereis „ Trumbull. Trans. Conn. Acad., iii, p. 265, pis. 42— 44.
1879. Alitta „ Webster. Trans. Albany Inst., ix, p. 35.

„ Nereis ,, Tauber. Annul. Danic, p. 100.

„ „ (Alitta) virens, Verrill. Check List, U.S. Comm. P. & F., p. 8.

„ „ virens, idem. Invert. Yiny. Sound. Ibid., p. 591, pi. xi, f. 47 — 50.

1881. Alitta virens, Leslie and Herdman. Proc P. Pliys. Soc, Bdinb., vol. vi, p. 274.

1883. Nereis „ Levinsen. Yidensk. Meddel. Foren. Kjobenli., p. 233.

1884. „ „ Webster and Benedict. Rep. U.S. Comm. F. & F., p. 717.

1891. „ (Alitta) virens, Hornel. Trans. Biol. Soc. Liverp., vol. v, p. 243.

1892. „ virens (eyes), Andrews. Journ. Morph., p. 177, pi. ix, f. 3.

1893. „ „ Levinsen. Yidensk. Ud. ' Hauclis.,' p. 329.

1901. „ „ Johnson. Proc. Bost. Soc. Nat. Hist., vol. xxix, No. 18, p. 398.
„ „ „ Whiteaves. Geol. Surv. Canada, No. 722, p. 81.

1902. „ „ Mcintosh. Ann. Nat. Hist., ser. 7, vol. x, p. 257.

Habitat. — Small specimens from 10 in. to a foot occur in great numbers along with
Nephthys in muddy sand on the north side of the pole at the West Hocks, St. Andrews.
They are found at various parts of the coast — both east and west — in similar localities.
Larger specimens are procured to the east of the Step-rock in the same line of sand. Very
large epitokous examples, again, are tossed on the West Sands at St. Andrews after storms
in October and April, especially the latter month, as well as in May (E. M.). Shore of
the Mersey estuary (Benham). Aberdour (Leslie and Herdman).

Norway (Sars). East coast of N. America (Ehlers). Virginian coast (Webster).
Puget Sound, Pacific (Johnson).

Head (Plate LXI, fig. 2) of the typical outline, with two short tentacles anteriorly.
Eyes arranged in a trapezoid behind the anterior triangle ; anterior pair wider apart ;
often with lenses. The longest tentacular cirrus is the dorsal of the second pair. Colour,
bluish-purple.

Body of 100 — 173 segments or more, from 1 to 3 ft. in length, and If in. in
breadth across the feet, slightly diminished anteriorly, whilst posteriorly it gradually
tapers, the feet being proportionally broader, and terminates in an anal segment and

332 NEREIS (ALITTA) VIRENS.

aperture which has a lax, fringed border, except interiorly, where a smooth portion gives
origin to two long cirri. Each of the latter has a dark basal region of a brownish-purple
hue on the ventral surface, and is thus distinguished from the adjoining parts.
Peristomial segment about twice the breadth of the succeeding. The colour of the dorsum
is of various shades of iridescent bluish-purple, the same tint extending to the shorter
anterior feet. The finest specimens have the great lamellae coloured greenish-purple.
Others vary according to the prevailing tint of the specimen. The lamellae are also finely
tinted and iridescent, and as they gracefully fold over each other like the slates on a roof
the effect is striking. The dorsal cirrus, which is larger anteriorly than in the feet with
foliaceous lamellae, is of a deep green. The ventral surface is pale iridescent pinkish,
becoming deeper posteriorly, and with a pinkish belt across the front of each segment.
In others this surface is iridescent bluish, with the belts at the anterior part of each
segment of a deeper hue. In contraction the tints are darker.

Ehlers (1868) observes that in the atokous form the vent has inferiorly two long cirri,
whereas in the epitokous form the vent has a ring of short papillae.

The proboscis (Plate LXI, figs. 2 and 2 a) in extrusion is greyish-blue, somewhat
lighter than the hue of the body, and similarly iridescent. Dorsally, in front of the palpi,
and on the basal ring, one or two paragnathi (VI) occur on the eminence on each side,
with an intermediate point (V), whilst on the maxillary (distal) segment a group (II) —
somewhat triangular in outline and with the apex in front — lies on each side behind the
jaws, with two points (I) longitudinally arranged between them. Ventrally (Plate LXI,
fig. 2 a) the basal segment has a somewhat irregular double row (VII and VIII) of
paragnathi on the folds, and they leave but a brief interval between them and the dorsal
points in front of the palpi. On the maxillary segment a long, somewhat irregular double
row (IV) proceeds backward as a longitudinal band from each jaw, and in the middle is
a transverse belt of points (III), the larger in a single row behind, and the smaller dotted
in front. The maxillae have six teeth. The smaller examples between tide-marks have, as
a rule, the paragnathi more complete than the large epitokous forms stranded on the beach.

From the posterior part of the pharynx pass two elongated caeca, which have
internally a coating of blackish pigment-granules, the smaller showing active molecular
motion when separated. The walls of the caeca are tough, and longitudinally plaited.

The coelomic bodies are transparent structures of an elliptical form, the longer
diameter having an average of T1 f 5 - - of an inch (O02 mm.), many being much shorter and a
few longer and larger. Their transverse diameter also varies (Plate LXI, fig. 3). Most of
these bodies are faintly striated longitudinally, and similar markings occur in the circular
corpuscles. They are protoplasmic, and yield when they encounter obstacles.

Circulation. —The blood must be in considerable quantity in large specimens.

The ultimate fibrils of the ventral muscle presented peculiar fibrillation as if from
fine lines crossing each other obliquely. In the centre of the fibril is a granular streak,
though in some this could not be made out. Acetic acid and calcium chloride obliterated
the streak. A contracted fibril is seen in Plate LXI, fig. 4. Occasionally the streak in
the fibril assumes the form of a granular band (Plate LXI, fig. 5), the central region
representing the contracted fibril, and the transparent sarcolemma or other muscular
element projects externally.

NEREIS (ALITTA) VIKENS. 333

The first foot (Plate LXXIII, fig. 2) has (in spirit) a short and somewhat thick dorsal
cirrus, the tip of which does not reach that of the dorsal lobe. The latter is lanceolate,
slightly contracted at the base in lateral view, and tapering to a blunt tip. The
setigerous process is attached to the anterior face of the second lobe so that only the tips
of the bristles project beyond it, and nothing is seen of the process itself from behind.
These bristles all have slender, tapering, finely spinous tips (Plate LXXXL, fig. 7, a,b, c)
and slightly camerated shafts. The middle lobe is tongue-shaped in lateral view, and its
blunt tip projects further than that of the dorsal lobe. The ventral lobe is closely
united at its base with the middle lobe, and its vertical diameter is greater in the free
region. Its blunt tip scarcely projects as far as that of the former. The ventral cirrus
arises close beneath, and its tip does not reach, by a larger interval than in the case of
the dorsal, the tip of the ventral lobe.

The third foot, as in other Nereids, has two setigerous lobes, and the lamellae,
especially the dorsal, are somewhat longer. The fourth foot shows a decided increase in
the dorsal lamella and in the space between it and the body, and these points continue to
increase until at the tenth foot (Plate LXXIII, fig. 2 a) — by the development of these
parts in breadth and depth — a large foliaceous lobe is formed with the dorsal cirrus
amalgamated and chiefly visible on the posterior border. The superior setigerous lobe
forms an ovate process in front of the bristles, the tongue-shaped middle lobe, which
projects considerably further, being fused with it inferiorly. A gap exists between the
foregoing and the inferior setigerous lobe, which has a free lanceolate lamella at its upper
and posterior edge. In front of this the setigerous lobe proper is bifid, with a small
upper and a larger inferior papilla from which the tip of the spine projects in front of
the bristles. The groups of bristles correspond to the ordinary types, but all have
simple tapering tips, the inferior group (corresponding to the heterogomph series)
having decidedly shorter tips. All the tips of the bristles are finely serrated and have
cameras in the shafts, except a small portion at the base and distal end. The ventral
cirrus arises from an elevation on the ventral surface and does not reach the middle of
the ventral lobe.

The development of the superior lamella continues behind the tenth foot, and the
dorsal cirrus becomes more and more lamellar in position, so that in the region of the
thirty-seventh foot (Plate LXXIII, fig. 2 b) the maximum is reached. It is true that at
the fifty-seventh foot the lamella is proportionally larger, but the foot generally is
considerably less. The second lamella is also thin and foliaceous (lanceolate), especially
after the anterior third. In life large pale wart-like papillae occur on the edge of the
dorsal lamella. The dorsal and ventral cirri show no blood-vessels, and no cilia, but
palpocils occur at the tip of both.

In front of the tail (twelfth bristled foot from the tip) the dorsal lamella still retains
its characteristic foliaceous form, the cirrus is longer and more slender, though its tip
does not reach that of the lamella. It arises from a slight notch of the dorsal edge of the
lamella about its middle. The foot is thinner and more flattened, so that the setigerous
papillae are brought into the vertical line of the edge, and thus do not possess the more
complex arrangement they have in front. The tips of the lobes below the dorsal are also
more pointed. The ventral cirrus remains short.

334 NEREIS (ALITTA) VIRENS.

J. T. Cunningham 1 describes the segmental organ as a somewhat spherical mass
composed apparently of a convoluted tube ending in an efferent duct which opens on the
ventral surface. The internal aperture is funnel-shaped and fimbriated and seems to
open on the anterior face of the septum. He thinks the generative elements are not
discharged through this organ, but by dehiscence through the body-wall.

Reproduction.— In those procured in May the females discharged from the broken
surfaces (posterior) a cloud of grass-green ova. They measured from -^ to T -^ 5 - - of
an inch (O03 - 0'02 mm.), and contained numerous granules that by transmitted light
have a pale brownish or greenish aspect, the larger masses especially showing the former
tint. On the bottom of the vessel the ova formed a deep grass-green layer. The addition
of acetic acid did not affect their colour.

The males when placed in spirit about the end of April (21st) emitted streams of
spermatozoa from the bases of the feet. In the fresh specimen the sperms maintain
an active wriggling motion. Amongst these are also some bodies longer than the head
of the ordinary form, and even more active in movement. The masses of sperms in the
ccelom assumed a greyish hue.

The digestive apparatus and the whole organs of the animal at this season (April
and May) seem to be modified for the furtherance of reproduction.

In October the small ova occur in masses at the bases of the feet, extending into the
lamellae. To the naked eye they are pale, but under the microscope slightly greenish.
Moreover, in the bottom tow-nets toward the end of the month numerous faintly yellow
or greenish-yellow larvas in the trochophore stages are found, with a diameter of about
•228 mm. At the same period an early mesotroch stage is met with. Neither tentacular
cirri nor eyes are present, and the snout is short, whilst only traces of the feet in the
form of lateral papillge are observed. A little pinkish-white pigment occurs at the
ciliated ring. The next stage presents three bristled feet on each side, though neither
eyes nor tentacular cirri are present. Two short tentacles spring from the anterior
border of the snout, and two rounded anal cirri occur posteriorly.

Toward the end of October a considerable number of a later stage appeared in the
bottom-net, and they were readily recognized by the pale pink pigment in the form of a
ring behind the preoral belt of cilia, and also by similar pigment posteriorly on each side
of the purplish streak of the alimentary canal (Plate L, fig. 12). The pigment on the
head forms a flattened horseshoe, thus readily distinguishing this form, and there are two
large eyes, besides two short tentacles, which seem to spring from the ventral edge of the
snout, and traces of another process on each side. The preoral ring of cilia is powerful,
and seems to be the chief locomotor apparatus. Behind it three long cirri project —
probably the representatives of the tentacular cirri, which in somewhat younger stages
are considerably shorter. Fourteen or fifteen feet are present, each having a prominent
dorsal and a long ventral lobule with a cirrus. Simple bristles occur dorsally. Posteriorly
the pygidium has two processes like flattened papillse, probably the precursors of the anal
cirri, and the region has rose-pink pigment. In swimming the tentacular cirri behind the
preoral ring of cilia are closely adpressed to the body.

An older stage, of the same date (Plate L, fig. 13), having sixteen bristled segments
1 ' Quart. Journ. Micros. Sci./ N.S., vol. 28, p. 256, PI. xviii, figs. 15 and 16, 1888.

NEREIS (ALITTA) VIUENS. 335

presents a tentacle and a second process (palpns ?) on each side of the snout, whilst the
preoral ring of cilia is narrower and the body considerably more elongate. The activity
of the ciliated ring is noteworthy, and in addition to its locomotor functions it may aid in
obtaining nourishment. Four tentacular cirri of different lengths, viz., the dorsal and
ventral of each pair, occur behind the ciliated ring. A long proboscis now separates the
mouth from the granular alimentary canal which commences at the seventh bristled foot.
The two anal processes are larger. The dorsal lamellae of the feet are lanceolate and
richly ciliated on the edge which is next the body.

Habits. — In confinement a muscular wave occurs from before backward in healthy
animals, and the species is fairly hardy, the elaborate coloured drawing (Plate LI II)
having been made from those kept for a considerable time fifty miles from the sea-coast.
When dying, or when placed in fresh water, they eject the proboscis with the jaws at the
tip, and decomposition takes place rapidly with considerable odour. Hornel states that
they secrete much mucus from an enormous number ol t abides in the lobes of the feet.
This species is often used as bait for plaice and other fishes. About the year 1890 the
local fishermen began to dig for them, having found for many years that those stranded
on the beach after storms were valuable in this respect.

Packard (1868) considered his specimens, which were 17 ins. in length, enormous, in
short, "these are princes amongst worms, ranking above the smaller forms by their superior
size and organization and their rich imperial dress." If this able and genial naturalist
had seen those fully 3 ft. in length at St. Andrews his words would have been even more
eloquent.

Ehlers (1868) points out that Nereis (Alitta) Brandti, the Nereis virens of Grube,
a closely allied species, frequents the Pacific shores of America and the sea of Ochotzk.

A Japanese form, Nereis {Alitta) oxypoda, Marenzeller, 1 from Yokohama, shows great
development of the dorsum of the foot between the cirrus and the body, so that the cirrus
lies in a hollow between two foliaceous lobes.

Grube's Nereis virens, 2 from the shores of Eastern Siberia, comes very near this
form, though Malmgren considered it a distinct species. AVhether the differences
indicated, such as the form of the great dorsal lamella, are due to environment and other
factors is still an open question.

J. T. Cunningham (1888) 3 states that he cannot confirm the account of the nerve-
cords formerly given. 4 In many sections three or four neural canals are seen, which are
not quite symmetrical; these are sections through interganglionic transverse commissures.
In the cords between successive ganglia there is seen to be a single pair of canals, one of
which is often divided into two. The pair occupies an exactly similar position to that
of the neural canals in Sigalion." He adds, "Probably he" (Mel.) "examined sections
of the ganglia in which several canals are often seen. But these have not a constant
relative position, and are, I believe, due to the subdivision of the two canals which are
seen in the separated cords." A re-examination of the careful sections, unstained and
made by the hand, does not lead, however, to any change of the view stated in 1877.

1 ' Suctjap. Annel./ p. 12, Taf. ii, fig. 3, 1879.

3 Op. cit., p. 6, Taf. i, figs. 2, 4, 5 and 6, 1858.

8 'Quart. Journ. Micros. Sci./ n.s., vol. 28, p. 270.

4 ' Proc. Eoy. Soc. Edin./ 1877.

336 NEREIS (NEREILEPAS) FUCATA.

Cunningham 1 denies that the neural canals are colossal nerve-fibres. He thinks they
are supporting structures to prevent the nerve-cords being bent at sharp angles. The
canals attain greatest development in worms which are extremely long; and further,
in these the nerve-cords are not separated from the epidermis, and therefore are more
exposed to danger. The neural canals have a position in relation to the nerve-cords
and ventral blood-vessel which is similar to that of the notochord in relation to neuro-
chorcl and aorta.

H. Parlin Johnson (1901) considers the Alitta Brandti of Malmgren and Ehlers, and
the Nereis foliata, Baircl, the same species, and observes that the Pacific examples grow
to the length of 50 cm. with a diameter of 25 mm. He had not seen an epitokous form.

Such types as the Pseudonereis gallapagensis of Kinberg from the shores of Peru
and the Galapagos Islands show the same leaf -like dorsal cirrus, and probably the same
vascularity. 2

9. Neeeis (Neketlepas) fucata, Be Blainville. Plate LII, fg. 6; Plate LXI, figs. 6, 6 a
—head; Plate LXXIII, figs. 3-3 c— feet; Plate LXXXI, figs. 8-8 b— bristles.

Specific Characters. — Head of the typical shape, with two subulate tentacles in front.
Eyes black, comparatively large, the anterior pair somewhat wider apart and occasionally
larger. Tentacular cirri of moderate length. Palpi white, only the distal process being
pale. Body 4 — 9 ins. in length, and segments 95 to 110. Peristomial segment wider
than the succeeding. The slender tail has two long cirri. The general hue is buff,
enlivened by two white streaks on the dorsum, with the red dorsal vessel between, and a
dotted white line along each side. The median white lines begin at the base of the
tentacles, enlarge before reaching the eyes, between which they pass, and again contract
on the first segment. The last segment has a conspicuous white patch. Ventral surface
pinkish buff. The proboscis has dorsally on its maxillary (distal) segment in extrusion a
group of somewhat large paragnathi (II) on each side at the base of the jaw. In the
space between them a single tooth (I) sometimes occurs. On the ventral surface two
groups of paragnathi (IV) also are found, the long axis being oblique, whereas that of the
dorsal series is more nearly transverse. Between them in the middle line posteriorly are
a few paragnathi (III) arranged transversely. The basal division of the organ has on the
eminence in front of each palpus a group (VI) of five or six small paragnathi. Ventrally
a row of larger paragnathi in groups of two to four (VII and VIII) occurs in front of the
area, and there are groups of minute ones behind, a long interval existing between the
dorsal groups and these. The strongly curved maxillse have from thirteen to sixteen teeth.

First foot chiefly noteworthy for the large size of the dorsal cirrus. Whilst the
dorsal lobe in this is bluntly rounded, it gradually increases in size from the tenth to the
twenty-seventh, where it forms a great dorsal lamella, pointed externally and rounded
internally, the dorsal cirrus projecting from the upper border. Superior setigerous lobe
amalgamated with that beneath it, the bristles passing out in front. The second lobe is
ovate or ovato-lanceolate. Inferior setigerous lobe bifid, the longer and smaller papilla

1 Op. cit., p. 275.

3 C. Gravier, ' Arch. Zool. exper./ t. x, p. 629, pi. xvi, figs. 15—20, 1909.

NEREIS (NEREILEPAS) FTJCATA. 337

anterior. Ventral lobe tongue-shaped. Ventral cirrus extends a little beyond the tip of
the foregoing lobe. In the superior division of the foot the bristles have long, tapering,
spinous tips, and at the lower edge of the fascicle are a few with stouter shafts and shorter
tips. In the inferior division the upper series consists of those like the foregoing, then
intermediate forms, and inferiorly a dense group of bristles with short tips (heterogomph).
The change in the feet of the epitokous female begins in the twenty-first foot as a
slight enlargement at the inner border of the ventral cirrus. A similar process appears
at the inner base of the dorsal cirrus of the twenty-sixth or twenty- seventh foot. These
lameilaB increase to the sixtieth foot and again diminish. Coincident with these a small
lamella occupies the posterior surface of the inferior setigerous process. In one specimen
the bristles retained the normal form, yet the ova were nearly ripe. Ehlers observes that
the change in the feet of the epitokous male begins in the seventeenth foot.

Synonyms.

1820. Lycoris fucata, Savigir^. Syst. Annel., p. 31.
1825. Nereis „ De Blainville. Diet. Sc. Nat., t. xxxiv, p. 436.
1828. Nereilepas fucata, idem. Ibid., t. lvii, p. 468.
„ „ ,, idem. Ibid., t. lvii, p. 469.

1833. Nereis „ Audouin and Edwards. Ann. Sc. Nat., torn. xxix ; p. 210.

1834. „ „ idem. Annel., p. 188.

1839. „ bilineata, Johnston. Ann. Nat. Hist., iii, p. 295, pi. vi, f. 4.

1840. „ fucata, idem. Ibid., v, p. 175, fig. 7.

1851. „ „ Grube. Fam. Annel., pp. 49 and 127.

1855. „ bilineata, Thompson. Ann. Nat. Hist., 2 ser., vol. xvi, p. 78.

1865. „ „ Carrington. Annel. Southport, p. 8.

„ „ imbecillis, Johnston. Cat. Worms Brit. Mus., p. 156 (excl. syn. Grube, fide Malmgren).

„ Nereilepas fucata, idem. Ibid., p. 158.

„ Nereis „ De Quatrefages. Annel., i, p. 547.

1867. Nereilepas „ Malmgren. Annul. Polych., p. 53, Tab. iii, f. 18.

„ „ ,, Parfitt. Trans. Devon. Assoc., ii, p. 22 (sep. copy).

1868. Nereis „ Ehlers. Borstenw., ii, p. 546, Taf. xxi, f. 41 — 44 ; Taf. xxii, f. 1—4.
1870. „ „ Grube. Arch. f. Naturges., Bd. xxxvi, p. 307.

„ „ bilineata, idem. Ibid., p. 304.

1873. „ fucata, Yerrill. Invert. Yiny. Sound, U.S. Comm. F. & F., p. 591.

1874. Nereilepis fucata, Mcintosh. Ann. Nat. Hist., ser. 4, vol. xiv, p. 199.
„ „ „ Malm. Goteb. vet. o. vitt. Handl., xiv, p. 84.

1875. „ „ Mcintosh. Invert, and Fishes St. Andrews, p. 123.
1879. „ ,, Tauber. Annul. Danic, p. 99.

1883. Nereis „ Levinsen. Vidensk. Medell. Foren. Kjobenh., p. 233.

1890. „ „ Malaquin. Annel. Boulom, p. 30.

„ Nereilepas „ idem. Rev. biol. Nord France, t. ii, p. 2 (fide De St. Joseph).

1891. Nereis (Nereilepas) fucata, Hornel. Trans. Biol. Soc. Liverp., vol. v, p. 242.
1895. Nereilepas fucata, Coupin. La Nature, l e semestre, p. 29 (fide De St. Joseph).
1898. Nereis fucata, De St. Joseph. Ann. Sc, Nat., 8 e ser., t. v, p. 300, pi. xvi, f. 83—87.
1902. Nereilepas fucata, Mcintosh. Ann. Nat. Hist., ser. 7, vol. x,, p. 257.

1904. Nereis „ Allen. Journ. M. B. A., n.s., vol. vii, p. 224.

1906. Nereilepas „ Eisig. Fauna u. Fl. Neap., xxviii, p. 158 et seq.

100

338 NEREIS (NEREILEPAS) FUCATA.

1908. Nereis fucata, Ehlers. Deutsch. Tiefsee-Exped., p. 71.

1909. Nereilepas fucata Elwes. Journ. M. B. A., n.s., vol. viii, p. 351.

„ „ „ Lo Bianco. Mittli. Zool. Stat. Neap., Bd. xix, p. 582.

Forma epitoca.

1771. ? Die faserige Nereide, 0. F. Miiller. Yon Wiirmern, p. 144, Tab. viii.

1776. Nereis fimbriata, idem. Proclr. Zool. Danic, No. 2627.

1791. „ „ Linn. Syst. Nat. (Gmelin), ed. 13, i, pt. 6, p. 3115.

1833. „ podophylla, Audonin and Edwards. Ann. Sc. Nat., t. xxix, p. 211, pi. xiii, f. 13.

1834. „ „ idem. Annel., p. 189, pi. iv a, f. 13.
1865. Nereilepas fimbriatus, De Quatrefages. Annel., t. i, p. 559.

,, ,, margaritaceus, idem. Ibid., p. 562.

„ Heteronereis margaritacea, Johnston. Cat. Worms Brit. Mns., p. 166.

,, „ renalis, idem. Ibid., p. 163.

„ „ glaucopis, Malmgren. Nord. Hafs- Annul., p. 110, Taf. xi, f. 16, 16 a.

1867. „ „ idem. Annul. Polych., p. 60, Taf. iv, f. 26, 27.

1869. Nereis fucata, Grube. Sehles. Qesell., 1868—9, p. 113, sep. copy, p. 5.
1874. Heteronereis glaucopis, Malm. Groteb. vet. o. vitt. Handl., xiv, p. 84.
1888. Nereis fucata, Wiren. Svensk. Yet. Akad. Handl., Bd. xiv, afcl. iv, No. 5, p. 1—14, Taf. i— iii.

3, ,, „ B inquilina, idem. Ibid., p. 9, Taf. i — iii.

Habitat. — Generally distributed round the shores of Britain, including Ireland, from
the eastern to the western borders. Abundant in deep water off St. Andrews Bay, and
frequently commensalistic with Pa gurus Bemhardus in Buccinum, both the large and the
smaller (littoral) varieties, and in Fusus Islandicus. Part of the annelid is inserted within
the shell, and the anterior region lies along the lip. Large examples are also thrown on
the beach after storms (E. M.). Dr. Johnston found his specimens in the same or allied shells
with Hermit-crabs. Stomach of cod and haddock, St. Andrews (E. M.). Large examples
from the deep water off Montrose (Dr. Howden). In 80 fathoms eighteen miles west of
Skellig, Ireland (J. Gr. Jeffreys). Torquay (Elwes).

Shores of France (Audouin and Edwards). North Sea (Ehlers, Levinsen, etc.).
Eastern American shores (Verrill).

Head (Plate LXI, fig. 6) of the typical shape, terminated anteriorly by two subulate
tentacles which are about the length of the head. Eyes black, comparatively large, the
anterior pair somewhat wider apart, and often larger than the posterior. The palpi are
normal, and the tentacular cirri of moderate length.

Body 4 — 8 or 9 ins. in length, slightly narrowed anteriorly, and diminished to a
slender tail terminated by two long cirri posteriorly. Segments from ninety-five to one
hundred and ten. The peristomial segment is wider than the succeeding. The general
hue is buff, enlivened by two white streaks along the dorsum with the red dorsal blood-
vessel between, and a clotted white line along each side. The two median white lines
begin at the base of the tentacles, enlarge before reaching the eyes, between which they
pass, and again contract behind them on the first segment. They continue to the tip of
the tail, the last segment having a conspicuous white patch. On the arch of each segment
a white streak passes outward toward the white dorsal process of each foot. In the
anterior feet the white pigment is less developed, and it is rare for the pigment on the
arch of the foot to unite with the central white lines. The palpi are white, only the

NEREIS (NEREILEPAS) FUCATA.

339

distal process being pale or slightly translucent. The tentacles and cirri are pale. In
some the yellowish buff is best marked over the bases of the feet anteriorly, leaving a pale
belt on each side of the central white line. The ventral surface is pinkish buff.

The proboscis (Plate LXI, fig. 6) has dorsally in front of each palpus a group (VI)
of five or six small paragnathi. Upon the maxillary segment near the base of the jaws
is on each side a group (II) of larger, pointed paragnathi, the groups separated by a wide
interval, in which (I) a single tooth sometimes occurs. Ventrally (Plate LXI, fig. 6 a) the
basal segment has a row of larger paragnathi (VII and VIII) in front in groups of two
to four, and a series of minute ones behind, a long interval occurring between the dorsal
groups of this division of the organ. Distally are two lateral groups (IV) of distinct
paragnathi, the long axis being oblique, whereas the long axis of the corresponding dorsal
series (II) is more nearly transverse. Between them in the middle line posteriorly are a
few paragnathi (III) arranged transversely. The maxillae, which are strongly curved at
the tip, have from thirteen to sixteen teeth. 1 Paragnathi II and IV are often the most
conspicuous in size.

Fig. 75. — Transverse section of the body-wall of Nereilepas fucata. Letters as before.

In connection with the food of this species, De Haan 2 records that Lycoris fucata
devoured Teredo.

In transverse section the arrangement of the ventral longitudinal muscles and their
reflection is similar to that of Nereis pelagica. The nerve-cords show a median neural
canal, and two lateral canals which are situated near the external border of each nerve in
the interganglionic areas (Fig. 75).

The segmental organs 3 agree generally with the type which has already been alluded
to on p. 250.

The first foot (Plate LXXIII, fig. 3) is chiefly remarkable for the large size of the
dorsal cirrus. The dorsal lobe is bluntly rounded, the setigerous lobe pointed, and the
ventral lobe is bevelled to a blunt tip. The ventral cirrus extends considerably beyond it.
There are two black spines passing to the apex of the setigerous region. The superior
group of bristles consists chiefly of those with long tapering tips and spinous edges
(Plate LXXXI, fig. 8), but there is at least one with a short terminal piece slightly
1 Levinsen gives a resume of teeth according to series I, II, etc., ' Viclensk. Meddel./ 1883, and
Tauber (op. cit.) notices the variability of the paragnathi.
3 'Zool. Record/ 1866, p. 586.
3 Vide Goodrich, ' Quart. Journ. Micros. Sci./ vol. xxxiv.

340 NEREIS (NEREILEPAS) FUCATA.

curved at the tip as in the next series (heterogomph), sometimes two or three. The
inferior group presents bristles with somewhat shorter tapering and spinous tips superiorly
(Plate LXXXI, fig. 8 a), then a few having medium tips with a very slight curve, and
inferiorly those with shorter tips with a curved probe-point (Plate LXXXI, fig. 8 b).

The double bristle-tuft appears in the third foot as in allied forms. At the tenth
foot (Plate LXXIII, fig. 3 a) the outline trends from above downward and inward. The
dorsal cirrus is long and slender, stretching far beyond the tip of the conical superior lobe.
The superior setigerous lobe is represented only by the bristles. The lobe beneath it is
irregularly conical and considerably less than the superior. The inferior setigerous lobe
is bifid and has two strong fascicles of bristles. The ventral lobe is less than either of
the preceding and conical at the tip. The ventral cirrus does not reach the extremity of
the latter. The bristles of the superior lobe have long tapering tips; those of the inferior
are in two groups — the upper having bristles with long tips above and with short tips
below. The inferior group has only those with short tips (epitokous female).

Towards the enlarged end of the shaft the camerated region of the bristles is
narrowed, and the markings are less regular. At the base of the shaft the cameras become
indistinct, and the diameter of the bristle is less. The spines are rather brittle. At the
base they are pellucid, then brownish, and finally black distally. The inferior spine is
the stronger.

Reproduction. — Two females having the body distended with ova of moderate size
came from deep water off St. Andrews Bay in January (E. M.). Others with large ova
were tossed on shore after storms the same month. One in March had smaller ova. In
October the reproductive elements are but slightly developed. Prof. Garstang 1 and
Dr. Allen give May as the breeding season at Plymouth ; whereas Lo Bianco has found
mature forms in October and November as well as from April to July. In the ripe
condition the ova extend forward to the anterior region ; whilst the coloration of the
posterior end of the body in some is lost by the prominence of the internal mass of
greyish ova (De St. Joseph).

In an incipient epitokous female from Cornwall, kindly sent by the late Dr. Baird, a
slight enlargement is observed at the inner border of the ventral cirrus of the twenty-
first foot. This increases at the thirtieth to form a small lamella adherent to the inner
border of the cirrus. This lamella continues to the sixtieth and again diminishes,
disappearing about the seventieth foot. In the same way a slight enlargement appears at
the inner border of the base of the dorsal cirrus about the twenty-sixth or twenty- seventh
foot. Behind this it by-and-by forms an elevated crest, and continues to the same
region as the former. Coincident with these a small lamella occupies the posterior surface
of the inferior setigerous process, its margin in lateral view being visible above and below
the lobe, but very little of it is seen at its tip. Traces of this continue beyond the
seventieth foot, and then disappear. The bristles retain the normal form in this specimen,
no swimming bristles being present. The ova are ripe or nearly so.

In an epitokous female the dorsal lobe of the foot gradually increases in size after
the tenth, until at the twenty-seventh (Plate LXXIII, fig. 3 6) the upper region of the
foot, including that between the dorsal cirrus and the body and the dorsal lobe, forms a

1 ' Journ. M. B. A./ vol. iii, p. 225.

NEREIS (NBREILBPAS) FITCATA. 341

great lamella pointed externally (apex of dorsal lobe) and rounded internally, with the
dorsal cirrus projecting from the upper border. The setigerous lobe is amalgamated
with that beneath it, the bristles, which have cylindrical shafts regularly camerated, and
long, tapering tips, passing outward from the base in front. The second lobe is some-
what ovate or ovato-lanceolate. The inferior setigerous lobe is bifid, the longer and
smaller papilla being anterior. The superior bristles have long tapering tips delicately
serrated. The inferior group consists of those with long tapering tips (somewhat shorter
than the upper series) superiorly, then intermediate forms, and inferiorly a dense group
of those with short falcate tips as in the figure, the edge being beset with spines. The
ventral lobe is tongue-shaped, and the ventral cirrus, which extends a little beyond the
tip of the lobe above, arises from an eminence of its own.

The huge superior lamella somewhat increases in size proportionally, especially in its
vertical diameter, from the foregoing to the fifty-seventh foot (Plate LXXIII, fig. 3 c)
but, except a slight increase in the size of the middle lobe, the foot does not essentially
alter. The apex of the dorsal lobe, however, becomes slightly more bulbous than in the
twenty-seventh, and about the sixty-eighth or seventieth the foot presents an opaque
brownish glandular aspect. This brownish glandular condition becomes more marked in
the succeeding segments to the tip of the tail. About the same region (seventieth foot)
a tendency to a similar thickening occurs in the middle lobe of the foot, and at the
ninety-fourth foot the brownish coloration marks the contents. The terminal segments
thus have a brownish touch at the tip of each of these lobes, and the middle lobe has
slightly increased in size. The ventral cirrus is also longer, reaching beyond the tip of
the ventral lobe.

In the fully developed epitokous form both divisions of the feet glisten with the
swimming bristles, which have broad, sabre-shaped tips.

Habits. — Young forms occur in large numbers in the crevices of stones and shells
from deep water and on Filigrana. The young thus follow a different habit from those
adults which become commensalistic with the hermit-crab in the shells of mollusks.
W. Thompson mentions one at Weymouth which constructed a free tube inside the shell of
the whelk. The motions of the annelid are very graceful.

In a young example about If ins. long procured with many others on a mass of
Filigrana implexa near the Bell Rock, the dorsal edge of the fourth foot on the right was
fused with a calcareous growth, apparently of a young Gellepova, which had fixed itself
on the dorsum near the base of the foot, which thus by-and-by had become involved in
the growth.

In sickly examples the caudal segments long retain vitality whilst the anterior region
is blanched in death, the opposite condition of that occurring for instance in Polydora, in
which the anterior region is active whilst the posterior is decayed. It is used as a bait
for whiting.

This is the Nereis imbecillis (Mus. Leach) from Sandgate, Kent, in the British
Museum.

The commensalistic habit of this species has not resulted in much structural change,
and in this respect it diverges from the Lycastis (Anoplonereis Ilerrmanni of Griard, 1 ), which
1 'Compt. Rend./ August 21, 1882, p. 389; and 'Ann. Nat. Hist./ ser. 5, vol. x, p. 330.

342 NEREIS (NEREILEPAS) FUCATA.

has an altered superior division of the foot (absence of the superior ligula) with simple
capillary bristles, and the jaws have disappeared, whilst there is a third median tentacle.
This species was found by the late Prof. Griard 1 in the tunnels of Balanoglossus. A
survey of other commensals amongst the Polychsets leads to a similar conclusion as regards
structural alteration. Moreover, the groups in which this habit has been developed are few,
the most conspicuous being the family of the Polynoidas which haunt the tubes of
Terebellids and Clrdetopterids, the ambnlacral grooves of Echinoderms, grooves in
mollusks, and occur on Hydrocorallines or within the framework of Venns's Flower Basket.
The Hesionids have few examples, the best known being Ophiodromus flexuosus on
Astropecten and Luidia. Syllids, again, are common in sponges, the unique Syllis ramosa
permeating the canals with its branches; Eisig 2 excludes Polydora in the group " Endo-
Commensalen " since it perforates oyster-shells so extensively, but it also does the same
in various rocks, and in such a case is no more a commensal than Oliona.

In Montagu's MS. volume of drawings (1808) in the Linnean Society this species is
represented in Plate LI, fig. o.

Chamisso and Eysenharclt 3 (1819) describe from Unalascha a Nereis heteropoda which
appears to resemble Nereilepas fucata very closely, though the authors place the change in
the form of the foot at the forty-third. They may, however, refer only to the full
development of the altered foot.

In accordance with their views Audouin and Mime Edwards (1834) considered the
enlarged dorsal lamella as specially branchial. Their Nereis podophylla is a female
epitokous form of the present species.

The Nereilepas fused of CErsted (1843) is probably the same form.

Ehlers (1868) did good service in rectifying the synonymy of this species.

Claparede (1868) adopted Nereilepas as a sub-genus, following Johnston and Malmgren
rather than CErsted, Kinberg, and De Quatrefages, though Stratonice, Malmgren, was
included.

He describes two species of the genus from Naples, viz., N. parallelo gramma (the
A T . pulsatoria of Grube), and N. caudata of Delle Chiaje. Both species seem to be in
need of revision and accurate comparison with N. fucata. Claparede saw vibratile
cilia on the lamellas of the latter species leading by grooves to apertures out of which he
supposed the sexual elements escaped. This needs confirmation. In Delle Chiaje's
figure of what appears to be a foot (Plate Oil, fig. 1 5 4 ) the structure is inverted.

Wiren (1888) gives a careful account of this form in the atokous and epitokous
conditions, indicating the changes in the feet and body-wall respectively. He terms the
epitokous form var. inquilina. The diminution of the muscular layers in this condition is
marked, as in other annelids under the same circumstances. It occurs, as in Britain, in
shells covered with Hydractinia and inhabited by hermit-crabs.

1 < Compt. Rend./ August 21, 1882, p. 389; and ' Ann. Nat. Hist./ 5th ser., vol. x, p. 330 (as
Anoplonereis Herrmanni) .

2 < Fauna u. Fl. Neap./ xxviii, p. 167.

3 'De Anim. Quibusd./ etc., Fascic. ii, p. 349, Taf. xxiv, fig. 2, 1821.

4 'Descriz. Anim. Invert./ vol. iii, p. 96, Tav. cii, figs. 10 and 15, and < Mem./ vol. ii, pp. 403 and
426, Tav. xxviii, figs. 100, 115.

EUNICIDiE. 343

Hornel (1891) states that the worm takes possession of the shell of the mollusk first,
and that it tolerates the intrusion of the hermit-crab for benefits subsequently received.
This is uncertain. It is more probable that the same affinity as in other commensalistic
forms holds.

De St. Joseph (1898) considered Wiren's view that the posterior region of the body
was furnished with thinner longitudinal muscles to fit it for its peculiar habitat was
untenable, and that such was only a step to the Heteronereid condition.

At Naples Lo Bianco 1 (1909) finds that this species especially frequents Prosobranchs
inhabited by Eupagurus Prideauxii.

Family XI. — EuNiciniE.

Head somewhat triangular or semi-oval, with lobate palpi more or less united, rarely
lateral tentacles, or none. Tentacles subulate or elongate posteriorly; sometimes none.
Eyes two or four or absent. Nuchal organs in the form of ciliated grooves (Bacovitza).
Body long, vermiform, often slender, somewhat rounded, iridescent, with numerous short
segments, ending in two or four anal cirri. Peristomial segment bi-annular, with two
tentacular (nuchal) cirri or none. The body-wall is characterized by its great muscu-
larity, and the nerve-cords are comparatively large, and lie in the interval between the
ventral attachments of the oblique muscles. Externally are the basement-tissue, the
hypoderm, and the cuticle. The changes made by the ripe sexual elements in the male
are indicated in Fig. 76. Proboscis with paired or unpaired horny or calcareous plates;
one superior maxilla with hooked tip on each side, simple or toothed ; great dental
plates hooked anteriorly, toothed, truncate or convex, besides smaller anterior and lateral
plates. Mandibles 2 chisel-shaped. Feet simple or double, bearing dorsal (often with
bristles) and ventral cirri, one or more spines, and one or two fascicles of bristles ; often
without cirri, but with a setigerous lobe. Bristles simple or simple and compound ; the
former sometimes slender, winged, the latter with hooked tips. Uncinate spines
posteriorly.

Branchia3 springing from the dorsal cirrus in the form of simple styles or pectiniform
or sub-fascicular, and occasionally as spiral tufts, or absent. Blood has haemoglobin. 3

Goodrich states that the segmental organs are always provided in the adult with
large funnels opening into the coelom, and they perform the function both of excretory
and genital ducts. They were misunderstood by the earlier observers, such as Ehlers.
Fage 4 considers they have a close analogy with those of the Syllidae, though they differ

1 The recent loss of this able observer at the Naples Zoological Station must be deplored not
only by the staff, but by every marine zoologist.

2 In the figure in the ' Cambridge Nat. Hist/ (p. 270, fig. 140, l) these organs appear to have
been inverted.

s Ray Lankester, c Proc. Roy. Soc./ No. 140, p. 2, 1873.

4 < Ann. Sc. Nat./ 9 e ser., pp. 347—352, figs. 36 and 37, pi. vii, f. 32—36.

344 EUNICID^E.

in so far as the opening into the ccelom is by a large and permanent genital funnel,
instead of by a small nephrostome as in the family mentioned. They transmit the genital
products.

The sexes occasionally exhibit, as in Eunice norvegica, considerable differences — for
instance, in the branchiae.

The larva passes through two ringed prototroch, and through metatrochophore and
nectochaete stages (Hacker).

Some secrete tubes — hyaline, papyraceous, or strengthened with fragments of shells,
stones, or sea-weeds, and they may be slightly branched ; whilst others frequent tunnels
of sandy mud in clefts and fissures of rocks, or under tangle-roots and the crusts of
Lithothamnion. Mr. Crossland 1 observed Eunice siciliensis in the Red Sea in long, tortuous
holes, lined with parchment-like material. He thinks the importance of this form is
noteworthy in breaking clown coral, for it not only eats considerable quantities, it may be
for the parasitic algae found in stony corals, but its tunnel enables Ghone to gain an entrance.
He is inclined to think the dental apparatus is the boring apparatus as in Ly si dice, but
such has not yet been proved.

Audouin and Milne Edwards (1834) observe that in the Eunicidae the jaws attained
the maximum development, and that some forms (Lumbrinereidae) had a symmetrical
arrangement (four on each side), whilst others show irregularity in this respect (Eunice,
Aglaura). They point out the uniramous feet and the presence or absence of branchia3,
which form a more or less pectinate fringe or a spiral. They divide the group into
Branchiate and Abranchiate, the former including Eunice, Onuphis, and Diopatra, the
latter Lysiclice, Lumbrinereis, Aglaura, and (Enone. They term the mandibles the inferior
jaws. The maxillae of this work are their superior pair; the second pair are their
maxillae ; the third pair (from behind) correspond to the anterior plates, whilst their
supernumerary plate is the azygos plate. They arrange the species according to the
presence or absence of the tentacular cirri.

Grrube, 2 after Savigny, grouped in his early work the genus Eunice along with Diopatra,
Onuphis, Lumbriconereis, Aglaura, and (Enone under his third family, Eunicea.

Dr. Thos. Williams 3 states that the segmental organs are large in Eunice, and the
branchiae closely connected with them. The generative products are contained in delicate
areolar tissue. The ciliated segmental tube and its appended vascular tuft are not easily
seen, though more saccular and prominent than in Nereis. He figures, in Eunice gigantea,
the organ as loop-like in connection with a blood-vessel. He observes that the female is
nearly twice as large as the male at the reproductive season.

Kinberg 4 gave as the general characters of the Eunicea (Sav.) coadnate mandibles
and longitudinal maxillae, of which the first lamina is present. The Eunicea proper, again,
have seven to nine maxillae, part ii edentate ; five tentacles, and neither antennas nor palpi.
The buccal segment is double. He arranged the genera according to the form of the

1 I am much indebted to my friend for many useful notes from the Reel Sea.

2 <Fam. Annel./p. 44, 1851.

3 'Philos. Trans./ p. 127, pi. viii, fig. 20, 1858.

4 'Ofvers. af k. Vet.-Akad. Fork/ 1864, No. 10, pp. 559 and 561.

EUNICIDAE. 345

cephalic lobe, the presence or absence of tentacular cirri, and the condition of the
branchiae.

Dr. George Johnston (1865), following Audouin and Edwards, grouped the Eunicidae
according to the presence or absence of branchiae, dividing those devoid of gills into those
with antennae and those in which these were absent or rudimentary.

The same author 1 thought that, following the Amphinomaceae, the Euniceae, his third
family, made the connection between the first-named and the Nereids less abrupt. He
included seven species of Eunice m the British Fauna, viz., Eunice norvegica, E. annuli-
cornis, E. antennata, E. Harassii, E. gigantea, E. sanguined, and E. margaritacea. Two of
these, only casually alluded to by Dr. Williams, may be withdrawn, and probably refer to
well-known forms. The first two also are probably synonymous, and the last refers to
Marpliysa smiguinea. Besides these, Onuphis conchylega and Hyalinorcia tubicola, Lysidice
ninetta and Arabella tricolor, represent the other groups.

De Quatrefages (18G5) followed Schmarda in separating Savigny's family of the
Eunicidae into two, viz., the Eunicidae and Lumbrinereiclae, the one characterized by
branchiae, the other by their general absence. He thought the Eunicidae were the most
elevated type of Annelids and contained some of the largest forms. He divided the
family into two groups according to the number of the tentacles (his antennae), Eunice
and Marpliysa having five, whilst Diopatra and Onuphis have seven, a sub-division in each
being made by the absence or presence of tentacular cirri (his tentacles). He terms the
maxillae the superior maxillae ; the great dental plates are his inferior maxillae ; the
other pieces are comprehended under his two pairs of denticles. The mandibles are his
labrum.

Ehlers (1868) describes the Eunicea, Grube (char, auct.), as having an elongated
segmented body, cephalic lobe rounded, either with or without tentacles and palpi, or
neck-pads ; mostly with eyes. Foot generally single, or double, without appendages or
lobes ; dorsal and ventral cirri and branchiae with compound and simple bristles, or with
only simple bristles. Four anal cirri under the vent. Proboscis with jaws, the upper of
several joints, and with two plate-like lower jaws.

He 2 divided the family Eunicea into two great groups, viz., the Eunicea labidognatha
and the E. prionognatha. The former has the interposing pieces of the upper jaw
heterogeneous. The pieces following the " Trager," and the untoothed pincers (Zange),
become surrounded by a half circle of smaller pieces. The feet single, with several
bundles of bristles.

This he subdivided into those in which the two halves of the upper jaws have an
unequal number of pieces — one more in the left than in the right. Cephalic segment
with tentacles (E. labidognatha tentaculata). Under the second division of this group —
with five long posterior and two stumpy anterior tentacles are Diopatra, Aud. & Edw.,
with two tentacular cirri, and Onuphis devoid of them. Under the third division with five
tentacles and branchiae are Eunice, Cuvier, with two tentacular cirri, and Marpliysa, De
Quatrefages, in which these are absent. In the fourth division is Nicidon, with five tentacles
and no branchiae. In the fifth division occurs Lysidice, Savigny, devoid of branchiae.

1 'Cat. Worms Brit. Mus./ p. 129, 1865.
3 c Borstenwurmer/ ii, p. 280, 1868.

101

346 EUNICID^E.

Under the sixth division are Nematonereis, Schmarda, with dorsal cirri, and Blainmllea,
De Quatrefages, without them.

In the second series of this group both halves of the jaws have an equal number of
pieces; no true cephalic tentacles {E. labidog natha nuclei). It comprises Ninoe, Kinberg,
with branchiae, and Luvibriconerei.% Blainville, without them.

In the second great group (the Eunicea prionognatha) he makes two divisions, viz.,
first, the E. prionognatJta monocopa, in which the feet are single and the bristles simple.
In the first section the dorsal cirri are stunted or absent ; the first pair of teeth with
pincer-shaped hooks at the tip and alike. Aracoda, Schmarda, has five pairs of teeth,
whilst in Arabella, Grube, the first pair of teeth are unlike. In these all the teeth are
similar. In the group following the first pair of teeth is devoid of pincer-shaped ends. In
this he places Notocirrus, Schmarda, with stunted dorsal cirri. In the second division the
pinnse are bifid, with simple and compound bristles, and under this is Stawocephahis, Grube.
Claparede, in his 'Supplement' (1870), takes up the subject of the teeth of the
Eunicidas, and compares his nomenclature with that of Ehlers.

Ehlers : Claparede :

Trager . . . Support (most posterior piece).

Zange . . . Pince.

Zahn .... Deut, or piece dentaire.
Sageplatten . . . Plaques en scie

or or

Reibplatten . . . Plaques en rape (Claparede would

call latter paragnathes).
Maxillaire inferieure (for lower
jaw).
He is of opinion that while he may have minimized their importance, Ehlers has exagge-
rated it — in classification. He thinks that Ehlers' Labidognathes and Prionognathes pass
gradually into each other. Whilst the jaws may be utilized in a certain measure for tribes,
their employment is difficult in constituting genera, for the same form of jaw occurs in
different genera. The importance of the shape of each piece is less than Ehlers supposes.
Claparede found great variations in the jaws of the same species. He further observes that
the group (Eunicea) has large tubular fibres on the dorsal surface of the nerve-area.

At a later period Grube, in his ' Philippine Annelids,' describes the Eunicea (and he
includes Staurocephalus) as having a subtriangular or semi-oval head, with lobate palpi (his
subtentacles) more or less united, rarely lateral tentacles or none. Tentacles subulate
or elongate posteriorly, sometimes none. Eyes two or four or absent. Body long,
vermiform, often slender, sometimes rounded, iridescent, with numerous short segments,
ending in two or four anal cirri. Buccal segment biannular, two tentacular cirri or none.
Proboscis with paired or unpaired horny or calcareous plates. One superior maxilla with
hooked tips on each side, simple or toothed ; inferior laminae hooked anteriorly, toothed,
truncate or convex. Branchiae simple styles or pectiniform or subfascicular, occasion-
ally spiral, springing from the dorsal cirrus, or absent. Feet simple, dorsal and ventral
cirri, one spine or more ; one or two fascicles of bristles, often without cirri, but with a
setigerous lobe. Bristles simple, or along with compound bristles ; sometimes slender,

EUNICIML

347

scalp riform ; apex truncate and pectiniform. Inferior setae or spines posteriorly uncinate.
Some make tubes — hyaline, or with fragments of shells, stones, and sea-weeds.

He divided them into three main groups, viz., (1) the Labidognatha, with unequal
pieces in the jaw-apparatus : (2) the Lumbriconereidea, which he again subdivided into
those with leaf -like dorsal cirri, and those without dorsal cirri ; and (3) the Stauvo-
cephalidea.

Grube l subdivides Eunice as follows :

m t . . . f Frontal border 4-lobed . . S.-ff. Eriphyle, Kbg.

Two tentacular cirri . fe ** %

[ „ „ 2-lobed . . b.-g. Leodice, s.st. (xr.

No tentacular cirri . ,, ,, 2-lobed or entire . S.-g. Marphysa, Sav.

In the same paper he subdivides the sub-genus Leodice (1) according to the presence
of the branchiae in the middle region of the body and their absence posteriorly, and
(2) those species in which the branchiae are continued to the end of the body or nearly
so. He adopts the same arrangement with Marphysa. He appends Lysidice, Amphiro,
and Nematonereis without special heading, distinguishing Lysidice according to the
position of the median tentacle.

The Eunicidae form the first family of Levinsen's Euniciformia vera, the first division
of his Euniciformia (1883). Benham makes it the ninth family of his Nereidiformia.
Both link on the Glycericlas, the former as a division, and the latter as merely the
succeeding family of the sub-order.

Pruvot (1885) considered the appendages on the second segment of the body in
certain forms as homologous with the dorsal cirri. He gave a careful description of the
nervous system of several types of the group.

The nuchal organs are in the form of ciliated grooves, and may be protected by the
border of the first segment (Racovitza).

Ehlers, in his ' Florida-Anneliden ' (p. 67), groups the tentacle-bearing Euniciclaa as
follows : —

No. of

Without tentacular cirri.

With tentacular cirri.

tentacles.

Without branchiae.

With branchiae.

Without branchiae. With branchiae.

7

Onuphis
(Paronuphis)

Onuphis

Diopatra
(Paradiopatra)

Heptaceras

Rhamphobrachium

Diopatra

5

Paramarphysa

Marphysa

Nikidion

Eunice

o

O

Lysidice

Amphiro

—

—

1

Nematonereis

—

—

■ — ■

1 ' Jahresber. der schl. Gresellsch. fur vaterl. Cultur/ 1877, t. lv, p. 98.

348

EUNICIDAE.

One dorsal cirrus and
one ventral cirrus

He discusses the supposed jaws of Eunicidae found by Dr. Hmde in the Silurian
strata, and points out that no mandibles have been described. He mentions the Rotifera
as having similar structures. He also adheres to his former classification of the Eunicidae
as a whole.

Grravier 1 classifies the Eunicea as follows : —

Eunices — three tribes.
' Superior jaw formed of pieces, few in number
(generally three)
Superior jaw formed of small pieces (teeth),
numerous, and arranged longitudinally .
No ventral cirrus . Dorsal cirrus rudimentary or foliaceous

Grravier again divides Eunice into—
No branchiae . .

f Reduced to one filament more or less divided

' 1 Arborescent or pectiniform

[ Seven antennae ; no tentacular cirri, branchiae

cirriform ....

Less than seven antennae; with tentacular

cirri ....

Five antennae

With branchiae

With branchiae

Eunicidae.

Staurocephaliclae.
Lumbriconereidae.

Nicidon.
S.-g. Eriphyle.

S.-g. Leodice.

Hyalince'cia.

No tentacular cirri -

Eunice.

Marphysa (Nau-

phanta, Nausicsea,

Macduffia).
Ampl dro.

Nicidon.
Lysidice.

Nematonereis.

Three antennae

' With tentacular cirri : Five antennae
No branchiae . . Without „ ,, Three antennae

„ ,, ,, One antenna

Instead of giving numbers to the parts of the dental apparatus it has been thought
more simple to give names.

Metatrachophore and Nectochaete larvae of Eunicidae from Naples have been described
by Hiicker, 2 the former having two eyes and two pairs of bristle-bundles, the latter four
eyes and three pairs of bristle-bundles. He finds the Eunicid larvae generally have a
two-ringed prototroch. In the Plankton Expedition 3 he met with another Nectochaete
stage with two bristle-bearing feet on each side.

Though, so far as known, no British Eunicid is phosphorescent, Mr. Cyril Crossland
found that many foreign species exhibit it when killed in the dark. " The phenomenon
is especially well shown, under the influence of mechanical stimulus, and in dim light, by
Eunice afra, Peters, at Suez. The animal is strongly phosphorescent during removal
from the mud, blue light being given out from points just below each parapodium (mucus-
secreting pads of the ventral cirri), and a considerable amount of mucus is secreted at the
same time, and this remains luminous." He mentions also that Mr. Stanley Gardiner

1 C. Grravier, ' Nouv. Arch. Museum/ Paris, 4 ser., t. iii, p. 217, 1900.

2 'Zeitschr. f. wiss. Zoo!./ Bd. lxii, p. 78, 1896.

3 'Die pelagischen Polychast./ etc., p. 9, Taf. 1, fig. 2, 1898.

EUNICIDJ5. 349

found the same form phosphorescent in the Maldives. Eunice siciliensis, which some
consider a borer, is stated by Mr. Crossland also to be phosphorescent.

The distribution of the BunicidaB is world-wide, though they are apparently most
plentiful in the warmer seas. They range from tide-marks to 1240 fathoms.

Their food consists of both vegetable and animal structures — chiefly invertebrate —
with sandy mud and its contents.

The majority of them secrete tubes, and some of these are branched. Others frequent
holes in the limestone rocks, e. g., Eunice siciliensis, and some on this account have been
credited with the power of boring in solid, hard substances as do the Eunicids found in the
telegraph cables, but, as elsewhere stated, 1 there is as yet no direct proof that they do
so. Mr. Crossland has recently found both Eunice and Lysidice common in the coral
blocks of the Reel Sea, and perhaps may be able to clear up the ambiguity on this point.

Under this family fall the most conspicuous representatives of the "Palolos" of
foreign seas. These have already been alluded to in the ' Challenger ' volume 2 as well as
subsequently, 3 and it need only be added in connection with the Atlantic " Palolo " that
an explanation of the remarkable absence of heads is afforded by the recent observations
of A. Gr. Mayer, 1 who found that the ripe examples of Eunice fucata turn tail outwards
in their tunnels in the coral and limestone rock and detach the tail-end containing the
reproductive elements. These swarm at the surface, and shed, as the sun rises, their ova
or sperms by rupture or otherwise, whilst the shrivelled remnants fall to the bottom and
perish. The adults in the tunnels regenerate their tails. In this form the males are
orange posteriorly, the females greenish. Moreover Lysidice oeelef the " wawo " of
Amboina, in the Malay Archipelago, leaving its retreat, swarms on the second and third
nights after full moon in March and April.

The group of the Eunicidaa is one in which representatives of two divisions (E.
labidognatha and E. labidognatha nuda) are found in a fossil condition in the Solenhofen
slates, the genera being termed respectively Eunicites and Lumbriconereites® In both
instances the form of the dental apparatus considerably differs from that of the living
types. These fossil forms range back to the Silurian rocks of the United States, Canada,
Eno-land (Wenlock and Ludlow formations), Scotland, and Scandinavia, and a great
variety of minute jaws has been described and figured by Dr. George Jennings Hinde. 7
Comparatively few examples of the maxillse occur, the majority consisting of the great
dental plates, or of the antero-lateral plates. Not a single example of the mandibles
has been found, so that Mr. Hinde considers these ancient forms were devoid of them,

1 ' Challenger/ vol. xii, Annelida, p. 262.
3 Ibid., p. 257, 1885.

3 ( Ann. Nat. Hist./ ser. 7, vol. xv, p. 33, 1905.

4 Carnegie Instit., Washington, vol. i, pp. 107 — 112, pi. i, 1908. A similar condition is described
by Percy Moore in Eunice paloloides at San Diego (' Proc. Acad. Nat. Sc. Pliilad./ June, 1909).
Recently (April, 1910) Treaclwell lias found that moonlight is not necessary for the swarming of
Eunice furcata.

5 R. Horst, 'Mus. Harlem/ pp. 105—108, 1905.

6 Elders, l Ueb. foss. Wiirmer/ Cassel, 1869, and ' Zeitschr. f. wiss. Zool./ xviii, p. 421, 1868.

7 'Quart. Journ. Geol. Soc./ August, 1880, p. 368, pi. xiv; and ' Bihang till k. Svenska Vet.-
Akad. Handl./ Bd. 7, No. 5, p. 1, pis. i— iii, 1882.

350 STAITROCEPHALID^.

and thus differed materially both from those described by Ehlers and from the living
types. The persistence of these annelids (if the interpretation put on these remains
is sound) is remarkable, as well as their adherence to a size akin to those of modern
representatives. In some the jaws more nearly resemble those of the genus Eunice
(Eunicites), in others they approach (Enone (CEnonites), in a third they are near Arabella
(ArabellUes), and in a fourth have the f acies of those of Lumbrieonereis (Lumbriconereites) .
It is noteworthy that in the Silurian beds the jaws have been found singly — a condition
possibly due, according to Dr. Hinde, to the abundance in the same beds of Ostracoda,
which may have preyed on the soft parts, and thus rendered them more readily
separated by the movement of the water. The locking together, however, of the parts
in certain forms would obviate such dispersion. Various irregular track-like markings
were observed by Dr. Hinde on the rock- surfaces where the jaws occurred, and may have
been due to the living annelids. Few instances of the occurrence of such fossils have
been recorded between the Silurian and the present period. As indicated elsewhere,
Ehlers is inclined to connect the foregoing structures with other phyla of the animal
kingdom such as the Rotifers.

In this group are some of the giant Polychasts. Thus De Quatrefages found Eunice
Rousseaui from Banyuls 332 mm. in length and 16 mm. in breadth. On the other hand
Ophryotroclia puerilis, Claparede and Metschnikoff, is only a few millimetres in length, and
the male is unknown. This form has been found in the body-cavity of Gucumaria planci
at Naples, by Monticelli, 1 and Labroro stratus parasiticus by St. Joseph, in the body-
cavity of Eusyllis monilicorms, S. prolifera, Pionosyllis lamelligera, and Grubea clavata.
This shows a reduction of the maxillary apparatus, especially of the superior jaw. The
same author found others amongst algas (Rytiphlasa), so that parasitism may be temporary.

Oligognathus Bouellide, again, was found by Spengel in the general cavity of
Bonellia, and in this form the inferior jaw is much reduced. 2

Hsematocleptes terebellidis, Wiren 3 , occurs in the blood-lacuna in the wall of the
stomach of Terebellides Stroemii. The maxillary apparatus is also reduced.

Euniceas appear essentially to be dwellers in cracks and crevices, some especially
haunting muddy fissures (e.g., Marphysa sanguinea.) The bristles of this species and its
great power enable it to adhere so firmly to its tube either by the anterior or posterior
end that rupture ensues before extraction. Small twigs on the stem and branches of the
Antipathidas are modified to form a tube for these annelids (Marphysa and others).

STAUEOOEPHALIDiE.

Eunicea prionognatha dicopa (Ehlers, 1868).

In this group the dorsal longitudinal muscles are lappet-shaped in transverse section,
the broad ends being inferior. The ventral longitudinal muscles are massive with convex
borders inferiorly. The oblique muscles pass below the nerve-cords, which have a single

1 ' Monitore Zoolog. Italiana/ Fierenze, 1892, No. 12, p. 250.

2 f Mitth. Zool. Stat. Neap./ 3 e Bd., 1882, pp. 15—52, Taf. ii— iv.

3 ' Bihang till K. Sv. Akad. Handl./ Bd. xi, No. 12, 2 pis., 1886.

STAUROCEPHALID^. 351

neural canal median in position between the cords, dorsal at the ganglia, and occasionally a
small one is seen at the side as if from a branch. The cords are comparatively large.

Grube first established the genus Staurocephalus for the common species 8. rubro-
vittatus from Trieste. He afterwards constituted the genus Anisoceros for certain foreign
species, which further examination proved to fall under the former genus, and conse-
quently this lapsed. In his final publication on the group (1879) he arranged them
according to the presence or absence of a terminal segment to the subtentacle and
the proportionate length of the latter, the condition of the dorsal cirrus, tentacles, and
anterior segments, the eyes and other features being of service in arranging the
sub-divisions.

Kinberg 1 placed this group in the fourth division of his Eunicea, and distinguished
it as having the mandibles split, the maxillary series short, and the posterior appendages
of the maxillae short. The family itself (Staurocephalea) was characterized by the
numerous maxillae, the fourth series being toothed and touching laterally.

De Quatrefages (1865) included Prionognathus as one of the genera of his Syllids,
the type being P. ciliatus, Keferstein, whilst Staurocephalus was placed under genera
and species of uncertain position. He thought that the presence of a fine bristle in the
dorsal cirrus of Staurocephalus rubrovittatus indicated a bifid condition of the foot, but
Claparede argues that the same occurs in Sacconereis helgolandica, Fritz Miiller, in
Sigambra Orubei, in Eunice tsenia, and in Psamathe, and therefore thinks its importance
exaggerated.

Claparede (1868) observes that Grube, in his Annulata (Erstediana, constituted the
genus Anisoceras for several foreign species which he considered different from Stauro-
cephalus already established. But after a study of Anisoceras rubrovittatus he changed
the title, making Anisoceras a sub-genus. Unaware of Grube's observations, Keferstein in
1862 made a new genus, Prionognathus, for the same type. Fritz Miiller again pointed
out the close relationship of the genus with the Eunicidas, and Keferstein's view
corresponded, though he linked them also with the abnormal Syllideans.

The same author (1868) found a dorsal, two lateral and a ventral vessel in 8. Ghiaji,
and complex plexuses in the feet, cirri, and tentacles. The lateral and dorsal vessels send
forward, the ventral backward currents.

Levinsen (1883), after Kinberg, grouped the Staurocephalidas as the last of the
first section of his Euniciformia, viz., the E. vera, those preceding being the Eunicidas,
Onuphidas, and Lumbrinereiclas. It may be presumed that Benham includes the group
under his Eunicidas, the tenth family of his Nereidiformia, having in front the Amphino-
midas and in rear the Glyceridas, a classification which evoked the criticism of Miss
Florence Buchanan, who rightly objected to the author separating Polydora from the
Spionidas as an independent family while assimilating the Staurocephalidas with the
Eunicidas and placing Spinther, Euphrosyne, and Amphinome under one family.

Gravier (1900) considers, after Pruvot and Racovitza, that the Staurocephalidas are a
more primitive type than the Lumbriconereidas.

Claparede and Mecznikow 2 (1869) describe a young stage of Staurocephalus Ghiaji,
1 Ofvers. Kongl. vet.-Akad. Forlmndl., 1864, p. 574.
3 'Zeitsclir. f. wiss. Zool./ Bd.'xix, p. 186, Taf. xv, fig. 3.

352 STAITROCEPHALUS.

Clprd., from Naples, measuring 08 mm., of nine segments, four of which had cirri
(branchise). The snout is hoof-shaped, with two eyes, the tentacles arising immediately
behind them. Nuchal grooves occur at the posterior edge on each side, and a dental
apparatus is developed. The foot is fairly formed and the dorsal cirrus is articulated.

Genus LXI. — Staurocephalus, Grille, 1855.
Anisoceras, Grube. Prionognathus, Keferstein.

Head rounded, quadrangular or pentagonal, with two articulated tentacles, two
palpi, and four eyes. Nuchal organs primitive, ciliated surfaces in direct contact with
the exterior (Racovitza). Two segments devoid of appendages. Body with compara-
tively few segments. Pygidium with two shorter and two longer cirri.

Upper jaw with two rows of jaw-plates, the wide halves of the mandible diverging
posteriorly into narrower processes.

Foot long, bifid, the upper division with simple bristles, the lower with compound
bristles. Dorsal cirri unequal with a short terminal process. Ventral cirrus arising
from the foot, short and simple.

In transverse section the body-wall (Fig. 76) has somewhat thin dorsal muscles,

n.c. o.jii.

Fig. 76. — Transverse section of a nearly ripe male of Staurocephalus rubrovittatus. The ccelom is much distended.
d. Alimentary canal, n.c. Nerve-cord. o.m. Oblique muscles.

but the ventral are more massive in proportion. The ventral nerve-area is large, and the
oblique muscles pass downward to be inserted at its sides. At the reproductive season,
as in the section figured, the sexual elements largely distend the ccelom, and the muscular
layers undergo more or less atrophy.

One of the most interesting features in the genus is the occurrence of pelagic
"swarming" in a species (Staurocejohalus gregarious) from the Tortugas Islands, Florida,
as described by A. Gr. Mayer. 1 The entire worms leave the coral rocks, swim at the
surface, and discharge the sexual products soon after sunrise, e.g., on July 19th, when the
last quarter of the moon falls on July 18th. The author also gives figures of the eggs
and larvas.

G rube's first group contains those in which the palpi (his sub-tentacles) have no
terminal segment. It includes the common British species.

1 A. G. Mayer, < Bull. Mus. Comp. Zool./ Harvard Coll., vol. xxxvi, No. 1, 1900.

STAUROCEPHALUS RUBROVITTATUS. 353

1. Staueocephalus rubeovittatus, Grubs, 1855. Plate LV, fig. 1; Plate LXT, figs. 7-
7 c— teeth ; Plate LXXIII, figs. 4-4 a— feet ; Plate LXXXI, figs. 9 and 9 a— -bristles.

Specific Characters. — Head somewhat rectangular, with four brownish-black eyes,
the anterior considerably larger than the posterior and wider apart. Two short clavate
(four-ringed, Grube) tentacles arise between the eyes of each side, and extend nearly to
the anterior edge of the snout. Two curved or sickle-shaped thick palpi (sub-tentacles,
Grube) spring below the snout. A slight fillet occurs on each side of the head
posteriorly. Head and appendages pale.

Body somewhat fusiform, with fifty-three or more bristled segments, tapered
from the anterior third forward, and from rather more than the posterior third backward
to the tail, which ends in two short cirri, two longer being in front of them (Ehlers).
First two segments devoid of feet. The first is fine rose-red, the second is paler. Two
bars of the same colour on each segment. Ground-tint of the body is flesh or cream-
coloured. The mouth has two lips. The dorsal dental apparatus consists of a double
row of horny teeth which are pinnate, the median fang of the dorsal series in front
being prolonged into a slender process. The lower row is the stronger, is dark brown,
and has about twenty to twenty-one teeth. 1 The upper row has ten to twelve teeth
which are well developed. The mandibles are club or wedge-shaped, the broad anterior
edge being coarsely crenate. The foot has a stout dorsal cirrus with a terminal articu-
lation which projects beyond the tip of the setigerous region. A slender spine occupies
its middle. Setigerous lobe large, dorsal ridge ciliated, and the free border split
superiorly into two conical processes, and with convex edges inferiorly. The ventral
lobe projects beyond the adjoining part of the setigerous lobe. A single translucent
spine is present. The upper bristles are transparent, gently widening upward and
forming a flattened blade with a serrated edge. Lower bristles equally translucent,
curved, dilating at the distal end of the shaft, which is bevelled, and articulates with
a bifid terminal piece, the secondary process being separated by a considerable interval
from the terminal hook.

Synonyms.

1855. Staurocephalus rubrovittatus, Grube. Arch. f. Naturges., xxi, p. 97, Taf. iii, f. 9.

1860. „ „ idem. Ibid., xxvi, p. 78.

1861. „ „ idem. Ausfhig nach Triest, pp. 24 and 140, pi. 1, f. 10 and 11.

1864. „ ,, idem. Die Insel Lussin, p. 80.

1865. „ erucseformis, Malmgren. Nord. Hafs-Annul., p. 184.
J} „ rubrovittatus, De Quatrefages. Annel., ii, p. 83.

1866. Dorvillea lobata, Parfitt. Zoologist, 2nd ser., vol. i, p. 113.

1867. „ „ idem. Trans. Devon Assoc, ii, p. 23 (sep. copy).

„ Staurocephalus erucseformis, Malmgren. Annul. Polych., p. 62, Taf. viii, f. 50.

1868. Staurocephalus erucseformis, Ehlers. Borstenw., p. 424, pi. xviii, f. 1 — 16.

1875. „ rubrovittatus, Marion and Bobretzky. Ann. Sc. nat., 6 e ser., tome ii, p. 10.

1879. „ erucseformis, Langerhans. Zeitschr. f. wiss. Zool., xxiii, p. 299.

1885. „ rubrovittatus, Pruvot. Arch. Zool. exper., 2 e ser., t. iii, p. 274, pi. xi, f. 4;

pi. xii, f. 10—13.

1 Grube gives twenty-eight and twenty-one respectively.

102

354 STAUROCEPHALUS RITBROVITTATITS.

1885. Staurocephalus rubrovittatus, Oarus. Fauna Medit., p. 217.

1888. „ „ De St. Joseph. Ann. So. Nat., 7 e ser., t. v, p. 235, pi. x, f. 99.

1895. m> 9J Pruvot and Eacovitza. Annel. Banyuls, in Arch. Zool. exper., 3 e

ser., vol. iii, p. 349, pi. xv.
1902. „ „ Mcintosh. Ann. Nat. Hist., ser. 7, vol. x, p. 257.

}i y} „ Marenzeller. Polych. Grund., p. 15.

1904. „ „ Allen. Journ. M. B. A., n.s., vol. vii, p. 226.

1905. „ „ G-raeffe. Arbeit. Zool. Stat. Triest, xv, p. 322.

1906. „ „ De St. Joseph. Ann. Sc Nat., 9 e ser., t. iii, p. 215.

Habitat.- — The under surfaces of stones near low-water mark, and the larger stones
in pools at Herm (A. and R. M.), Devonshire (Col. Montagu), Plymouth (Allen).

Extends to the Mediterranean and the continental side of the Channel. Grube first
procured it in the Adriatic, Pruvot found it common in tufts of Gystoseira at Roscoff.
Apparently the same form is mentioned by Malmgren from deep water in Fin mark.

Head (Plate LV, fig. 1) somewhat rectangular, with four brownish-black eyes, the
anterior pair considerably larger (though hidden by the tentacles in the figure) than the
posterior, and wider apart. Two short clavate tentacles spring between the eyes of each
side, and extend nearly to the anterior edge of the snout. 1 Two curved or sickle-
shaped thick palpi arise below the snout and project on each side like horns. A slight
ciliated fillet is indicated on each side of the head posteriorly. 2 Head and appendages
pale, with palpocils on the anterior border (Pruvot and Racovitza).

Body of fifty-three or fifty-four bristled segments, and fully an inch long (15 — 29 mm.
Pruvot and Racovitza), somewhat fusiform, slightly tapered from the anterior third
forward, and from rather more than the posterior third backward, and terminating in a
moderately tapered tail with two short cirri and a dorsal anus. The first segment has
no foot and is broader than the second. It is tinted of a fine rose-red, the " organs
nucaux " of Pruvot and Racovitza, which are developments of the anterior border of
the first segment, are innervated by the posterior lobe of the cephalic ganglia.
The second is paler, and also is devoid of feet. Bands of the same rose-red hue
cross the body from the anterior to the posterior end, being fainter in front and behind.
Each segment has two bands which cross the anterior and posterior parts (with a pale
belt between), diminishing to a point before reaching the bases of the feet. The tint of
each posterior band is somewhat deeper. The feet and cirri, with the ground-tint of the
body, are flesh or cream-coloured.

The upper dental apparatus consists of two rows of dark brown horny teeth, the
inferior and inner (Plate LXI, fig. 7) being the stronger. This row commences anteriorly
in an irregularly differentiated horny band, which soon rises into distinct teeth separated
from each other, and of a complex nature. Twenty-one teeth on one side and twenty on
the other are visible in this part of the apparatus. These teeth (Plate LXI, fig. 7 a), like

1 Ehlers represents four articulations in his examples, hut those procured in Herm, whether
imperfect or otherwise, were as described and as Pruvot and Racovitza state. In S. Chiaji,
Claparede, there are six segments.

3 In the S. Chiaji of Claparede two ciliated pits, supposed to be sensory, occur at the posterior
border of the head; and two other vibratile pits are present inferiorly at the sides.

STAUROCBPHALUS RUBROVITTATUS. 355

those of the upper (outer) row, conform to the same type, though variously modified.
They are split at the base into two flattened plates which lie on each side of a lingual
band, and give attachment to the muscles moving the tooth. The median tooth is
powerful, and, in the drawing, is by no means acute, having apparently been in active
use. On each side are four smaller teeth, the last being minute and often obscured when
in situ. They extend along the ridge in a somewhat intricate manner, the teeth having
the longest processes being in front of the middle of the row, whilst the posterior teeth
are more compact and their processes smaller. The curve at the tip is also more decided.
The last tooth abuts on a long serrated horny band, the crenations or teeth being much
larger than those on the corresponding portion of the outer and upper row.

The outer and upper row has ten or twelve distinctly developed teeth (Plate LXI,
fig. 7 b). They also occupy a ridge clasped by the bifid base, each lamella of which is
somewhat expanded at the tip. A few anteriorly are small, but they soon show an
elongated median tooth which projects into the pharynx like a spine slightly bent
toward the tip. About five have the long median process, though several of those
succeeding have a tendency to the same condition. These teeth diminish posteriorly,
sometimes with an irregular mass or two, and then end in a long bar with a finely
denticulated edge.

This apparatus evidently is of importance in feeding, the more powerful lower row
performing the sections, whilst the more slender upper series probably aid in holding or
manipulating the morsels, the long horny processes especially being useful in this respect.

The external rows of rudimentary teeth indicated by Ehlers either were not present
or had been overlooked in the dissection. These small teeth form a single row beginning
externally rather in front of the middle, and continuing backward to the commencement
of the posterior bar, when two rows occur, a shorter inner and a longer outer.

The mandibles are club- or wedge-shaped, broad in front and tapering to a point
posteriorly. The broad anterior border (Plate LXI, fig 7 c) is coarsely crenate at the
outer edge, but very finely denticulated on its inner edge anteriorly. The two mandibles
project beyond the lower lip in some preparations.

The alimentary canal contains fragments of Crustacea, sand-grains, spicules of
sponges, and other debris.

The minute first foot presents three pointed lobes, a single pale spine and a group of
bristles. Two of the lobes are superior, one bearing the spine, and below it the bristles,
which have jointed bifid tips and dilated ends of the shafts with bevelled edges. The
other is the ventral cirrus.

The feet gradually increase in size, so that at the fourth or fifth there is dorsally a
thick cirrus, with a conical articulation at the tip (cirrophore and cirrostyle, Pruvot and
Racovitza), and internally two slender spines which pass almost to the tip of the organ.
Beneath are a trifid setigerous lobe— the pale spine passing between the upper divisions—
and also the simple bristles, which are slightly curved toward the somewhat blunt end
and finely serrated on the upper or convex edge. Along the lower edge of the furrow of
the trifid region is the dense group of the translucent jointed bristles, the shafts having
dilated and bevelled ends and bifid terminal pieces. The ventral lobe of this division is
broadly lanceolate.

356 STATTBOCEPHALtTS RUBROVITTATITS.

The typical condition of the foot is assumed about the tenth and subsequent feet
(Plate LXXIII, fig. 4) toward the middle of the body. Dorsally is the stout dorsal
cirrus with the terminal articulation, its tip projecting considerably beyond the soft parts
of the foot beneath. The slender spine occurs internally as in the foregoing foot.
Claparede 1 describes the basal segment as having four parallel blood-vessels, which do not
enter the terminal piece, but anastomose with each other. He considers this the homo-
logue of the short ceratophore of the majority of Annelids. " The setigerous lobe is lar^e
and strong, its dorsal ridge richly ciliated, and the free border split into two conical
processes superiorly and convex edges inferiorly. The ventral cirrus projects beyond
the adjoining part of the setigerous lobe. The bristles of the dorsal part of the setigerous
lobe (Plate LXXXI, fig. 9) are transparent and elongated, gently widening upward and
forming a flattened blade with a finely serrated edge, the tip being curved and ending in
a slightly tapered point. Some of the bristles have a broader and shorter flattened
region at the tip than that figured. The serrations are elevated distally. Pruvot and
Racovitza describe them as having an irregular border. The bristles of the inferior
group are more numerous, but equally transparent, the shaft dilating into a somewhat
large distal region, which is bevelled and carries the rather elongated and minutely
serrated terminal piece, which is bifid (Plate LXXXI, fig. 9 a), the terminal hook
being separated by a considerable interval from the secondary process, and thus charac-
teristically differing from such types as Stawroceplialus Budolphii, from Naples, a
form in which the secondary process passes distally like a knife-blade and touches the
terminal hook.

The presence of a spine in the dorsal cirrus, so called, raises the question as to
the composition of such a foot, De Quatrefages interpreting it as an indication of a
dorsal division, whilst Claparede thought differently.

The feet retain the same structure to the posterior end of the animal, but they
diminish in size as in front. Claparede showed that the feet discharge many bacillary
bodies from follicles.

In an example caught in a tow-net off Sark by Mrs. Collings in June, 1869, and
measuring about three quarters of an inch, the feet (Plate LXXIII, fig. 4 a) differed
considerably from those of the foregoing ripe male. Thus the dorsal cirrus was less
massive, the inferior processes of the setigerous region were more acute in lateral
view, and the bristles, both simple and compound, were more attenuated. The terminal
pieces of these, however, maintained the character of the species, the terminal hook
and the secondary process being separated by a considerable curved interval, though
the process was somewhat shorter (Plate LXXXI, fig. 9&).

Reproduction, — Most of the examples procured in July at Herm were males in which
the body-cavity was distended with sperms. These also invaded the feet, almost to the
tip, and a column of them passed along the centre of the dorsal cirrus. Pruvot and
Racovitza give July as the breeding period, and add that most of the young have a single
pigment-band on each segment. In transverse section the body of the male shows
greatly stretched muscular layers, whilst the alimentary canal forms an elliptical
central slit, apparently without food.

1 ( Annel. Chet. Nap./ i ; p. 118, pi. vii, fig. 2 b, f.

STAITROCEPHALUS PITBPOVITTATUS. 357

The segmental organs in this form are visible as transparent objects in the yonng
examples (Fage 1 ). The external aperture is placed above the glandular epidermic pad
at the base of the foot.

Habits. — It is active like a Hesione and appears to be tolerably hardy. De St. Joseph
states that it secretes much mucus. Eisig 2 describes the Neapolitan form, S. Rudolphii,
D. Ch., as swimming freely in an undulating manner, and performing various movements
even after decapitation.

It is possible that further examination may reveal a closer approach between this and
allied forms such as Delle Chiaje's species named by Claparede S. ChiajL Moreover the
Nereis Budolphii of Delle Chiaje 3 appears to be the present species. In the ' Descrizione '
it is termed Syllis Radolphiana. 4i

In Montagu's MS. volume of drawings (1808) in the Linnean Society is a figure of
this species (Plate XLVII, fig. 1), named Nereis pennata r and with" Dor inllea lobata"
pencilled opposite. The proboscis is thrust out and coloured pink. Only two eyes are
represented. It is clear, therefore, that Col. Montagu was the first naturalist who
procured the species.

This is one of the numerous species the accurate description of which science owes
to the indefatigable exertions — continued throughout a long life — of Prof. Ed. Grube, of
Breslau, who found it in 1855 in the Adriatic at Triest, Fiuma, and Cherso. An amended
description, generic and specific, was published a few years later (1860). He held the
foot to be uniramous, and the blood red. Grube also pointed out its differences from
Anisoceras vittata, rubra, and bioculata, Grube and CErsted.

Pruvot agrees with Ehlers in considering the inferior appendages of the head to be
palpi. He found the nervous system of this species conform to the type in the Eunicicla3.

Pruvot and Eacovitza 5 describe the first segment as having four red bands. More-
over they mention two varieties, viz., those with the two bands on each segment, which
are commonly dredged and which secrete a tube of mucus, and the variety with one band
which inhabits dead colonies of Dendrophyllia and Amphihelia and also occurs amongst
Serpuhe. The authors give an elaborate description of the structure of the foot and of
the proboscis and its armature.

The accurate comparison of the American forms of this group has yet to be carried
out, and the connection between such species as the Staurocephalus pallidus of Verrill,
and 8. sociabilis of Webster, from the American shores, and the British forms, may be
closer than is at present supposed.

The following fall under Grube's second main division in which the palpi are jointed,
whilst the dorsal cirri are very short and without an articulation, or absent.

1 < Ann. Sc. nat./ 9 e ser., hi, p. 349, 1906.
» f Fauna u. Fl. Neap./ xxviii, p. 211, 1906.

3 < Mem./ hi, pp. 166, 176, Tav. xliii, figs. 13, 14, 19.

4 T. iii, p. 95.

5 ' Arch. Zool. exper./ 3 e sei\, vol. hi, pp. 349—373, pi. xv, 1895.

358 STAUEOCEPHALUS KEFEESTEINI.

2. Staueocephalus Kefersteini, Mcintosh, 1869. Plate LV, fig. 2; Plate LXI, fig. 8—
teeth; Plate LXXIII, fig. 5— feet; Plate LXXXI, figs. 10-10 b— bristles.

Specific Characters. — Head bluntly conical; eyes two, black, at its posterior border.
A moniliform tentacle springs from the outer side of each eye ; palpi long with terminal
articulation. Body slender, elongate, terminating in two articulated cirri. First of the
two naked segments the broader. Colour orange, with pale head, tail, and feet. Proboscis
with a double row of teeth as in 8. rubrovittatus, but smaller. Lower and inner row of
dark brown curved teeth with a median and one or two lateral points, with lateral wings.
Interiorly it terminates in a long process with alternately large and small teeth. Outer
and upper row consists of amber-coloured teeth of similar but more slender form.
Toward the lower end more than one small tooth intervenes between the longer. Man-
dibles brownish-black, clavate, and with about five denticulations on the edge. Foot when
fully developed stout, with ovate dorsal lobe. Setigerous region with a single pale spine at
the base and a few stout, straight, translucent bristles, the shaft being dilated and then
bifid at the end, one end having a claw, the other being simply pointed. Below are a
few slender bristles with straight shafts and curved tapered tips which are finely serrate.
Beneath the spine are five or six bristles with curved shafts dilated at the end, and with
a few serrations, and terminal pieces which vary from above downward. The tips of the
latter are really bifid.

Synonyms.

1869. Staurocephalus Kefersteini, Mcintosh. Trans. E. Soc. Edin., xxv ; pt. ii ; p. 417, pi. xvi ; f. 11 a-a.
1879. , } , } Grube. Jahresb. Schles. Gresellsch., p. 112.

1895. ;; „ Pruvofc and Eacovitza. Arch. Zool. exper., 3° ser. ; vol. iii ; p. 372.

1902. „ „ Mcintosh. Ann. Nat. Hist. ser. 7, vol. x, p. 257.

Habitat. — Under stones near low-water mark on both eastern and western shores of
North Uist, Outer Hebrides. One occurred in an old tube of Serpula and another was
dredged in Loch Portan (an inland sea more or less brackish) in three to four fathoms.

Head (Plate LV, fig. 2) conical, snout rounded, forming the blunt apex of the cone.
Eyes two, of moderate size, black, situated toward the posterior border of the head.
A tentacle springs from the outer side of each eye, and is annulated or moniliform.
They seem to be about as long as the diameter of the head in life. The palpi arise from
the infero-lateral region of the head, have a short terminal articulation, marked off by a
constriction, and are proportionally long. They are often coiled and twisted in a grace-
ful manner, and both they and the tentacles are pale.

Body slender, elongated, composed of many segments, and terminating posteriorly
in two pale articulated cirri of moderate length and with two shorter cirri in front of
them, shorter than in Keferstein's S. ciliatus. In general outline it resembles a Syllis,
and is about an inch in length. Colour orange, paler at head and tail, the feet being also
pale. The orange band on the dorsum curves inward opposite each foot, and slightly
outward at each segment-junction. Two reddish lines proceed backward from the head
and appear to be due to vessels, as waves passed from behind forward. In the example
from Loch Portan a faintly marked dark bar occurred at the posterior part of each

STAUROCEPHALTTS CILIATUS. 359

segment on eacli side of the intestine. The first two segments are devoid of feet, and in
the preparation the anterior seems to be the broader.

The proboscis is armed with a double row of teeth as in Staurocephalus rubrovittatus,
but from the diminished size of the animal they are proportionally smaller. The lower
and inner row consists of dark brown strongly curved teeth with a median and one or two
lateral points, with the basal wings for support. Each terminates inferiorly in a long
process, which has alternately large and small teeth. The outer and upper row consists
of amber-coloured teeth of similar but more slender form and terminating inferiorly in a
similar process with smaller sharp teeth alternately larger and smaller. Towards the
lower end more than one small tooth intervenes between the larger.

The mandibles are clavate and of a brownish-black colour. The free edge is coarsely
denticulated (Plate LXT, fig. 8), about five teeth occupying the edge, the last or outer
with an accessory process.

The anterior feet are smaller and somewhat rudimentary, but when fully developed
(Plate LXXIII, fig- 5) have a stout form, and a somewhat ovate or ovato-lanceolate
dorsal lobe narrowed at its attachment and projecting above the large setigerous region.
This has superiorly a single pale spine, the point of which just touches the surface.
Above the spine are one or two stout, straight, translucent bristles, the shaft of which
dilates and is bifid at the end (Plate LXXXI, fig. 10), the longer limb of the fork slightly
flattened and with a claw at the tip. The other is simply pointed. Below these are
one or two slender bristles with straight shafts and finely tapered tips which are
definitely curved and finely serrated from a little below the bend — upward along the
tip on the convex side of the curve — which is usually directed dorsally (Plate LXXXI,
fig. 10 a). The bristles beneath the spine are generally five or six in number, translucent
with curved shafts, dilated at the end, and with a few serrations (the convexity being
directed dorsally), and terminal pieces which form a diminishing series from above down-
ward, the upper being the longest and the lower the shortest (Plate LXXXI, fig. 10/)).
The tip of the terminal piece, at first thought to have a simple hook, is really bifid, for
close to the terminal hook a minute secondary process is present. The setigerous region
is produced outward inferiorly into a prominent papilla at the origin of the inferior
bristles. The terminal pieces do not show serrations on the edge in the preparations, but
they may have had them when fresh. The ventral lobe arises nearer the body than the
dorsal, but has a similar shape, the base being narrowed at its attachment to the foot.

The example from which the drawings were made was a female with small eggs
showing nucleus and nucleolus in August.

Habits. — They often twist themselves into a circular ball, crawl about frequently on
their sides with a wriggling motion, and can also swim in a similar manner. They are
on the whole active and lively.

3. Staurocephalus ctliatus var. Pobertian/e. Plate LXI, figs. 9 and 9 a — teeth; Plate
LXXIII, fig. 6— foot; Plate LXXXI, figs. 11-11 6— bristles.
Specific Characters. — Head flattened and tapered anteriorly to a blunt snout. Eyes
two, large, black. Palpi arising laterally in front of the eyes and having a terminal arti-

360 STAUROCEPHALUS CILIATUS.

dilation. Tentacles moniliform, scarcely half as long, much smaller in diameter than the
palpi, and springing from the dorsum of the head behind the eyes. Body about one inch
long, pale lavender, a yellow belt bounded by a red line on each side along the centre,
slightly tapered in front and more definitely diminished posteriorly, where it ends in two
caudal cirri. Proboscis has in the lower row a dense series of curved dark brown
denticles with a median and two lateral teeth, the maximum development occurring
toward the posterior third, and terminating in a thickened dark brown band with
irregular denticles. Upper row begins anteriorly in a series of pale brown horny fila-
ments which merge into the long teeth of the row. Posteriorly they diminish in
length, but increase in robustness, and terminate in a band of small denticles. An
accessory row externally on each side. Mandibles shaped like a boomerang, with a
series of eight deeply cut teeth on the anterior edge, besides three separate denticles
externally. Feet generally uniform, distinguished by a well-developed dorsal cirrus,
which commences on the second foot, containing a slender spine which goes as far as the
articulation of the considerable terminal process. Setigerous region large, bevelled
superiorly, and trending to the prominent inferior border. The obovate ventral cirrus
projects beyond it, less, however, than the dorsal cirrus. Two kinds of bristles
superiorly, viz., a long slender translucent form with an abrupt curve below the serra-
tions and a finely tapered tip, and a stronger bristle or two with a straight shaft and a
dilated, bifid extremity, one of the legs of the fork being shorter, broader, and more
acute at the tip. The inferior bristles compound, with curved shafts, and dilated, bevelled,
and serrated distal ends. Terminal piece with a bifid tip.

Synonyms.

1862. Prionognathus eiliatus, Keferstein. Zeitschr. f. wiss. Zool., Bd. xii, p. 99, Taf. viii,. f. 13 — 19.

1885. Btauroce'phalus Chiajei, Cams. Fauna Medit., p. 217.

1902. „ (third), Mcintosh. Ann. Nat. Hist., ser. 7, vol. x, p. 257.

„ ,, (fourth), idem. Ibid., ser. 7, vol. x, p. 257.

1904. „ ciliatus, Allen. Journ. M. B. A., n.s., vol. vii, p. 226.

Habitat. — Under stones between tide-marks, St.- Peter Port, Guernsey, July, 1868
(W. 0. M.) ; Plymouth (Allen).

Head flattened and tapered anteriorly to a blunt snout. Byes two, black, large
and distinct. Palpi arising from the sides of the snout in front of the eyes and having
a terminal articulation. Tentacles, scarcely half the length and much smaller in diameter
than the palpi, springing from the dorsum of the head behind the eyes, and moniliform
throughout.

Body upwards of an inch long (25—30 mm., Keferstein), of a pale lavender hue,
marked along the centre of the dorsum by a yellow belt, bounded on each side by a red
line, probably from the blood-vessels. It is slightly tapered in front and more definitely
so posteriorly, where it ends in two caudal cirri. Segments numerous, the first and second
being devoid of feet. When preserved the feet of the anterior fifth are crowded, those
behind are less so.

Proboscis.— In this and allied forms the food first comes under the action of the
serrated edges of the mandibles, and is then passed inward (upward) to the double

STAUEOCEPHALUS CILIATUS. 361

row of hard horny points which guard each side of the pharyngeal slit. The elaborate
series of muscles which move these denticles, and their conformation, denote important
functions. In this species (Plate LXI, fig. 9) the lower row has a dense series of strongly
curved dark brown denticles with a median and two or three lateral teeth. They have
their maximum development toward the posterior third of the row, whereas in front they
form a comparatively even series of smaller and less dusky teeth. Posteriorly the row
terminates in a thickened dark brown band with irregular denticles. The upper row
commences anteriorly in what seems to be a series of pale brown horny filaments which
merge into the somewhat long teeth of the row, the curves of which, like those of the
lower row, are directed backward. Posteriorly the teeth diminish in length but increase
in robustness, and have toward the inferior end a brown hue as deep as that of the lower row.
The row terminates in a band of small denticles. Besides these rows a less developed
accessory row occurs to the outer side of them, and on one side a similar row on the
inner side. Ehlers shows a similar accessory row in Staurocepl talus rubrovittatvs.

The mandibles (Plate LXI, fig. 9 a) are shaped somewhat like a boomerang, and have
on the produced anterior edge a series of eight deeply-cut teeth, besides three separate
denticles at the outer edge.

The nerve-cords in the preparations are flattened in transverse section, but such may
be due to the methods of preparation.

The feet of this form are distinguished from those of the Hebridean species by the
presence of a well-developed dorsal cirrus, which appears on the second foot. At the
twentieth foot (Plate LXXIII, fig. 6) the parts have attained an average outline and
structure. Dorsally is the stout cirrus with a slender central spine proceeding as far as
the articulation of the terminal process, which is, proportionally, of considerable length.
The setigerous region is large, bevelled superiorly, the outline in lateral view trending to
the prominent inferior border whence the compound bristles arise. This prominent
region attains even greater development posteriorly. Inferiorly, at some distance from
the edge, the obovate ventral lobe projects outward beyond the setigerous region, less,
however, than does the dorsal cirrus.

Superiorly two bristles occur, viz. (Plate LXXXI, fig. 11), along, slender, translucent
bristle with a somewhat abrupt curve below the serrations and a finely tapered tip, on
the slender terminal region of which the serrations become indistinct. They are directed
distally, and are most distinct inferiorly. Accompanying this is a stronger translucent
bristle (or two) with a straight shaft and a dilated, bifid extremity (Plate LXXXI, fig. 11 a),
one of the legs of the fork being shorter, broader, and more acute at the tip, the other less
expanded and longer. Traces of two spines occur on the edge beneath. The inferior
group consists of compound bristles (Plate LXXXI, fig. lib and b ! ) with translucent shafts,
curved, with the convexity upward, and dilated and bevelled distal ends which are also
serrated. The terminal piece, as usual, varies in length, diminishing from the upper to
the lower bristles, one of the shorter being represented in the figure. The tip is bifid, a
secondary process passing forward nearly parallel with the axis of the terminal piece to
the distal hook, and thus differing from such as have the secondary process projecting at
a larger angle.

103

362 STAUROCEPHALUS CILIATUS.

The feet do not alter materially in character posteriorly, except that the lower part
of the setigerons process at the exit of the inferior bristles is more prominent.

Reproduction, — The specimen is a female, with comparatively early ova (July). On
the other hand an example from Plymouth, kindly sent by Dr. Allen, had fairly developed
ova in June.

This species approaches Keferstein's Prionognathus ciliatus, and it may be that the
differences are due to imperfections in description and figures. The additional pair of
small eyes behind the tentacles is a marked distinction, though variation may occur in this
respect, yet in colour, structure of feet, bristles, and other parts, it closely corresponds.
The description of the teeth, however, differs, and he figures the ventral bristles as having
a simple instead of a bifid tip. Moreover, Grube places weight on the occurrence of only
two eyes, which is the condition, for instance, in his 8. bioculatus from Costa Rica.

This appears to be distinguished from Claparede's S. Ghiaji by the presence of only
two eyes, by the much longer palpi, and the shorter tentacles, by the absence of the dorsal
peristomial papilla, by the absence of the elongated spines on the simple bristles, by the
absence of boldly serrate compound bristles, and in other particulars.

So far as can be observed the Staurocephalus pallidus of Langerhans 1 approaches
closely to S. ciliatus, Keferstein, and this conclusion is borne out by an examination of an
example from Plymouth, kindly sent by Dr. Allen. The example has two very distinct
eyes at the inner border of the tentacles.

Claparede and Mecznikow 2 describe and figure a young example of the allied
Staurocephalus Ghiaji, 0*8 mm. long, with two projecting lateral lappets of the pro-
stomium, and a pair of unjointed tentacles with palpocils at the tip. The dental appa-
ratus is present, and there are two dorsal organs on the peristomial segment. The feet
are fairly formed with a jointed dorsal cirrus.

Another form, which may be termed provisionally Staurocephalus B, requires a brief
notice : —

Habitat. — Procured in a trammel net off Fermain Bay, Guernsey, in July, 1868.
The net was placed at a considerable depth — near the bottom.

Head furnished with four red eyes, and probably having the typical structure, though
such was not observed in the example, which was injured.

Body elongated as in Staurocephalus Kefersteiui, and of a pale yellow hue, brownish
from the intestine, posteriorly. A line of white pigment passes along the middle of the
dorsum from the proboscis backward. Posteriorly the line becomes double from an
intervening streak and disappears towards the tip of the tail.

The feet closely agree with those of Staurocephalus Kefersteiui in regard to general
outline, the proportions of the dorsal and ventral lobes (the latter being somewhat smaller
and more elongate), and the structure of the bristles. The compound series inferiorly
show the same very minute secondary tooth below the distal hook. The bifid bristles
of the upper part of the setigerons process are identical, the only difference observed
being the absence of the curve in the slender tapering bristles of the upper series,

1 ' Zeitschr. f. wiss. Zoo!./ xxxiii, p. 300, Taf. xvi, fig. 35, 1869.

2 Ibid., xix, sep. abd., p. 24, Taf. xv, fig. 3.

STAUEOCEPHALUS PALLIDUS. 363

and the absence of serrations. The great length, straightness, and slenderness of the
dorsal bristles may be connected with a sexual change associated with the largely
developed ova.

The example is a female laden with large ova, each of which has its longer diameter
exceeding the vertical diameter of the foot.

Unfortunately the head and anterior region were injured, so that considerable doubt
must remain. It differs from Staurocephahis pallidus, Langerhans, in the presence of
four eyes, and in the structure of the bristles.

4. Staubooephai/us pallidtjs, Langerhans, 1879. Plate LXXXVII, fig. 9— head,
9 a— foot; Plate LXXXVI, figs. 10, 10 a, and 10 b— bristles.

Specific Characters. — Head with only two eyes ; tentacles nine to ten, ringed, and
longer than the two-jointed palpi. No nuchal grooves. Body 1 cm. long, with fifty
segments. First two without feet and third without a dorsal cirrus. Anal segment with
two longer jointed dorsal and two smaller simple ventral cirri. Lower jaw with from seven
to ten teeth on either side, and three to five free denticles. Upper jaw as in other species,
only with two rows of teeth without accessory jaws. The plate of the upper row has
about twenty-five teeth, simple in front but posteriorly denticulated. That of the lower jaw
is a thin toothed leaf, having smooth upper but denticulated lower teeth. Foot as long as
breadth of body ; dorsal bristles simple, serrated, or with a distal process ; in the fourth
segment simple, serrated; twelfth segment with long simple tips above the serrated edge ;
in the thirteenth segment stout bifid forms. Inferior division has compound bristles, the
terminal process having a hook distally and a process beneath (Langerhans).

Synonyms.
1879. Staurocephalus pallidum, Langerhans. Zeitschr. f . wiss. Zool., Bd. xxxiii, p. 300, Taf . xvi, f . 35.
1904. „ „ Allen. Journ. M. B. A., n.s., vol. viii, p. 226.

Habitat. — Dredged on Asia shore, Plymouth (Allen).

Head (Plate LXXXVII, fig. 9) with only two eyes ; tentacles with nine double rings,
longer than the bi-articulate palpi. No nuchal grooves observed.

Body about 1 cm. in length (Langerhans), and having fifty-nine segments. The first
two segments are devoid of bristles, and the third has no dorsal cirrus, which, as a two-
jointed structure, appears on the fourth. Anal segment with four cirri, two longer jointed
dorsal cirri, and two smaller simple cirri ventrally.

Lower jaws armed with seven to ten teeth on either side, and three to five denticles.
Upper jaw as in other species, only with two rows of teeth, without secondary jaws.
The plate of the upper row is compact, with about twenty-five teeth which are more
simple in front, but posteriorly denticulated. The plate of the lower jaw is a thin,
toothed leaf, with smooth teeth in the upper but denticulated teeth in the lower row.

Foot as long as the breadth of the body, with the bristles of the upper row simple, or
with a distal process and serrated edge beneath ; in the fourth segment simple, serrated ;
at the twelfth segment with long simple tips above the serrated region ; at the thirteenth
segment the bristle is bifid, with a few serrations on the curved limb. In the inferior

364 OPHRYOTROCHA PUERILIS.

division compound bristles, the terminal process having a hook distally and a process
beneath it (Langerhans).

Langerhans thinks that this form agrees with Delle Chiaje's rubra and vittata in
the upper bristles, but possesses only two eyes.

The foot (Plate LXXXVII, fig. 9 a) differs from that of 8. rubrovittatus in the
inconspicuous condition of the two upper processes, but it more nearly approaches that of
8. Kefersteini.

The dorsal bristles (Plate LXXXVI, fig. 10) are translucent, long, slender and finely
tapered, with serrations on the edge below the long smooth tip, whilst the bifid forms
(Ibid., fig. 10 a) have a longer and a shorter limb, about half a dozen serrations occurring
on the edge below the latter. The ventral series of falcate bristles (Ibid., fig. 10 b and b')
has the terminal piece long and slender as well as bifid.

The exact position of the Plymouth example is uncertain. It may be immature.
The figures of Langerhans certainly differ from the structures observed in this form,
but they may not be accurate.

Genus LXII. — Ophryoteocha, Glaparede and Mecznikotv, 1869.

Small forms with horse-shoe-shapecl head ; eyes four ; two short dorsal tentacles in
front of the eyes, and two similar tentacles ventrally. Body comparatively short, first
two segments devoid of feet ; two clavate cirri posteriorly. Proboscis armed with a
complex upper dental apparatus, and an /-shaped mandible. Foot resembling that of
Staurocephalus, with simple upper bristles and compound lower bristles. Oviparous.

1. Ophryotrocha puerilis, Glaparede and Mecm., 1869. Plate LV, fig. 9 — head and
anterior region; Plate LVI, figs. 4 and 5; Plate LXI, figs. 10 and 10 a — young
and teeth; Plate LXXIII, fig. 7— foot ; Plate LXXXI1, figs. 1 and 1 a— bristles.

Specific Characters. — Head horse-shoe shaped, with palpocils anteriorly. Eyes
variable, generally four, two situated externally in a dimple behind the dorsal tentacle,
and two smaller on the summit of the head just in front of the nuchal fold. Occasionally
a pigment-spot or two in front of the latter. In front of each anterior eye a small
tentacle of two segments, the basal shorter than the distal. A pair of tentacles of similar
structure on the ventral surface. Body nearly half an inch long, of more than thirty
segments, slightly tapered at each end, pale and translucent. First two segments
devoid of feet. Two clavate cirri posteriorly. Proboscis with dark brown or blackish,
strongly curved maxilte, and with an acute posterior process. Six denticles beneath and
in front, each hoof-shaped, with the free edge denticulated, and diminishing in size
from before backward. Mandible elongated and curved like the letter /, cutting
edge bifid, each cusp being arcuate. Foot resembles that of 8taurocephalus, with four
conical terminal processes. At the upper end is the conical dorsal lobe. Beneath is a
conical process for the spine, one margin of the foot trending obliquely into the setigerous

OPHRYOTROCHA PUERILIS. 365

region, whilst the other leads to the ventral lobe (cirrus) nearer the body. Dorsally are
three translucent bristles with slightly flattened tips, which end in a hook. Below the
spine are about six compound bristles, the ends of the curved shafts dilated and bevelled
for the terminal piece with a simple (?) tip.

Synonyms.
1869. Ophryotrocha puerilis, Claparede and Mecznikow. Zeitschr. f. wiss. Zool., Bel. xix, p. 184,

Taf. xiii, f. 2.
1878. „ Glaparedii, Studer. Arch. £. Naturges., xliv, p. 119, pi. v, f. 11.

1884. „ minimus, Langerhans. Zeitschr. f. wiss. Zool., Bd. xl, p. 257, Taf. xv, f. 16.

1885. Staurocephalus Siberti, Mcintosh. Ann. Nat. Hist., ser. 5, vol. xvi, p. 482, pi. xiii, f. 5—8.
„ Ophryotrocha pueri lis, Carus. Fauna Medit., p. 217.

1886. „ ., Yiguier. Arch. Zool. exper., 2 e ser., t. iv, p. 417, pi. xxv, f. 11—17.

„ Paractius littoralis, Marenzeller. Die Osterr. polarstat. Jan Meyen, III, Zool. Wiirmer, p. 21.

1888. Ophryotrocha puerilis, De St. Joseph. Ann. Sc. nat., 7 ser., T. v, p. 239, pi. x, f. 96—98.

„ Paractius mutabilis, idem. Ibid., p. 240, pi. x, f. 103 — 112.

1890. „ „ Malacuiin. Rev. Biol, du Nord. (Annel. Boulon), T. hi, p. 381.

„ Ophryotrocha puerilis, Griard. Bullet. Sc. Fr. et Belg., T. xxii, p. 77.

„ „ „ idem. Ibid., T. xxii, p. 77.

1892. „ „ Monticelli. Monit. Zool. Italiano, III, p. 250 (fide aut.).

1893. „ „ Bonnier. Bullet. Sc. Fr. et Belg., T. xxv, p. 198, pis. 1—4.
}} „ „ idem. Oompt. Rend. Acad. Sc, T. cxvi, p. 524.

1895. „ „ (development), Korschelt. Zeitschr. f. wiss. Zool., Bd. lx, p. 543, pis.

27—34.
idem. Ibid., Bd. lx, pp. 543, 689.
1902. „ „ Mcintosh. Ann. Nat. Hist., ser. 7, vol. x, p. 257.

1904. „ „ Allen. Journ. M. B. A., n.s., vol. vii, p. 226.

1905. „ „ Graeffe. Arbeit. Zool. Stat. Triest., xv, p. 322.

Habitat.— When at Whitstable in June, 1884, a small Ennicid was noticed in an
aquarium belonging to Mr. Sibert Saunders, who kindly forwarded examples to the St.
Andrews Marine Laboratory. The specimens had been procured from the oyster-
beds at Whitstable, so that in all probability the species haunts the Algaa and other
growths attached to the shells of the oysters. Plymouth (Grarstang and Allen).

Shores of France (Griard and Bonnier) amongst Bryozoa and Ascidians, as well as on
the tubes of Sabellaria. Body-cavity of Oucuma/rla plauci at Naples (Monticelli).

Studer 1 found a very similar species in Kerguelen.

Head (Plate LXI, fig. 10, and Plate LV, fig. 9) horse-shoe shaped, with a series of
rather long palpocils on the whole of the anterior edge. These organs have broad bases,
taper to a fine point, and apparently are of great tactile sensibility. Their motions on the
edge of the snout are so lively that it seems to be ciliated. The number of eyes is variable,
two being situated externally in a dimple behind the dorsal tentacle. They are simple
pigment-spots, blackish by reflected and pale brownish by transmitted light. The snout
often shows a slight furrow nearly opposite each eye, apparently the second ciliated crown
of Bonnier. On the summit of the head just in front of the nuchal fold is a small pair
of eyes placed near each other, and occasionally a pigment-spot or two in front of these.

1 ' Archiv f. Naturges.,' 1878, p. 119, Taf. v, fig. 11.

366 OPHRYOTROCHA PUERILIS.

In front of each anterior eye is a small tentacle of two segments, the basal shorter than
the distal, which is bluntly rounded and furnished with palpocils similar to those on the
anterior arch. A pair of tentacles having the same structure occupy a corresponding
position on the ventral surface of the snout, but they are more external in position so that
they project distinctly on each side. Claparede and Mecznikow describe a ciliated groove
between the head and the following segment.

Body from 8 or 9 mm. to nearly half an inch in length, with more than thirty bristled
segments. It is little tapered anteriorly, and only slightly so posteriorly. It is pale and
semi-translucent, the internal organs, viz., the blackish dental apparatus and the straw-
coloured or greenish alimentary canal, being visible from the exterior. Two segments
behind the head are devoid of feet. The feet project outward somewhat stiffly.
Posteriorly are two clavate cirri provided with palpocils which proceed from a caudal
segment devoid of feet.

The armature of the proboscis (Plate LXI, fig. 10 a) consists of dark-brown or blackish,
strongly curved, and sharp-pointed maxillae, behind which an acute posterior process
projects. Six denticles, which probably represent the dental plates of allied forms, occur
beneath and in front of these, each being shaped somewhat like a hoof, the free edge
anteriorly being finely denticulated. They diminish in size from before backward, the
last having a long slender process which reaches the posterior border of the maxillas.
Young examples have a smaller number of denticles. The mandibles are elongated and
curved like the letter f, a wing-like border occurring close behind the anterior region.
The cutting edge is bifid, and both cusps are minutely crenate.

"When fully formed the foot (Plate LXXIII, fig. 7), like that of Staurocephalus Kefer-
steini, has no dorsal cirrus, but projects somewhat stiffly outward with four conical
terminal processes. At the upper end is the conical dorsal lobe, which rises from the
dorsal margin of the foot by a long stiff slope, whilst the distal border is shorter and
more nearly vertical. Beneath is the blunt or expanded conical process for the long
translucent spine, one margin of the foot then trending obliquely outwards into the
elongated process for the compound bristles of the region, whilst the other leads to the
ventral lobe, which is less prominent than the other processes, that is, is nearer the body,
as usual in the group, and has a long ventral and a shorter distal slope. The bristles
also differ from those of any other species observed. Dorsally above the spine are about
three elongated translucent bristles (Plate LXXXII, fig. 1), with slightly flattened tips
(in one short) which end in a short point slightly hooked. No serrations are visible on
the edge in the preparations. Below the spine and little separated from the upper series
are about six compound bristles equally translucent. The shafts are curved toward the
end, which is also dilated and bevelled for the terminal piece. The two upper bristles
are as long as the dorsal, but the rest diminish in length interiorly. All, however, are
proportionally long. The terminal pieces appear to be simple, but it is sometimes very
difficult to observe a minute secondary process if close to the point (Plate LXXXII, fig. 1 a).

Reproduction. — All the examples observed, and they were under observation for several
months, had large ova internally, their longer diameter being nearly equal to the vertical
diameter of the foot. Mr. Garstang thought the breeding season of the species was in
August, but Dr. Allen has since found it spawning in the tanks in May.

OPHRYOTROCHA PUERILIS. 367

At Naples Lo Bianco 1 states the ova are deposited from January to May, so that in
our country it is somewhat later, but it evidently extends over several months.

Habits. — It is very hardy in confinement and breeds freely. It crawls actively on
the seaweeds and zoophytes or along the glass.

The description and figures of Claparede and Mecznikow had been overlooked when
the first account of the British examples w r as drawn up, but comparatively little alteration
is necessary in regard to either. The figure of the maxillse and mandibles given by these
authors, indeed, is not characteristic of our forms, and probably considerable variation
exists, though the amalgamation of the anterior denticles is different from anything seen
in the British forms, which have a row of six separate processes. The lateral view of the
foot also (fig. 2f) has been from an injured or imperfectly preserved example. The older
authors' views of the body and head, however, are good, and their account and figures of
the young are also accurate. They describe and figure an eyeless larva 0*3 mm. long
with five ciliated rings, tactile cilia on the prostomium, and a median caudal cirrus. The
dental apparatus shows the mandibles and rudimentary maxillae (the joint authors' labrum).
Then two lateral caudal cirri are developed by-and-by, and finally feet. When 2 # 5 mm.
long two black eyes appear and the ova are ripe. The authors do not allude to the male.

De St. Joseph (1888) described and figured this species under the name of Paractius
•mutabilis, and had found it in considerable numbers at Dinard. He, however, pointed
out that it probably fell under the same genus as Staurocephalus minimus, Lang., 8. Siberti
Mcintosh, and Ojpliryotrocha puerilis. In the same contribution he figured a young example
as Ophryotrocha puerilis, the condition of the dental apparatus apparently having misled
him.

A careful account of this species was given in 1893 by Bonnier from specimens
preserved at Wimereux, where it had formerly been found by the distinguished founder
of the Laboratory, Prof. Griarcl. Bonnier rightly gives a wide margin for variation in this
almost cosmopolitan species, and the dental apparatus especially shows divergences. Thus
in the British examples the lateral denticles are much less prominently denticulated than
in those from Wimereux. Moreover the terminal lobes of the feet differ slightly. Bonnier
shows three or four of the posterior denticles as boldly denticulated, whereas in ours only
one (the most posterior) presented this structure. Bonnier criticises the groups of Ehlers,
viz., the Labidognatha and Prionognatha, averring that in its young condition (four to
fifteen segments) Ophryotrocha is like one of the Prionognatha, whereas in its adult state
it resembles a Labidognath. The classification therefore holds only in the adult condition.
This author also considers Paractius, Levinsen, from Greenland, as synonymous. It
presents the larval character of a band of cilia around each segment.

Hacker (1896) mentions a larval form with " intertrochalen " ciliated rings.

Czwiklitzer 2 lately removed the head-segment in one, with the result that degenerative
processes followed for six weeks, viz., loss of cirri, bristles, and most of the feet, but a week
later there was complete regeneration of the lost parts. If the first segment is removed,
the second may take on the character of the head. When the first and second segments

1 ( Mitth. Zool. Stat. Neap./ xiii, p. 489, and ibid., xix, p. 583.

3 ' Arch. Entwickelnng./ xix, p. 140, and figs., 1905 ; and < Journ. R. Micr. Soc./ Feb., 1906, p. 28.

368

EUNICEA LABIDOGNATHA NUDA.

Minor injuries to the first segment are rapidly

are removed no regeneration ensues,
repaired.

Another experiment was made by Braem, 1 who halved a female example with ripe
eggs. The anterior region with the head and thirteen segments regenerated seven
segments in three weeks. Moreover, the female elements had disappeared, and were
replaced by male. The author thought that in consequence of the injury the indifferent
germ-cells had developed into male cells.

EUNICEA LABIDOGNATHA NUDA.

Both halves of the jaws have an equal number of pieces. No true cephalic tentacles.
Some with and some without branchiae.

The body-wall in the Lumbrinereidse (Fig. 75) is characterized by the toughness and
iridescence of the cuticle, the comparative thinness of the hypoclerm and the circular layer,

Fig. 77. — Transverse section through the body-wall of Lumbriconereis fragilis. The massive condition of the
dorsal and ventral longitudinal muscles is apparent. The oblique muscles are attached above the
nerve-cords.

and the massiveness of the dorsal and ventral longitudinal muscles, which occupy the
greater part of the area in transverse section. Moreover, the marked dissimilarity which
usually distinguishes the dorsal from the ventral longitudinal muscles is in this group not
apparent. Both form compact masses of fibres, slightly diminished towards the middle line
clorsally and ventrally. The ventral muscles form the thickest and shortest masses,
though, as mentioned, the usual disproportion is not pronounced. In consequence of the
great development of the ventral muscles the nerve-cords are pushed inward, their con-
nection with the superficial parts being by a pedicle. As a rule a conspicuous neural
canal occurs superiorly, and it is often as large as the great ventral vessel above it. To
the dorsal edge of the investment above the neural canal the oblique muscles are attached.
Fibres from the circular muscular layer seem to pass across the gap at the pedicle. In
certain forms, as in Lumbriconereis impatiens, a smaller neural canal exists in each trunk
below the larger, and indications of a second (branch) occasionally appears. In some the

1 ' Anat, Anzeiger/ xxxiii, pp. 19—27, 2 figs., 1908.

EUNICEA LABIDOGNATHA NUDA. 369

deeply stained nerve-cells not only form an external investment, but go downward
almost to the outer end of the pedicle.

In the mid-dorsal line a mobile band passes from the circular layer to the muscular
walls of the dorsal blood-vessel, and thereafter is continued to the arch of the alimentary
canal, the nuclei of the cells of which are mostly external.

The form mentioned by Donati 1 as living in a cavity or tunnel in a sponge is probably
one of this family (Lumbriconereidae), from its general structure and iridescence. The
figures are, however, too indefinite for accurate identification.

Pallas described an example under the title Nereis ebranchiata, and De Blainville
placed the Lumbricus fragilis of 0. F. Miiller under the genus Scoletoma.

Grube, 2 after Savigny, made Lumbriconereis one of "the genera of the Euniceae, the
others being Diopatra, Onuphis, Eunice, Lysidice, Aglaura, and (Enone.

The Lumbriconereicla constitute the fifth family of Kinberg's 3 Eunicea, characterized
by eight maxilla), part Ii edentate, tentacles tubercle-like or none, branchiae absent. He
gives only two families, viz., Eranno, in which the tentacles are two and tuberculiform,
and Lumbriconereis, in which they are either obsolete or absent.

Claparede and Mecznikow (1868) 4 give an account of the development of what they
considered a Lumbriconereis or an allied form. The youngest stage is spherical, with a
long tuft of cilia in front, two reddish eyes with lenses, a general covering of cilia, with
a gap posteriorly, and then a broad but short caudal tuft. The alimentary canal and its
glandular coat are differentiated. The following day the outline is pear-shaped, and the
posterior ring of cilia is better developed. On the fourth day greater elongation ensues,
and the proboscis is apparent, whilst the broad anterior coating of cilia and the caudal
ring are more conspicuous. Then a few larval bristles appear on each side posteriorly,
and this region soon shows two distinct tufts of bristles elevated on rudimentary feet, and
a pair of short caudal cirri on cirrophores, whilst the proboscis is armed with what appear
to be maxillae, great dental plates, and mandibles in an early condition.

Cams (1885) makes the Lumbriconereidae a sub-family of the Euniciclae.

Hacker (1896) quotes Fewkes as stating that Lumbriconereis lays eggs in galleries,
and has non-pelagic young. Fewkes says that from each pigment-spot, cephalic or
caudal, a number of delicate threads pass toward the centre of the larva, and that they
are ultimately lost in the larval body.

In his 'Annelids of the Shores of Dinard,' De St. Joseph, 5 while criticising Grube's
classification, which he considers to be a combination of the systems of Kinberg and
Ehlers, follows a similar arrangement.

The veteran student of the Annelids, Prof. Grube, G published, the last year of his life
(1879), a careful general account of the Lumbriconereiclea, primarily subdividing them
according to the absence or presence of dorsal cirri, the bristles and teeth being of

1 ' Delia Storia Naturale Marina dellAdriatico/ Venezia, 1750, p. 59, Tav. viii, figs. G, h, j, k.

2 f Fam. der Annel./ p. 293, 1851.

3 < Ofvers. af-K. Vet.-Aead./ 1864, No. 10, p. 567.

4 'Zeitschr. f. wiss. Zool./ xix, p. 182, Taf. xv, figs. 1 — In.

5 < Ann. Sc. nat./ 7 e ser., t. v, p. 192, 1888.

e < Jahresber. Scliles. Gesellsch./ 1879, p. 78.

10-1

370

EUNICBA LABIDOGNATHA NUDA.

service in the subdivisions. In arranging the representatives of the genus Lumbrico-
nereis he made use of the structure of the jaw-plates and the shape of the head, or in
sections b and o the condition of the bristles. He gives Lumbrlco nereis four jaw-plates,
the next genus Larymna having five plates. Under the section b, just mentioned, there
are large winged bristles, and the maxillae -have elongated, slender posterior appendages.
The mandibles (lamina ventral is) are horny, black, bilobed in front, the short limbs
united or split. Under this head come Drilonereis and Arabella with four pairs of jaws,
and Avacoda and Maclovia with five pairs of jaws. Unlike Kinberg, he regarded the
biannulate buccal region as a single segment.

The Lumbrinereidae formed one of the families of Levinsen's Euniciformia vera
(1883) following the Onuphidae, a position in keeping with their structure. Benham
includes the group under his Eunicidae, a family of the Nereidiformia.

One of the most recent classifications of the group is that of Gravier, who has done
excellent work, especially amongst the foreign Annelids :

Capillary bristles [Digital branchiae fixed to setigerous lobe

[Four pairs of jaws .
No branchiae - Five pairs of jaws .

i

f [One pair of jaws (hooked)

CD

^

accompanied by
hooks or com-
pound bristles

Only capillary
winged bristles

Four pairs of jaws

Four or five pairs
of jaws ; two
first
achetous

segments

One pair of jaws not
hooked, two filiform sup-
ports

Jaws of the third and
fourth pairs reduced to
hooks

Ninoe, Kinberg.
Lumbvie oner els.
La rymna (including
Zygolobus, Grube) .
La i rt i id a, Kinberg.
Noto cirrus,

Schmarda.

o

CD
O

.3

Three antennae in-
serted on pos-
terior part of
prostomium

Jaws of the third and
fourth pairs denticulated m
plates ; jaws of second
pair unequal
Five pairs of jaws First segment achetous .
Supports of upper
jaw long and fili-

foi

mi

Four eyes, six pairs of jaws
Two eyes, five pairs of jaws

Drilonereis, Clapa-
recle (De St.
Joseph, char,
emend) .

Arabella, Grube
(De St. Joseph,
char. emend.,
Maclovia).

A racoda, Schmarda.

Danymene, Kin-
berg.

Halla, A. Costa.

Support of upper jaw short — in form of a plate Lysaretea, Kinberg.

Three antennae covered by the prostomium ; nucal organ forming Aglaur ides, &&vigny

S-i

m

o

^

w.

en

CD

Pi

-4^

CD

• r— 1

m

s<

o3

s

c3

o

P-i

o

two voluminous organs behind antennae

iNo antennae, no nuchal organs

f Lower jaw much developed, parasitic or

commensal on Syllideans
Lower jaw much reduced, a parasite or com-
- mensal of Bonellia
Parasitic or commensal in Terebellides Stroernii Edema tody pies,

Wiren

Four eyes

No eyes

and Ehlers.
CEnone, Savigny.
Labrorostratus, De

St. Joseph.
Oligoglia fit us, Spen-
gel.

LUMBRICONEKEIS. 371

In this group are instances of the somewhat rare endo -parasitism of Polychaeta,
indeed, the majority of the examples appear to pertain to it, and, further, they are
parasitic on Vermidia of one kind or other, chiefly Polychaeta. Thus the Amphinomids
Hipponoe Gaudichandii and H. Mulleri inhabit Lejpas anatifera; and the Ctenophore
Gydippe densa, Forsk., is the home of Alciopina parasitica ; whilst Ophvwricola cynips is
found in a cleep-sea Ophiuroid (Ophioglypha tumulosa). The Lumbriconereid forms,
again, are Oligognathus Bonellix, which haunts the Gephyrean Bonellia ; Hxmatocleptes
terrebellidis occurs in a blood-sinus of the gut of Terebellides stroemii; Labrorostratus
parasiticus is found in the body-cavity of various Syllicls, viz. : — Odoiitosyllis ctenostoma,
Eusyllis moiiilicornis, Syllis prolifera, Pionosyllis lamelligera, and Grubea clavata; and
another Lumbriconereid in the same cavity in Marphysa sanguinca. Lastly, Monticelli
met with Ophryotrocha puerilis in the body-cavity of Oucumaria Planci at Naples.

The Lumbrinereids are common in mud, indeed some, such as the red form mentioned
by Mr. Crossland, are the only inhabitants of the mud which forms the bottom between
the coral reefs in Red Sea harbours.

Genus LXIII. — Lumbbiconebeis, De Blainville, 1828 (char, emend.).

Head more or less conical, without tentacles or palpi; a neck-fillet with retractile
papillas. Mouth with two lips. Body elongated, distinctly segmented, little tapered
anteriorly, but more distinctly so posteriorly, where it terminates in four anal cirri.
Two smooth segments behind the head. Maxillge smooth with basal processes; maxillary
plates symmetrical, toothed, and with a broad horny ledge. Mandibles connate in the
broad middle region anteriorly, and diminishing posteriorly. Foot with a long terminal
flap, simple and compound bristles, in the posterior segments (and in the young in all)
simple hooks. Body-wall as in the type (see p. 368). The circulation is highly deve-
loped, and the vascularity of the feet is characteristic. The segmental organs, repro-
duction, and development are all in need of further investigation.

Kinberg 1 made two divisions of the genus Lumbrico nereis, viz., (1) those having
maxilla Illi unidentate, and (2) those with the same part bidentate.

Ehlers (1868) points out the substitution by Grube of the term Lumbriconereis for
Lumbrinereis, Blainville; and amalgamates Scoletoma, Blainville, Zygolobus, Grube, and
Eranno, Kinberg, with the genus, whilst Araeoda, Schmarcla, and JSfotocirrus, Schmarda,
are also partially connected with it.

The species range from tide-marks to very great depths, e.g., in the ' Challenger ' to
2225 fathoms.

A form {Liimbricoiiereis) occurs in the fresh waters of Trinite 2 (Trinidad?).

Gravier 3 mentions Miss F. Buchanan's observations on the regeneration of the
anterior region in Lumbriconereis impatiens and in Holla. Regeneration of the tail is
more frequent.

1 ' Ofversigt af-K. vet.-Akad. Forliancll./ 1864, pp. 568 and 569.

3 Benham, ' Camb. Nat. Hist./ ii, 1897.

3 < Ann. Sc, nat,, Zool./ 9 C ser., t, ix, p. 136, 1909.

372 LUMBRICONEREIS FRAGILIS.

Claparede and Mecznikow (1863) gave the early stages of the development of a species
of Lumbriconereis which they found at Naples in November and December, and though
the precise relations of the form are unknown, it may be mentioned that the youngest
stage is a rounded trochophore having two reddish eyes with lenses, a central digestive
region, a long anterior whip of cilia, and a broad ciliated ring behind the eyes, as Avell as
a posterior tuft. The following day the shape is more elongate (pear-shaped). On the
fourth day the long anterior whip has disappeared, but the broad zone behind is in full
development, as also is the posterior cincture. The digestive tract shows an anterior
(pharyngeal) and a posterior region, whilst a pair of bristle-bundles appears in front of the
anal ring. Two bristle-tufts occur on the eleventh day, and the anal ring of cilia has
disappeared. The prostomial region is larger. At a later date the young annelid shows
two well-marked setigerous segments, a conical prostomium with a lateral eye-speck on each
side, the pharynx is sunk in the digestive tract, and two short anal cirri occur posteriorly.
The outline is elongate ovoid. The authors were uncertain whether the form belonged to
Eunice, Lumbriconereis, or Notoeirrm. The figures of the bristles are so small that little
help is obtained from them.

1. LuMBiucoNEREis EiiAGiLis, 0. F. Milller, 1788. Plate LXII, figs. 1 and la — teeth;
Plate LXXIII, figs. 8-8 c— feet ; Plate LXXXII, figs. 2-2 b— bristles.

Specific Characters. — Head conical and smooth, with a dimple posteriorly leading
into two nuchal pits with papillae. Body 5 — 10 or more inches in length, ending in four
caudal cirri. First segment longer than the second. Colour iridescent brownish.
Mouth opens behind two massive lips separated by a median fissure, with a curved and
crenate hind lip. Maxillae boldly curved, continued posteriorly into two spear-shaped
processes. Great dental plates with four teeth (five to six, Ehlers). Inner lateral plate
has a horny tooth internally, and so with the outer. Horny patch behind these fits against
the tooth, and externally a horny bar passes towards the maxilla. Mandibles anchylosed,
with dense whitish calcareous edges. Body- wall (Fig. 77) massively muscular, the
great thickness of the ventral longitudinal muscles being conspicuous. Foot with simple
winged bristles as far as the twenty-first, when winged hooks appear. Anterior lobe
bluntly rounded, the posterior with a nearly horizontal dorsal edge, a bluntly conical
point, and then, after fusing with the lower edge of the anterior lobe, sloping inward and
backward to the body. The space between the lobes greater in the anterior foot. Feet
increase in length posteriorly, and the hind lobe becomes pointed. Hook with a shaft
like that of the bristles, but after the curve and expansion it contracts to a neck. The
terminal hook has a serrated crown and is guarded by a wing on each side. Spines four
in number, black. The tapering winged bristles disappear before the sixtieth foot, the
posterior feet having only the winged hooks and two black spines.

LUMBRICONEREIS FRAGILIS.

Synonyms.

1776 Lumbricus fragilis, 0. F. Mailer. Zool. Dan. Prodr., p. 216, No. 2611.
1788. „ „ idem. Zool. Danica, i, p. 22, Tat', xxii, £. 1—3.

1791. „ „ Linnaeus. Syst. Nat. (Gmelin), ed. 13, i, pt. vi, p. 3086.

1825. Nereis frag His, De Blainville. Diet. Sc. Nat., xxxiv, p. 454.

1827. Lumbricus frag His, Bruguiere. Encycl. Meth., Tab. liv, f. 15a.

1828. Scoletoma fragilis, De Blainville. Diet. Sc. Nat., lvii, p. 492.
1833. Lumbriner els fragilis, x\.ud. & Edw. Ann. Sc. nat., t. xxviii, p. 244.
1835. „ „ Sars. Beskriv. og Iagtt., p. 48.

„ ,, „ idem. Nyt Mag. f. Naturvid, vi, p. 209.

1843. ,, „ (Ersted. Annul. Danic. Consp., i, p. 15, f. 1 and 2.

„ „ „ idem. Reg. Marinis, p. 78.

1844-5. „ „ idem. Kroyer's Nat. Ticls. (Dr^bak), p. 408.

1851. Lumbriconereis ,, Grube. Fain. Annel., pp. 45 and 124.
1857. Lumbrinereis ,, Danielssen. Reise, p. 50, and 1858, p. 116.

1864. Lumbriconereis borealis, Kinberg. Annul. Nova, Ofvers. a£-K. vet.-Akad. Fork., No. 10, p. 568.

1865. „ fragilis, De Quatrefages. Annel., ii, p. 365.

1867. ,, borealis, Malmgren. Annul. Polych., p. 63, Taf. xiv, f. 83.

1868. „ fragilis, Ehlers. Borstenw., ii, p. 395.

1869. Lumbrinereis fragilis, Mcintosh. Rep. Brit. Assoc, 1868, p. 337.

1873. „ ,, Sars. Brdrag Christ. Fauna, p. 22.

,, „ „ Verrill. Invert. Yiny. Sound, U.S. Comm. F. and F., p. 594.

1874. Lumbriconereis fragilis, Mcintosh. Ann. Nat. Hist., ser. iv, vol. xiv, p. 199.
,, „ „ Smith and Harger. Trans. Conn. Acad., iii, p. 16.

„ Lumb vineries fragilis, Malm. Goteb. vet. o. vitt. Handl., xiv, p. 85.

1875. Lumbriconereis fragilis, Ehlers. Zeitschr. f. wiss. Zool., xxv, p. 53.

,, „ ,, Mcintosh. Invert, and Fishes St. Andrews, p. 123.

„ ,, „ Mobius. Jahresb. Comm. deutsch. Meere, p. 167.

1878. „ „ Mcintosh. Trans. Linn. Soc., Zool. I, p. 503.

1879. Lumbrinereis fragilis, Tauber. Annul. Danic, p. 101.

„ „ „ Hansen. Nyt Mag. f. Naturvid., xxiv, pp. 5 and 268.

1881. Lumbriconereis fragilis, tlorst. Niederl. Arch. Zool., Snppl. Bd. i, p. 12.

1883. Lumbrinereis frag His, Wiren. Choet. Vega Exped., p. 402.

„ „ ,, Levinsen. Yidensk. Medclel. Foren. Kjobenh., p. 224.

1884. „ „ Webster and Benedict. Rep. U.S. Comm. F. and F., p. 720.
1886. Lumbriconereis fragilis, Harvey Gibson. Proc. Lit. and Phil. Soc. Liverp., vol. xi, p. 154.

,, Lumbrinereis fragilis, Levinsen. Kara.-Hav. Ledorme, p. 8.
1889. „ ,, Marenzeller. Arch. f. Naturges., Iv, p. 129.

1891. „ „ Hornel. Trans. Biol. Soc. Liverp., vol. v, p. 241.

1893. ,, ,, Levinsen. Yidensk. Ud. "Hauchs," p. 331.

1901. „ „ Whiteaves. Geol. Surv. Canada, No. 722, p. 80.

1902. „ ,, Marenzeller. Polych. Grund., p. 15.

1903. Lumbriconereis fragilis, Mcintosh. Ann. Nat. Hist., ser. 7, vol. xi, p. 560.
„ „ „ idem. Ibid., vol. xii, pp. 158 and 164.

1908. „ „ Ehlers. Deutsch. Tiefsee Exped., p. 95.

Habitat. — Found at various stations off the British shores in the ' Porcuoine '
expedition of 1869, at depths ranging from 15 to 1380 fathoms; 30 to 50 fathoms off

374 LUMBPICONEREIS FRAGILIS.

Valentia and in Dingle Bay and other parts on the W. and S.W. coasts of Ireland (J. G.
Jeffreys). Procured also in the 'Knight Errant' at Station 3; Connemara (A. G.
Moore). Most observers have, from the days of 0. F. Muller, found it on muddy
ground.

Nova Zembla (Theel) ; Siberia and Behrings Strait (Wiren) ; Canada (dredged by
Dr. Whiteaves). It seems to be northern, and does not reach the Mediterranean.

Head forms a smooth conical process with a dimple posteriorly, which leads into two
pits (nuchal), into which two small papillae or bosses project from the first segment.

Body elongate, little diminished anteriorly, but more tapered posteriorly, and ending
in four caudal cirri. The first two segments are devoid of feet, and the first is broader
than the second. The mouth opens inferiorly behind two massive tips (homologues of
palpi ? ) separated by a median fissure. The hind lip is curved and crenate.

The dental apparatus (Plate LXII, fig. 1) presents a pair of boldly curved maxillse,
the base being continued posteriorly into a spear-shaped process, and each of the great
dental plates has four teeth. The inner antero-lateral plate has a prominent tooth
internally and so has the outer in most cases. A horny patch occurs behind these and fits
against the teeth, and internally a horny bar passes toward the great dental plate. From
the ventral surface the broad lamina of the great dental plate has a similar pale area in
front, so that the surface is boldly chequered. The mandibles (Plate LXII, fig. 1 a) are
anchylosed, and have dense, whitish calcareous edges. The fine brown lines run from
side to side continuously. The appearances are somewhat the same as in L. Pettigrewi of
the ' Challenger.'

In this form the nerve-cords in section (Fig. 77) often show a symmetrical arrange-
ment of the neuroglia, the median band passing upward, splitting and surrounding the
great dorsal neural canal, which has coagulable contents. The pedicle is shorter than in
some species. Additional smaller neural canals occur below the larger, but whether this
is due to branches or to the methods of preparation is uncertain.

The first foot has a considerable tongue-shaped lobe (Plate LXXIII, fig. 8) projecting
posteriorly, five black spines, and only simple somewhat short winged bristles of the same
type, such as exist on the feet behind it. These bristles form two groups, an upper
series of longer, and a ventral series of shorter forms with slightly broader wings. The
following feet to the twentieth retain this structure except that the posterior lobe
becomes a little longer. The anterior lobe is bluntly rounded in lateral view, whilst the
posterior has a nearly horizontal dorsal border, a bluntly conical point, and then, after
fusing with the inferior edge of the anterior lobe, the outline trends inward and backward
with a long curve to the body. The space between the two lobes is greater in the
anterior than in the posterior feet, thus affording room for the larger setigerous process
anteriorly. Posteriorly, and especially toward the tail, the feet considerably increase in
length, and the posterior lobe becomes pointed.

At the tenth foot two groups occur, a dorsal of strong curved bristles with the wings
on the convex or inferior edge and a comparatively short tapering tip, and a more
numerous lower group with less elongated tips, the edge of the wings often uppermost.
The foot is supported by five spines, pale at the base, black throughout the rest of their
extent. This structure occurs in the succeeding feet as far as the twentieth, the only

LTJMBBICONEREIS FRAGILIS. 375

change being the somewhat different arrangement of the bristles, which often form
groups of two and three.

The twenty-first foot usually shows the first long winged hook. This has a shaft
similar to that of the bristles, but after the bend and expansion, it narrows to a neck,
with a terminal hook having a serrated crown. A narrow wing on each side guards the
hook and stretches from the dilatation to the tip. This hook appears below the spines,
which are four in number. The spines at the thirtieth foot are three in number, but the
structure of the foot is little altered except by the presence of the long winged hooks.

At the fiftieth foot or thereabouts (Plate LXXIII, fig. 8 b), only a single bristle is
present, or occasionally two of the tapering winged type (Plate LXXXII, fig. 2) remain,
whilst the strong winged hooks are prominent. These (Plate LXXXII, fig. 2 a) have a
considerably stouter shaft than the bristle of the same foot, and the shaft is curved distally,
the winged region being comparatively short and obliquely striated with fine lines. A
constriction forming a neck occurs below the terminal hook which has a crown of minute
serrations, directed obliquely forward to the hook. The developing hook has a long
filiform cap or process distally (Plate LXXXII, fig. 2 b) — apparently a translucent exten-
sion of the wing. When seen antero-posteriorly, the double wing is observed to curve
inward at the tip on each side and to be attached to the central stem of the hook. Two
black spines occur in these and the succeeding feet.

The tapering winged bristles disappear before the sixtieth foot, so that the posterior
feet only have the long winged hooks with the comparatively short wing and the two
black spines supporting the foot (Plate LXXIII, fig. 8 c).

In looking at the feet from above in spirit-preparations a tendency to a " probe-point "
at the tip of the posterior lobe is apparent, especially anteriorly. A blood-vessel passes
round the posterior lobe of the foot, a little within the margin.

Habits. — An active form. It frequently twists itself into a knot, and occasionally
is preserved in this condition.

In a variety from Grodhavn Harbour, Greenland, dredged by the ' Valorous,' the tips
of the long bristles are remarkably attenuated and have a double curve. Moreover, the
posterior lobe of the foot anteriorly is less attenuate, that is, has a broader cone in lateral
view. By-and-by, as at the fifty-seventh foot, the attenuation of the long bristles becomes
extreme, the tip forming a long, delicate, hair-like process, and the wings are very narrow.
The shafts are brownish. The winged hooks have blackish-brown and very long shafts,
with a short dilated terminal region and a narrow wing. Both lobes of the feet posteriorly
form narrow cones. Whether this is a marked variety remains to be seen. The spines
are similar to those of L. fragilis.

Langerhans l describes L. fragilis from Madeira, but his figures of the hooks give
rise to some doubt as to the species referred to by him.

Two species described in the 'Challenger' volume 2 from the Amierican coast, viz.,
Lumbriconereis punctata and L. Ehlersii var. tenuisetis, approach this form in many
respects.

A form (B) procured in Herm in 1868 is distinguished from the ordinary Lumbrico-

1 'Zeitschr. f. wiss. Zool./ Bd. xxxiii, 1879, p. 297, fig. 30.
3 Pp. 252 and 253, etc.

376 LUMBRICONEKEIS LATREILLII.

nereis fragilis by the much more distinctly segmented body, each segment having a
greater antero- posterior diameter, and the feet are thus wider apart. The contour of
the foot also diverges, yet it is difficult to express the change in words, though the
posterior flap is larger in L. fragilis. In addition to the long winged bristles others are
peculiarly jointed, for instance, from the first in the tenth foot, and the structure of the
posterior hooks also differs, as likewise do the spines which are not black.

A variety of L. fragilis, from the south west of Ireland, off the Blasquet, has in the
tenth foot the middle group of bristles slightly blunted, so as to form a stage in its
approach to the foregoing.

2. Lumbriconekeis Lateeillii, Audouin and Edwards, 1834. Plate LIV, fig. 6; Plate
LXXIII, figs. 9 and 9 a— feet ; Plate LXXIV, fig. 12— head ; Plate LXXXII, figs.
3-3 c — bristles.

Specific Characters. — Head a short blunt cone, with a median band attached to the
succeeding segment in the centre posteriorly, and at each side of the latter a dimple
receiving a process from the first segment. Two papillae in front of the mouth are
transversely elongated. Body similar to that of L. fragilis, with four anal cirri. Maxilla?
strongly curved and rather short, the basal plates trending to a short posterior lozenge.
Great dental plates with four teeth ; each anterolateral with one tooth ; a horny patch
in front, and a horny bar outside the anterior half of the maxilla?. Mandibles have
spatlmlate anterior ends anchylosed in a wide angle, and two short roots. First foot has
three pale spines, a short, bluntly conical posterior flap, winged tapering bristles superiorly
and inferiorly, and among the latter one or more winged jointed hooks, the ends of which
have four or five spines. Typical foot (as far as seventeenth) has a nearly horizontal
edge to the posterior lamella, whilst the ventral edge is long and sinuous. Spines (four)
pale. Tapering winged bristles superiorly ; articulated hooks, with the distal end of the
shaft slightly curved backward and dilated, and with a narrow wing on each side. End
of shaft bevelled to a sharp point, and with a posterior fillet. Terminal piece of consider-
able length, diminishing distally, constricted at the neck of the hook, which is small, with
several sharp spikes above it. One or two winged bristles occur below the spines at the
ventral border of the foot. Behind the foregoing simple winged hooks take the place of
the jointed. The foot also has a more abrupt inferior curve, so that its depth is
increased. Posterior feet have the posterior lobe prominent, and the hooks have the
primary fang large.

Synonyms.

1834. Lumbrinereis Latreillii, Aud. and Edw. Annel., p. 168, pi. iii b, f. 13 — 15.
1840. Lumbriconereis Nardonw, Grube. Actin. Echin. und Wiirmer, p. 79.
1851. „ „ idem. Fam. AnneL, pp. 45 and 124.

1862. „ tingens, Keferstein. Zeitschr. f. wiss. Zool., Bch xii, p. 102, Taf. ix, f. 1—9.

1863. „ Eclwardsii, Claparede. Beobacht., p. 58, Taf. xiv, f. 14—22,

1864. „ ,, Grube. Insel Lussin, p. 79.
„ Zy globus Edwardsii, Claparede. Glannres Zoot., p. 114.

1865. „ „ De Quatrefages. Annel., i, p. 366.

LUMBRICONEREIS LATREILLIL 377

1868. Zygolobus tingens, Ehlers. Borstenw., ii, p. 391, Taf. xvii, f. 11—14.

„ Lumbriconereis Nardonis, idem. Ibid., ii, p. 381, pi. xvi, f. 23—30.

>» » :, Claparede. Annel. Nap., p. 147, pi. ix, f. 3.

1870. Zygolobus Latreillii, Grube. Arch. f. Xaturges., xxxvi, p. 302.

1875. Lumbriconereis Latreillii, Marion and Bobretsky. Ann. Sc. nat., 6 e ser., t. ii, p. 15.

„ (Under L. fragilis) Ehlers. Zeitschr. f. wiss. Zool., xxv, p. 54, Taf. iii, f. 22, 23.

„ Lumbriconereis Latreillii, De St. Joseph. Ann. 8c. nat., 6 e ser., t. ii, p. 15.

18 ?9. „ Nardonis, Grube. Jahresb. Schles. Gesells. (for 1878), p. 90.

„ „ tingens, idem. Ibid., p. 91.

1885. „ Nardonis, Cams. Fauna Medit., p. 215.

1888 - „ tingens, De St. Joseph. Ann. Sc. nat , 7 e ser., t. v, p. 21-2, pi. viii, f. 62— 64.

1890. „ „ Malaquin. Annel. Boulon. in Rev. Biol. Nord. Fr., ii, p. 180.

1896. „ Latreillii, Roule. Camp. < Caudan/ p. 449.

„ ,, Nardonis. Polych. deutsch. Meere, p. 14.

1898. „ Latreillii, De St. Joseph. Ann. Sc. nat., 8 e ser., t. v, p. 276.

1903. „ Nardonis, Mcintosh. Ann. Nat. Hist., ser. 7, vol. xi, p. 561.

1904. „ Latreillii, Allen. Journ. M. B. A., n.s., vol. vii, p. 226.

1905. „ Nardonis, Graeffe. Arbeit. Zool. Stat. Triest, xv, p. 321.

Habitat. — Not uncommon under stones between tide-marks, St. Peter Port, Guernsey,
and in Herm. Deep sea fishing off St. Andrews Bay, and in the stomach of the flounder
(E. M.). Plymouth (Allen). Loch Slyne, Co. Cork, K.I.A. Bxped., 1886.

Naples (Claparede). Kaila Bay, Japan Sea, by Capt. St, John, 1874 (W. C. M.) ; 400
metres off France (Roule). Eighty-one fathoms off Cape Finisterre, ' Porcupine,' 1870.

The head forms a short, blunt cone with a median band in the centre posteriorly
attached to the succeeding somite, and at each side of the latter a dimple into which
a projecting process of the first segment fits. The under surface in the preparations
generally has a median furrow.

Body similar in appearance to that of L. fragilis, being slightly diminished anteriorly,
and tapered posteriorly, ending in four cirri, which appear (in reproduced tails at any rate)
to arise by the bifurcation of two — one at each side of the vent — and there is nothing to
contra-indicate this in the fully developed tail.

The first segment is slightly wider than the second. The two papillae in front of the
mouth are transversely elongated. The general aspect of the segments and feet resembles
that in L. fragilis, the posterior flap of the foot projecting outward, and being slightly
probe-pointed when seen from above.

The dental apparatus l has superiorly a pair of strongly curved and rather short
maxillae, the basal plates of which trend to a short posterior lozenge (Plate LXV, fig. 12). 2
The great dental plates have each four teeth, and the antero-lateral a tooth each. A
horny patch lies to the outer side of the antero-lateral, and a horny bar outside the anterior
half of the maxillae. The mandibles (Plate LXV, fig. 12 a) have spatulate anterior ends,
anchylosed in a wide V, and two short roots.

The pedicle of the nerve-cords is somewhat long.

The first foot has a short, bluntly conical posterior flap and three pale spines. It

1 In a female of moderate size.

3 In the description of Plate LXV, fig. 12, Nardonis has been entered for Latreillii.

105

378 LUMBRICONEKEIS LATREILLII.

bears superiorly a series of winged tapering bristles with moderately elongated tips.
Similar bristles occur below the spines, and amongst them one or more winged,
jointed hooks, the ends of which have four or five spines behind the small hook. The
wing is narrow. The occurrence of these in the first foot, together with the pale spines,
aids in the discrimination of the species.

At the tenth foot (Plate LXXITI, fig. 9) the outline shows a nearly horizontal dorsal
edge to the posterior lamella, whilst the ventral edge is long and sinuous. Above the four
pale spines the larger tapering winged bristles (three or four in number) occur (Plate
LXXXII, fig. 3). Beneath the spines are about four articulated hooks (Plate LXXXII,
fig. 3/;). The distal end of the shaft of these is slightly curved backward (i.e., from the
point of the hook), dilated, and, moreover, has a narrow wing on each side. The end
of the shaft is bevelled to a sharp point and has a posterior fillet, thus keeping to the
plan of structure seen in the SyllidaB, NereidaB, and other forms. To this edge is attached
the terminal piece, which is of considerable length, flattened, and diminished clistally,
then constricted at the base of the neck of the hook. The latter is small, with a crown of
several sharp spikes above it. The wings are somewhat narrow, so that the lines of the
distal region are nearly parallel. Minute striaB slant from above downward and backward
over the wings, and a fillet occurs at the neck of the hook. Besides the articulated hooks
one or two tapering winged bristles appear below the spines at the ventral border.

This form of foot (Plate LXXTII, fig. 9a) with the compound hooks occurs as far as
the seventeenth foot or thereabout, but variations exist. Then simple winged hooks take
the place of the jointed. The foot also has a more abrupt inferior curve, so that greater
depth is given in lateral view.

The thirtieth foot has three or four pale spines, a few rather slender simple tapering
bristles superiorly, and the stout winged hooks. At the thirty-fifth foot only a single
slender bristle is present, and the same at the fortieth, but this type disappears before the
fiftieth foot.

In the posterior feet the hind lobe remains prominent, and the hooks (Plate LXXXII,
fg. 3 c) are distinguished by the large size of the primary fang.

Reproduction, — An example from St. Peter Port, Guernsey, was distended with large
ova in July, and the appearances denoted that the functions of the alimentary canal must
have been considerably interfered with.

Claparede thought that this species was closely related to L. Grubiana of Port-
Vendres, yet the hooks are different.

The Lmnbriconereis oxychs&ta, of Gravier, 1 from Djibouti, on the Keel Sea, appears to
be allied in a very close manner with this common form. That so few representatives of
Southern Europe should be found in the Red Sea is a noteworthy feature.

Mr. Crossland (in lit.) thought that the Lumbriconereis japonica, Marenzeller, of the
' Challenger,' 2 might be this species (L. Johnstoni in that work), to which it is closely
allied ; but the posterior appendages of the maxillary apparatus are shorter in the
example procured by the £ Challenger,' and, as mentioned in the British form, 3 " the

1 < Nouv. Arch. Mus. tVHist, Nat./ 4 e ser., t. ii, p. 275, pi. xiv, figs. 96—98.

2 P. 243, pi. xxxvi, figs. 13 — 15; xviiA, fig. 17 ; and xviiiA, fig. 1.

3 Op. cit., p. 245.

LTJMBBICONEEEIS IMPATIENS. 379

dorsal winged bristles are longer and more attenuate, as are also the jointed hooks, the
crowns of the latter, moreover, being less distinctly serrated. The bristles are more
numerous in the Japanese species, and the posterior lamellae more acute superiorly. The
spines in the foreign forms are black, whereas in the British they are pale." It is possible,
however, that intermediate forms may yet occur which will render it necessary to modify
the views here expressed.

3. Lumbhioonebbis IMPATIENS, Glaparecle, 1868. Plate LXII, fig. 2 — mandibles; Plate
LXV, figs. 8 and 8 a— teeth ; Plate LXXIII, figs. 10-10 6— feet; Plate LXXXII,
figs. 4-4 d — bristles.

Specific Characters. — Head forming a blunt cone, with conspicuous ciliated organs
posteriorly ; body typical, but the bristles are somewhat longer than in L. fragilis. Tail
terminating in four short cirri. MaxillaB strong, with curves which keep to the hori-
zontal. Great dental plates each with four teeth ; right anterior plate has three small
teeth, whereas the corresponding plate on the left has only an unbroken edge. The other
plate has a single tooth. An accessory horny plate and a curved band passing toward
the great dental plate complete the upper system. The conformation ventrally resembles
that of L. fragilis. The mandibles form a long, narrow wedge on each side, and are
fused along one-third of the inner border ; marked by five curved brown lines anteriorly
and by madder-brown longitudinal lines on the reverse side.

First foot is distinguished by the occurrence of long, narrow, un jointed winged
hooks and of black spines. Feet in front of seventh show an irregularly conical posterior
lobe, somewhat like that of L. gracilis, and a narrow anterior lobe. The long winged
hooks occur below the spines, and their flattened terminal pieces have nearly parallel
sides. These hooks become modified in progress backward, so that at the twentieth foot
they are considerably shorter. The dorsal winged tapering bristles form two groups at
the thirtieth foot, and they by-and-by diminish, disappearing in the posterior feet. At
the fiftieth foot the winged hook has a short tip, and the chief fang is larger ; three black

spines.

The upper neural canal is often very large, with the usual coagulable contents, and
a smaller median canal occurs below it, besides a lateral, which, however, varies in
appearance as if from branching (Fig. 78).

Synonyms.

1825. Lumbricus fragilis, Delle Cliiaje. Mem. Anim. s. Vert.,ii, pp. 409 and 428, Tav. xxviii, f. 8 — 20.
1841. Lumbrinereis fragilis, idem. Descriz. e. not., iii, p. 83, Tav. ci, f. 8 — 11 and 14 — 20.
1868. Lumbriconereis impatiens, Claparede. Annel. Nap., p. 145, pi. ix, f. 2.
1868. „ breviceps, Ehlers. Borstenw., ii, p. 388.

1885. „ „ Cams. Fauna Medit., p. 215.

n impatiens, Pruvot. Arch. Zool. exper., 2 e ser., t. iii, p. 279, pi. xi, f. 5, and

pi. xiv, f. 1 — 3.
1893, } v F. Buchanan. Quart. Journ. Micr. Sci., n.s., vol. 34, p. 537.

1898. „ De St. Joseph. Ann. Sc. nat., 8 e ser., t. v, p. 279.

380

LUMBRICONEREIS IMPATIENS

1903. Lumbriconereis assimilis, Mcintosh. Ann. Nat. Hist., ser. 7, vol. xi, p. 561.
,, „ „ (near), idem. Ibid., vol. xii, p. 158.

1904. „ impatient, Allen. Journ. M. B. A., N.S., vol. vii, p. 226.
1906. „ „ De St. Joseph. Ann. Sc. nat., 9 e ser., t. iii, p. 161.

Habitat. — Stomach of flounder, St. Andrews, and from the deeper water of the Bay
(E. M.). Dredged twenty-five miles off North Unst, Shetland, in ninety fathoms, by Dr.
Grwyn Jeffreys in July, 1868 ; Nymph Bank, South-west Ireland, in two and a half
fathoms, Royal Irish Academy's Expedition, 1886. In the centre of tubes of Panthalis
CErstedl, in fifty to seventy fathoms, west of Peel, Isle of Man (Herdman). Plymouth
(Allen).

Shores of France, at Banyuls, in crevices of corallines (Pruvot). Naples and the
Mediterranean.

Fig. 78. — Transverse section through a ripe female of Lumbriconereis impaiiens, the ccelom being occupied by-
masses of ova.

Head forming a blunt cone as usual in the group, two nuchal organs with vibratile
cilia occurring posteriorly.

Body up to 50 cm. (De St. Joseph) and o mm. in diameter, the first segment broader
than the second, and with numerous rings of nearly equal breadth. The feet generally
do not offer diagnostic features in a superficial view, though the bristles are somewhat
longer. Delle Chiaje describes it as reddish-brown with fine iridescence, and very fragile,
while De St. Joseph gives it a pale rose colour. The segments number between three and
four hundred in a large example. Posteriorly it ends in four anal cirri — two dorsal and two
ventral. The great development of the dorsal and ventral longitudinal muscles and the
position of the nerve-cords are well seen in this species (Fig. 78).

The dental apparatus (Plate LXV, figs. 8 and 8 a) presents a pair of strong maxilla?,
the curves of which keep to the horizontal, and end posteriorly in a pointed lozenge-shaped
process. The great dental plates have each four teeth. The adjoining right anterior
plate has three small teeth, whereas the corresponding plate on the left has only an
unbroken edge. The other plate has a single tooth. An accessory horny patch and a
curved band toward the great dental plates complete the upper system. The conformation

LUMBBICONEREIS IMPATIBNS. 381

ventrally resembles that of L. fragills. The mandibles (Plate LXV, fig. 8 a) form a long
narrow wedge on each side, and they are fused along a third of their inner border. A
wedge-shaped gap occurs in the centre anteriorly. They are prettily marked on one side
by five curved lines of brown anteriorly, and by similar brown lines arranged longitudinally
on the reverse side. Considerable variation, however, occurs in the aspect of these organs
— probably in connection with regeneration, as may be observed by comparing the fore-
going figure with another example (Plate LXII, fig. 2).

De St. Joseph found a Monocystis in the intestine of one at Dinard.

The first foot is readily distinguished from allied forms by the occurrence of long
hooks with rather narrow wings, and of black spines. The feet in front of the seventh,
indeed, show an irregularly conical posterior lobe somewhat like that of L. frag His, and a
narrow anterior lobe. There are three black spines. Dorsally are two groups of winged
bristles, which have moderately tapered tips. Below the spines is a series of long winged
unjointed hooks, the flattened tip of which has parallel sides (Plate LXXXII, fig. 4 a), the
drawing in this case being from the first foot and showing a differentiation at the end of
the shaft, indicating the morphology of the tip, viz., as a modification of a jointed bristle.
The distal end has a series (four) of small hooks, the first and last of which are somewhat
larger, but not much. On the front (the convex border) of the distal end of the shaft are
indications of minute serrations.

The winged hooks (Plate LXXXII, fig. 4) with the elongated tips continue to the tenth
foot, though even in this there are signs of modification (Plate LXXIII, fig. 10). 1 The
dorsal bristles are split into two fascicles and the tips are similar to those in front. More-
over, the wings are traceable almost to the tip. The upper group has longer, the under
shorter tips. The ventral series of winged bristles (below the hooks) resemble those
with shorter tips in the dorsal. In lateral view the posterior lobe of the foot is perhaps
more massive than in L. fragills. Four black spines are present.

At the twentieth foot the structure of the winged bristles is the same, and especially
as regards the continuation of the wings on the filiform tip. The flattened distal region
of the winged hooks is now considerably shorter, though in antero-posterior view they
retain the slight bend seen in front.

The thirtieth foot has the dorsal bristles in two groups, an upper of stouter winged
bristles (Plate LXXXII, fig. 4 b) with a marked curve toward the tip, and beneath these a
larger group of more slender bristles (Plate LXXXII, fig. 4 c), the tips being very attenuate.
The median series of hooks has now much shorter tips, but the wings remain narrow, and
their slightly bevelled condition opposite the point of the hook is still evident. The fang
is comparatively small, and above it is a crown of four or five minute teeth (in lateral
view). Inferiorly is a group of stout tapering winged bristles like the upper series of the
dorsal. The foot has three strong black spines.

At the fifty-seventh foot only a single bristle of each kind remains above the hooks,
which are stouter and now show considerable abbreviation of the tip (Plate LXXXII,
fig. 4cZ). The chief fang is larger. The strongest winged hook is above the three black
spines. The tapering winged bristles generally disappear about the sixtieth foot, though a
single bristle below the two powerf ul dorsal hooks is found occasionally considerably beyond
1 The artist has omitted the winged hooks in the figure.

382 LUMBRICONEREIS IMPATIENS.

this foot, whilst the hooks at the eightieth have a larger first fang, a shorter winged region,
and increased strength, This bristle would appear to belong to the lower dorsal group,
though the tip is now short. From the 70th to the 180th foot (Plate LXXI1I, fig. 10 a) l
only two black spines occur in each foot.

The bristles on the whole approach those of Zygolobus, though differing in arrangement,
and especially anteriorly.

Fage 2 notes that the segmental organ of this form is green, and that the genital
funnel is of large dimensions.

A small form dredged in considerable numbers off S.W. Ireland at log 55 in
twenty-three to thirty-eight fathoms in 1886 shows certain features resembling the fore-
going. At the fifteenth foot some of the winged hooks have very slender tips, the points
(hooks) being very minute, but the spines are pale.

Reproduction. — An example procured in Shetland by Dr. Gwyn Jeffreys in July had
well-developed ova.

This species attains a comparatively large size in the south. Thus De St. Joseph
records one of 34 cm. from St. Vaast (P. Fauvel), and those from Plymouth cannot be
much short of two feet.

In Delle Chiaje's ' Memorie ' (1825) the maxillas are represented with only four teeth,
as observed in the British examples. Two of the figures are omitted in the ' Descrizione.'
He calls the species " Tremolino," and besides the dental apparatus, he describes and
figures the alimentary canal, circulation, and nervous system.

This form differs from Lumbriconereis tingens, Keferstein, in having the long narrow
hooks anteriorly unjointed. It appears to correspond with Lumbriconereis impatiens of
Claparede from the Mediterranean, a form long previously mentioned by Delle Chiaje as
Lumbricus fragilis (non 0. F. Midler), as above noted.

Claparede (1868) cleared up certain ambiguities in Grube's description, especially in
the structure of the hooks anteriorly and posteriorly.

Ehlers (1868) observes that his L. breviceps has bristles similar to those of L.
impatiens.

Cams, again (1885), thought this form synonymous with Grube's Zygolobus lauren-
tianus, which it can scarcely be.

Pruvot (1885) represents two somewhat clavate lobes of the inferior region of the
cerebral mass which pass to the ciliated sacs at the posterior part of the head. He
regards the lateral elevations at the mouth as the homologues of the palpi.

Miss Florence Buchanan 3 (1893) describes spirals of different lengths occurring in
this species from Naples. She likewise mentions that in this annelid and in Halla
regeneration of head and tail occurs.

Lo Bianco 4 found this species mature at Naples from May to September.

1 The artist has omitted the long hooks in this figure.

2 ' Ann. Sc. Nat./ 9 e ser., Ill, p. 380, fig. 36, 1906.
8 ' Quart. Journ. Micr. Sci./ n.s., vol. 34, p. 537.

* f Mitth. Zool. Stat. Neap./ xiii, p. 487, 1899.

LUMBRICONEREIS HIBERNICA. 383

4. Lumbriconereis hibernica, n.s. Plate LXII, figs. 3 and 3 a — lie ad ; Plate LXXIV,
figs. 1 and la — feet; Plate LXXXII, figs, b-b d — bristles.

Specific Characters. — Head forms a somewhat narrow cone with a pointed apex.
Body typical; the bristles, however, approach those of L. impatiens in length. Posterior
lobes of feet not elongated. Maxillae normal; great dental plates with four teeth. Two
lateral plates simple. Mandibles with somewhat elongated anterior plates, the edges of
which are crenate. First foot with three or four pale spines, and two groups of tapering
winged bristles. The tenth foot has four pale spines and a short conical posterior lobe.
Dorsally are winged tapering bristles, whilst below the spines are characteristically
tapered simple hooks — indicating how easily such could merge into a tapering bristle. The
tips of these gradually shorten, so that at the twentieth they are, proportionally, only
half the length. At the thirtieth foot they have short tips and broad wings, and this
condition continues to the sixtieth, in which, as in the thirtieth, the lobes of the feet are
nearly equal.

Synonym.
1903. Lumbriconereis hibemica, Mcintosh. Ann. Nat. Hist., ser. 7, vol. xi, p. 561.

Habitat. — Dredged in ninety fathoms, twenty-five miles west of the Blasquet, S. W\
Ireland, May, 1869 (Dr. Grwyn Jeffreys) ; in the same region (log 55) in twenty-three to
thirty-eight fathoms, P. I. A. Exped., 1886.

Head (Plate LXII, fig. 3) forms a somewhat narrow cone with a pointed apex, and a
slight median groove on the ventral surface.

Body of the normal outline, tapered for a short distance anteriorly, and more
distinctly so posteriorly. The first segment is rather broader than the second, and the
glistening bristles are more conspicuous than usual, in this respect approaching Lumbrico-
nereis impatiens. So far as can be observed in the examples the feet do not present the
elongated posterior lobe so characteristic of several forms, but none are complete
posteriorly. The segments after the twentieth have a tendency to increase in breadth, and
the posterior are about twice the breadth of the anterior. Reproduction of the posterior
end takes place readily, a miniature tail sprouting from the last segment, and showing
the four cirri at the tip.

The great size of the dorsal longitudinal muscles and the arrangement of the oblique
muscles, which pass below the nerve-cord, are noteworthy (Fig. 79).

Proboscis.— In an extruded organ (Plate LXII, fig. 3 a) the maxillae occupy the
anterior and central region of the bilobecl organ, their action being lateral, that is, each
working in and out from its own side. The great dental plates are behind them, their
four prominent teeth projecting on each side of the central space in front and slanting
inward to the posterior part of the plates. The two lateral plates extend outward on
each side, so that an elongated transverse dental series is formed in this position. The
mandibles with their double curves and median fissure project a short distance behind,
the cutting edge being bare.

So far as can be observed the food would be seized by the maxilla? and lacerated by

384

LUMBBICONEREIS HIBERNICA.

the scissor-like action of the great dental plates, aided by the chisel-like effect of the
mandibles, which would not only sever pieces held by the maxillae, but would also press
them against the great dental plates. The muscles attached to all these parts are powerful.
The action of the parts when inverted would not be very different. When the proboscis
is thus thrust out, the head is bent up at an angle, and in semi-extrusion the same
tendency is noticed.

The neural canal is small and indistinct, and situated in an upper median segment of
the ovoid area in transverse section. The area is surrounded by a dense series of nuclei.
The oblique muscles are attached to the upper and outer edges of the area, The
hypoderm is comparatively thick and the pedicle long.

The first foot has a small triangular posterior lobe and two groups of tapering
winged bristles. One in the centre has a somewhat short tip, and thus may indicate the
condition in the following feet, The spines are pale and three or four in number, and

Fig. 79. — Transverse section of the body-wall of Liimbriconereis hibernica.

thus the form is readily discriminated from the allied species L. vm^atiems. The second
foot has five pale spines.

The tenth foot has four pale spines and a short conical posterior lobe, and a shorter
curved anterior lip. Dprsally is a group of tapering winged bristles (Plate LXXXII,
fig. 5), with curved and finely attenuate tips. Below the spines is a group of character-
istically tapered simple hooks (Plate LXXXII, fig. 5 a), which indicate how easily such
forms could merge into a tapering bristle. The terminal hook is minute, with a series of
coronal serrations. When viewed antero-posteriorly, the double frill of the wings is
observed at the tip (Plate LXXXII, fig. 5 /;). Below are one or two tapering winged
bristles.

The tips of the simple winged hooks gradually shorten, so that at the twentieth foot
they are proportionally about half the length. The tapering bristles above and below
them are similar to those in front. There are four pale spines.

At the thirtieth foot the lobes (anterior and posterior) are more nearly equal, and
there are four pale spines. The second or inferior group of dorsal bristles has finely
tapered and very long tips, as in certain allied species, and the hooks (below the spines)
have now short tips with broader wings (Plate LXXXII, fig. 5 c) and long shafts. The

LUMBPICONEREIS GRACILIS. 385

main fang is well marked, and there are three or four spikes above it. A single winged
and finely tapered ventral bristle is usually present.

The condition remains much the same (Plate LXXXII, fig. 5 d) as the foregoing to
the sixtieth foot, beyond which none of the specimens were of service. The differentiation
into an upper stouter and a lower more slender set of dorsal bristles continues as far as
this foot. The lobes of the feet are nearly equal in this region. Only in the anterior
third does the posterior lobe stand backward and outward, and even there the foot has a
slightly truncate appearance.

This form seems to be different from any of the preceding.

5. Lumbrioonereis gracilis, Ehlers, 1868. Plate LXII, figs. 4 and 4 a— head ; Plate

LXXXII, figs. 6 and 6 a — bristles.

Specific Characters. — Head forms a blunt cone. Body small, elongated, and finely
iridescent, tapering posteriorly to a slender tail. The colour is reddish or brownish-
orange (Pruvot and Racovitza). Proboscis with a pair of curved maxillse which articulate
posteriorly with a process having a short anterior and a long tapered posterior region.
Each great dental plate has four teeth. The antero-lateral plates are two in number on
each side, each with a tooth. Mandibles anchylosed in front, the edge presenting a
double crescent, an inner and an outer. The external edge has a pigmented shelf or
process. First foot has a small triangular posterior lobe and about four pale spines and
tapering simple winged bristles, the adjacent end of the shaft likewise having two wings.
The jointed hooks have a short terminal piece with a crown of small hooks and two wings,
and broad wings occur on the distal end of the shaft. This hook continues to the fifteenth
foot and then disappears, its place being taken by simple winged hooks which by-and-by
become stout, the chief fang being strong, whilst above it are three or four erect spines
on the crown. As a rule two pale spines are present in each foot, and the latter is
bi-papillose from the tenth backward.

Synonyms.

1868. Luml>riconereis gracilis, Ehlers. Borstenw., p. 893, Taf. xvii, f. 6 — 10.

1879. ,, „ Langerhans. Zeitsclir. f. wiss. ZooL, xxxhi, p. 298, f. 31.

1885. „ „ Cams. Fauna Medit., p. 215.

1903. „ „ Mcintosh. Ann. Nat, Hist,, ser. 7, vol. xi, p. 561.

Habitat. — First procured on muddy ground mixed with clay (eight to nine fathoms)
in Lochmaddy, August, 1865. Coast of Kerry (A. G. Moore). S.W. Ireland, log 23,
in thirty-five to thirty-seven fathoms, II. I. Acad. Exped., 1885 ; in Bantry Bay, five
and a half fathoms ; and in Loch Slyne, Co. Cork, in 1886. Dredged in the ' Porcupine '
Expedition of 1869 in 370 fathoms off Ireland. Dredged in ninety fathoms twenty-five
miles west of the Blasquet, S.W. Ireland, by Dr. Gwyn Jeffreys, May 25th, 1869, and by
the same naturalist in one hundred fathoms in St. Magnus Bay. In the stomach of the
haddock, St. Andrews (E. M.).

106

386

LUMBRICONEREIS GRACILIS.

Ranges to the Mediterranean, Adriatic (Ehlers), Madeira (Langerhans), and to
Norway.

Dredged in the e Porcupine ' Expedition of 1870 in eighty-one fathoms off Cape
Finisterre.

Head (Plate LXII, fig. 4) forms a blunt cone, sometimes in lateral view almost ovoid,
as Ehlers says. A median dorsal process with backward curve on each side occurs
posteriorly.

Body small, elongated, and beautifully iridescent, tapering posteriorly to a slender
tail, which in the only example of the region was devoid of cirri. Ehlers had also only
incomplete examples. Few of the specimens had the bristles in a satisfactory condition,
probably from friction in the net of fine gauze inside the dredge.

Proboscis. — The dental apparatus (Plate LXII, fig. 4 a) has, superiorly, a pair of
curved maxillae which are articulated posteriorly to a process which has a short anterior
region and a larger tapered posterior part as in L. ting ens. Each of the great dental plates

Fig. 80. — Transverse section of Lumbriconereis gracilis.

has four strong teeth. The anterior or antero-lateral plates are two in number on each
side and appear to have only one tooth in each, but as the posterior on one side has a broad
edge, this, as Ehlers states, may often be double ; indeed, this is well seen in one from
Lochmaddy. As in allied forms their surface is granular.

The mandibles are anchylosed in front, but the edge has a V-shaped fissure, beyond
which on each side the tooth presents a double crescent, viz., an inner and a larger outer.
The external edge has a pigmented shelf or process.

The body- wall is typical in transverse section. The pedicle of the nerve-cords is short
and the neural canal of large size. It is placed superiorly at the ganglia, but passes down-
ward between the cords in the intervals. The ganglia are surrounded by a zone of deeply
stained cells, which often form a mass inferiorly. The oblique muscles are fixed to the
upper and outer borders of the investment of the neural canal.

The first foot has a small triangular posterior lobe, and a series (four) of pale spines.
It bears tapering winged bristles and also jointed hooks, which in this and the second foot
have a short terminal piece with a crown of small hooks and two wings, the adjacent end
of the shaft likewise having two wings. The second foot has five spines.

LUMBRICONEREIS G. 387

The tenth foot (Plate LXXXII, fig. 6) is bi-papillose, a small conical papilla occurring
in front and a larger obliquely conical papilla behind. Between them the fascicle of
bristles projects. The tapering bristles have the usual wings, and the point is moderately
attenuate. The winged hooks are jointed and have comparatively broad wings on the
distal end of the shaft. In some a slight incurvation of the distal wings is evident, as in
the figure. The foot has two or three pale spines.

In a mounted example dredged in Torbay, apparently referable to this species and
forwarded by Major Elwes, the posterior region of the head and every subsequent
segment is, in the transparent preparations, studded with large brownish-red, clearly
defined pigment-cells, so far as can be observed, lodged in the hypoderm. The entire
animal, with the exception of the anterior region of the snout, the segment- junctions, and
the last five segments, thus has the aspect of being studded with eye-spots. Even the
pygidium has a spot on each side at its anterior border.

The jointed hooks continue to the fifteenth foot and then disappear, their place being
taken by simple winged hooks which at the thirtieth foot (Plate LXXXII, fig. 6 a) have a
stout shaft and rather broad wings, whilst the terminal hook in those best developed is
strong, the crown behind it having three or four erect spines in lateral view. Some of
the dorsal bristles (inferior dorsal) are very attenuate towards the tip, as shown in the
figure. Two pale spines occurred in the Norwegian example from which the thirtieth foot
was drawn.

Several fragments, viz., from the deeper water off St. Andrews, from a depth of one
hundred fathoms in St. Magnus Bay, Shetland, and from the western coast of the Isle of
Man, approach this species.

The L. coccinea of Renier, Xardo, and Grube comes near this species.

Pruvot and Racovitza 1 term the maxilla of this and previous communications the
mandible, the great dental plate maxilla 1, and the two plates in front respectively
maxilla 2 and maxilla 3. The mandibles are called "labre." They give a careful
account of Lumbriconereis coccinea, a form which comes very near L. gracilis, and their
remarks on the dental apparatus are specially interesting.

6. Lumbriooneeeis G. Fragment. Plate LXII, fig. 5— head ; Plate LXXIV, figs. 2 and 3
—feet ; Plate LXXXII, figs. 7 and 7 a— bristles.

Specific Characters, — Head and body typical. The armature of the proboscis differs
from that of L. gracilis in the marked shoulder externally on the maxillae, and in the
broader posterior appendages.

At the tenth foot the anterior border (seen in profile) forms a blunt cone, beyond
which the posterior lobe appears as an irregular flap. Bristles translucent simple winged
forms. Hooks, so far as the fragmentary example shows, with curved shafts dilated
distally to the commencement of the wings, then tapered to the neck, the tip having a
short main fang and a series of spikes above it.

Habitat. Procured in Trawl 2, on board the 'Knight Errant' in 516 fathoms,

August 24th, 1882.

Arch. Zool. exper./ 3 e ser., t. iii, pp. 374—384.

i <

388 LUMBRICONEREIS ACUTIFRONS.

Head of the typical conical outline (Plate LXXII, fig. 5) with a slight median
inflection (backward) of the peristomial border posteriorly. The mouth has the concavity
of its posterior lip directed backward, and two rounded papillas lie in the fossa anteriorly.

Body somewhat like that of L. bre wipes, the segments with the exception of the first
two being very narrow and the posterior lobe of the foot small. The specimen is
incomplete and most of the bristles are absent.

Proboscis similarly armed to L. gracilis, the only apparent difference being the more
distinct shoulder at the posterior and outer border of the maxillas and the proportionally
broad and short posterior processes which have a boldly convex, almost semicircular
outer border. The great dental plates have fourth teeth, and in front of the two anterior
plates is a third granular bar on one side. An accessory granular horny belt also lies
to the exterior of the great plates. The mandibles unfortunately were lost.

At the tenth foot (Plate LXXXII, fig. 7) the anterior border forms a blunt cone
beyond which the posterior lobe appears as an irregular flap. The bristles consist only
of one kind, viz., tapering, winged, translucent forms, but it has to be borne in mind that
the specimen had lost most of the bristles and hooks. The spines are pale. In some of
the feet of this form a small papilla appears above the foot, in profile, as in Plate LXXIV,
fig. 2.

The general form of the elongated hooks from the middle of the body is shown in
Plate LXXXII, fig. 7 a.

7. Lumbeiconereis acuhfrons, Mcintosh, 1903. Plate LXXXVII, fig. 7 — head; Plate

LXXXVI, figs. 7-7 b— bristles.

Specific Characters. — Head tapers to a delicate extremity from a base of the normal
breadth, and is rounded. Body little diminished anteriorly, and it remains of uniform
diameter throughout the fragment (f in.). Dental apparatus is translucent madder-
brown ; maxillse somewhat broad posteriorly, with a concave border, but taper in front to
strongly curved and sharp points. Their posterior appendages are constricted behind the
articulation, then expand into somewhat long processes with a straight inner and a convex
outer edge, the whole appendage being thus unusually long. The great dental plates
appear to have six strong recurved teeth, each of which is connected by a canal with a
brown band externally. A single plate with a tooth in front of these. Mandibles trans-
lucent, anchylosed in front, then split into the oblique cutting edge.

Feet project evenly, and from above present a cylindrical setigerous process, behind
which is a short subulate papilla — sometimes probe-pointed — and largest in the anterior
region of the body. Dorsal bristles translucent and brittle, the tips russet-brown,
tapering distally to a fine point, in some considerably prolonged and winged. A few are
more slender, with a hair-like tip. Hooks translucent, with the shaft curved backward
distally and gradually dilating to the short neck. Main fang small, with several small teeth
on the crown above it.

Synonym.

1903. Lumbriconereu- acutifrons, Mcintosh. Ann. Nat. Hist., ser. 7, vol. xii, p. 146, pi. xii, f.

29—32.

LUMBPICONEREIS ACUTIFPONS. 389

This form, dredged in the 'Porcupine' Expedition of 1870, may be briefly
alluded to here, for, in the absence of a note of its locality, it may yet be procured in
the neighbouring waters.

The single example is small and imperfect, but is distinguished by the attenuated
conical snout (Plate LXXXVII, fig. 7), which from a base of the normal breadth tapers
to a delicate extremity. In lateral view it is even thicker at the tip than when seen from
above, for the dorso-ventral flattening is less marked than in ordinary types.

The body is very little diminished anteriorly, and remains of the same diameter as
far as it goes, the total length of the fragment being f in. The dental apparatus is trans-
lucent madder-brown by transmitted light. The maxillae are somewhat broad posteriorly,
with a concave border, but taper in front to strongly curved and sharp points. Their
posterior appendages are constricted after the articulation, then expand into somewhat
long processes, with a straight inner and a convex outer edge, the whole appendage being
thus unusually long. The great dental plates appear to have six strong, recurved teeth,
each of which is connected by a canal with a brown band externally. These correspond
respectively with the central canal of the tooth and the layer of odontoblasts of the dental
matrix of Pruvot and Racovitza, 1 as shown in their account of Lumbriconereis coccinea.
In front the preparation presented only a single plate with a tooth. The translucent
mandibles are anchylosed in front, then split into the oblique cutting edge which is capped
externally with brown. Nearly parallel lines pass from these blades into the shafts, where
they converge. The apparatus, however, had been injured.

In glancing along the lateral region of the body the feet are seen to project evenly, for
the basal part forms, when observed from the dorsum, a cylindrical process, the setigerous
lobe constituting the anterior or median region of the tip, whilst posteriorly is a short
subulate papilla or lobe, sometimes probe-pointed, and which is largest in the anterior feet.
The bristles are translucent and brittle, so that comparatively few remain on the specimen.
The spines, of which four or more occur in the anterior feet, are also pale and translucent,
the tips only being russet-brown. The dorsal bristles (Plate LXXXVI, fig. 7) are slightly
curved at the end of the shaft, the winged tip dilating in the usual manner and then
tapering distally to a fine point, which in some is considerably prolonged. A few bristles,
again, are of a more slender structure, dilating little at the end of the shaft, and termi-
nating in a very attenuate hair-like tip (Plate LXXXVI, fig. 7 a). The hooks likewise are
translucent, and appear to be absent from the anterior feet, though, as mentioned, they
might have been removed. The shaft is curved backward toward the tip, and gradually
dilates to the commencement of the short neck of the hook (Plate LXXXVI, fig. 7 b).
The main fang in this region of the body is small, and the crown above it has several small
teeth.

The species is peculiar, and probably came from a considerable depth.

Delle Chiaje 2 gives figures of the dorsal and the ventral surface of a form with
a similar sharp snout, but neither description nor name has been found, and the arrange-
ment of the segments appears to differ. It is, at any rate, a near approach to the foregoing
species from the ' Porcupine.'

1 ' Arch. Zool. exper./ 3 e ser., t. hi, p. 380, text-fig. 3.
3 'Descriz. Anim. Invert./ Tav. 178, figs. 4 and 5.

390 LUMBRICONEREIS (ZYGOLOBUS) LAURENTIANUS.

Genus LXIV. Zygolobus, Grube, 1863.

Head a blunt (rounded) cone. The anterior margin of the buccal segment bears two
bosses which dorsally overlap the cephalic lobe. Proboscis, body, and bristles as in
LumbriconereiSj with which this probably should be united.

Ehlers insisted that Grube's genus Zygolobus should be included in Lumbriconereis, the
two swellings of the first segment, in his opinion, not sufficing for distinction.

Lumbriconereis (Zygolobus) laueentianus, Grube, or n.s. ? 1863. Plate LXII, figs. 6-6 b —
head and teeth; Plate LXXXII, figs. 8-8 c —bristles.

Specific Characters. — Head a short cone, with two prominent dorsal bosses in the
middle line posteriorly. Body 7 — 9 ins. in length, and fully 3 mm. in breadth, with the
general aspect of a Lumbriconereid, and finely iridescent. Foot with a well-marked conical
setigerous lobe having black spines (four or five in number), strong winged simple bristles
with a marked curve at the distal end of the shaft above the spines, a more slender
group with a different curve distally below the spines, and long winged hooks which
anteriorly (twentieth foot) have a narrow and elongated distal region, but posteriorly this
becomes shorter and more rounded in outline. The distal end has a narrow neck for the
hook, the chief fang of which is of moderate size, and the crown above it has four or five
minute spines.

The Liimbriconereis mucronata of Ehlers, 1 from the mouth of the Congo, has an
elongated prostomium, but this is more lanceolate than acutely tapered, and the hooks of
the two diverge.

Synonyms.

1863. Zygolobus laurentianus } Grube. Arch. f. Naturges., xxix, p. 40, Taf. ix, f. 3 and 3a.
1865. „ „ De Quatrefages. Annel., i, p. 382, and ii, p. 652.

1874. Liimbriconereis laurentiana, Mcintosh. Ann. Nat. Hist., ser. 4, vol. xiv, p. 199.

1875. „ „ idem. Invert, and Fishes St. Andrews, p. 123.
1885. „ breviceps, Cams. Fauna Medit., p. 215.

Habitat. — Procured in the stomach of cod and haddock from St. Andrews Bay
(E. M.).

Prof. Lorenz sent it to Prof. Grube from St. Martino in Lussin Piccolo, in the
Adriatic.

Head (Plate LXII, fig. 6) a short cone, and distinguished by the two prominent
bosses in the middle line posteriorly.

Body probably 6 or 7 ins. or more in length and fully 3 mm. in breadth in the prepara-
tion. The general aspect is that of Liimbriconereis, and it has a finely iridescent cuticle.
So far as the preparation shows, a single segment about twice the breadth of the others
follows the head, but there are indications of two segments, as Grube says, the broader

1 ' Deutsch. Tiefsee Exped./ p. 95, Taf. xii, figs. 9—13.

LUMBRICONEREIS (ZYGOLOBUS) LAURENTIANITS. 391

being posterior. At the anterior border of the peristomial segment are two globular
papillae of considerable size, one on each side of the middle line of the dorsum. Grube
considers them to be tentacular organs.

The proboscis, the teeth of which were not examined by Grube, is armed with a pair
of much curved, dark brown maxillae, the points of which curve dorsally in a more
pronounced manner than usual in the group. Looked at from above there is, therefore,
a ventral curve of the blades and a dorsal curve of the tips. Their bases are articulated
posteriorly to processes which slope to a constriction, then to a fusiform and pointed
region behind (Plate LXII, fig. 6 a). The great dental (or maxillary) plates have a
small dark brown area directed vertically (dorsally) and bearing four teeth on each side.
In lateral view from the outside each of these brown areas is triangular, whereas internally
a rectangular process passes ventrally in the middle line. The anterior plates form a
symmetrical pair on each side, the front plate being the larger, irregularly triangular, and
with a single curved tooth directed dorsally. The second is also somewhat triangular, with
a single tooth directed dorsally. External to the anterior plate is a horny comma-shaped
patch, and on the right a horny bar curves backward from a line passing outward from
the anterior border of the great dental plate and joins the pale edge of the great maxillary
plate. It is incomplete on the left. The mandibles (Plate LXII, fig. 6 b) are pale, some-
what pointed behind, expanded and evenly rounded in front, with the exception of a small
external notch, as viewed from above. On this surface they have a transverse row of black
dots behind the anterior edge, and parallel lines running backward. On the ventral
surface the pale Tellina -shaped anterior area is marked by parallel lines, which follow the
outline of the double area from side to side.

The first foot presents a small setigerous lobe with three black spines, and a group of
simple tapering winged bristles. The posterior lobe is large and ovate or broadly conical,
for it is difficult to say what the original condition was in a specimen so softened.

The type of foot (Plate LXXIV, fig. 3, from the middle of the body) with the black
spines appears to be similar in the segments behind as far as the thirtieth or fortieth. So
far as the preparations go, two groups of bristles appear to be present, e.g., in the tenth
foot, a dorsal group of stronger, and another below the spine of bristles with shorter and
more slender tips and shafts. Three or four black spines occur in the tenth foot. At
the twentieth there are four or five spines, and the groups of bristles are similar. The
dorsal bristles (Plate LXXXII, fig. 8) have a bold curve at the commencement of the
wings, which are finely striated from the edge downward and backward. The group
below the spines has more slender shafts, and the curve of the tip is bow-like (Plate
LXXXII, fig. 8 a). One or two long winged hooks also appear in the twentieth foot
(Plate LXXXII, figs. 8 b and 8 c) below the spines. The tip is dilated beyond the shaft,
and the diameter of the terminal region is nearly uniform with rather narrow wings, the
striae on which, like those of the bristles, are directed from above downward and back-
ward. It becomes narrow at the tip to form the neck of the hook, the main fang of
which is of moderate size, with a crest of four or five minute spines above it.

The type of the narrow winged hook is also seen in Lambriconerels impatiens,
though no nuchal papillae are present.

The anterior end of a small example was procured in the stomach of a haddock in

392 DRILONEREIS.

St. Andrews Bay by E. M. in 1869. In this specimen the two globular papillae are situated
very close together. The condition of the feet is unsatisfactory, but the setigerous
region seemed to be bilobed and rather truncated at the tip. The simple winged bristles
appeared to be the same as in that from the stomach of the cod. The winged hooks had
shorter tips at the twentieth foot, and the first fang was hardly distinguishable in size
from the small hooks above it. No black spines existed in this example, but it was much
smaller than the other. The anterior hooks have a smaller main fang than in the
posterior segments, indeed, anteriorly it is scarcely to be distinguished from those
above it.

Delle Chiaje * gives a figure of a greenish form with a head somewhat like Zygolobus,
but it has black specks at the bases of the feet. Its relationship with this form, however,
is uncertain.

Grube's 2 example was 9 ins. long. He did not examine the proboscis, and his figures
of the bristle and hook are too small for accurate work, and, indeed, he appears to have
seen only the posterior hooks, since the contour of the terminal region is described and
figured as rounded.

Claparede's Lumbriconereis Edwardsii (' Beobach. ii. Anat.,' 1863) approaches this
species very closely, only it has compound bristles, which this has not.

The L. tingens of Keferstein, however, has simple bristles, but they differ from those
of the British species.

From the remarks of Claparede in his ' Glanures,' 8 it would seem that the bristles
and hooks of Z. laurentianns, Grube, are simple, and the dental armature resembles that
of the author's Z. Edwardsii. In the other species, termed by Claparede Z. Grubianus,
which has similar coloration to the former, each segment, however, having transverse
brown stride, the inferior hooks are articulated.

B. EUNICEA PEIONOGNATHA.

B. 1. Eunice a prionognatha monocopa.

Ehlers (1868) thus describes the group : Head naked, two segments without feet.
Foot devoid of dorsal and ventral cirri, with simple bristles, the broad terminal wing of
which is serrate. Upper jaw long and dagger-like. Five pairs of jaw-plates, accessory
jaws, the first pair with hook at tip. Mandible shorter than the foregoing, each half
being wide.

Genus LX.V. — Drilonebeis, Claparede, 1870.

Head and body as in Lumbriconereis. Proboscis with four pairs of symmetrical
teeth: tooth I hook-like; tooth II declining; two maxillas with long, slender, posterior
appendages, great dental plates denticulated ; mandibles wedge-shaped, and of the con-

1 f Descriz./ Tav. 155, fig. 14, 1841.

3 < Arch. f. Naturges./ 1863, p. 40, Taf. iv, fig. 3 and 3a.

3 P. 115, pl.iv, fig. 2.

DBILONEBEIS LONGA. 393

sistence of those of Arabella. Foot with a small dorsal lobe; winged bristles and stout
spine-like bristles, both hooked and compound bristles being absent.

1. Dbiloneb.uis longa, Webster, 1879, var. Elisabeth^. 1 Plate LXII, figs. 7, la and lb —
head and teeth ; Plate LXX1V, fig. 4- foot ; Plate LXXXIII, figs. 1 and 1 a—
bristles.

Specific Characters. — Head a short blunt cone, with four eyes in a transverse line at
its posterior border and a lunulate central mark. Body 4 or more inches long, of the
shape of Lumbrlconereis, and easily recognized by its finer and more persistent iridescence.
Proboscis has powerful black, much-curved maxillae with a broad base, having about
three teeth on the inner edge, articulating posteriorly with the very long black processes.
Great dental plates elongate, black and nearly rhomboidal. Each has from six to eight
teeth along the inner edge. Antero-lateral plates, three, small, each with a single long,
sharp fang, the posterior and larger having in addition a second small tooth at the base
of the larger fang. Mandibles wedge-shaped, horny, dark brown, the anterior edge
rounded and rough. Typical foot has a small dorsal lobe sloping outward and upward, a
short setigerous lobe with four or five ordinary spines and a larger stout spine below the
others, and a broadly lanceolate inferior lobe directed upward. Bristles simple winged
tapering forms with oblique striae on the wings.

Synonyms.

1879. Drilonereis longa, Webster. Trans. Albany Inst., vol. ix, p. 40, pi. vii, f. 84 — 88.

1885. Laranda „ Mcintosh. Annel. ' Challenger/ p. 237, pi. xxx, f. 10; pi. xxxvii, f. 5.

1903. Drilonereis Elisabeths, Mcintosh. Ann. Nat. Hist., ser. 7, vol. xi, p. 562.

Habitat. — First procured at St. Andrews in the stomach of a haddock, in 1869, and
afterwards in the stomach of the cod (E. M.). S.W. Ireland in forty fathoms; log 29,
E. Irish Acad., 1885.

Shores of Virginia and New Jersey (Webster).

Head (Plate LXII, fig. 7), forming a short blunt cone, finely iridescent like the rest
of the animal, and with the four eyes at its posterior border, in a transverse line. These
have been omitted in the figure.

Body 4 or more inches in length, tapering slightly anteriorly and more distinctly so
posteriorly. It is easily recognized amongst the other members of the group by its finer
and more persistent iridescence. The colour on the dorsum consists of four longitudinal

bands.

The first two segments are devoid of feet as in the other members of the group,
and a lunular median mark occurs in the line of the segment-junction between them.

Proboscis. — The dental apparatus (Plate LXII, fig. 7 a) consists of a pair of powerful
black maxillse, massive at the base posteriorly and strongly curved in front. The
broad base formed by these organs differs from that in most of the group, and especially
from the condition in Lumbriconereis and Eunice. The two halves touch in the middle
fine posteriorly and articulate with the V-shaped end of the very long posterior black

1 Named after the lady who found it.

107

394 ARABELLA.

processes. Moreover, the inner edge of the base is furnished with teeth, three of which
are visible in front. The great dental plates are elongate, black, and nearly rhomboidal.
They have about six recurved teeth along the inner edge besides a considerably larger
first fang. The small antero -lateral plates are three in number, each with a long sharp
fang, the largest, in addition, having a small tooth at its base, and thus the arrange-
ment materially differs from that of A. tricolor, which has in each plate five or six
teeth. The mandibles (Plate LXII, fig. 7 b) are dark brown, horny, and wedge-shaped.
The broad anterior edge is rounded and roughened. A slight slope occurs on the inner
edge of each, running to the point of contact of the halves.

The first foot forms a simple conical lobe, without marked indication of the process
for the bristles as in the second and subsequent feet. It bears a group of simple tapering
winged bristles. Very soon a dorsal process or papilla appears, so that at the sixth foot
it forms a prominent structure in certain views (Plate LXXIV, fig. 4). The typical foot
has a similar outline, viz., a small dorsal lobe sloping outward and upward, a short
setigerous lobe projecting straight outward, and a broadly lanceolate inferior lobe directed
upward and backward, its longer outline being inferior. Four or five spines support the
foot anteriorly (tenth) ; five occur at the thirth- seventh, and posteriorly four or five are
present, in addition to a large stout spine, apparently below the others, the tip of which
(enclosed in the tissues) is pointed (Plate LXXXIII, fig. 1) and solid. It is a brittle,
chitinous structure like the teeth and gives way when compressed. The great strength
of this spine in contrast with the others is noteworthy, and apparently its function is
similar to the great hooks of the Eunicidae. The bristles (Plate LXXXIII, fig. 1 a) are
simple — tapered and winged — with moderately long points, and have the usual oblique
striae on the wings. The dorsal are somewhat stronger than the ventral. In glancing
along the feet anteriorly the prominent lobe is seen to be ventro-posterior in position.

Habits. — This species seems to inhabit sand or sandy mud at the bottom, yet the
number of specimens procured in the stomach of the haddock by my mother seems to
indicate that it occasionally leaves its retreats to crawl about on the bottom, or even to
swim in screw- coils over it, though it may also be ejected by storms.

This form seems to be closely allied to the Drilonereis (Laranda) longa of Webster,
from the Virginian coast and New Jersey. Discrepancies may be due to imperfections in
description. The example procured by the s Challenger ' was obtained off the American
coast between Halifax and New York. The British form differs in the larger size of the
dorsal process and in the presence of eyes, but it has been thought unnecessary to
constitute a new species.

Genus LXVI. — Arabella, Grube, 1851.

Conical head devoid of appendages, but with eyes. Body with two footless segments
anteriorly, 1 and finely iridescent. Terminal segment bilobate. Four anal cirri. Proboscis
with two strong maxillae ; a great dental plate and three others on each side, the two
anterior having long, sharp teeth. Mandibles short with wide anterior plates. The

1 Grube terms the buccal segment biannulate, the second produced mferiorly to the mouth.

ARABELLA IRICOLOR. 395

dental apparatus is symmetrical. Foot with a small dorsal lobe sloping outward and
upward, a short setigerous division, and a lanceolate inferior lobe. Besides the ordinary
spines is a specially strong pointed spine of a brittle nature. Bristles simple, winged, and
tapering.

Grube (1879) apparently makes three genera of this, viz., Arabella, Grube ; Aracoda,
Schmarda ; and Maclovia, Grube ; the first being distinguished by the presence of four
pairs of jaws, whilst the others have five, thus forming two distinct groups. Both groups
are stated to have jaw I hook-like, but Arabella has the base serrated, all the jaws
toothed, jaw II asymmetrical (Ungleich), two maxillae, and a rudimentary dorsal cirrus.
On the other hand Grube gives to Aracoda of the second group a single buccal segment,
two maxillae (Kiefertrager), and according to Schmarda jaw II is symmetrical; whilst
Maclovia, Grube, has jaw II asymmetrical, as in Arabella, and three maxillae (Kiefertrager).
Considerable confusion, therefore, was introduced into this group by the distinguished
naturalist, probably from relying on erroneous descriptions.

Ehlers, in his ' Florida Anneliden ' (1887), follows Grube's diagnosis, and places under
Aracoda those in which the bases of the maxillae are dentate.

Arabella iricolor, Montagu, 1804. Plate LIV, fig. 4; Plate LXII, figs. 8-8 c — head and
teeth ; Plate LXX1V, figs. 5-5 d— feet ; Plate LXXXIII, figs. 2 and 2 a— bristles.

Specific Characters. — Head somewhat flattened, bluntly conical, and with four eyes,
one on each side of the middle line, and very slightly in front of the others, which are
external and less conspicuous. Body 8 ins. to a foot long, firm, rounded, often thrown
into a spiral, and of an iridescent pinkish colour with a tinge of reddish from the blood-
vessels. It tapers posteriorly to a tail, which has four short cirri — two somewhat longer
dorsally, and two ventrally. First two segments evenly rounded, but the feet cause a
lateral dimple in the others. First segment narrower than the second dorsally. Proboscis
with maxillae strongly curved and massive at the base, which is articulated with two
tapering processes, the narrow posterior end joined to two long parallel blackish rods,
which gradually diminish posteriorly and end in slight enlargements. The great dental
plates have nine or ten recurved teeth. First antero-lateral plate has five teeth, the two
anterior far surpassing the rest in size. The posterior has the same number of teeth, the
first and third being largest. Mandibles cleaver-shaped, black, with a pale tip and an
oblique cutting edge of considerable length. The posterior process commences after an
abrupt shelf.

The foot has a small dorsal lobe above the setigerous process and a larger lower
lobe. At the tenth foot the row of bristles passes from above downward and forward —
below the upper and in front of the posterior lobe. This continues to the tail, where
the line of the bristles is less oblique and the setigerous lobe more prominent. The
bristles are of two kinds : (1) a dorsal series with a gentle curve and long, finely tapered tips
with narrow wings, and (2) a series with shorter tips which have a more abrupt curve at
the end of the shaft, the free edge of the curve having about five serrations. Anteriorly
the foot has from five to seven spines, and by-ancl-by the dorsal region has a group (four

396 ARABELLA IrllCOLOR.

or five) of small spines which pass to the base of the cirrus. Posteriorly this dorsal group
increases in size.

Synonyms.

1804. Nereis tricolor, Montagu. Trans. Linn. Soc, vii, p. 82.

1807. „ „ Turton. Brit. Fauna, p. 135.

1812. „ „ Pennant ; Brit. Zool., ed. 4, iv, p. 94.

1828. Lumbricus Saint- Hilaire, Delle Chiaje. Memorie, iii, pp. 170, 178, Tav. xlii, f. 4, 11, 16.

1836-7. CEnone metadata, M. Edwards. Regne Anim., pi. xi, f. 4.

1840. Lumbricoiiereis quadristriata, Grube. Actiu. Ecliinod., etc., p. 79.

1841. Lumbrinereis S. -Hilairii, Delle Chiaje. Descriz., iii, p. 83 ; v, p. 97, Tav. 96, f. 4, 11, 16.
1851. ,, (Arabella) quadristriata, Grube. Fam. Annel., pp. 45 and 124.

1861. Aracoda ceendoea, Schmarda. Neue Wirb. Thiere., I ij, p. 115, Tab. xxxii, f. 253.

1864. Lumbricoiiereis quadristriata, Grrube. Insel Lussin, p. 80.

1865. Lumbrinereis maculata, De Quatrefages. Annel., i, p. 365.
„ „ quadristriata, idem. Ibid., i, p. 366.

„ „ gigantea, De Quatrefages. Ibid., i, p. 360.

„ Nereis iricolor, idem. Ibid., p. 383.

,, „ tricolor, Johnston. Cat. Worms Brit. Mus., pp. 142 and 341.

„ „ iricolor, idem. Ibid., p. 280.

1867. „ ,, Parfitt. Trans. Devon. Assoc., iL P- 2*3 (sep. copy).

1868. Arabella quadristriata, Fillers. Borstenw., pp. 399 and 405, Taf. xvii, f. 15 — 24.
„ Lumbricoiiereis (Zygolobus) quadristriata, Claparede. Grlanures, p. 116, pi. iv, f. 5.
„ Notocirrus Hilairii, idem. Annel. Nap., cl, pi. ix, f. 4.

1871. Maclovia gigautea, Grube. Jahresb. Schles. Ges., p. 58.

1872. „ „ idem. Mitth. u. St. Malo, etc., p. 86.

1875. Notocirrus tricolor, Fillers. Zeitschr. f. wiss. ZooL, xxv, p. 56, Taf. iii, f. 33.
„ Arabella quadristriata, De St. Joseph. Ann. Sc. nat., 6 e ser., t. ii, p. 16.

1878. Lumbricoiiereis iricolor, Grube. Jahresb. Schles. Ges., 1878 ; Breslau, 1879, p. 87.

1879. Maclovia gigantea, idem. Ibid., p. 104.

1881. Arabella Hilairii, Langerhans. Canar. Annel. in Nova Acta Leop., xlii, p. 112.
1885. Notocirrus capensis, Mcintosh. Annel. ' Challenger/ p. 236, pi. xxxvii, f. 3 and 4; pi. xviiiA,

f. 15.
,, „ Hilairii, Carus. Fauna Medit., p. 216.

1888. Maclovia gigantea, De St. Joseph. Ann. Sc. nat., 7 e ser., t. v, p. 30, pi. ix, f. 92 — 95.
1898. „ ,, idem. Ibid., 8 e ser., t. v, p. 282.

1900. „ iricolor, Willey. Journ. M. B. A., n.s., vol. vi, i, p. 98.
1903. Arabella „ Mcintosh. Ann. Nat. Hist., ser. 7, vol. xi, p. 563.

1905. Notocirrus hilarii, Graeffe. Arbeit. Zool. Stat. Triest, xv, p. 322.

1906. Arabella St. Hilarii, -De St. Joseph. Ann. Sc. nat., 9 e ser., t. iii, p. 214, pi. iv, f. 82.
„ Maclovia gigantea, Bolm. Ibid., 9 e ser., t. iii, p. 101.

1909. Aracoda iricolor, Banham. Polych. Subantarctic Is., N. Zealand, p. 245.

Habitat. — Abundant under stones between tide-marks at St. Peter Port, Guernsey,
and at Herm, whence it was first sent to me by Sir Ray Lankester. It is also common on
the southern shores of England. It seems for the most part to frequent the tidal region
both at home and abroad. West Coast of Ireland. Off Galway in fifteen to twenty
fathoms, 'Porcupine,' 1869. Bantry Harbour, H.I.A. Exped., 1885.

Shores of France (De St. Joseph) ; Naples (Delle Chiaje) ; Canaries (Langerhans) ;

ARABELLA TRICOLOR,

397

South Africa ; Japan ; Juan Fernandez ; Straits of Magellan. May yet be found on the
American shores.

Head slightly flattened from above downward, bluntly conical, and of the same
pinkish iridescent hue as the rest of the body. It is distinctly separated from the peri-
stomial segment both dorsally and ventrally. At the posterior border it bears four eyes,
one on each side of the middle line, and very slightly in front of the others, which are
lateral and less conspicuous. The under surface of the snout (Plate LXII, fig. 8) has a
longitudinal furrow, probably in connection with the eversion of the proboscis. The
mouth opens just behind the prostomium. Two well-marked furrows pass backward
from the oral border on each side of the median line, and there is a shorter lateral one.

Body 8 — 10 ins. long, and with 178—200 or more segments, firm, rounded, often
thrown into a spiral. The first two segments are evenly rounded, whereas in the
succeeding the feet cause a dimple at the side of each. The first segment is narrower

nc vm

Fig. 81. — Transverse section of Arabella iricolor, about the anterior third. The nerve-cord has been pushed
inward by the largely developed ventral longitudinal muscles. Letters as before.

than the second dorsally, but broader ventrally, and proportionally deeper anteriorly.
The body is slightly tapered in front, and more distinctly so posteriorly toward the tail,
which has the anus at the tip, a kind of funnel being formed by the four short cirri, the
tips of which are nearly on a level (Plate LXII, fig. 8 d) } The general colour is
iridescent pinkish, with a tinge of reddish here and there from the blood-vessels.

The proboscis is armed with a series of five brownish-black teeth (Plate LXII, figs.
8 6?, 8 b). The maxilla are short, strongly curved, and massive at the base, the inner
edge of which is denticulated (eight and nine points). Behind are two tapering pro-
cesses with an enlargement immediately behind the maxillas, then a contraction, followed
by another enlargement with a spur, the narrow end being continuous with two long
parallel blackish rods, which gradually diminish posteriorly and end in slight enlarge-
ments. The great dental plates have each eight or ten recurved teeth. The antero-
lateral plates are situated on each side of the middle line in front of the maxillas. The
posterior plate has four to five well-marked recurved fangs, the first and the third being
1 Reference to this figure has been omitted in the description of the plate.

398 ARABELLA TRICOLOR.

larger. The plate in front has the same number of teeth, but the anterior far surpasses
the rest in size. The small anterior tooth has a single large curved fang. The mandibles
(Plate LXII, fig. 8 c) are cleaver-shaped, black, with a pale tip which forms an oblique
cutting edge sloping outward from the point of attachment. The body of each process is
long, and the short and narrow posterior process commences after an abrupt shelf.

In transverse section (Fig. 81) the cuticle is dense, and the hypoderm proportion-
ally well developed. The longitudinal muscles are massive, the dorsal exceeding the
ventral in size, only a chink separating them in the mid-dorsal line, and the dorsal vessel
is thus (in contraction) pushed far inward. The oblique muscles are powerful, and are
attached to the upper and outer border of the nerve-area, the vertical bands from the
alimentary canal being inserted at the same points. The nerve-area has a long pedicle
furnished with many blood-vessels, and in contraction is somewhat ovoid, no neural canal
being visible.

The first foot (Plate LXXIV, fig. 5) is of considerable size, and consists of a small
dorsal lobe above the setigerous process, and a large lower lobe (posterior and tongue-
shaped in lateral view) which has a slight slant backward. The row of bristles in this
foot is less oblique than in the subsequent feet, being situated between the dorsal cirrus
or lobe above and the large ventral lobe beneath. At the tenth foot (Plate LXXIV,
fig. 5 a) the bristle-row passes from above downward and forward below the upper lobe,
and in front of the posterior lobe. This arrangement continues throughout the greater
part of the body, only posteriorly at the tail the line of the bristles is somewhat less
oblique, and the spinigerous lobe more prominent. The tenth foot has only bristles of
one kind, viz., tapering winged bristles, and while the dorsal lobe or cirrus is of consider-
able size the large posterior lobe is less than it becomes in the succeeding feet. The
points of many of the developing bristles project from the foot at the base of the
long bristles, which appear to be brittle. The bristles (Plate LXXXIII, figs. 2 and 2 a)
are of two kinds, (1) a dorsal series with a gentle curve and rather long, finely tapered
tips with narrow wings, and (2) a series with shorter tips which have a more abrupt
curve at the end of the shaft, the wings opposite the curve being expanded and provided
with about five serrations.

The foot has from five to seven spines anteriorly, and by-and-by the dorsal region
has a group of small spines which go to the base of the cirrus. In the posterior feet this
dorsal group of spines is considerably larger, though they do not attain the size of the
inferior series. There are four or five in the upper group.

The foot in the middle of the body (Plate LXXIV, fig. 5 b) diverges little from the
tenth, except in the form of the dorsal cirrus, which becomes a mere papilla. The long
posterior lobe curves upward behind the setigerous process, but is not longer than in
front. The hypodermic cells are regularly arranged, and the whole process is highly
vascular.

The posterior feet (Plate LXXIV, fig. 5 c), again, present a dorsal cirrus somewhat
smaller and more tapered than in front, whilst the curved posterior lobe is slightly
shorter than in the middle of the body.

The feet continue prominent, though the body tapers to the tail, two or three imme-
diately in front of the pygidium being minute. Dorsally over the vent are two cirri,

ARABELLA IRICOLOR. 399

thicker than those beneath, and having a distinct cirrophore or enlargement at the base,
from which the cirrus comes off at a slight angle as it slopes outward, the tip in the
preparations being blunt. The ventral cirri are somewhat more slender than the dorsal
and a little shorter. By separating the sides of the vent the dorsal and ventral cirri of
each side are found to be attached to the same basal flap. The caudal segments would
appear to be frequently regenerated.

Reproduction. — A dark-coloured specimen in a firm tube of small stones and shell-
fragments under a stone in a littoral pool at Herm in July was filled with ova, which
escaped apparently by pores (?) near the feet, especially towards the posterior half.

Habits. — It is active and vigorous when stimulated, and is fond of twisting its body
into coils. It tinges spirit of a deep pinkish hue, then of a fine purple. In some instances
the animal is preserved in a firm spiral like a wire-spring.

It is difficult to diagnose the precise form mentioned and figured by Donati l from
an Alcyonium in the Adriatic. It may be either this form or a Lumbriconereis.

This is one of the striking British forms procured on the shores of Devon by the
indefatigable Col. Montagu, who in 1804 described an example as thick as a raven's
quill in diameter and 3 feet long. It is the Nereis iricolor of his MS. volume of drawings
(Plate XXXII, fig. 3, 1808) by his niece, Miss Dorville.

In the illustrated edition of the 'Regne Anim.' (183G — 7) four striao are distinctly shown,
two stronger lateral, and two less marked on each side of the middle line of the dorsum.

Grube (1840) gave this form only two eyes.

There is some doubt as to whether Delle Chiaje's Lumbricus Saint Hilaire (1842) is
this species, for the four longitudinal series of spots, the two conspicuous eyes, and the
figure of the dental apparatus differ, yet it can hardly be anything else.

The Lumbriconereis lonjissima of Grube 2 from Valparaiso is a form very nearly
related to this species, if it is not the same.

The Aracoda coerulea of Schmarda 3 from the coast of Chili, Juan Fernandez, and other
localities, seems to be very closely allied, if not identical, the slight changes in the dentition
and other parts being probably due to variation, and these remarks are rendered more
probable by the perusal of the description and figures of Ehlers. 4

Claparede (1868) associated Delle Chiaje's species with Grube's and subsequent
authors', and thought that the differences in coloration might be surmounted, some being
boldly marked with spots, others, as in those from the Channel Islands, being without
them. The very wide distribution of this form gives a margin for considerable variation.

Ehlers (1868) points out that Grube in his original description mentioned only two
eyes, and omitted the dorsal cirrus. He also corrects the synonymy.

The Notocirrus genicidatus of Claparede (1875), as described by Marion and
Bobretzky, 5 does not seem to differ materially from this species.

1 ' Delia Stoi-ia Nat. Mar./ p. 60, Tav. viii, figs, g— k, 1750.

3 f Annulat. CErsted in Naturhisfc. Foren. Vidensk. Meddel./ 1857, p. 158 (p. 1 sep. copy).

3 < Xeue Wirb. Thiere/ ii, p. 115, 1861.

4 ' Polychajt. Magell. u. Chilen./ 1901, p. 143, Taf. xix, figs. 1—6; and 'Zool. Jahrb./ snppl.
" Fauna Cliilen/' ii, 1901, p. 263.

5 ' Ann. Sc. nat./ 6 e sei\, t. ii, p. 15, pi. i, fig. 2.

400 XOTOCIRRUS SCOTICTTS.

Grube in 1871 instituted tlie genus Maclovia for De Quatref age's Lumbriconereis
gigantea, and in his later publication (1879) adheres to the genus, distinguishing it from
Arabella by the presence in the latter of only four pairs of jaws. In Maclovia, again,
Grube assigned four large basal teeth to the first pair of jaws (Kieferpaares), whereas in
such forms as Verrill's Arabella opalina the four basal teeth of the first pair are small.

The Arabella opalina of Verrill, which Treadwell 1 also found at Porto Rico, seems to
be a closely related form common on the American coast, where formerly Leidy termed
it Lumbriconereis splendid a, 2 Webster and Benedict found it at low water in sand. 3

The note on the synonymy of this species by Dr. Willey, 1900, did good service in
drawing attention to the confusion on the subject, especially in connection with the
supposition that Notocirrus scoticus, McL, was the same species. The two forms are
readily discriminated by the naked eye.

Genus LXVII. — Notocirrus, Schmarda, 1861. Char, emend.

Head rounded and devoid of appendages. Eyes present or absent. Body firm, of
moderate length, deeply segmented so as to be more or less moniliform. Maxillae with
long fangs ; four pairs of jaw- plates, the first with uniformly toothed edge, the second
dissimilar ; very long posterior processes. Mandibles pointed posteriorly. Foot with a
stumpy rounded dorsal and ventral cirrus. Bristles simple, with short tips and broad
wings.

Ehlers 4 describes this genus as having a rounded naked head. Foot with a stumpy
rounded dorsal and ventral cirrus, simple bristles with sparsely toothed wings. Upper
jaw with long fangs, four pairs of jaw-plates, the first with uniformly toothed edge
and without end hook, the second dissimilar.

Claparede (1868) observed that De Quatrefages considered the absence of eyes
diagnostic, yet two species he described were furnished with four. Then he did not
think the presence of a dorsal cirrus of much differential weight in connection with
Lumbriconereis. On the other hand, the cirrus or tubercle sometimes representing it
always had a group of small spines. These are absent in the Lumbriconereids. He
evidently included Arabella as a chief type, and thus his views differ from those in
the present work.

1. Notocirrus scoticus, Mcintosh, 1869. Plate LXII, figs. 9-9 I — body and teeth; Plate
LXXIV, figs. 6 and 6a— feet; Plate LXXXIII, fig. 3— bristles.

Specific Characters.— Bead acutely conical with two distinct eyes at its posterior
border. Body about 3 ins. long, of a general reddish-brown colour, and less iridescent

1 < Journ. Acad. Nat. Sc. Philad./ vol. iii, p. 147.

2 < Eep. U. S. Comm. F. and F./ 1884, p. 721 ; and Webster, < 32nd Ann. Rep. X. York Mas./ p.
116, 1879.

3 < Eep. IT. S. Comm. F. and F./ 1901, p. 199.

4 'Borstenw./ ii, p. 406, 1868.

NOTOCIRRUS SCOTICUS. 401

than Lumbriconereis fragilis. It has a moniliform or deeply segmented appearance, the
enlargements occurring at the feet. It diminishes to a tapering tail posteriorly. The
typical foot has a small dorsal lobe (cirrus) in which is a single vascular loop. Setigerous
region beneath is supported by two strong spines, and the brittle bristles have compara-
tively short, broadly winged tips, boldly serrated at the edges. A small lobe or papilla
also appears ventrally.

This genus was included under the eighth family of Kinberg's Eunicea (Laidea), 1
and was characterized by the presence of an entire cephalic lobe, devoid of eyes.
Branchiee (cirri dorsuales) and ventral cirri mamilliform.

Synonyms,

1869. Notocirrus scoticus, Mcintosh. Trans. Hoy. Soc. Edinb., vol. xxv, p. 417, pi. xvi, f. 17,, a } b.

idem. Rep. Brit. Assoc. (1868), p. 337.
1903. „ „ idem. Ann. Nat. Hist., ser. 7, vol. xi ; p. 564.

Habitat. — Dredged in tenacious grey clay in nine fathoms in Ardmaddy Bay, Loch-

Fig. 82. — Transverse section of Notocirrus scoticus about the anterior third.

maddy, North Uist, in 1865 (W. C. M.), and subsequently in several parts of the Hebridean
seas by that veteran naturalist, the late Dr. Gwyn Jeffreys. Also in 100 fathoms in St.
Magnus Bay (J. G. J.). Irish Sea off Peel, July, 1879 (W. C. M.). It was met with in
the ' Porcupine ' Expedition of 1869 in 80 — 110 fathoms in muddy sand with pebbles.

Head (Plate LXII, fig. 9) acutely conical, with two distinct eyes at its posterior border,
where it is slightly contracted. The eyes are rather close to each other on each side of
the middle line. It is somewhat flattened from above downward in lateral view. Ventrally
a median depression goes forward from the mouth toward the tip of the snout.

Body about 3 ins. in length, of a general reddish or reddish-brown colour, less
iridescent than Lumbriconereis fragilis. It is slender, tapered a little at the anterior end,
which has the segments more compactly arranged and generally thicker, diminishing
posteriorly. The greater part of the body has a moniliform or deeply segmented
appearance, the enlargements occurring at the feet. The cuticle is tough and minutely
punctate, from the presence of the apertures of what appear to be minute tubes, which,
when the cuticle is torn off, project internally as slender structures like hairs.

1 ' Ofvers. Kongl. vet. Akad. Forhancll./ 1864, p. 571.

108

402 NOTOCIRRUS SCOTICUS.

The first two segments are scarcely so broad as the succeeding, and with the following
six or seven are more rounded (as in Lumbriconereis) than those behind, which present a
deeper sulcus at the segment-junction. Behind the thirtieth foot the body becomes
moniliform, the lateral body- wall projecting outward like a cone with the foot at the tip.
The anterior segments, moreover, have longer bristles, and the tips of these are more
finely tapered than in the region behind.

Only one example had a tail, and that had recently been regenerated. No distinct
papillae occurred at the tip, though the minute size of the part and the absence of feet on
the segments showed that it was only in process of growth.

Proboscis. — In Notocirrus, as in Cirrobranchia, Ehlers, a considerable modification of
the dental apparatus has occurred (Plate LXII, fig. 9 a). The maxillae are no longer
prominent and might easily be overlooked. They form small toothed processes in the
usual position, with two to four curved fangs and an enlarged base, the thicker edge of
which touches the enlarged anterior ends of the posterior appendages. Some would
consider them merely continuations of the series of dental plates which lies in front of them,
but in all probabitity the foregoing is the more satisfactory interpretation, unless it is to
be supposed that the maxillas are entirely absent. The posterior appendages have an
enlargement at the articulation, and a spur a short distance downward, and then form
two long tapering processes. In front of the maxilla? are three dental plates. The posterior
is the largest, and has a dark blackish-brown colour, with several large teeth in front,
and an inner long process, which passes backward to the base of the maxillae, bearing
smaller recurved teeth. Externally about the anterior hook of the maxillae it has a well
marked process or shoulder. The plate in front also has a posterior process or shoulder,
and six or seven teeth on its inner edge. The anterior plate is still smaller and shows six
well marked teeth. Each of these plates is supported by a thin plate, which passes
inward and ventrally.

The mandibles (Plate LXII, fig. 9 b) form two dark brown processes with a crenate
anterior edge which is slightly bevelled. Behind is a constriction externally, the outline
curving outward again and then inward to the slightly tapered posterior end. The two
limbs (that is, the posterior processes) enclose a wedge-shaped space between them, and the
edge along which they touch in front is nearly straight. It is marked by a slight shoulder
posteriorly.

The typical foot (Plate LXXIV, fig. 6) has a small dorsal lobe (cirrus P) in which is
a single vascular loop. The setigerous region beneath is supported by a pair of strong
spines, the points of which often project beyond the surface. The bristles (Plate
LXXXIII, fig. 3) are very brittle, and have comparatively short tips with wings,
which are broad and boldly serrated at the edges throughout the greater part of their
length. Foreign particles thus adhere very readily to them and obscure their structure.
A small lobe or papilla (Plate LXXIV, fig. 6 a) also exists ventrally, so that the conical
foot has two, a dorsal and a ventral, as in Arabella tricolor.

The Gordius piisilliis of Delle Chiaje l would seem to refer to this or a very closely
allied species.

1 < Mem./ iv, pp. 177, 19G ; Tav lxiv, figs. 8 and 9, 1829 ; and ' Descriz./ iii, p. 82; v, p. 97, 1841.

ONUPHIDID^B. 403

2. NOTOCIKRUS, Sp.

Synonym.
1903. Notocirrus sp., Mcintosh. Ann. Nat. Hist., ser. 7 ', vol. xi, p. 565.

Another small and very active species was procured under a stone in a tide-pool at
Herm, but unfortunately it has been lost. The head was smoothly rounded in front, and
of a brighter reddish-orange than the rest of the body, which was dark orange, with the
dorsal blood-vessel in the centre. The segments appeared to be minutely granular as if
punctured. The tail terminated in two longer and two shorter papillae.

Sub-Family Onuphidhle.

Kinberg 1 made the Onuphaea the first family of his Eunicea, characterized by having
seven maxillae, part Ii edentate, tentacles five, antennae two, palpi two, and a single
buccal segment. He grouped the genera according to the condition of the branchiae,
which are plumose in Diopatra, pectiniform or cirriform in Onuphis.

No example of the genus Diopatra, a southern type, has hitherto been met with in

Fig. 83. — General arrangement of the muscles in the body-wall of Hyalinoecia tubicola, O. F. M. x 21. The
vertical bands of fibres passing by the sides of the nerve-cords to the sides of the alimentary canal are
diminishing. The single large neural canal toward the ventral edge of the nerve-cord is conspicuous.
The oblique muscles arise inferiorly at the sides of the nerve-cord.

British waters, though it may yet be found in the neighbouring parts of the Atlantic.
As Claparede points out, the existence of eyes is not general.

Grrube 2 followed Savigny in grouping the Onuphidae under the Eunicea.

The Onuphidae follow the Eunicidae in Levinsen's classification (1883), and apparently
with good reason, and Benham likewise includes the group under the Eunicidae.

Hacker (1896) 3 quotes Wilson as alluding to a stage of Diopatra cuprea with five
segments, and this is a non-pelagic larva, bred in the gallery where the eggs are laid.
He gives a detailed account of the histology of the pelagic and other larvae.

1 < Ofvers. af. K. Vet.-Akacl. Forh./ 1864, No. 10, p. 559.

2 < Fam. der Annel./ p. 44, 1851.

3 'Zeitschr. f. wiss. Zool./ Bd. Ixii, p. 98.

404 ONUPHIS BRITANNICA.

Remarkable tubes are formed by several foreign examples of the group, such as those
of Onuphis sombreriana of the ' Challenger,' which utilizes the long glassy spicules of
vitreous sponges, and 0. Willemoesii, which secretes a finely rounded tube bristled with
long spines of similar material. The Onuphididae are distinguished by their bathymetrical
distribution, for almost all occur in water of considerable depth, and many range to very
great depths, e.g., 2225 fathoms. Not a few of the Eunicids and Lumbriconereids, on
the other hand, frequent shallow water.

In this group reproduction of lost parts has been noted several times in Diopatra by
De Quatrefages, Ehlers, Fauvel, and others.

Genus LXVIII. — Onuphis, Audouin and Edwards, 1834.

Head bearing two eyes and seven tentacles, arranged in three rows — two in front,
two in the middle, and three in rear. Buccal segment with two tentacular cirri. Dental
apparatus highly developed. Branchiae pectinate or simple. 1 Bristles in front large
falcate, bifid forms ; then the foot has simple winged and tapering upper bristles, with a
few brush-shaped forms, and interiorly powerful bifid winged hooks.

1. Onuphis bpjtannica, Mcintosh. Plate LI, figs. 6 and 6a; Plate LXIII, figs. 7-7 d —
head, teeth, tube; Plate LXV, ^.g. 13 — double maxillae; Plate LXXV, fig. 5 —
foot ; Plate LXXXIV, figs. 3-3 c— bristles.

Specific Characters. — Head typical, with well-marked black eyes, situated external
to the base of the long lateral tentacles. First segment about half the breadth of the
succeeding, and having two subulate tentacular cirri which arise from the centre of the
segment. First pair of feet project forward almost to the anterior border of the snout,
and bear falcate bristles (bifid tips) furnished with wings, whilst dorsally are simple
tapering bristles. At the third foot are the latter and brush-shaped bristles dorsally, and
ventrally the falcate bristles. The filiform branchiae commence from the eighth to
tenth foot, in which latter the ventral cirrus is in the form of a boss or pad, and continues
to the tip of the tail. In the same foot the setigerous region is bluntly conical, with a
stout papilla projecting in front of the bristles at the tip, and a long subulate papilla
behind them. Dorsally are simple bristles, finely tapered at the tip, but with broad
wings, and a few brush-shaped bristles. Beneath is a single long bifid hook with wings.
At the twentieth foot the branchia absorbs the entire basal region, the small, slender
cirrus appearing as a process of this organ. The bristles are slightly modified posteriorly,
but the hooks retain almost the same structure. Tubes formed of tough secretion,
strengthened by numerous pieces of bivalve shells.

Synonyms.

1870. Nothria conchylega, Baird. Journ. Linn. Soc, vol. x, p. 356 (partim).
1903. Onuphis britannica, Mcintosh. Ann. Nat. Hist., ser. 7, vol. xi, p. 555.

1 In the ( Camb. Nat. Hist./ ii, p. 319, 1896, the branchiae are described as pectinate (Benham).

ONUPHIS BEITANNICA. 405

1904. Onwphis conchilega, Allen. Journ. M. B. A., n.s. vol. vii, p. 225.

? 1905. „ conchylega, Willey. Rep. Ceylon Pearl Fisheries, iv, p. 276.

Habitat.— Dredged by Dr. Gwyn Jeffreys in 100 fathoms, St. Magnus Bay, and 90
fathoms off North Unst, Shetland, in 1867. Dredged thirty to fifty miles W. of Valentia
in 90 — 125 fathoms and in Donegal Bay (J. Gr. J.) ; in ninety-two fathoms on Adventure
Bank in sandy mud, 'Porcupine,' 1870. Ceylon ? Dr. Willey identifies the Ceylonese
species with that at Plymouth, so that in all probability they agree. S.W. Ireland,
Stat. I, 1885, log 3, forty-five fathoms; Stat, VII, ditto, twenty-two fathoms, 1890
(12th May) ; S.W. Ireland, ditto, Stat. I, log 3; eighty fathoms, P. Irish Acad. Expect.,
1885, Plymouth, as 0. conchilega, (Allen).

Head (Plate LXIII, fig. 7) with two small lobate frontal tentacles on each side of the
middle line, and five long tentacles, the median, as in Eunice, being longest and the inner
lateral much exceeding the outer in length. Each arises from a ringed cirrophore. The
median is most posterior, and the longer lateral springs behind the shorter. A well-
marked black eye occurs to the exterior of the base of the long lateral, and occasionally a
black speck lies in front of the median. On the ventral surface are two flattened palpi,
separated by a median fissure, and projecting laterally rather than anteriorly.

The body is tapered in front and more distinctly so posteriorly, and has forty-two or
more bristled segments, and therefore is much shorter than in Hyalinoecia, The peri-
stomial segment is about half the breadth of the succeeding, and bears dorsally two
subulate tentacular cirri which arise from the centre of the segment, and at the base of
each externally is a black speck. A similar black speck occurs on the next and is distinct
on the following eight or nine segments — gradually fading after the commencement of
the branchiae. The segment following is broad and carries the large first pair of feet,
which project in the preparations forward almost to the anterior border of the snout.
The next feet are also large but they gradually diminish. The body is somewhat
rounded dorsally, especially in front, and has a deep ventral groove. The general
colour, in life, is pale brownish with an iridescent lustre, the pigment being darkest in
front and along the median line, which is also marked by the dorsal vessel. A tinge
of yellow occurs at the tail, and the branchiae are red. The head and tentacles are
yellowish. The body terminates posteriorly in two caudal cirri arising beneath a
dilated anus.

Proboscis. — The dental apparatus (Plate LXIII, fig. 7 a) is pale with a slight tinge
of chocolate, also with dark chocolate bands on the edges of the maxillae and dental
plates and in the form of an English cross at the posterior appendages. The maxilla?
are broad posteriorly and narrow in front, with a sharp tip. The right great dental
plate has eleven or twelve teeth, the first being most prominent and hooked. The
left has ten teeth, but from a hiatus one appears to have been broken. The left
azygos has ten teeth. The right anterior curved plate has ten or eleven teeth> and
the left about the same number. The posterior appendages taper backward from a
somewhat broad base so that they are more or less triangular. The mandibles (Plate
LXIII, fig. 7 b) have long tapering shafts, and the somewhat narrow dental edge has
a few points. A dark spot occurs at the shoulder externally, and in some an angular

406 ONUPHIS BRITANNICA.

or stellate black mark is found toward the inner edge of each dental plate. Certain
differences exist between the foregoing and that figured in the ' Challenger '—it may
be from difference in age. In one example the maxillas had been reproduced (Plate
LXV, fig. 13), both sets being visible in the preparation. No sign of duplication in
the posterior appendages is present.

The first foot is long and slightly tapered — with a bulbous extremity — bearing a
subulate dorsal cirrus about its middle dorsally, and having toward its tip dorsally a
long and rather stout papilla. A stout subulate process (ventral cirrus) occurs near
the base ventrally. The foot is supported by a series of strong yellow spines, and,
though the tip of only one bristle projects beyond the skin, others are included in the
tissues. These consist of large falcate bristles with bifid tips, furnished with wings
(Plate LXXXIV, fig. 3). In the second foot two of these project beyond the tip of the
foot, and the dorsal cirrus is longer, its tip reaching beyond that of the foot. The
terminal papilla is also longer. This foot is smaller than the first.

The third foot is shorter, but is likewise directed forward and outward, and it differs
from the foregoing in the longer dorsal cirrus, and in the short lobate condition of the
ventral cirrus, which is thus homologous with the ventral pads which occur at the bases
of the feet behind. The foot is considerably diminished in size, the dorsal cirrus is very
long, and the distal papilla extends far beyond the tip. Dorsally the setigerous lobe bears
simple and brush-shaped bristles, and below these the tenth foot has dorsally the subulate
dorsal cirrus, from the inner base of which the first branchia arises as a somewhat broad
crenate process which varies in length according to the example; some having short
processes, others having a branchia about as long as the dorsal cirrus and occasionally a
spur appears on it toward the tapering distal end. The setigerous region is bluntly
conical with a stout papilla projecting in front of the bristles anteriorly, whilst posteriorly
is a longer subulate process or papilla resembling a cirrus. The ventral cirrus is in
the form of a pad inferiorly. Dorsally and ventrally are stout bristles with rather short
and finely tapered tips which have broad wings inferiorly. A few brush-shaped forms
are also present, and a single long hook with a bifid tip.

The branchias thus commencing on the tenth foot continue as single filaments almost
to the tip of the tail. Whilst these organs form long thick processes, the dorsal cirri
diminish in size from before backward, the disproportion between the posterior branchiae
and their minute cirri being great, the dorsal cirrus, moreover, moving further up the
branchia in each case.

The twentieth foot (Plate LXXV, fig. 5) presents dorsally the large branchia which
has absorbed the whole basal region, so that the small, slender cirrus appears as a process
of the branchia. The setigerous region forms a short cone with a bulbous tip. The
bristles are the same as in front, viz., dorsally and ventrally stout simple bristles
(Plate LXXXIV, fig. 3 a) with a broad wing on the short and finely tapered tip, a few
brush-like forms (Plate LXXXIV, fig. 3 b), and the powerful bifid hooks with wings
(Plate LXXXIV, fig. 3 c).

The structure of the foot alters little posteriorly except in the proportion of its
parts. The brush-shaped bristles are often prominent from the absence of the tips of
the winged forms.

ONUPHIS BRBVIBRAOHIATA. 407

The tubes (Plate LXIII, figs. 7 c and 7 d) are formed of the tough translucent
secretion of the glands, strengthened by pieces of shell, or sometimes attached to an entire
valve of Pecten. Special attention has been paid to the nature of the protective internal
valves of this species by Mr. Arnold Watson 1 (his Onuphis conchilega), and he figures two
arrow-like va]ves at one end, but whether anterior or posterior is uncertain. The species
appears to carry about its tube caterpillar-fashion (Watson). In the Zetlandic examples
such bivalves as Nucida, Leda, and Lucina are frequent.

This species has certain resemblances to the Diojpatra socialis of Ehlers (1875) from
the c Porcupine ' Expedition of 1869, but, amongst other features, it diverges in the
position of the tentacular cirri ; the simple nature of the branchiae throughout, whereas
in D. socialis they are simple, bifid, or trifid ; by the fact that they arise on the tenth
segment; and in the nature of the tube, which in D. socialis is secretion and mud.

The relationship of Nothria conchyphila of Verrill (1885) remains uncertain, but it
" constructs flat free tubes, about 2 ins. long, out of broken bivalves, and often occurs in
vast numbers in the warm zone," viz., off the northern coast of the United States. The
critical points are unfortunately not shown in the figures ; indeed, the Hyalinoecia artifex
of this author likewise closely approaches the British form.

2. Onuphis blievibraohiata, Elders, 1875. Plate LXIII, figs. 8 and 8 a — teeth; figs. 10
and 10 a— head ; Plate LXXV, figs. 6 and 6 a— feet ; Plate LXXXIV, figs. 4-4 c—
bristles.

Specific Characters. — Head with two rounded and flattened frontal tentacles.
Tentacles short, arising from a ringed cirrophore. Tentacular cirri slightly fusiform
from a constriction at the base. Palpi form two flattened bosses ventrally. Body
resembling that in Hyalinoecia. Dental apparatus has strongly curved maxillae, which are
sharp at the point and rather abruptly swollen a little behind the middle. The posterior
appendages are half-spoon-shaped, with a constriction at the base. Great dental plates
have about ten teeth, and the azygos plate has the same number. Right anterior curved
plate has five teeth ; left broken, but appears to have more. Mandibles with an oblique
smooth edge anteriorly, and an external projection. Branchiae commence on the twelfth
foot, are simple as far as the sixteenth foot, the seventeenth having a branched branchial
process, the twentieth a simple base attached to the cirrus, and the rest being dichotomously
divided. The thirtieth branchia has three divisions, and then the tip is dichotomously
divided. First three feet much larger than the succeeding, more or less ventral in
position, flattened and directed outward. Ventral cirrus of the first shifted, so that it
adjoins the mouth. The tenth foot has a subulate dorsal cirrus with a ventral hiatus
followed by a swelling from which it tapers to the point. It is supported by slender
spines, the blunt points piercing the tip, and a short flap projects posteriorly. Dorsal
bristles long, translucent, slightly curved, and finely tapered— with narrow serrated wings.
Ventral bristles translucent, shafts enlarged and bevelled distally for articulation with the
long tapering terminal piece. The latter kind of bristles disappear at the twentieth foot,
1 ' Trans. Liverp. Biol. Soc./ vol. xvii, p. 311, pi., fig. 8.

408 ONUPHIS BREVIBKACHIATA.

their places being taken by two powerful yellowish hooks with bifid, winged tips. The
brush-shaped bristles offer no peculiarity;,

Synonyms.

1875. Diopatra brevibrachiata, Elilers. Zeitschr. f. wiss. Zool., Bd. xxv, pp. 21 and 49, Taf. hi, figs.

11—21.
1903. Onuphis {Diopatra) brevibrachiata, Mcintosh. Ann. Nat. Hist., ser. 7, vol. xii, p. 133, pi. x,

figs. 5 — 10.

Habitat, — Dredged in the ' Porcupine' Expedition of 1869 at Station 36, 21st June,
48° 50' N., 11° 7' W., and therefore in the British area, at a depth of 725 fathoms on a
bottom of sandy mud.

Also procured in the ' Porcupine ' Expedition of 1 870, probably between Stations 46
and 47, in 358 fathoms, and east of Cape cle Gatte, six miles from shore, in 60 to 160
fathoms, and west of the Strait of Gibraltar, on a bottom of fine grey mud, and in water
varying from 286 to 322 fathoms in depth.

Head (Plate LXIII, figs. 10 and 10 a [after Ehlers] ) with two frontal tentacles
(Stummelfiihler) arising close together in the median line, and forming two rounded and
somewhat flattened lamella in spirit. The five tentacles have the normal position, but they
are short — much shorter than in Diopatra neapolitana — though they also have an enlarged,
ringed cirrophore. The tentacular cirri arise from the anterior border of the first
segment, are wide apart, and have a slightly fusiform outline due to a constriction at the
base. The palpi form two flattened bosses on the ventral surface.

Body somewhat resembling that in Hyalinoecia, but readily distinguished by the
presence of the tentacular cirri and the condition of the branchiae, which lean to the type
seen in Eunice, The length is uncertain, as no complete example occurs. Ehlers gives
24 mm. and forty-four segments, with a breadth 3-4 mm. The first segment of the body
is about the breadth of the succeeding. The dental apparatus (Plate LXIII, fig. 8) is
pale, a dark brown touch occurring at the tip of the posterior appendages of the maxillae,
whilst at the junction of the several parts is a triradiate band. The tips of the maxillae
and the teeth of the great dental plates are slightly brownish. The maxillae are strongly
curved, sharp-pointed, and rather abruptly swollen a little behind the middle. The
posterior appendages are half-spoon-shaped, with a constriction at the base. Each of the
great dental plates has about ten teeth, and the azygos plate the same number. The
right anterior curved plate has five ; the left appears to have a larger number, but is
broken. The mandibles (Plate LXIII, fig. 8 a) have an oblique smooth edge anteriorly,
and at the junction of this with the shaft, externally, is a projection.

The branchiae commence on the twelfth foot as simple filaments attached to the
dorsal cirrus a little above its base. As far as the sixteenth foot they remain simple, long
filaments. The seventeenth foot has a branched branchial process, and at the twentieth
it has a single stalk above the cirrus, and the terminal part is clichotomously divided — a
feature of the species. At the thirtieth foot three divisions occur above the cirrus, which
they considerably exceed in length, and then the tip is clichotomously divided into two
still longer processes. The specimen terminated at the thirty-fifth foot, and on this, so
far as could be seen in the injured preparation, the number of gill-filaments was not less.

ONITPHIS BREVIBRACHIATA. 409

The first three feet are much larger than the succeeding, flattened laterally and
directed outward. The first foot has a large and rather thick dorsal cirrus, which is bent
downward, whilst into its base pass three spines. A bluntly conical setigerous region
follows, having a short tapering papilla at the tip. No bristles are present. The ventral
cirrus is of considerable size, somewhat fusiform in outline, and shifted so that it adjoins
the mouth. The next two feet (second and third) are similar in general structure, the
size slightly increasing, and the ventral cirrus of the third foot is both larger and more
flattened. In none do bristles project beyond the skin, though they occur internally below
the spines. The fourth foot, like the foregoing, is more or less ventral in position. Its
dorsal cirrus is large and subulate, the setigerous region has short bristles projecting from
its edge, and the ventral cirrus is a lanceolate or conical lobe immediately beneath it.

At the fifth foot conspicuous glistening bristles project from the setigerous region,
which has a posterior flap. The dorsal cirrus is somewhat less, though still prominent, and
the ventral cirrus is visible inferiorly as a short process. The dorsal bristles are trans-
lucent and finely tapered, with narrow wings and serrated edges, as in the feet behind.
The ventral are equally translucent, and have their shafts slightly curved and dilated at the
bevelled end, from which a long tapering blade extends distally. The ventral cirrus is
by-and-by included in the low glandular pad (behind the foregoing region), and the dorsal
cirri diminish in size.

The tenth foot (Plate LXXV, fig. 6) has a subulate dorsal cirrus with a ventral
" bite," followed by a swelling, from which it tapers to the point. It is supported by a
group of slender spines. The setigerous region has two strong spines, the blunt points of
which pierce the tissues, and a short flap projects posteriorly. Superiorly is a group of
long, translucent, very slightly curved, finely tapered bristles (Plate LXXXIV, fig. 4)
with narrow wings, which are serrated at the edge. The ventral bristles (Plate LXXXIV,
fig. 4 a) are equally translucent, and have slightly curved shafts — enlarged and bevelled
at the ends for articulation with the long tapering terminal piece, which at the base is
fully the breadth of the enlarged end of the shaft.

At the twentieth foot the jointed bristles have disappeared, and their places are taken
by two powerful yellowish hooks with bifid winged tips (Plate LXXXIV, fig. 4 b), which
in outline resemble miniature pliers. The dorsal bristles have shorter tips with a more
distinct bend at the end of the shaft. The two brown spines have increased in strength.
The thirtieth foot (Plate LXXV, fig. 6 a) presents little change on the foregoing, except
in the branchia3 and the slightly increased slenderness of the dorsal bristles. The brush-
shaped bristles (Plate LXXXIV, fig. 4 c) offer little peculiarity. From the posterior end
of the fragment projected a series of fibres, apparently formed by the glands of the
annelid, and enveloped in a translucent matrix. The secretion is probably allied to that
of Panthalis. The example from Cape de Gatte had the same tough fibres projecting
from its broken end, and Loxosomse occurred on many of the branchial processes.

This species approaches the Diopatra brevibr adnata of Ehlers, though no spiral line
appeared on any branchise present. Ehlers states that the first segment is broader than
the succeeding, but in this form little difference between them is observed. The number
of teeth in the dental plates is greater. It differs from Diopatra simplex, Grube (^izOnuphis
Panceri, Claparecle), in the form of the anterior feet and other particulars.

109

410 ONUPHIS CONCHYLEGA.

3. Onuphis conohylega, Sars, 1835. Plate LXIII, fig. 9 in tube; Plate LXIV, figs. 1, 1 a
—teeth; Plate LXXV, fig. 7— foot; Plate LXXXIV, figs. 5-5 c— bristles.

Specific Characters.— Head resembling that of Onuphis britannica, but differing in the
shorter and more rounded frontal tentacles with pigment behind them. The eyes are
somewhat less. Tentacles with ringed cirrophores (in spirit). The first segment is
narrow, and carries the tentacular cirri at its anterior border. The proboscis agrees in
colour with that of Onuphis britannica, the T-shaped dark bands being conspicuous at the
posterior appendages, as well as a dark patch at the posterior end of each great dental
plate. An incurvation of the inner edge of each maxilla occurs, connected by the
dark mass of pigment with the great dental plate. Right great dental plate has
nine teeth ; the left eight ; the azygos plate has nine or ten teeth. The left anterior
curved plate has nine or ten teeth ; the right has twelve. The usual dark brown
base occurs at each of the latter. A single isolated denticle lies outside each. The
mandibles have an oblique anterior edge with a single denticle, and long tapering
shafts. First pair of feet somewhat shorter than in Onuphis britannica, and the terminal
flap is nearly circular, the dorsal cirrus is subulate, the posterior papilla nearly cylindrical,
and the ventral cirrus (on each side of the mouth) has a slight constriction in the middle.
The stout falcate bristles are strongly hooked at the tip, and beneath is a small secondary
process or riclge. The tenth foot has a long subulate dorsal cirrus, a minute button-like
anterior papilla, and a long posterior papilla at the tip. The ventral cirrus forms a boss.
Dorsally are stout simple bristles, with the tip bent at an angle, and winged as well as
finely tapered. A single bifid hook beneath. The branchiae in some begin on the eleventh,
though generally on the twelfth foot, and each continues as a simple filament almost to
the tip of the tail. Marenzeller 1 describes in the examples from Spitzbergen two or more
divisions, but they never exceed four, the northern forms thus differing from the southern
in this respect.

Synonyms.

1835. Onuphis conchylega, Sars. Beskr. og Iaktt., p. 61, Tab. x, f. 28a — e.

,, Diopatra Eschrichti, Grube. Fam. Annel., pp. 43 and 123.
1843. Onuphis Eschrichtii, CErsted. Gronl. Annul. Dors., p. 20, Tab. iii, f. 33—41, 45.
1851. ,, conchylega, Sars. Nyt Mag. f. Naturvid., vi, p. 209.

„ „ „ Grrube. Fam. Annel., pp. 44 and 123.

1853. „ „ Sars. Nyt Mag. f. Naturvid., vii, pp. 386 and 391.

„ „ Eschrichtii, Stimpson. Invert. Gr. Manan, p. 33.

1857. „ „ Danielssen. Reise, p. 59.

1858. „ „ idem. Ibid., p. 116.

1859. „ conchylega, Sars. Ibid., p. 14 (sep. copy).

1860. „ „ Packard. Canad. Nat. and Greol., vol. vii, p. 403.

1863. „ conchilega, Stimpson. Ann. Greenl. in Proc. Nat. Sc. Pliilad., p. 140.
1865. Diopatra Eschrichtii, De Quatrefages. Annel., i, p. 349.
„ Nothria conchylega, Johnston. Cat. Worms Brit. Mus., pp. 138 and 341.

1 < Archiv f. Naturges./ lv, p. 129, 1889.

ONUPHIS CONCHYLEGA. 41]

1866-69. Onuphis Eschrichtii, Packard. Mem. Bost. Soc. Nat. Hist., i, p. 294.
1867. Nothria conchylega, Malmgren. Annul. Polych., p. 66.

1873. „ ,, M. Sars. Bidrag Christ. Fauna, iii, p. 16 (sep. copy).

1874. „ „ Malm. Goteb. vet. o. vitt. Hand!., xiv, p. 85.

,, „ ,, Smith and Harger. Trans. Conn. Acad., iii, pp. 11, 16, 20.

1875. „ „ Mobius. Jahresb. Comm. deutsch. Meere, p. 168.

1876. „ „ Mcintosh. Proc. Roy. Soc, xxv, p. 217.

1878. Onuphis Escltrichti, Grrube. Schlesisch. Gesellsch., 20th June, 1877 (sep. abdr.), p. U.
„ Nothria conchylega, Marenzeller. Denksch. k. Akad. Wien., Bel. xxxv, p. 396.

„ Onuphis „ Mcintosh. Trans. Linn. Soc, Zool., i, p. 503.

1879. „ „ Hansen. Nyt Mag. f. Naturvid., xxiv, p. 268.
„ ,, hyperborea, idem. Ibid., p. 5, Tab. iv, f. 1 — 9."

„ „ conchylega, Theel. K. Sv. Akacl. Handl., Bd. 16, No. 3, p. 44.

„ Nothria „ Tauber. Annul. Danic, p. 102.

1881. Onuphis „ Horst. Niederl. Arch. Zool., Suppl. Bd. i, p. 14.

1882. „ hyperborea, Hansen. Norsk. Norcl. -Exped., vii, p. 32, pi. iv, f. 5 — 13.

1883. „ conchylega, Wir hi. Cheetop. ' Vega ; Exped., p. 403.

„ „ „ Levinsen. Yidensk. Meddel. Foren. Kjoben., p. 226.

„ „ hyperborea, idem. Ibid.

1885. Nothria conchylega, Mcintosh. ' Challenger ' Annel., p. 305.
1889. Diopatra „ Marenzeller. Arch. f. Naturges., Iv, p. 129.

1891. Onuphis conchilega, Hornel. Trans. Biol. Soc. Liverp., vol. v, p. 240.
1893. Nothria conchylega, Levinsen. Yidensk. Ud. " Hanchs,^ p. 331.
1901. „ „ Whiteaves. Geol. Surv. Canada, No. 722, p. 79.

1903. Onuphis „ Mcintosh. Ann. Nat, Hist., ser. 7, vol. xi, p. 555.

„ „ „ idem. Ibid., vol. xxii, pp. 149 and 164.

1906. „ eschrichtii, Augener. Ball. Mus. Comp. Zool., vol. xliii, p. 141.

1908. Diopatra conchylega, Bhlers. Deutsch. Tiefsee Exped., p. 79.

1909. Nothria „ Moore. Proc U.S. Nation. Mus., vol. 37, p. 138.

Habitat. — In vast numbers in 608 fathoms in the Atlantic as dredged by H.M.
ship 'Triton/ Station 9, 23rd August, 1882; off Tynemouth in forty fathoms (Prof. G. S.
Brady). Frequently procured in the deeper parts of the North Sea by the Fishery Board
Ship " Goldseeker."

Ranges to the Norwegian shores, to Greenland, Finmark (Canon Norman, CErsted,
Mcintosh), Nova Zembla (Theel), and the American coast. Numerous examples from
depths ranging from 98 to 325 fathoms off Florida (Bhlers), Eastern shores of America
(Verrill).

Head (Plate LXIII, fig. 9) generally resembling that of Onuphis britannica, but
differing in the shorter and more rounded frontal tentacles, which likewise have pigment
behind them. The eyes are somewhat less and lie to the exterior of the bases of the long
or posterior lateral tentacles. The palpi are similar. The tentacles have ringed cirro-
phores in the preparations.

Body (Plate LXIII, fig. 9) of the same general outline as in 0. britannica. The first
or peristomial segment is narrow and bears the tentacular cirri 1 towards its anterior
border — one on each side dorsally. They are somewhat short. The proboscis and its

1 Grrube states that 0. conchylega has no tentacular cirri.

412 ONUPHIS CONCHYLEGA.

armature (Plate LXIV, fig. 1) resemble in colour and general aspect that of Onuplds
britannica. The T-shaped dark bands are conspicuous at the posterior appendages, and
a dark patch occurs at the posterior end of each great dental plate. The aspect and
curvature of the maxillaa are the same. A little in front of the posterior appendages an
incurvation of the inner edge of each occurs, connected by the dark mass with the
base of the great dental plate. The right great dental plate has nine teeth and a
posterior ridge; the left has eight and a ridge. The azygos plate (left) has nine or
ten teeth. The curved anterior plate on the left has nine or ten teeth; that on the
right has about twelve. Each of these plates has a dark brown base. A single isolated
denticle occurs beyond (anterior to) each curved process. The mandibles (Plate LXIV,
fig. 1 a) have a single denticle on the oblique edge in front and long tapering shafts.
The outer edge is pointed. The segment following the peristomial is about the same
breadth and carries the first pair of feet, which are shorter than in 0. britannica, have a
nearly circular flap at the tip, though this varies in different specimens, and a little
within it arises the subulate dorsal cirrus. Posteriorly is a blunt and nearly
cylindrical papilla, and ventrally, on each side of the under lip, is a short, thick
ventral cirrus with a slight constriction at the middle, as if a cirrophoral differentiation
existed. The foot forms a short cone supported by three tapering spines, and with
a series of long and strong falcate bristles (Plate LXXXIV, fig. 5), the tip of which is
boldly hooked, with a small secondary process or ridge a short distance beneath. From
the edge of the hook below the ridge, wings extend forward to the tip. In the de-
veloping hook the part first formed is the tip, and it is then seen that the secondary
process attains no larger size than what is figured and described (Plate LXXXIV, fig. 5 a).
In many examples, however, the tip forms a rounded blunt hook without a trace, or
only with a very slight trace of the secondary process, and devoid of wings, a
condition probably due to attrition. The tip as a rule remains boldly hooked. The
structure of the foregoing bristle or hook is fundamentally different from that of Onuphis
britannica.

The second foot forms a still shorter conical process, also usually directed forward,
and having a rounded flap projecting from the tip. The dorsal cirrus is subulate, con-
siderably longer than in the first foot, and arises from the middle of the dorsum. The
papilla from the posterior edge of the tip of the foot is long and subulate. The ventral
cirrus has nearly the same form and size as in the first foot, the constriction indicating
the ceratophore being clearly marked. Dorsally are short simple bristles which have only
a trace of wings, and brush-shaped forms, the long and stout hooks beneath being the
conspicuous appendages. Each hook has the structure already described, the tips of
those projecting being blunt from wear. The third foot has still further diminished,
though the dorsal cirrus is longer. The terminal button-like flap is smaller, the posterior
papilla is long, but the ventral cirrus is a short ovate lobe attached to the base of the foot
ventrally. The bristles and hooks are the same.

The tenth foot has a long subulate dorsal cirrus arising from the base (Plate LXXV,
fig. 7), a short conical setigerous region, with a minute button-like papilla at the tip, and
a long posterior papilla. The ventral cirrus now forms a rounded boss which is becoming
less and less prominent. The setigerous region has stout simple bristles with the tip bent

ONUPHIS QUADEICUSPIS. 413

at an angle and winged, as well as finely tapered (Plate LXXXIV, fig. 5 6). Brush-shaped
bristles occur amongst them. A single bifid hook (Plate LXXXIV, fig. 5 c) is present.

A trace of a branchial process appears in some on the eleventh foot, but as a rule
only on the twelfth foot. It is continued backward, as in the former species, almost to
the tip of the tail. It is a simple process throughout; only occasionally is an accessory
filament found on it. On the whole it is a considerably smaller process than in Onuphis
britannica. The dorsal cirrus diminishes from before backward.

The tube (Plate LXIII, fig. 9) generally consists of coarse gravel and minute stones
fixed to a lining of the secretion. More rarely it is formed of fragments of shells and
stones. Dr. Johnston found the glairy secretion so tenacious that the threads could be
drawn out a foot without breaking.

Several forms, though differing in certain features, approach this species, such as
Onuphis abranchiata, 0. pycnobranchiata, and 0. macrobranohiata of the ' Challenger.'

Hansen (1879) gave a description of this form as a new species in the account of
the Annelids of the Norwegian North Atlantic Expedition in 1876, and mentioned it sub-
sequently in the next year's Expedition. His figures are somewhat indefinite in detail,
but, like the description, give no new feature.

Horst's figure 1 of the jaws of the species is good, though his example has asymmetrical
posterior processes to the maxillse.

Ehlers (1887), in his ' Florida Anneliden,' adopts (Ersted's title under the genus
Diopatra as D. Eschrlchtii, yet admits that it is identical with the unpins coiichylega of
Sars.

Hornel (1891) mentions this species as common off the Liverpool bar, but whether
his form pertains to 0. britannica or 0. coiichylega is uncertain, though in all probability it
is the former.

The Nothvia opalina of Verrill 2 is probably a closely allied form, if not one of the
foregoing species.

4. Onuphis quadricuspis, M. Sars, 1871. Plate LXIV, figs. 2 and 2a — teeth; Plate

LXXV, fig. 8— foot.

Specific Characters. — Read nearly typical, but the median is considerably shorter and
somewhat more slender than the adjoining tentacles. Frontal processes (antennas, Sars ;
palpi, Ehlers) ovate, small. No eyes. Tentacular cirri two, at the anterior border of the
buccal segment. Body elongate, minute, pale bluish iridescent, with two adjacent
brownish bands in the mid-dorsal region. Segments about 150 (Sars), flattened, with the
exception of the first five. It is little tapered anteriorly, but more distinctly so posteriorly,
where, it ends in four cirri — two longer dorsal and two shorter ventral. Branchige begin as
simple processes on the ninth foot, by-ancl-by become bifid, then in three to five divisions
or pectinate, and finally in the posterior part of the body again simple (Sars). Proboscis
with the maxillge much curved with a marked constriction behind the point of junction,

1 'Niederl. Arch. Zool./ Suppl. Bd. i, 1881.

3 « Proc. Amer. Assoc. Sc./ 1873, p. 381, pi. 4, f. 4, 1874.

414 ONUPHIS QUADRICUSPIS.

where a bar of pigment occurs. The posterior processes are pointed. Left great dental
plate has six teeth ; the right has eight. The left lateral paired plate has six, the unpaired
six. The right lateral has seven teeth. The mandibles have curved shafts and the cutting-
edge a denticulation or two, somewhat symmetrically arranged. Tube smooth, a delicate
hyaline secretion coated with greyish sticky mud or muddy sand.

Synonyms.

1871. Onuphis quadricuspix , M. Sars. Vidensk.-Selsk. Forhandl., 1871, p. 4 (sep. copy).
1873. „ „ a. 0. Sars. Bidrag K. Christ. Fauna, p. 16, Tab. xv, f. 7—19.

1875. Diopatra socialis, Ehlers. Zeitschr. f. wiss. ZooL, Bd. xxv, p. 46, Taf. iii, f. 5 — 10.

y, „ quadricuspis, Mobius. Jahresb. Comm. deutsch. Meere, p. 168.

1878. Onuphis „ Grrube. Schles. Gesellsch., Breslau, 1878 (sep. abdr.), p. 10.

1879. Diopatra

1885. Nothria
1903. Onuphis

Tauber. Annul. Danic, p. 103.
Hansen. Nyfc Mag. f. Naturvid., xxiv, p. 268.
Mcintosh. ' Challenger ' Annel., p. 332,
idem. Ann. Nat. Hist., ser. 7, vol. xii, p. 149.

Habitat. — Dredged at various stations during the ' Porcupine ' Expedition of 1869,
viz., Station VI, 808 fathoms, Station VII, 725 fathoms, and Station VIII, 650 fathoms,
on a bottom of muddy sand or sticky mud (Ehlers). Dredged by the ' Knight Errant ' at
Station VI, August 11, 1880, lat. 59° 37' N., long. 7° 19' W., in 530 fathoms; bottom
temperature 46°*5, surface temperature 57 o, ; sea-bottom ooze.

Norway, Drobach (M. Sars). Extends to Canada (Dr. Whiteaves).

Head 1 more or less typical, but the median is considerably shorter than the adjoining
tentacles, all of which have a ringed cirrophore — often of considerable length.

Body minute, about 1 mm. in diameter, but apparently typical in structure. The
buccal segment is broad and bears the subulate tentacular cirri.

The maxillae (Plate LXIV, fig. 2) are much curved, and there is a marked constriction
of the posterior processes immediately behind the point of junction. The processes are
pointed posteriorly, and the tip of each is dark brown and a bar of pigment also exists
behind the maxilla3. The left great dental plate has six teeth (seven, Ehlers), the right
eight. The left lateral paired plate has six (four, Ehlers), the unpaired six (seven, Ehlers).
The right lateral has seven. The mandibles (Plate LXIV, fig. 2 a) have curved shafts,
and the cutting edge presents a denticulation or two, somewhat symmetrically arranged.
The entire apparatus is soft. The dentition was not minutely examined by Sars.

The branchiae begin as simple processes on the ninth foot (Plate LXXV, fig. 8), in
which, as in other respects, it agrees with a specimen dredged by Canon Norman off
Bergen, Norway, and with the descriptions and figures of M. Sars, as given from his
unpublished manuscripts by G. O. Sars, and also of E. Grube. The specimen appears to
be young, and has only about twenty-five segments. The branchiae, therefore, have not
attained full development. The highest number of divisions is four, as Prof. Grube also
states, whereas five are present in the Norwegian form. Ehlers, again, indicates consider-
able variation in the origin of the branchiae in the example (his Diopatra socialis) from

1 From the description in the l Challenger ' volume.

ONITPHIS FRAGOSA. 415

deep water, one having these on every foot, whilst in others they range from the sixth to
the eighth backward.

The first foot has dorsal and ventral cirri and a rather longer and larger median
process behind the setigerons lobe. A few slender spines go to the base of the dorsal
cirrus, whilst the setigerons lobe bears a tuft of rather strong bristles with slender,
minutely-hooked tips, probably bifid, but the preparation is indistinct on this head.
The wings are prolonged to a filamentous process at the tip, and occasionally they
separate.

At the tenth foot simple winged bristles occur, and two stout bifid hooks (Plate
LXXV, fig. 8, ninth foot), the shaft being enlarged in the middle and convex forward,
then sloping gently backward to the neck. The main fang is very strong, its inferior
edge forming more than a right angle with the shaft, and a strong tooth curved slightly
forward projects from the crown. The twentieth foot presents similar bristles and
hooks, and, in addition, three spines of varying size, but with the tips prolonged as
filiform processes. The brush-shaped bristles of the upper division offer no peculiarity.

The posterior feet agree for the most part with the last-mentioned, each having
three spines with the filiform tips, two long hooks, and a tuft of bristles with narrow
wings.

The tube is composed internally of a delicate hyaline secretion coated either with
sticky mud or greyish muddy sand, and it is rather friable.

This is one of the many discoveries made by M. Sars (1871) in the Norwegian
waters, and described from his manuscripts after his death by his talented son. It
apparently haunts the deeper waters up to 300 fathoms off Lofoten, and to a greater
depth off the British shores.

5. Onuphis fragosa, Ehlers, 1887. Plate LXIV, figs. 3-3 b— head, teeth, tube ; Plate
LXXV, figs. 9 and 9 a— feet; Plate LXXXIV, figs. 6-6 b— bristles.

Specific Characters. — Head comparatively small, with slender tentacles of moderate
length, the median being shorter than the adjoining lateral, and all the posterior tentacles
having long, ringed cirrophores. The anterior form rounded bosses. Body somewhat
rounded in front, flattened throughout the rest of its extent. First segment narrower
than the succeeding, and carries the comparatively short, subulate tentacular cirri at its
anterior border. Proboscis, yellowish with dark touches. Maxillge strongly curved
anteriorly and acute. A dark band between them and the posterior appendages, which
are spathulate, with a notch at the end. Eight great dental plate has nine teeth ; left has
nine. Azygos plate nine. The left anterior curved plate has eight, the right nine or ten.
The mandibles are small in front with narrow tips in the centre, then a band of pigment,
and laterally the sloped petaloid blade. First foot bears a series of translucent bristles
with pseudo-articulations at the tips, which are minutely bifid and furnished with
long pointed wings. A few simple bristles with tapering tips (without evident wings)
occur dorsally. The feet decrease in prominence from the first to the fifth, and thereafter
they still further diminish. The dorsal cirrus after the first six or seven becomes very

416 ONITPHIS FRAGOSA.

small, forming at the tenth foot a short subulate process. Besides the spines only short
simple bristles, with tapering winged tips, are present. The dorsal bristles are more
elongate at the twentieth foot, and beneath are two strong bifid hooks. Tube of greyish
sticky mud. Branchias absent. Tube either of pure secretion like a quill or secretion
strengthened with gravel and shell-sand.

Synonyms.

1887. Diopatra {Paradiopatra) fragosa, Elders. Florida Annel., p. 75, Taf. xx, f. 7—14; Taf. xxi,

f. 1—4.
1903. OnupUs „ „ Mcintosh. Ann. Nat. Hist., ser. 1, vol. xi, p. 556.

Dredged in 370 fathoms in the ' Porcupine ' Expedition of 1869 in sticky mud off
the coast of Ireland.

Norway (A. M. Norman) ; off the coast of Florida (Ehlers).

Head (Plate LXIV, fig. 3) comparatively small, with slender tentacles of moderate
length, the median being shorter than the adjoining lateral, whilst the shorter anterior
lateral have finely tapered tips. No eyes are visible in the preparations. The frontal
tentacles are ovoid with narrow pedicles. The palpi form prominent rounded lobes on
the ventral surface^ All the tentacles have long ringed cirrophores.

Body somewhat rounded in front, flattened throughout the rest of its extent. The
first segment has less breadth than the succeeding, and carries laterally, at the anterior
border, the tentacular cirri, which are subulate and comparatively short. The general
colour of the proboscis (Plate LXIV, fig. 3 a) is yellowish, with dark touches here and
there. The maxillas are acute and strongly curved anteriorly. A dark band lies
between them and the posterior appendages, which are outlined in blackish-brown, and
spathulate, a median notch occurring posteriorly. The right great dental plate has nine
teeth; the left has nine. The azygos plate on the left has nine. The left anterior
curved plate has about eight ; the right nine or ten. The mandibles are small,
terminating in front in two rather narrow central tips with a band of dark brown
pigment externally between them and the expanded and sloped external blade.

The first foot (Plate LXXV, fig. 9) is short and is directed slightly forward, the
dorsal and ventral cirri and the terminal papilla being of moderate length. It bears
ventrally a series of translucent bristles (Plate LXXXIV, fig. 6), the curved tips of which
are minutely bifid, with long, pointed wings extending beyond, whilst at the junction of
the tip with the shaft is a pseudo- articulation, as in allied forms, though this is not shown
by Ehlers. Dorsally are a few simple tapering bristles (often only one) in which no wing
is evident.

The feet diminish in prominence from the first to the fifth, and thereafter they still
further decrease so that they* are by no means conspicuous. So far as can be observed
no branchial process is present, even the dorsal cirrus, after the first six or seven,
becoming very small.

The tenth foot has a short flatly conical setigerous region, which has, besides the
spines, only short bristles with tapering winged tips. The ventral cirrus is now adnate,
forming a rounded swelling beneath. The dorsal cirrus is a short subulate process.

HYALINCECIA SICULA. 417

At the twentieth foot the dorsal cirrus is still the same. Behind the dorsal bristles
is a minute flat papilla, the remnant of the longer process anteriorly. The translucent
dorsal bristles are considerably longer with traces of wings at their finely tapered tips
(Plate LXXXIV, fig. 6 a). Below are two strong bifid hooks (Plate LXXXIV, fig. 6b),
the inferior hook marking off the setigerous region from the long elevated ventral pad
formed by the metamorphosed ventral cirrus.

The species secretes a thin but tough tube (Plate LXIV, fig. 3 b) which is coated
with the greyish sticky mud amongst which it dwells.

This form approaches the Diopatra (Paradiopatra) fragosa of Ehlers in the absence
of branchiae, and in the aspect of the head, tentacles, and body. The armature of the
proboscis is also similar. Ehlers 1 describes and figures the tube, however, as formed
of fragments of shells set on edge after the manner of tubes of Owenia filiformis.

Genus LXIX. — Hyalincecja, Malmgren, 1867.

Head with ovate frontal lobes, five smooth filiform tentacles arising from ringed
cirrophores. Eyes two, or absent in the preparations. Palpi in the form of two bulbous
pads, separated by a median fissure ventrally. No tentacular cirri. Body long, flattened,
slightly tapered anteriorly, more distinctly tapered posteriorly, and ending in two long
anal cirri, with the anus on the dorsum of the last segment (pygidium). Proboscis with
maxillge enlarged posteriorly, and all the parts well developed. Mandibles have long
tapering shafts and leaf -like notched anterior edges. First foot is large and slopes forward,
and has a disc-like process at the tip. It carries falcate bristles. Ventral cirrus soon
becomes pad-like. Winged and brush- shaped bristles occur dorsally in the anterior feet,
then strong bifid hooks appear below them with a few winged bristles inferiorly.

1. Hyalinoecia sicula, Be Quatrefages, 1865. Plate LXIV, figs. 4-4 c — head, teeth, and
tube; Plate LXXV, fig. 10— foot; Plate LXXXIV, figs. 7-7 d— bristles.

Specific Characters. — Head as in Hyalinoecia tubicola. The small black eyes lie
outside the base of the long lateral tentacles. All the tentacles have ringed cirrophores.
Body characterized in spirit by two parallel bands of brown from a transverse belt behind
the head, and by a brown spot between each foot from the fifth backward. Maxillas
strongly curved and sharp-pointed ; the posterior appendages elongated and conical. The
right great dental plate has nine or ten teeth ; the left has eight ; the azygos plate has
eight. The curved anterior plates seem to have few teeth — four or five on the right,
fewer on the left. Mandibles with long, narrow, and tapering shafts, and a blunt, leaf-like
anterior plate with a few notches. Branchiae commence on the fifth foot, and extend as
a simple process beyond the fiftieth foot. The bristles of the anterior feet {e.g., the first)
have the end of the shaft slightly dilated, and the bevelled region has spikes. The
moderately elongated terminal piece is sloped at the base and bifid at the tip, the

i < Florida Annel./ p. 76, Taf . xxi ; f . 4.

110

418 HYALINCECIA SICULA.

prominent processes being guarded by wings. The simple bristles have very narrow
wings. The powerful posterior hooks somewhat resemble the bifid forms of Eunice and
of other Oiruphids. The tube is long and narrow, formed of an inner lining of secretion,
more or less strengthened by minute particles of shells, fragments of echinoderms, and
similar structures.

Synonyms.

1865. Onuphis sicula, De Quatrefages. Annel., p. 352.

!869. „ „ Mcintosh. Rep. Brit. Assoc. (1868), p. 337.

„ Hyalinoecia sicula, idem. Trans. Roy. Soc. Edinb., vol. xxv, p. 418, pi. xvi ; f. 3, 36, 3c.
1870. „ bilineata, Baird. Journ. Linn. Soc, vol. x, p. 358.

1901. Onuphis sicula, Whiteaves. Geol. Sarv. Canada, No. 722, p. 80.
1903. „ „ Mcintosh. Ann. Nat. Hist., ser. 7, vol. xi, p. 557.

Habitat— In ninety fathoms off North Unst, Shetland (J. G. Jeffreys) ; Plymouth
(Allen); Coast of Cornwall (Baird); Connemara, Galway (A. G. Moore). ' Porcupine '
Expedition of 1869, off the coast of Ireland in sticky mud at 370 fathoms; Nymph Bank,
S.W. Ireland, in 52J fathoms, Eoyal Irish Academy's Expedition, 1886. 'Porcupine'
Expedition of 1870, at Station VIII, Lat. 48° 26' N., Long. 9° 44' W., in 358 fathoms and
a bottom temperature of 50° F., and at various stations in the southern waters without
and within the Mediterranean, generally on sand or muddy sand. Also obtained by the
' Knight Errant ' at Station III.

Shores of Italy, Sicily, and Palermo (De Quatrefages). 'Porcupine' Expedition,
1870, east of Cape de Gatte, six miles from shore in 60 — 160 fathoms. Gulf of St.
Lawrence (collected by Whiteaves).

Head (Plate LXIV, fig. 4) with three elongated tentacles — a median and two lateral
as in Hyalinoecia hibicola. The small black eyes lie to the exterior of the base of the long
lateral. All the tentacles have ringed bases, and in the preparations (spirit) they often
present crenations or pseudo-articulations.

Body characterized in spirit by two parallel bands of brown which course along the
iridescent dorsum from a transverse belt of the same colour immediately behind the head,
and by a brown spot between each foot from the fifth backward.

Proboscis. — Maxillae (Plate LXIV, fig. 4 a) strongly curved and sharp-pointed. The
posterior appendages are elongated and conical. The right great dental plate has nine or
ten teeth. The left great dental plate has eight teeth; the azygos plate has eight. The
curved anterior plates seem to have few teeth, four or five on the right, fewer on the left,
but they may have been broken. The mandibles (Plate LXIV, fig. 4 b) have long, narrow,
slightly curved and tapering shafts, and a blunt leaf-like anterior region with a few
notches. The branchiae commence as a simple filament on the dorsal cirrus of the fifth
foot, and appear to extend far backward (beyond the fiftieth foot), but both they and the
cirrus are considerably diminished in size, the branchia, at first the smaller, becoming the
larger and continuing so to the end of the fragments.

In the structure of the bristles of the anterior feet a marked difference distinguishes
this species from Hyalinoecia tubicola and 0. coiichylega, since instead of the large un-
jointed winged hooks characteristic of these, jointed or compound forms, after the fashion

HYALINCECIA TUBICOLA. 419

of those of Eunice, occur (Plate LXXXIV, figs. 7 and 7 a). The end of the shaft is
slightly dilated and the bevelled region has spikes. The moderately elongated distal
piece is sloped at the base and bifid at the tip, both the terminal and the secondary
processes being prominent, and they are guarded by wings. The bristles (Plate
LXXXIV, figs. 7 b and 7 c) are slender and have very narrow wings, whereas in the fully
developed forms of the two species above mentioned they have the shape of Valentin's
knife.

Posteriorly the jointed hooks are supplanted by two simple forms which resemble
those so frequently found in Eunice, wings being present in the perfect examples, but
they are sometimes abraded. The bristles are also shorter and some are characteristically
curved at the point. None of the brush-shaped bristles were observed. The hooks in
the posterior region (Plate LXXXIV, fig. 7 d) have a powerful main fang and a smaller
one on the crown.

This species inhabits a tube (Plate LXIV, fig. 4 c) composed of secretion strengthened
by gravel and shell-fragments, yet De Quatrefages thought that its tube resembled that
of Hyalinoecia tubicola.

Ehlers (1868) confused this species with Hyalinoecia tubicola, from which it differs in.
essential features, such as the arrangement of the branchiae, the presence of eyes, the
structure of the bristles, and the pigment.

Dr. Baird (1870) described this form as new under the name of Hyalinoecia Ulineata,
his examples having been dredged off the coast of Cornwall. He noticed that it had a
pellucid, soft tube of a thin horny texture, but he does not allude to any extraneous
materials.

Grube (1878) considered De Quatrefage's species as identical with Hyalinoecia
tubicola.

2. Hyalincecia tubicola, 0. F. Mutter, 1788. Plate LIV, figs. 5, 5 a — head, and 5 b— teeth ;
Plate LXIV, figs. 5-5 b— head, teeth, tube; Plate LXV, fig. 15— mouth; Plate
LXXV, fig. 11— foot; Plate LXXXIV, figs. 8-8 d— bristles.'

Specific Characters. — Head typical, the tentacles arising from ringed bases, and being
long and smooth. No eyes. The frontal lobes (palps, Pruvot; tentacles, Grube) are ovate.
The palpi form two bulbous pads separated by a median furrow ventrally. Body long, of
about eighty-five bristled segments, pale brownish, iridescent, with the double red line along
the dorsum, and the bright red double line in each branchia. The first segment is narrower
than the succeeding. The second segment is considerably broader and carries the first
feet, which are large, slope forward, and have a disk-like process at the tip. No tentacular
cirri. The falcate bristles are strongly hooked at the tip, and beneath is a prominent
secondary process resembling that in Onujohis britannica, and winged. At the sixth and
seventh feet the ventral cirrus is pad-like. Proboscis with maxillae enlarged posteriorly
so as to leave a short hooked anterior region. Posterior appendages bluntly conical in
outline. A dark brown border between these runs forward to a transverse bar between
them and the maxillae. Eight great dental plate with thirteen teeth ; left with thirteen
(sixteen, Ehlers). Left anterior curved plate with seven teeth; right with eight.
Mandibles with a slightly denticulated edge anteriorly and long tapering shafts.

420 HYALINCECIA TUBICOLA.

At the tenth foot is a proportionally longer fusiform dorsal cirrus. The setigerous
region has dorsally one or two long and rather stout bristles with winged and tapering
tips, besides a group of brush-shaped forms. Below is a series of similar winged bristles
with broader wings and shorter tips. The branchiae commence about the twenty-third
or twenty-sixth foot as simple filaments, and continue almost to the posterior end (absent
in the last seven, Langerhans).

At the thirtieth foot the dorsal cirrus has diminished to a slender subulate organ, the
branchia being very much larger and longer. The short blunt setigerous region has dorsally
a group of three longer winged bristles with rather short tips. Brush-like bristles occur
amongst them. Below are two strong bifid hooks. Then follow a few bristles with shorter
tips and broader wings. A ventral pad represents the cirrus. Posteriorly the body
diminishes to a somewhat flattened tail, with the anus on the dorsum and two long finely
tapered caudal cirri. The tube is firm, horny, and quill-like, devoid of extraneous
structures. It tapers towards one end, and has apertures at both.

Synonyms.

1766. Nereis tubicola, 0. F. Miiller. Zool. Danic. Prod., p. 217, No. 2625.
1788. „ „ idem. Zool. Danica, i, p. 18, Tab. xviii, f. 1 — 6.

1806. „ „ Turton's Linnaeus, iv, p. 87.

1820. Leodice „ Savigny. Syst. ArmeL, p. 52.

1827. 'Nereis „ Bruguiere. Encycl. Meth., Vers, i, p. 134, Tab. Iv, f. 7—12.

1828. Nereidonta tubicola (Nereitube), De Blainville. Art. Vers in Diet. Sc Nat., t. lvii, p. 447.
„ Spio filicornis, Delle Chiaje. Mem., vol. iii, pp. 173, 176, Tav. xlv, f. 6, 7.

1833. Nereis tubicola, Aud. & Edw. Ann. Sc. nat., t. xxvii, p. 228.

1834. Onuphis „ idem. Ann el., p. 154.

„ Eunice „ Fleming. Encycl. Brit., 7th ed., vii, p. 219.

1835. Onuphis „ Sars. Beskr. og Iaktt., p. 48.

1838. „ „ G-rube. Zur Anat. Kiemenw., p. 45, Tab. ii, f. 10.

1840. Nereis „ idem. Actin. Echin., etc., p. 82.

1844. „ „ W. Thompson. Rep. Brit. Assoc, 1843, pt. 2, p. 76.
1844-5. Onuphis tubicola, (Ersted. Kroyer's Nat. Tids. (Drobak), p. 405.

1845. Nereis tubicola, Johnston. Ann. Nat. Hist., xvi, p. 6, with figs.
,, „ „ Peach. Rep. Brit. Assoc, 1844, pt. 2, p. 64.

1847. „ „ idem. Ray. Soc Rep. Zool., 1843-4, p. 508.

1851. Onuphis „ Grrube. Fam. Annel., pp. 44 and 123.

1852. Eunice „ Landsborongh. Excurs. Arran, p. 49.

1853. Onuphis „ Sars. Nyt Mag. f. Naturvid., vii, pp. 386 and 391.
1859. „ „ idem. Reise, p. 14 (sep.).

„ „ „ Danielssen. Ibid., 1858, p. 115.

1864. „ „ Grrube. Insel Lussin, p. 79.

1865. Northia „ Johnston. Cat. Worms Brit. Mus., pp. 136 and 341.
,, Onuphis tubicola, De Quatrefages. Annel., i, p. 351.

1867. Hyalinoecia tubicola, Malmgren. Annul. Polych., p. 67.

„ Northia tubicola, Parfitt. Trans. Devon Assoc, ii, p. 20 (sep. copy).

1868. Onuphis „ Ehlers. Borstenw., ii, p. 297, Taf. xiii, f. 1—14.

1869. Hyalinoecia tubicola, Baird. Journ. Linn. Soc, x, p. 357.

1873. „ „ K. Kupffer. Jahresb. Comm. deutsch. Meere, 1871, p. 150.

1880.
1882.
1883.

1885.

HYALINGECIA TUBICOLA. 421

1873. Hyalinoecia tubicola, Sars. Bidrag Christ. Fauna, p. 16.

1874. „ „ Mcintosh. Ann. Nat. Hist., ser. 4, vol. xiv, p. 200.
>} ), „ Malm. Goteb. vet. o. vitt. Handl., xiv, p. 85.

1875. Onuphis tubicola, Mobius. Jahresb. Comm. deutsch. Meere, p. 168.

„ Hyalinoecia tubicola, Mcintosh. Invert, and Fishes St. Andrews, p. 123.
„ Onuphis „ Marion and Bobretzky. Ann. Sc. nat., 6 e ser., t. ii, p. 10.

1879. Hyalinoecia tubicola, Tauber. Annul. Danic., p. 103.

,, Langerhans. Zeitschr. f. wiss. Zool., Bd. xxxiii, p. 291, pi. xv, f. 26.
„ Marion. Ann. Sc. nat., 6 e ser., viii, Art. 7, p. 17.
artifex, Verrill. Proc. U. S. Nat. Mus., hi, p. 357 (fide Augener).
tubicola, Schmiedeberg. Mitth. Zool. Stat. Neap., t. iii, p. 373.
„ Wiren. Chsetop. Vega Exped., p. 403.
„ Levinsen. Yidensk. Meddel. Foren. Kjobenh., p. 227.
„ Mcintosh. ' Challenger ' Annel., p. 335.
„ Cams. Fauna Medit., i, p. 209.

„ Pruvot. Arch. Zool. exper., 2 e ser., t. iii, p. 256, pi. xiii, f. 1 — 5.
1887. Onuphis gracilis, Ehlers. Florida Annel., p. 78.
1893. „ „ Levinsen. Yidensk. Ud. "Hauchs," p. 331.

3 ) ,, „ Marenzeller. Zool. Ergebn., ii, p. 9.

1896. „ „ Roule. Camp. ' Caudan/ p. 445.

1898. Hyalinoecia tubicola, De St. Joseph. Ann. Sc. nat., 8 e ser., t. v, p. 241.

1903. „ „ Mcintosh. Ann. Nat. Hist., ser. 7, vol. xi, p. 557.
„ ,, „ idem. Ibid., vol. xii, p. 164.

1904. „ „ Moore. Proc. Acad. Nat. Sc. Philad., vol. x, p. 144.

1906. „ „ De St. Joseph. Ann. Sc. nat., 9 e ser., t. iii, p. 199, pi. iv, f. 75.

„ „ „ Augener. Bull. Mus. Comp. Zool., xlviii, p. 135.

„ „ „ Eisig. Fauna u. Fl. Neap., xxviii, p. 215.

1908. Onuphis tubicola, Ehlers. Deutsch. Tiefsee Exped., p. 83.

1909. „ „ Benham. Polych. Subantarctic Is., N. Zealand, p. 245.

Habitat. — In various parts of the deeper waters of the British seas, as off Plymouth.
Off the Hebrides, Dr. J. Gr. Jeffreys, July, 1866. Off Balta, Shetland, in fifty fathoms,
Dr. J. Gr. Jeffreys, 1867 (July). Ninety to one hundred fathoms, in the 'Porcupine'
Expedition of 1869. In eighty fathoms eighteen miles W. of Skellig (J. Gr. J.) ; seventy
fathoms in Dingle Bay ; twenty-five fathoms off Valencia, S.W. Ireland ; ninety to one
hundred fathoms twenty-five to thirty miles W. of the Blasquet, S.W. Ireland, 26th May,

1870 (j. a. J.).

A conspicuous feature in the collection of Annelids made by the Fishery Board's ship
6 Groldseeker ' from the Firth of Forth northwards in the North Sea during the Inter-
national Investigations. It occurred at more than thirty stations and generally at
considerable depths. Many of the tubes show numerous and somewhat regular opalescent
curves from internal membranous valves.

In various parts of the Atlantic and Mediterranean during the ' Porcupine ' Expe-
dition of 1870, and in the former case at considerable depths, e.g., 795 fathoms. Marion
thought it was both rare and small at Marseilles. Shores of France (De St. Joseph).
Siberia (Wiren). Christiania Fjord and other parts of Norway (M. Sars). Off Bergen,
Norway (Canon Norman). Off Fayal, Azores, coast of Buenos Ayres, off New Zealand,

422 HYALIXCECIA TUBICOLA.

Torres Strait, and Canary Islands (< Challenger '). Japan (J. P. Moore). West Indies
(Augener). Bast Indies and Africa (Ehlers).

The wide distribution of this species and its varieties is noteworthy.
Head (Plate L1V, fig. 5 a) agrees generally with that in 0. conchylega, the tentacles
arising from ringed bases, and being long and smooth. No eyes. The anterior processes
(frontal palpi, Pruvot, and palpi, Claparede) are somewhat pyriform and slightly
marked by transverse wrinkles in life, but in spirit ovate. The palpi form two bulbous
pads with a median furrow ventrally.

Body typical. The first segment is narrower than the succeeding, and is smoothly
rounded from side to side. The mouth-parts (Plate LXV, fig. 15) present two bulbous
processes at each side of the aperture, with the mandibles posteriorly.

Proboscis (Plate LXIV, fig. ha). — The maxillae have a rather short hooked region
anteriorly, the enlarged region extending far forward. The tips are sharp. The posterior
processes are bluntly conical. A dark brown border to each runs forward and joins a
transverse bar between them and the maxillae, and from the centre a line passes obliquely
outward on each side to a dark spot at the end of the great dental plate. The right
great dental plate has teeth throughout nearly its entire extent, but they vary in number
apparently according to the size and age of the specimen. In the examples from the
' Challenger ' there are fourteen teeth. A British one had thirteen teeth ; the left great
dental plate has about thirteen. The left anterior curved plate has four, and the right
anterior curved plate four. The mandibles have a slightly denticulated edge anteriorly,
and long tapering shafts. The second segment is considerably broader and bears the
first feet, which are proportionally massive organs directed forward with a disk-like
process at the tip. The dorsal cirrus is somewhat short and stout, and arises from the
mid-dorsal region of the foot. The posterior papilla springs from the tip of the
foot behind the disk-like flap. It is subulate. The ventral cirrus, again, arises
near the base of the foot ventrally. The setigerous region is supported by three
spines, and a series of strong, bifid winged falcate bristles (Plate LXXXIV, fig. 8).
The tip is strongly hooked, and beneath the hook is a prominent secondary process,
closely resembling that in Onuphis britanniea, though the attachment of the wing is nearer
that process.

The second foot is considerably shorter and only slightly inclined forward, and as a
consequence the cirri and papilla are nearer each other. It bears both simple bristles
with a winged and tapering tip, and bifid hooks. The ventral cirrus forms a stout
subulate process, that of the next foot (fourth), however, being shorter and its tip blunt.
In the following three feet it gradually diminishes into the pad at the ventral edge of the
foot. The lower bristles of the dorsal series in the anterior region, e.g., the eighth foot
(Plate LXXXIV, fig. 8 d), have shorter tips and broader wings.

At the tenth foot (Plate LXXV, fig. 11) is dorsally the proportionally large and
somewhat fusiform dorsal cirrus — tapering, however, more toward the tip. The seti-
gerous region has dorsally one or two long and rather stout bristles, with winged and
tapering tips (Plate LXXXIV, fig. 8 a), besides a group of brush-shaped forms (Plate
LXXXIV, fig. Sb). Below is a group of similar bristles with broader wings and shorter
tips. The hooks do not seem to be prominent in these feet. The papilla at the posterior

HYALINCECIA TUBICOLA. 423

border of the tip is lanceolate, whilst the terminal process of the setigerous region is
small. At the twentieth foot the dorsal cirrus is more elongate; the posterior papilla is
shorter and more ovate, but the bristles have the same character. The setigerous region
is about the same length as at the tenth.

The branchiae commence from the twenty-third to the twenty-sixth foot as a simple
filament, and continue almost to the posterior end.

At the thirtieth foot the dorsal cirrus has diminished to a slender subulate organ,
whilst from the inner edge of its base arises a branchia more than twice its length and
bulk, and slightly fusiform in outline. The setigerous region is considerably shorter,
forming a short blunt cone with a trace of a papilla at the tip (and it disappears in the
next foot). Dorsally (above the spines) is a group of three longer winged bristles with
rather short tips. Brush-shaped bristles occur along with these. Then follow, below,
two strong bifid hooks with wings (Plate LXXXIV, fig. 8 c). Ventrally are about three
bristles with shorter tips and broader wings than those of the dorsum. The ventral pad
is the only representative of the ventral cirrus. The branchiae diminish in size posteriorly,
as also does the dorsal cirrus, but the arrangement of the bristles and hooks remains the
same.

Posteriorly the body diminishes to a somewhat flattened tail, the last segment of
which is ovoid, with the anus on the dorsum and two long caudal cirri from the hind edge.

In. an example procured in Norway by Canon Norman a subulate tentacular cirrus
of some length sprang from the middle of the first segment nearly in a line with the long
lateral tentacle of the right side. Such would seem to indicate that the absence or presence
of these organs is not a matter of fundamental importance, though it may be a convenience
in classification. In one of the British specimens a tail is being regenerated, and this
is tipped with three long and slender styles, one of which bifurcates near the tip.

Fage 1 observes that the ectodermic portion of the segmental organ of this form is
particularly well developed.

The tube (Plate LXIV, fig. 5 b) is a translucent horny structure, which the animal
drags after it like the larval Phryganea (Pruvot) 2 or other aquatic form. Schmiedeberg 3
has found a peculiar substance in it, which he calls onuphin, as well as an albuminoid
material.

Reproduction. — De St. Joseph gives the diameter of the eggs in ripe examples from
St. Raphael at 0'24 mm., with a porous zona r-acliata.

The original description by 0. F. Miiller (1788) is as definite as his figures, and he
specially points out the singular nature of its pellucid tube.

The tube of this form is represented in Plate LI, fig. 4, of Montagu's MS. drawings
by Miss Dorville (1808) in the Linnean Society's collection.

Grrube gave a brief description of the species from Mediterranean examples in one of
his early papers. 4 He thought the dorsal longitudinal muscles smaller than in Eunice.

1 < Ann. Sc. nat./ 9 e ser v t. in, p. 350, fig. 37 (genital funnel).

2 c Arch. Zool. exper./ 2 e ser., hi, p. 256.

3 'Mitth. Zool. Stat. Neap./ in, p. 373, 1882.

4 l Anat. u Physiol./ Kiemenw., p. 45, 1838.

42 4 HTALINCECIA TUBICOLA.

He mentions the ovaries in their correct position, and gives an account of other structural
features.

Peach (< Rep. Zool./ Ray Soc, 1847) thinks he had observed this form swimming on
the surface of the sea, but Edward Forbes was of opinion that this was merely accidental.
A dried form in a tube, or a tube with air in it, might float, but though active on the
bottom there is no proof up to date that it is pelagic.

Dr. Johnston (1865), founding on the absence of pectinate branchise, placed this form
under his genus Nothria, He made what he calls a vain attempt to form the tube into
a pen.

Ehlers (1868) included the Onuphis sicula of De Quatrefages under this species, a
view not held in the present work. Tauber followed him.

Schmiedeberg (1882), as indicated, made a chemical examination of the tube and
found it analogous to chitin. It would be interesting to compare his results especially
with regard to "onuphin'' with those recently obtained by Prof. Irvine 1 at St. Andrews
on a polarimetric method of identifying chitin in similar secretions in Crustacea, Mollusca,
and Fishes. It may be that onuphin has some relation to mucin.

Pruvot 2 gave an account of the nervous system of this species in which the cerebral
mass is large and pierced by the bases of the posterior lateral tentacles. The anterior
lobes give origin to the stomato-gastric system of nerves, and not the oesophageal con-
nectives. He shows a single central neural canal in the ventral cord. In the British
examples the neural canal is nearer the ventral than the dorsal border. He could not find
the ciliated pockets of Semper at the posterior part of the head. He found in the foot
two ductless glands connected with the bases of the dorsal and ventral cirri, and a smaller
third gland having a similar relation to the cirrus (or papilla) near the exit of the bristles
at the tip.

Arnold Watson 3 carefully describes the valves formed in the tube of this species, the
anterior being V-shaped, and the posterior zig-zag. The inrush of sea- water automati-
cally closes the valves. The worm carries about its tube in its movements on the surface
of the muddy ground it inhabits. From his experiments this observer found that the
annelids could renew the valves when cut off and also of course add to their tubes.

Eisig 4 gives a detailed account of the movements of this species in its tube and when
free. In its tube its motion is what he calls geometroid, and when swimming in the free
condition he terms its action mastigo-helicoidal. It can secrete a new tube in a day.

1 'Trans. Chem. Soc./ 1909, vol. xcv, p. 565.

2 ' Arch. Zool. exper./ 2 e ser., iii, p. 256, 1885.

3 ' Trans. Liverp. Biol. Soc./ vol. xvii, p. 306, etc., figs. 1 — 4, 1903 ; and i Rep. Brit. Assoc./ Belfast,
1902, p. 652.

4 ' Fauna n. Fl. Neap./ xxviii, p. 263, 1906.

EUNICE FASCIATA.

425

Genus LXX. — Eunice, Ouvier, 1817.

Head with five tentacles, cirrophore devoid of rings (Cams), two bulbous palpi,
the second of the first two footless segments with tentacular cirri. Foot with dorsal and
ventral cirri, simple dorsal bristles, compound ventral bristles, and branched or comb-
like branchiae. Four anal cirri. Upper jaw with teeth and fang, one plate more on the
left than on the right side. (Auclouin and Milne Edwards give seven plates.) Anal
segment with two cirri. 1

This genus corresponds with Leodice; Leoclice simplices, Savigny; Nereidonta, Be
Blainville; and Eunice, De Quatrefages. Some, such as Grube, make a sub-genus
Leodice, Savigny, characterized by having the margin of the head bilobate and with
two tentacular cirri.

The body-wall is less massive than in Lumhriconereis, the dorsal longitudinal muscles
especially being less compact. The nerve-area is situated immediately within the circular

Fig. 84. — Transverse section of Marphysa sanguinea, Mont. The great size of the ventral longitudinal
muscles is conspicuous. The dorsal blood-vessel (above the alimentary canal) has been cut along with
a branch on each side.

coat and the basement-membrane, and forms a more or less ovoid area of firm consistence
and with a neural canal toward the outer or ventral border. Jourdan 2 declines to
consider this a colossal nerve-fibre either structurally or functionally. The oblique
muscles pass clown laterally, some of the fibres reaching the basement membrane.

1. Eunice fasoiata, Eisso, 1826. Plate LIV, fig. 1; Plate LXII, figs. 10 and 10 a
—teeth; Plate LXXIV, figs. 7-7 6— feet; Plate LXXXIII, figs. 4 and 4 a— bristles.

Specific Characters. — Head comparatively small, bilobed anteriorly, with five
tentacles of a dull yellow colour with olive-brown bars which are five in number on
the median. Slight constrictions occur at the bars, and minute white grains cover the

1 Benham (' Camb. Nat. Hist/) states that four are present.
3 ' Ann. Sc. nat./ 7 e ser., t. ii, p. 262, 1887.

Ill

426 EUNICE FASCIATA.

surface. Two white spots behind the median tentacle and one in front of it. Eyes
at the base of the inner (posterior) paired tentacles, Body 6 to 9 ins. long, pale
brownish red, with a dorsal streak from the blood-vessel and variegated with regular
white markings. Anal cirri are enlivened with white grains and olive-brown bars.
Second segment equal to about two-and-a-half of the succeeding, with a shield-shaped
white spot in the centre of the anterior border, and two at each side. The next (also
a footless segment) bears the tentacular cirri, and is marked by a central white spot and
one on each side. The tentacular cirri have four olive-brown rings toward the distal
half. The third body-segment is the type of the succeeding, having a white spot in the
centre and three or four on each side, so that in series three longitudinal rows occur on
each side anteriorly. The dorsal cirri are whitish.

Proboscis has the maxillae moderately curved, and of a chocolate-brown hue, with a
dark edge to the flange posteriorly in front of the spatulate appendages. The left great
dental plate has seven teeth, the right six. The right anterior plate has six teeth, and a
dark brown chitinous process supports it. The accessory patch on the outer side has a
single tooth. The left anterior (curved) plate has six teeth and a horny inner edge;
accessory outer plate a single tooth. The left azygos plate has seven or eight teeth.
Mandibles have the anterior edge sloped outward with a tooth at the inner third, and
are marked by concentric lines. First branchial process (a simple filament) occurs on
the fourth bristled segment; next three segments have two, three, and four divisions
of the branchial process, the eighth foot has eight divisions and the tenth about
twelve, so that the semi-pinnate arrangement is marked. The maximum development
of the branchiae takes place in the anterior third of the body. At the fortieth foot
the processes have diminished to nine, at the fiftieth to five, and at the seventieth
foot to three. Gradually diminishing in number they are represented in the caudal
region by a single filament as in front. At the tenth foot the branchial divisions are ten,
and the dorsal cirrus extends beyond the bristles. The dorsal bristles are winged and
tapered, and there are brush-shaped bristles at their bases. The jointed bristles have
shafts curved and dilated at the end, and a short terminal piece bifid at the tip and
with guards. The setigerous region has three yellow spines. A powerful ventral hook
with wings appears before the thirtieth foot, and continues to the posterior end.

Synonyms.

1826. Leodice fasciala, Risso. Hist. Nat. Prod, de FEurope, t. iv, p. 421.

1833. Eunice Harassii, Aud. and Edw. Ann. Sc. nat., t. xxviii, p. 215, pi. xi, f. 5 — 7, 10, and 11.

1834. „ ,, idem. Ann el., p. 141, pi. iii, f. 5— 7, 10, and 11.

1838. „ „ Grube. Anat. n. Physiol. Kiemenw., p. 35, Taf. ii, f. 1—9.

1840. „ „ idem. Actin. Echin. u. Wiirm., p. 83.

1851. „ „ idem. Fam. Annel., pp. 44 and 123.

1864. „ ,, Claparede. Grlannres Zoot., p. 118, pi. ii, f. 5.
„ „ „ Grube. Insel Lnssin, p. 79.

1865. „ „ Johnston. Cat. Worms Brit. Mus., pp. 132 and 340.
„ „ „ De Qnatrefages. Hist. Annel., i, p. 307.

,, „ fasciata, idem. Ibid., p. 330.

EUNICE FASCIATA. 427

1868. Eunice Harassii, Ehlers. Borstenw., ii, p. 312, Taf. xiii, f. 15 — 21; Taf. xiv ; Taf. xv, f. 1 — 3.
1870. „ „ Baird. Journ. Linn. Soc., vol. x, p. 346.

„ „ ,, Grube. Arch. f. Naturges., Bd. xxxvi, p. 294.

1874. „ Claparediij Marenzeller. Adriat. AnneJL, p. 57.

1875. „ „ Ehlers. Zeitschr. f. wiss. Zool., xxv ; p. 52.
1879. ,, „ Langerhans. Ibid., xxxiii, p. 294.

1890. „ Harassii, Malaquin. Annel. Boulon., p. 24.

1891. „ „ Hornel. Trans. Biol. Soc. Liverp. ; vol. v, p. 239 (?).
1893. „ „ Marenzeller. Zool. Ergebn., ii, p. 9.

1895. „ „ Pruvot and Racovitza. Arch. Zool. exper., 3 e ser., t. iii, p. 384, pi. xvii,

f. 48—58; and pi. xviii, f. 69.

1896. „ „ Roule. Camp. ' Oaudan/ p. 445.

1903. ,, fasciata, Mcintosh. Ann. Nat. Hist., ser. 7, vol. xi, p. 557.
„ „ ,, idem. Ibid v ser. 7 ' , vol. xii ; p. 136.

1905. „ Harassii, Graeffe. Arbeit. Zool. Stat. Trieste xv, p. 321 (?).

„ „ tor quota (syn. Harassii, Aud. and Edw.) ; idem. Ibid., p. 321 (?).

1906. „ Harassii, De St. Joseph. Ann. Sc. nat., 9 e ser v t. iii, p. 201, pi. iv, f. 76 (juvenis).
,, ,, ,, Bohn. Ibid., 9 e ser., t. iii ; p. 101.

Habitat. — Dredged in fifteen fathoms off Fermain Bay, Guernsey, on shell-gravel, and
also procured between tide-marks, St. Peter Port (W. C. M.). Off the Hebrides and off
Inverary; in 160 fathoms, fifty-five miles Avest of Valentia, Ireland (J. G. Jeffreys). In
seventy-five fathoms on mud with stones and mussel-shells — ' Porcupine,' 1869 —
Station XXXIV; and in 80 — 110 fathoms on muddy sand with pebbles in the same
Expedition. Very fine examples are procured between tide-marks, Herm. S.W. Ireland,
log 56 (H. I. A. Expedition). Jersey (Hornell).

St. Vincent, Cape Verde Islands ('Challenger'). St. Malo, Isles de Chausey, Dinard,
and other shores of France (De Quatrefages, De St. Joseph). Off Cape Guardia, and in
eighty-one fathoms off Cape Finisterre, ' Porcupine ' Expedition, 1870.

Head bilobed anteriorly from the great palpi, comparatively small and crowded with
the five tentacles, the median being the longest. The ground colour of the tentacles
is dull yellow, with olive-brown bars. These organs do not taper much and seem to be
proportionally thick, slight constrictions occurring at the olive-brown rings, which are
five in number on the median tentacle, the most distinct being at the terminal ring.
Minute white grains also occur on the tentacles. Behind the median tentacle are two
white spots on the head and one in front of it. A few others occur near the base of the
lateral. A well-marked eye lies externally at the base of the inner lateral tentacle and
behind the shorter external tentacle. 1 The prominent posterior edge of the cephalic
segment is dark brown, and the slope of the ridge light brown, the whole defining the
region very distinctly.

Body 6 to 9 ins. in length, rounded in extension, flattened in contraction, and of a
fine pale brownish-red colour, with a dorsal streak from the blood-vessel, and variegated
with regular white markings. It is slightly tapered anteriorly, and distinctly so pos-

1 A careful account of the minute structure of the eye of this form and of other annelids is given
by Von Graber, ' Arch, f . Micros. Anat,/ xvii, p. 243, pis. 28—30.

42 8 EUNICE FASCIATA.

teriorly, the slender tail terminating in the amis, beneath which are two long cirri
variegated with white grains and olive-brown bars.

The second segment is equal to about two and a-half of the succeeding, and has a
slightly projecting collar at each side anteriorly. A shield-shaped white spot occurs in
the centre of the anterior border, a smaller spot on each side, and another to the outer
side and a little in front of the last. A short subdivision (forming the footless
segment of some) bears the tentacular cirri posteriorly, and it has a considerable white
spot in the centre posteriorly and one on each side, the tentacular cirri springing from
the anterior border of these two. The spots and the tentacular cirri are pure white,
and the latter have about four olive-brown rings towards the distal half. Another white
patch occurs on the side beneath. The third segment is the type of the succeeding,
having a white spot in the centre of the dorsum, and three or four on each side (not on
the bases of the feet), so that three longitudinal rows occur on the dorsum anteriorly. In
this segment one or two whitish touches appear at the anterior border, and the speck in
the centre is pale. The rows of spots continue to the tail, and the lateral, which are less
white, also do so, though the general speckling of the dorsum renders them less marked.
The dorsal cirri are whitish and add considerably to the elegant aspect of the animal.
The ventral surface is uniformly iridescent pinkish throughout.

A pale specimen between 8 and 9 ins. in length, procured between tide-marks in
Herm, shows only the anterior region of a faint reddish-brown. The rest is dull whitish
or greyish, with a reddish-brown streak from the intestine and blood-vessel. The whitish
marks throughout are also very indistinct.

Proboscis. — The maxillaa (Plate LXII, fig. 10) are strong, moderately curved, and
of a chocolate hue, with a dark edge to the flange posteriorly, the appendages in the
rear being spatulate. The left maxillary plate has seven teeth, the right six. The
curved plate in front of the right has six teeth, and a dark brown chitinous process
supports it anteriorly. The accessory patch on the outer side has a single tooth. On the
left the curved plate has six distinct teeth and a horny inner edge not evidently den-
ticulated. The accessory outer plate has a simple conical edge. The left azygos plate
has seven or eight teeth. The mandibles (Plate LXII, fig. 10 a) have the anterior edge
sloped forward and outward, with a tooth at the inner third, and are slightly marked by
concentric lines. The limbs are pointed posteriorly. Occasionally a shelf-like flap
occurs below the cutting edge, as in the figure.

The first branchial filament is simple and springs from the fourth bristled segment,
though slight variations are met with. It is attached to the inner base of the dorsal
cirrus. The next three segments have respectively two, three, and four divisions of the
branchial process, whilst the eighth foot has about eight divisions and the tenth about a
dozen. Some of the segments have branchias with fifteen divisions, forming a long
bright red semi-pinnate process of great beauty. Their maximum development takes
place in the anterior third of the body, so that at the fortieth foot they have diminished
to nine, at the fiftieth to five, and at the seventieth to three. Gradually diminishing they
are represented in the caudal region by a single filament as in front.

The first foot has a long and rather thick dorsal cirrus, and a ventral lobe or cirrus
of a massive conical outline. The setigerous lobe is bluntly conical, with two pale spines.

EUNICE FASCIATA. 429

It bears dorsally a small tuft of tapering, simple bristles, and below the spines a series
of jointed bristles, the shafts of which are curved and dilated toward the tip, which is
bevelled and marked by oblique strise. The terminal piece is bifid, with guards at each
side, as in the succeeding feet.

At the tenth foot (Plate LXXIV, fig. 7) the divisions of the branchios are ten in
number, the dorsal cirrus extends beyond the bristles, and there are three spines. The
superior bristles have the same type as in front (Plate LXXXIII, fig. 4). The brush-
shaped bristles at the base of these show no peculiarity. The inferior bristles agree with
those in front (Plate LXXXIII, fig. 4 a), though the terminal piece is less elongated. They
form a dense group.

The thirtieth foot has three spines, the tips of w4iich project beyond the setigerous
region. The bristles are as in front. The ventral cirrus is a short process, which
projects beyond the great ventral pad. The fortieth foot has also three spines, besides
a powerful ventral hook (with wings) issuing just above the ventral cirrus, which now
is elongated. The fiftieth is similar. At the seventieth foot the setigerous region has
diminished in size, and has two spines. The great inferior hook is well marked, and the
ventral cirrus (lobe) is longer.

The ninetieth foot (Plate LXXIV, fig. 7 a) has three spines, the middle one being the
longest, and the tips of all being distinctly curved. The great hook with its crest projects
below the ventral cirrus. Near the tip of the tail (Plate LXXIII, fig. 7 b) two sharp
spines occur in the somewhat deep setigerous region. The jointed ventral bristles are
comparatively large, and the great hook projects inferiorly. The. ventral cirrus is bent
upward and is comparatively long.

So far as can be made out the Leodice fasciata and L. punctata of Eisso l refer to this
form ; indeed, his description is fairly good.

Grrube (1838) in one of his early papers gave a careful account of the external and
internal structure of this species, with accompanying figures.

The head of a gorgeously coloured Eunice on Plate 142 of Delle Chiaje's ' Descri-
zione ' (fig. 7) may be that of this species, though the gill represented in fig. 9 resembles
that of E. aphroditois.

Claparede (1864) describes a well-marked variety of a vinous red colour with a white
band on the head, as well as on the first and fourth segments bearing feet, and with the
tentacles, and the tentacular and dorsal cirri, variegated with white and red.

It is doubtful if the Eunice Glapareclii of De Quatrefages 2 is other than a variety of
E. fasciata.

An elaborate structural account of the species is given by Ehlers (1868), and also a
description of the reproduction of the tail.

Pruvot (1885) examined the nervous system of Eunice torquata, a form which
approaches very closely to this species. It agrees generally with the arrangement in
Hi/alincecia tubicola, but the origin of the oesophageal connectives is double, and the large
neural canal is nearer the ventral border.

1 ' Hist. Nat. de TEurope/ t. iv, p. 421, 1826.

2 ' Annel./ ii, p. 652, 1865.

430 EUNICE FASCIATA.

Pruvot and Eacovitza (1896) x distinguish between this form and E. torquata, De
Quatrefages. Thus the colour of E. torquata is generally brown, and the sixth segment is
always coloured white on the dorsal surface. Median tentacle only as long as the first
four segments, whereas in E\ fas data it is as long as eight. All the appendages are
moniliform, with brown bars. Branchiae begin on the fifth segment and go to the posterior
end, with the exception of the last twenty-five in an example of 140 segments. Maximum
divisions of branchiae between seventeenth and thirty-eighth are about ten. Spatulate
bristles with a long filament at both edges, whereas in E. fasciata there is only a long
filament on one edge. Aciculi black and without valves— whereas those of E. fasciata
are brown, with two valves. Dental plates support a third which is the length of the
mandibular piece (this refers to the maxillae and the posterior piece).

Some of these distinctions are doubtful, but certainly the black spines are
characteristic.

Hornel (1891) shows a form which he refers with doubt to E. fasciata, with a
maximum of twenty divisions to the branchiae.

Gravier 2 records E. fasciata as a temporary commensal in Ostrea edulis from
Coutances on the shores of France. It does not seem to interfere with the mollusk, the
microscopic food of which would not offer much temptation to a form which habitually
selects Crustaceans, Annelids, Bryozoa, and Synapta3. Moreover Phyllodoce rubiginosa,
Nereis Dumerilii, and Lagisca extenuata were found under the same circumstances, thus
probably pointing to accidental commensalism.

A form dredged off the Hebrides (in the Minch) by Dr. Gwyn Jeffreys in July,
1866, shows certain marked differences from E. fasciata, though apparently resembling
it. Thus the branchia3 commence on the third foot, speedily increase in the number of
filaments, which in those best developed are fifteen, then gradually diminish, and cease at
the forty-seventh foot. The absence of branchiae throughout the rest of the body
posteriorly thus differs from the condition in E. fasciata. It agrees with E. fasciata in
the maximum number of the pinnae of the branchiae, but these go no further than the
forty-seventh foot.

The general form of the dental apparatus is similar, but the great dental plates have
seven teeth on the left, and eight on the right, the first on that side being occasionally
bifid. The azygos plate has eight teeth; the curved plate in front about the same
number, and only one continuation plate with a smooth edge is present beyond the latter
(absent on the left from rupture). On the right the anterior curved plate has about
twelve teeth, and a single flat plate with a smooth edge continues the curve.

The appendages to the maxillae form together a spathulate process. The mandibles
are similar to those of E. fasciata, with a tooth at the inner third of the anterior edge.

The foot likewise diverges from that of E. fasciata. At the tenth there are at most
ten divisions of the branchiae. The setigerous region is less prominent and the ventral
cirrus forms a smooth pad without the terminal process of E. fasciata. The dorsal
bristles do not differ much, but the jointed ventral have the first or proximal point of the
hook directed straight outward and not distally inclined as in E. fasciata, The edge of

1 'Arch. Zool. exper./ 3 e ser., t. iii ; pp. 384, 389, et seq.

2 f Bull. Mus. Hist. Nat. Paris/ vi, p. 415, 1900.

EUNICE V1TTATA. 431

the terminal piece is serrated and so is the edge of the enlarged end of the shaft. A
small process appears at the tip of the ventral cirrus at the twentieth and a larger at the
thirtieth and fiftieth, so that this may be a question of degree. The spines are yellow, as
in E. fasciata.

A similar form comes from the ' Porcupine ' Expedition of 1870, sixty-five miles
W. of Valentia, S.W. Ireland, at a depth of 160 fathoms. The branchiae had only eight
branches as the maximum number, but the specimen is smaller.

The same form with eight branches to the branchiae comes from an area twenty-five
miles N.N.E. from Unst, Shetland, in ninety-five fathoms, 1868 (Dr. Gwyn Jeffreys).

A Norwegian Eunicid dredged by Canon Norman approaches E. fasciata in some
respects, the branchiae commencing on the fourth foot and extending to the thirty-
seventh foot or thereabout. The number of pinnae on the larger branchiae appear to be
much less than in E. fasciata, being only six to eight, generally six. 1

So far as known E. fasciata does not harbour the crustacean entoparasites (Eunicicola
Clausii, Kurz.) found for instance in E. Glajjaredii.

2. Eunice vittata, Belle Ghiaje, 1829. Plate LXIII, figs. 3 and 3 a— teeth ; Plate LXXIV,
figs. 10-10 b— feet ; Plate LXXXIII, figs. 7-7 £— bristles.

Specific Characters. — Head pale, with an evenly rounded anterior margin of the fused
palpi, which, however, show a deep ventral furrow. Five long smooth tentacles. Eyes
large and black, in the normal position. Body slender, 2 — 3 ins. long. First segment
about thrice the breadth of the succeeding. The second bears two slender tentacular cirri
at its anterior border. Each in life has a white spot at its base. The anterior third or
fourth of the dorsum is pale brown, posteriorly it is pale fawn. The pale madder-brown
aspect anteriorly is due to the pigmentation of the general surface, and also to brownish
belts which cross the segment at the anterior border. A dark spot occurs at the base of
each foot. Branchiae commence on the third foot as a simple filament or two. On the
tenth foot there are four divisions, on the twentieth five, the same number being on the
thirtieth. The fortieth has four, the fiftieth two, and the sixtieth one, and so to the
eighty- seventh, the branchiae ceasing only on the last seventeen or eighteen segments.
Tenth foot has dorsally the cirrus with its supporting spines, the tips of which pass
into the cirrus beyond the origin of the branchiae, which show four nearly equal divisions.
Setigerous region bluntly conical, the points of two yellow spines projecting at the tip.
Superiorly are simple bristles slightly dilated at the commencement of the tapered,
serrated tip so as to give the organ the advantage of wings. Beneath the spines is a
dense tuft of compound bristles, the shafts being translucent, bevelled at the tip, and
marked by striae directed downward and backward. The terminal piece is rather broad
and short, is bifid, and has wings. Posteriorly the chief changes in the foot are the less
prominent condition of the setigerous region, the diminution in the number of the bristles,
and the presence of the powerful hook, the crown of which (above the fang) is often bifid.

1 Marenzeller's remarks show that considerable confusion exists between this species and
E. torquata, E. Claparedii, and allied forms (' Sitzb. k. Akad. Wiss. Wien/ Bd. lxix, p. 57).

432

EUNICE VITTATA.

1829.
1841.
1851.
1864.
1865.

))
1868.

1870.
1875.
1876.
1879.
1885.
1900.

1903.
1904.
1905.
1906.

1908.
1909.

Ner

tis

Synonyms.

vittata, Delle Chiaje. Mem., iv, pp. 176, 195, Tav. lxiv, f. 12 and 13.

idem. Descriz., iii, p. 94; v, p. 101, Tav. 166, f. 12 and 13.
minuta (juv.), Grube. Fain. Annel., pp. 44 and 123 (?).
vittata, idem. Insel Lussin, p. 79.
Rissoi, De Quatrefages. Annel., t. i, p. 315.
Laurillardi (partim), idem. Ibid., p. 314.
vittata, Claparede. Annel. Nap., p. 133, pi. vi, f. 3.
limosa, Elilers. Borstenw., ii, p. 348, Taf. xv, f. 15 — 22.
Tuhrocincta, idem. Ibid., p. 344.
vittata, Claparede. Suppl. Annel. Nap., p. 34.

„ Marion and Bobretzky. Ann. Sc. nat., 6 e ser., t. ii, p. 11.
limosa, Mcintosh. Proc. Koy. Soc,, xxv, p. 217.
vittata, Langerlians. Zeitsclir. f. wiss. ZooL, xxxiii, p. 293.

Cams. Fauna Medit., i, p. 210.

Fischli. Polych. Ternate (Moluccas), p. 104 (Abhand. Senek. Natur. Gesell.,
Bd. xxv).

Mcintosh. Ann. Nat. Hist., ser. 7, vol. xi, p. 558.

Moore. Proc. Acad. Nat. Sc. Philad., vol. x, p. 435.

(syn. Claparedii ?), Graeffe. Arbeit. Zool. Stat. Triest, xv, p. 321.

De St. Joseph. Ann. Sc. nat., 9 e ser., t. iii, p. 204.

Eisig. Fauna u. Fl. Neap., xxviii, p. 213.

Ehlers. Deutsch. Tiefsee Exped., p. 87.

Lo Bianco. Mitth. Zool. Stat. Neap., Bd. xix, p. 580.

Habitat. — Dredged off St. Peter Port, Guernsey, on shelly ground in fifteen fathoms
in a tube composed of broken shells and stones inside the valve of a large mussel, and in
cracks of leathery tube in another shell at the same depth. This form seems to frequent
water from fifteen to twenty fathoms or more in depth. ' Porcupine ' Expedition, 1870,
Stations XXVII, XXVIII, and XXVIII a. Galway (A. G. Moore); Polperro (W. Baird);
and Plymouth (C. S. Bate).

Shores of France (De St. Joseph). Madeira (Langerlians). Mediterranean (Delle
Chiaje, Grube, Claparede). Japan (Moore). Adventure Bank, e Porcupine,' 1870, ninety-
two fathoms. Cape Verde Islands (Ehlers).

Chiefly a southern form — widely distributed.

Head with an evenly rounded anterior margin of the fused palpi, which, however,
show a deep ventral furrow. It bears five long smooth tentacles, the median being the
longest, whilst the infero-lateral is little more than half the length of the superior. The
eyes are large and black, and they hold a similar position to those of E. fas data. The
head and tentacles are pale, and the latter are smooth under a lens.

Body slender, 2 — 3 ins. long, slightly tapered anteriorly, and forming a slender tail
posteriori}^. The first two segments are devoid of feet, the first being about thrice the
breadth of the succeeding. The second bears the two slender tentacular cirri at its
anterior border. In life each of these has a whitish spot at the base. The last six or
seven segments in front of the vent have very thin dorsal walls, so that even in spirit-
preparations the intestine is clearly visible. Such probably aids in respiration. Dorsum

EUNICE VITTATA. 433

for the anterior third or fourth of a pale brown colour, posteriorly of a pale flesh colour,
and so to the tip of the tail. The slightly madder-brown hue anteriorly is due to the
faint pigmentation of the general surface, but more particularly to brownish belts (red or
roseate, De St. Joseph), which cross the segments at their anterior border, and which
sharply define the segment-junctions. One of the striking characteristics is the presence
of a dark spot at the base of each foot, very distinct posteriorly. Ventral surface pale.

The branchiae commence on the third foot as a simple filament or as two filaments,
though in those from the ' Porcupine ' some show three branches on this foot. On the tenth
foot are four divisions, on the twentieth five, the same number being on the thirtieth.
The fortieth has four, the fiftieth two, and the sixtieth one. The latter number occurs on
the seventieth and the eighty-seventh, the branchiae ceasing only in the last seventeen
or eighteen segments. Variations, however, are frequent. Claparede mentions that they
are ciliated, a feature rarely present in the Eunicidse.

The body- wall in this form is less compact, both dorsal and ventral longitudinal
muscles being extended in transverse section. The nerve-area is within the circular coat,
having the basement-tissue, hypoderm, and cuticle externally. No neural canal is visible.
The oblique muscles are attached at the sides of the area.

Proboscis. — The maxillae (Plate LXIII, fig. 3) are well developed, and have no shoulder
at their posterior articulation. The left great dental plate has nine or ten teeth, the right
ten ; azygos plate seven to ten ; left anterior plate six or seven, and the right ten teeth.
On each side in front of these is a brown nodule. A dark brown pigment-band marks the
articulation between the maxillae and the posterior processes, and passes down each side
of the articulation between them, and another curved band bounds the hind border of the
dental plates a little in front of the articulation. The posterior processes are somewhat
lobate in shape, usually with bluntly rounded posterior edges, the specimen figured being
exceptionally narrow and pointed. The mandibles (Plate LXIII, fig. 3 a) have an oblique
cutting edge, often with a gap.

The first foot has a long dorsal cirrus, and a conical ventral lobe with two pale spines
and dorsal and ventral bristles. The second foot is similar to the first, only the setigerous
region is clearly differentiated from the ventral lobe (cirrus). The third foot has two pale
spines, and the setigerous region is more distinctly separated from the ventral lobe. The
dorsal cirrus is still long, and bears a single branchial filament, a condition Ehlers also found
in a large example, though he figures and describes the first branchia as having six
divisions. The condition, indeed, is variable, some of the British forms having one or
two, the latter being the number in the largest example (from Galway).

The tenth foot (Plate LXXIV, fig. 10) has dorsally the cirrus, which is slightly
enlarged beyond the origin of the branchia at its base, and which has a tuft of simple
bristles, the tips of which pass to the dorsal border of the cirrus beyond the branchia,
which has four nearly equal divisions. The setigerous region beneath is bluntly conical,
the points of the two yellow spines projecting at the tip. Above the spines is the tuft of
simple bristles (Plate LXXXIII, fig. 7), slightly dilated at the commencement of the tip,
and finely serrated, no brush-shaped bristles being visible in the preparations. Beneath
the spines is the dense tuft of compound bristles (Plate LXXXIII, fig. 7 a), the shafts
being translucent, bevelled at the tip, and marked by striae directed downward and back-

112

434 EUNICE NORVEGICA.

ward. The terminal piece is rather broad and short, is bifid, and has wings. The ventral
cirrus has a large granular swelling at its base, its own outline being conical. The foot
generally is vascular. The same type of structure remains at the twentieth foot (Plate
LXXIII, fig. 10 a), except that the blood-vessel going to the branchia is larger, and the
organ has five divisions, as has also the thirtieth foot, but the great hook has now appeared
below the jointed bristles. The basal enlargement at the ventral cirrus has disappeared
at the fortieth foot, and the setigerous region is less prominent, a feature still more
characteristic of the posterior region, where both kinds of bristles are fewer. In the
terminal segments the dorsal bristles are shorter, but the jointed, though few, are
prominent, as also are the great hooks (one in each foot). The crown of the hook
(above the fang) is often bifid (Plate LXXXIII, fig. 7 6).

Reproduction. — Lo Bianco l gives September as the month in which the Neapolitan
examples are mature.

Habits. — It is a vivacious and irritable species, and breaks if much interfered with.
Eisig 2 found this form very agile on the ground, paddling its way over the sand, and an
active swimmer. It also readily secretes a tube, and covers it with sand. When decapitated
the head is reproduced in about nine weeks.

Delle Chiaje, in his ( Memorie,' 1829, describes and figures what apparently is this
form as Nereis zonata and N. vittata, though the tints are brighter than in northern
waters, and the arrangement of the gills and other features are not very accurately
figured by the artist of the Italian naturalist.

Ehlers (1868) re-described this as a new species in the second volume of his ' Borsten-
wurmer ' from examples procured at Quarnero in the Adriatic.

Claparede (1868) found an example devoid of the third (smaller) transverse brown
band in the segment, and it would appear that this species is more definitely pigmented
in the Mediterranean than in the north. This author pointed out the identity of Delle
Chiaje's Nereis vittata with this species, though the figures are far from being
accurate in detail. The same author (Claparede) in his Supplement (1870) observes that
on the forty or fifty last segments, devoid of branchiae, he found a pigment spot at the
base of each foot, forming a minute eye with a lens. Similar spots occur in E. fascia ta,
E. rubrocincta, and Hyalinoecia rigida, but they are less superficial.

3. Eunice norvegioa, L., 1766 (= pennata, 0. F. M.). Plate LXIII, figs. 4 and 4 a — teeth ;
Plate LXXIV, figs. 11 and 11 a— feet; Plate LXXV, figs. 1 and 1 a— feet; Plate
LXXXIII, figs. 8 and 8 a, and 9-9 c for var. BA. — bristles.

Specific Characters. — Head with five long tentacles, which in spirit have an articula-
tion at the tip. Eyes two, of considerable size, black. Palpi fused, leaving a deep notch
in front and a deep groove ventrally, whilst superiorly they are pitted. Body 6 — 8 ins.
in length. First segment three or four times the breadth of the second, with a lateral
notch on each side ventrally. The second segment is even narrower than the succeeding.

1 'Mitth. Zoo]. Stat. Nap./ xiii, p. 486.

2 ' Fauna u. Fl. Neap./ xxviii, p. 213.

EUNICE NORVEGICA. 435

It bears the tentacular cirri. Colour pale brown, iridescent. Proboscis with strong
maxillae, the curve of which is rather flat, whilst the tips rise little above the horizontal.
A marked shoulder exists posteriorly, internal to which is a longitudinal ridge which
alternates with that of the opposite side and interlocks. The maxillae articulate with
spatulate processes posteriorly. Great dental plates have seven teeth— largest in front.
Left azygos plate has nine teeth, and the curved plate adjoining six teeth, and there are
also two separate brown denticles. Right anterior curved plate has ten small teeth, and
a larger and a smaller denticle continue the series as on the left. On the ventral surface
is a slender horny bar between the anterior curved plates and the great dental.
Mandibles with a denticulated anterior edge to the white oral plate; posterior limbs taper
to a blunt point, and are brownish grey with a darker internal edge. Branchiae com-
mence as short subulate processes on the fourth foot — as a rule. Three divisions occur
on the eighth foot. The twelfth foot has only two. Those following generally have two,
occasionally three, then diminishing to one ; they cease about the fortieth foot. The
highest number of branchial filaments noticed was four, but in some young specimens it
was three. The typical foot has dorsally the cirrus of moderate length, with the bifid or
trifid branchiae. The setigerous region is bluntly conical, supported by two black spines
(pale in young examples). Dorsal bristles, long and simple, flattened and widened before
tapering to the delicate tip ; also a series of brush-shaped forms. Ventral bristles have
a short, bifid, terminal piece, considerably narrower than the dilated end of the shaft.
The large posterior hooks have a powerful terminal fang and a strong spike above it.

Synonyms.

1767. Nereis norwegica, Linneus. Syst. Nat., ed. 12, i, pt. 2, p. 1086.

1768. „ madreporse pertusx, Ghmner. Trondhj. Selsk. Schrift., iv, pp. 41 — 51, Tab. 2, f. 6 — 12.

Ibid. Mus. Kircher, Class 12, p. 412, f. 8, 9, 10.
1776. „ pennata, 0. F. Miiller. Zool. Dame. Prod., p. 217, No. 2630.

„ „ pinnata, idem. Ibid., p. 217, No. 2629.

1788. „ pennata, idem. Zool. Danica, i, p. 30, Tab. xix, f. 1 — 3.

„ „ pinnata, idem. Ibid., p. 31, Tab. xix, f. 4 — 7.

1791. „ ,, Linn. Syst. Nat. (Gmelin), ed. 13, i, pt. 6, p. 3116.

1806. „ norvegica, Turton/s Linnasus, iv, p. 87.

„ „ pinnata, idem. Ibid.

1818. Leodice norvegica et N. pinnata, Lamarek. Anim. s. Vert., 1st edit., v, p. 323 ; 2nd edit, v

pp. 562, 563.
1820. „ norwegica, Savigny. Syst. Anim., p. 51.

1827. Leodice pinnata, Bruguiere. Encycl. Meth., Vers, i, p. 134, Tab. lvi, f. 1 — 4.

1828. Nereidonta norvegica, De Blainville. Diet. Sc. Nat., lvii, art. Vers., p. 476.
„ „ pinnata, idem. Ibid., p. 476.

1834. Eunice norvegica, Audouin and Edwards. Annel., p. 145.

1835. ,, „ Sars. Beskriv. og Iagtt., p. 48.
1836 — 7. ,, „ Cuvier. Regno Anim., iii, p. 100.

1844 — 5. „ norwegica, CErsted. Kroyer's Nat. Tids., Bd. i (Dr^bak), p. 406, pi. v, f. 13 15.

1851. „ „ Grube. Fam. Annel., pp. 44, 123.

„ „ norvegica, Sars. Nyt Mag. f. Naturvid., vi, p. 209.

436 EUNICE NORVEGICA.

1858—9. Eunice norvegica, Danielssen. Reise, 1857, p. 50; 1858, p. 116.
1865 - „ „ Johnston. Cat. Worms Brit. Mus., p. 131.

» ,, „ De Quatrefages. Aunel., i, p. 324.

» }, pinnata, idem. Ibid., p. 325.

» » annidicornis, Johnston. Cat. Worms Brit. Mus., p. 131.

1867. Leodice noruegica, Malmgren. Annul. Polych., p. 64.

1868. Eunice „ Ehlers. Borstenw.", ii, p. 347.

1869. Leodice „ Mcintosh. Rep. Brit. Assoc. (1868), p. 339.

1870. Eunice „ Baird. Journ. Linn. Soc, vol. x, p. 345.

1873. Leodice „ Sars. Bidrag Christ. Fauna, p. 22 (sep. copy).

1874. Leodice „ Malm. Goteb. vet. o. vitt. Hancll., xiv, p. 85.

1875. Eunice „ Mobius. Jahresb. Comm. deutsch. Meere, p. 168.
» jj „ Ehlers. Zeitsch. f. wiss. Zool., Bd. xxv, p. 53.

1879. Leodice „ Hansen. Nyt Mag. f. Naturvid., xxiv, p. 268.

,> „ ,, Tauber. Annul. Danic, p. 102.

1881. Eunice „ Horst. Niederl. Arch. Zool., Suppl. Bd. i, p. 13.

1883. Leodice „ Levinsen. Yidensk. Meddel. Foren. Kjobenh., p. 229.

1893. „ „ idem. Yidensk Ud. " Hauchs," p. 331.

1902. Eunice pennata, Marenzeller. Polychset. des Grundes, p. 16.

1903. „ „ Mcintosh. Ann. Nat. Hist., ser. 7, vol. xi, p. 558.
„ „ „ idem. Ibid., vol. xii, pp. 137 and 164.

Habitat. — Dredged off Inverary by Dr. Grwyn Jeffreys in July, 1866. One was in a
tube in a valve of Pecten. At Station XXXIV ' Porcupine ' Expedition of 1869, in 725
fathoms on mud and sand, and a variety in 539 fathoms in grey mud in the Expedition of
1870. Off Balta in fifty and ninety-five fathoms. Dredged fifty-five miles west of
Valentia, Ireland, in sixty fathoms (J. Gr. Jeffreys).

Amongst corals and mud, Norway (Canon Norman) ; Adria (Marenzeller) ; forty-five
fathoms off Cape Sagres, 'Porcupine,' 1870. Extends to Canada (coll. Dr. Whiteaves).

Head with five long tentacles, the median being longer than the others, and in spirit
presenting an articulation at the tip. 1 The adjoining pair have a tendency to further
crenulation at the tip, whilst the external pair, which are considerable shorter, have a
still more evident crenulation. In origin the median tentacle is most posterior ; the outer
lateral arise in front of the other three. The eyes are of moderate size, black, and lie in
a line behind the outer lateral tentacles, and to the exterior of the bases of the inner
lateral tentacles. The palpi are fused, leaving, however, a deep notch in front and a deep
groove ventrally. The palpi are excavated or pitted superiorly.

Body 6 — 8 ins. in length, little tapered anteriorly, but gradually diminishing to a
slender tail posteriorly, where it terminates in two long caudal cirri beneath the vent. In
the preparations it is rounded dorsally, and somewhat flattened ventrally. The first
segment is three or four times the breadth of the second, and has a well-marked lateral
notch on each side, and its front edge is concave from side to side ventrally. The second
segment is even narrower than the succeeding, and carries dorsally the two tentacular cirri,
which are slightly crenulate in spirit, and reach fully to the anterior border of the first
segment. Like the other processes they are tapered.

1 Muller's N'. pennata had non-articulate tentacles.

EUNICE NORVEGICA. 437

The proboscis has a strong pair of maxilla?, the curve of each of which is rather flat.
In lateral view the tips rise very little, so that the organs are nearly horizontal. Posteriorly
the outer angle of each maxilla is slightly elevated and projects as a prominent external
shoulder. Within this is a definite raised longitudinal ridge, which alternates with that of
the opposite side, so that the one fits into a pit superiorly, the other inf eriorly — thus locking
the jaws securely together, especially against vertical motion. Moreover, as the maxilla? fit
into a crescentic hollow of the great dental plate, additional security is given to the grasp of
food or living prey (Plate LXIII, fig. 4). Each maxilla articulates with a spatulate process
posteriorly. The great dental plates are powerful and each has seven teeth, the largest in
front. These teeth appear to have a scissor-like action, the left passing over the right.
Rather more than a third of the posterior edge is smooth. The first antero-lateral plate
on the left is further backward than that on the right, and forms what has been termed
an azygos plate with nine teeth. It is also less arcuate than that on the right. The
curved plate next it has six teeth, and there are two brown denticles — a larger and a
smaller — which continue the row, and each has a terminal denticle. The right anterior
curved plate has ten or more small teeth, and a larger and a smaller denticle continue the
series beyond it, as on the opposite side. On the ventral surface a slender horny bar
occurs on each side between the curved plates and the great dental, but it is disconnected,
and there is no horny bar going to the maxilla? as in other forms.

The mandibles (PI. LXIII, fig. 4 a) are powerful, the anterior oblique edge being
denticulated, and the oral plate (which, as usual, is most extensive ventrally) is white.
The posterior limbs taper to a blunt point. They are brownish-grey with a darker
internal edge. Slight movement occurs between the halves of the apparatus.

The branchia? commence as a short subulate process attached as a rule to the inner
base of the dorsal cirrus of the fourth foot, though it may be the fifth in some — indeed, there
is variation in this respect. About the eighth foot the branchia has three divisions and
the same on the right tenth foot, the left having only two. The eleventh has three on
one side and two on the other, whilst the twelfth has two on each side, but the right
dorsal cirrus is bifid. The branchise behind generally have two divisions, occasionally
three, then they diminish to one and cease at the fortieth foot, or occasionally there are
rudimentary or simple processes to the forty-third or forty-fourth. The highest number
of branchial filaments in each foot is four. In some young examples the highest is three.
0. F. Muller describes his E. pinnata as having branchia? from the seventh to the fortieth
foot, and the pinna? varying from six to eleven. Various abnormalities occur from injury
and partial reproduction.

All the parts of the foot undergo diminution at the fortieth, only a minute branchial
process being present on the base of the cirrus. The ventral cirrus is also less. The
spines are either black, or, in young forms, of varying degrees of yellow or brown. As a
rule they are gently tapered almost from the base to the tip, which is very slightly hooked
— that is, curved backward and then forward. The tips are often abraded.

Posteriorly are the blackish hooks (Plate LXXXIII, fig. 9 c), the shafts of which have
a more decided curve than the spines, the tip being bent backward and then forward as it
diminishes to the neck. The great fang is short and stout, its inferior border forming
more than a right angle with the neck, whilst the single short spike on the crown is

438 EUNICE NORVEGICA.

directed upward and slightly forward. The wings leave the neck a little above the point
of diminution and are often injured distally. They are of varying degrees of yellow or
brown in young specimens. There is little that is distinctive in these hooks in the group
to which this species belongs.

The first foot has a long dorsal cirrus, and a considerable ventral cirrus — the two
arising close together, so that but a small space remains for the two spines and
the dorsal and ventral bristles. The dorsal cirri of the first five feet, indeed, are
conspicuously longer, thereafter they diminish. The ventral cirri of five or six of the
feet in the same region are likewise proportionally long.

At the tenth foot (Plate LXXIV, fig. 1) the dorsal cirrus is still proportionally
long, and bears from its inner or dorsal base a single branchial process not half its
length. Occasionally there are two. The setigerous region is bluntly conical, and is
supported by two strong black spines (pale in young specimens). Superiorly are long
simple bristles (Plate LXXXIII, fig. 9) and a series of brush-like forms (Plate
LXXXI1I, fig. 9 a), while ventrally are the compound (Plate LXXXIII, fig. 9 b) or falcate
bristles.

At the twentieth foot the chief distinctions are the diminution in the dorsal cirrus,
the branchia being bifid (or it may be trifid), and the increase in the depth of the dorsal
cirrus, which has a thick cushion-like aspect.

In the examples brought from Norway by Canon Norman the tenth foot (Plate
LXXIV, fig. 11) has a branchia with four divisions. Twentieth foot (Plate LXXIV,
fig. 11a) has six divisions to the branchia. The falcate bristles of these are represented in
Plate LXXXIII, fig. 8, and a posterior hook in Plate LXXXIII, fig. 8 a. Teeth and
mandibles of this form are shown in Plate LXIII, figs. 4 and 4 a.

In a series of small examples dredged by Canon Norman in shell-gravel in Norway
the branchiae commence on the third foot. On the eighth foot the process is still simple.
The twelfth foot has four divisions, whilst the twentieth (Plate LXXIV, fig. 11a) has six
or seven. At the thirtieth foot are five, and at the thirty-ninth (last) a simple process.
The maximum number of filaments of the branchiae is nine or ten. They form tubes
amongst the shell gravel. Considerable variation, indeed, is present. The dental
apparatus of this small variety appears to agree with the type.

In a form of medium size in a very imperfect state of preservation, dredged in the
' Porcupine ' Expedition of 1870 at a depth of 539 fathoms in grey mud, certain
features diverge from the type above mentioned. Amongst these are the occurrence of
black spines, the comparatively feeble development of the branchiae, and the slight
articulations of the dorsal cirri. On the tenth foot the bifid branchia is little more than
a third the length of the dorsal cirrus, and it is slightly shorter at the twentieth foot,
one of the divisions being rudimentary. A single short filament only occurs on the
thirtieth foot, and it disappears before the fortieth. This form does not appear to lean
either to the Eunice tovquata of De Quatrefages or other type, and may require
separation. The material in hand, however, does not warrant this step.

On the third foot of an example a long process projected from the posterior aspect
of the setigerous region — evidently an abnormal papilla of the cutaneous tissues.

Pood. — In the intestine of examples from Norway were vast numbers of spicules

EUNICE FLOEIDANA. 439

of G-orgonidse, fragments of Crustacea, and fragments of shelly tubes of Serpulids
( Hydroides) .

Habits. — 0. F. Miiller mentions the occurrence of this species in " Madreporae," and
more recently Marenzeller observes that this species, like E. floridana, Pourtales, inhabits
coral-stocks of Loplwlielia prolifera and Amphihelia oculata. Such appears to be the habit
of certain Eunicids in all seas, the worms, as Dr. Alcock tells us, for instance in the stocks
of Lophohelia investigator is in the Indian Ocean, living in " tunnels formed by a crust of
scleroderm that connected two neighbouring branches." 1

Reproduction. — Those obtained in Norway by Canon Norman were loaded with ova
and sperms in July.

In the c Encycl. Meth.' (1827) Leodice pinnata, Lamk., is shown with few divisions
to the branchiae, whilst Leodice norvegica (idem, Vers, i, p. 134, Plate LVI, figs. 5 — 7)
has a larger number of divisions in the branchiae (six or seven), and the dorsal cirrus is
larger.

0. F. Miiller (1788) distinguished his N.pennata from N. pinnata by the presence of
whitish globules (ova ?) above and below the intestine, and by the pennate instead of
pinnate branchiae, which arise in the former on feet seven to forty, and in the latter from
feet five to twenty-six. Moreover, he figures the former with a dark speck at the base of
the dorsal cirrus, and a median line of dark specks along the dorsum.

(Ersted considered that the branchiae of the male E. norvegica were less divided (from
three to five pinnae) than those of the female, which had eight pinnse on the branchiae,
and he thus made E. pinnata of 0. F. Miiller the male, and E. pennata the female of the
species. In this connection Grube 2 observes that a female example with ova showed
from seven to thirteen processes in the branchiae, whilst one with minute eggs had four
divisions.

Dr. Johnston's description (1865) has been held as pertaining to this species, though
he states that he had not seen a specimen. The origin of the branchiae on the seventh
segment and the occurrence of from three to thirteen filaments in the branchiae are
differences. What the same author's E. antennata may be is uncertain.

Tauber places Eunice fasciata under this species, but he gives no definite reasons.
In E. fasciata, however, the branchiae continue to the end of the body, whereas in E.
norvegica they cease at the fortieth foot.

4. Eunice floridana, Pourtales, 1869. Plate LXV, figs. 7-7/; — head and teeth; Plate
LXXV, figs. 2 and 2 a— feet; Plate LXXXIII, figs. 10-10 b— bristles.

Specific Characters. — Head with the palps grooved dorsally. Eyes behind the cirro-
phore of the frontal (external) tentacle and exterior to the lateral tentacles. Median
tentacle as long as first ten or eleven segments. Tentacles slightly annulated. No other
process is so. Tentacular cirri stretching beyond the palps. Body of 110 to 150 or more

1 ' Zoological Gleanings/ 1901, p. 42, 1901.

2 < Schles. Gesellsch./ 20th June, 1877 (1878), p. 16 (sep. abdr.).

44 EUNICE FLORIDANA.

segments and from 120 to 190 mm. in length, of a pale rose colour dorsally, pale iridescent
ventrally. Two small anal cirri. Spines generally black throughout. Spatulate bristles
with a long filament on one edge. Compound, with stout terminal piece, and bifid. Hook
commencing about the thirtieth segment, and it is also black. Great dental plates with five
or six teeth. Azygos plate, five to six teeth ; anterior plates, four to eight teeth on left,
eight to ten on right. Posterior process of apparatus one third length of maxilla?. V
and VI anteriorly have each a tooth. Branchiaa commence on the seventh foot and
continue almost to the tip of the tail. The divisions generally do not exceed four, but
sometimes six to eight occur, and there are for the most part three anteriorly. Inhabits
parchment-like tubes often on Lophohelia and Awphihelia.

Synonyms.

1863-69. Marphysa florideana, Pourtales. Bull. Mus. Comp. Zool., vol. i, p. 108.

1887. Eunice floridana, Ehlers. Mem. Mus. Comp. Zool., xv, p. 88, pi. xxii, f. 1 7.

1893. „ philocorallia, F. Buchanan. Proc. K. Dubl. Soc, vol. viii, p. 173, pi. ix. f. 2 6; pi. x,

f. 7 — 9 ; and pi. xi.

„ ,, floridana, Marenzeller. Zool. Ergebn., xii, Polych. Grand., p. 7, Taf. ii, f. 5.

1895. „ florideana, Pruvot and Racovitza. Arch. Zool. exper., 3 e ser., t. iii, p. 395, pi. xvii, f.

59—62; pi. xviii, f. 62.

,, „ amphihelii, Eoule. Camp. c Caudan/ p. 447, pis. xix, xx, xxiii, and xxv, f . 1 — 3, 16—26.

1903. ,, philocorallia, Mcintosh. Ann. Nat. Hist., ser. 7, vol. xi, p. 559.

,> „ „ idem. Ibid., ser. 7, vol. xii, p. 140.

,, „ amphihelise, idem. Ibid., p. 164.

Habitat— Large examples dredged in the 'Porcupine' Expedition of 1869 in 173
fathoms between Galway and the Porcupine Bank. Amongst colonies of Lophohelia
prolifera at a depth of 200 fathoms, 50 miles off Bolus Head, Kerry (F. Buchanan).

Sand Key, Florida, in 100 fathoms (Pourtales). Gulf of Gasgoiue (Marion and
Roule). South of Cap St. Maira di Leuca (Marenzeller). Norway (Norman).

Head (Plate LXV, fig. 7) with the palpi separated by a notch in front and with a
deep hollow dorsally anterior to each frontal tentacle. These are about the diameter of
the head in spirit, and retain their crenations. They are little tapered in several examples,
but in one in which the left had been reproduced it was as subulate as a dorsal cirrus.
Behind these are the longer median and two lateral tentacles, the cirrophore of the former
being just posterior to the line from the notch between the palpi, whilst the latter tentacles
are slightly in front at each side. All bear traces of crenations. Behind the frontal and
exterior to the lateral tentacle is the eye, which is of moderate size.

Body 6 — 8 ins. long and about 4 mm. broad in front, very little tapered anteriorly,
but posteriorly diminishing to a slender tail with two cirri. Segments 100 — 150. First
segment about as broad as the three following, with a deep notch laterally opposite the
palpi, and a crenate border inferiorly. The second segment is shorter than the third and
bears the tentacular cirri, which vary in length, but often extend beyond the frontal
border of the first segment, and, Miss Buchanan observes, even beyond the palpi.

The armature of the proboscis (Plate LXV, figs. 7 a and 7 b) corresponds with the
description of Marenzeller. The posterior appendages of the maxillas are lobate. The

EUNICE FLORIDANA.

441

maxillary plates (II) have six teeth on the right, five on the left. The paired plates (IV)
have nine or ten on the right, seven or eight on the left; impaired (III) five teeth. The
two anterior (V and VI) have each a tooth. In small forms, as this author observes, the
teeth are fewer. The mandibles (Plate LXV, fig. 7 b) have expanded anterior plates and
long tapering shafts.

In transverse section (Fig. 85) the great muscularity of the species is noteworthy, and
the dorsal longitudinal muscles are specially large. A large neural canal exists inferiorly.

The typical foot (Plate LXXV, fig. 2) has an unjointed dorsal cirrus, which from the
seventh foot onward to the tip of the tail carries a branchia. The first branchia has one,
two, or three divisions, and as a rule the maximum number (which Marenzeller locates from
the sixteenth to the eighteenth) is four. The branchiae are less developed in small forms
(Plate LXXV, fig. 2 a). While, however, the foregoing description applies to many
examples, it does not suffice for all. Thus some well-developed specimens show greater
freedom in the branches of the branchiae, a feature exemplified even in the first, which in

Fig. 85. — Transverse section of the anterior third of Eunice floridana. The dorsal longitudinal muscles are

of great size.

one had been mutilated, and from the stump sprang a short tuft of six filaments. The
second, however, had only two of the usual aspect, but they rapidly increased, so that the
tenth foot had six, and some of the succeeding had seven filaments and possibly one or
two more might have been found in others, for Ehlers 1 figures nine. No apparent
difference could be detected in the branchiae of the two sexes.

The setigerous region is supported by two powerful black spines, the tips of which
generally pierce the skin. The dorsal bristles (Plate LXXXIII, fig. 10 c) are simple, with
finely tapered tips and narrow wings. The brush-shaped forms (Plate LXXXIII, fig. 10)
have a long process at one side. The compound (falcate) bristles (Plate LXXXIII, fig. 10 a)
are yellowish, dilated toward the distal end of the shaft, which is bevelled and has internal
striations. The terminal piece is somewhat short, with a bifid tip, and winged. The
ventral hooks (Plate LXXXIII, fig. 10 b) appear about the thirty-first foot and continue to
the posterior end. At first they are single, but by-and-by two project, and this till near
the tip of the tail, when again a single hook occurs. The greater part of the hook is
black, the proximal and distal ends alone being yellow. The tip has a strong fang at

1 c Florida Aimel./ pi. xxii, fig. 4.

113

442 MARPHYSA SANGUINEA.

right angles to the shaft, and a smaller hook on the crown. The commencement of the
wing is indicated by a constriction in front.

In front the ventral cirri are bulbous inferiorly — with a conical tip, but posteriorly
they are more elongate. The foot, indeed, in the posterior region of the body is some-
what more prominent, the spines are proportionally stronger, and the bristles more
slender and longer.

Pourtales gave no tentacular cirri in his original description, but such appears to have
been due to a misapprehension. He found it inhabiting "large deformed, paper-like
tubes, with lateral openings irregularly placed, though in general alternate, bordered by
laciniate and fimbriate flaps." He described the branclme as five- to seven-lobed, and
stated that they began on the seventh or ninth ring.

Miss Buchanan (1893) was right in pointing out the close resemblance of her
Eunice jpMlocorallia to E. floridana of Pourtales and Ehlers, the differences mentioned,
however, being only variations.

Marenzeller's account (1893) is both minute and careful. He figures a calcareous
tube with terminal and lateral openings attached to Lophohelia prolifera, and he also found
it on Lophohelia ocalata in the Museum at Vienna.

Roule (1896) does not allude to Miss Buchanan's species, and considers that E.
amphihelise most nearly approaches E. fasciata. He mentions that a note about this
form was first published by Filhol in " La Vie au fond des mers," ' Bibliotheque de la
Nature' (Paris, Masson).

Genus LXXI. — Marphysa, Be Quatrefages, 1865.

Head bilobed or rounded, with massive palpi. No tentacular cirri. Body typical,
terminating posteriorly in two cirri. Maxillse as in Eunice, but with a tendency to
elongation of the posterior processes. An azygos plate on the left. Foot with a dorsal
and a ventral cirrus. Knife-shaped bristles superiorly, compound falcate inferiorly.
Strong bifid hooks occur in the ventral region posteriorly.

Ehlers (1868) characterized the genus as having five tentacles and a pair of massive
palpi. No tentacular cirri on the second of the two footless segments. Foot with a
dorsal and a ventral cirrus, dorsally with simple knife-shaped bristles, and ventrally with
compound bristles. Branchiae simple or with a short stem from which project filaments
more or less elongate. Four anal cirri. Upper jaw with teeth and fang; an additional
plate in the left row of teeth.

1. Marphysa sanguinea, Montagu, 1807. Plate LIV, fig. 2 ; Plate LXIII, figs. 1 and 1 a
—teeth; Plate LXV, fig. 14— foot; Plate LXXIV, figs. 8 and 8 a— feet ; Plate
LXXXIII, figs. 5-5 b— bristles.

Specific Characters. — Head deeply bilobed, flattened, with five dull yellow and rather
long tentacles, which are slightly wrinkled. Eyes normal in position. Body 18 ins. to 2 ft.
long, rounded in extension, flattened in contraction, terminating posteriorly in two cirri.
Colour ash-grey or greenish brown, duskier in front, paler posteriorly, with fine irides-

MARPHYSA SANGUINEA. 443

cence. Dorsum minutely dotted anteriorly. Maxillse with a flattened curve and a
strong hook in front. A flange posteriorly indents the great dental plate, which on the
left has three teeth in front, and on the right four. The anterior crescentic plate on the
right has seven to eight teeth, the left five to six. The azygos plate on the left has six to
seven small teeth. The spathulate posterior appendages are dark brown, the ends of the
maxilla being pointed and pale chocolate brown. The mandibles have a dense porcel-
lanous plate ventrally, the front edge being bevelled to a blunt point externally and
having a black spot, whilst the limbs taper posteriorly and are marked by longitu-
dinal lines. The foot presents considerable changes in its progress from before backward,
the dorsal cirrus increasing in bulk, and the superior lobe increasing in depth. At the
tenth foot the superior lobe in profile is broadly conical, the posterior flap projecting
superiorly and sloping inward inferiorly. The branchise commence about the twenty-first
foot, and continue till within twelve to fifteen segments of the tip of the tail.

Synonyms.

1807. Nereis sanguinea, Montagu. Trans. Linn. Soc, xi, p. 26, pi. 3, f. 1.
1818. Leodice „ Lamarck. Anim. sans Vert., v, p. 323 ; 2nd edit., v, p. 563.

1820. „ opalina, Savigny. Syst. des Annel., p. 51.

1828. Nereidonta sanguinea, De Blainville. Diet. Sc. Nat., Art. Vers., t. lvii, p. 477, and Atlas, pi. 15, f. 2.
,, Leodice sanguinea, Stark. Elements, ii, p. 137.

„ Fleming. Encycl. Brit., 7th edit., xi, p. 219.

„ Audouin and Edwards. Ann. Sc. nat., t. xxviii, p. 220.

,, idem. Annel. Litt. de la France, p. 147.

„ (Juvier. Regne Anim., t. hi, p. 200.

„ (circulation), H. M. Edwards. Ann. Sc. nat., 2 e ser., p. 204, pi. 12, f. 2.

sanguinea, Gruhe. Fam. der Annel., pp. 44 and 123.
„ Leidy. Proc. Acad. Nat. Sc. Philad., p. 147.

„ Grube. Insel Lussin, p. 79.

,, Johnston. Cat. Worms Brit. Mus., pp. 134 and 340.

„ Marphysa „ De Quatrefages. Annel., i, p. 332.

„ „ Leidii, idem. Ibid., i, p. 337.

1867. Eunice sanguinea, Parfitt. Trans. Devon. Assoc, ii, p. 20 (sep. copy).

1868. „ „ Ehlers. Borstenw., ii, p. 360, Taf. xvi, f. 8—11.
1870. Marphysa ,, Baird. Journ. Linn. Soc, vol. x, p. 352.

,, „ „ Grube. Arch. f. Naturges., Bd. xxxvi, p. 298.

1873. ; , Leidyi, Verrill. Invert. Viny. Sound, U. S. Comm. F. and F., p. 593.

1875. ,, sanguinea, Marion and Bohretzky. Ann. Sc. nat., 6 e ser., t. ii, p. 12.

1879. „ ,, Webster. Trans. Albany Inst., ix, p. 36.

;; „ „ idem. 32nd Report N. York State Museum, p. 115.

1885. „ „ Cams. Fauna Medit., p. 212.

1887. „ „ Ehlers. Florida Annel., p. 97.

1888. „ „ De St. Joseph. Ann. Sc. nat., 7 e ser., t. v, p. 201.

1890. „ „ Malaquin. Annel. Boulon, p. 24 (Rev. Biol. Nord. Fr., t. ii, pp. 176 and

179).
1892. „ „ (eyes), Andrews. Journ. Morph., p. 180, pi. xi, f. 34, 35, 38, etc.

1901. „ „ Fauvel. Bull. Soc. Linn. Normand., 5 e ser., vol. v, p. 61.

1903. „ „ Mcintosh. Ann. Nat. Hist., ser. 7, vol. xi, p. 559.

)>

Eunice

1833.

)>

1834.

))

1836.

))

1838.

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1851.

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1853.

))

1864.

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1865.

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444 MARPHYSA SANGUINEA.

1903. Marphysa sanguined, Mcintosh. Mar. Invert. S. Africa, vol. iii, p. 44.

1904 - „ „ Allen. Jonrn. M. B. A., n.s., vii, p. 225.

1905 - „ „ Graeffe. Arbeit. Zool. Stat. Triest., xv, p. 321.

1906 - » „ Bonn. Ann. Sc. nat., 9 e ser., t. iii, p. 101.

Habitat— Abundant at St. Peter Port, Perelle Bay, and other parts of Guernsey,
and in similar places in Herm and the Gouliot Caves, Sark (W. C. M.). Polperro, Cornwall,
on a gravelly bottom (Laughrin). Coast of Devon (Montagu).

Mediterranean. Adriatic (Graeffe). American coast (Leidy). Shores of France
(De Qnatrefages, etc.). Vinyard Sound, United States (Verrill). Virginian coast and
New Jersey (Webster). South Africa (W. C. M.). Angra Peqaena (Marenzeller).

Head deeply bilobed and somewhat flattened. It bears five smooth dull yellow
tentacles of considerable length, the median being posterior and somewhat longer than
the others. The two lateral arise obliquely, one in front of the other a little exter-
nally and in front of the median. The tentacles generally present slight wrinkles. A
single eye on each side lies behind the outer tentacle and to the exterior of the inner.
The large flattened palpi form a continuation of the head anteriorly.

Body 18 ins. to 2 ft. long, somewhat rounded in extension, flattened in contraction,
slightly tapered anteriorly and finely tapered posteriorly to the tail, which ends in two
cirri springing from the ventral edge of the anus, as in Nereis. The peristomial segment
resembles in outline that of Nereis pelagica, though the dorsal and ventral wrinkles are
characteristic. Behind the foregoing is a narrow segment without appendages, and those
which follow are also narrow. In colour it is brilliantly iridescent, with a basis of ash-
grey or greenish-brown, the iridescence being most marked anteriorly, where the duskier
hue reflects the colours more conspicuously, so as to rival the sheen of tropical beetles.
In large examples the body is darker in the middle from a slight development of pigment,
gradually becoming pale posteriorly, especially toward the tip of the tail, which is often
greenish. As in Nereis, the dorsum anteriorly is minutely dotted.

Proboscis, — The maxillee (Plate LXIII, fig. 1) have a lateral flattening of the curve,
and a strong hook in front. A flange posteriorly indents the maxillary plate, which has
three teeth in front on the left and four on the right. The crescentic anterior plate on
the right has seven or eight teeth, the corresponding one on the left having fewer (five
to six), whilst the intermediate process on the left has six or seven small teeth. Some of
the teeth in the three plates just mentioned are not distinct. The posterior spatulate plates
are dark brown, the ends of the maxillge being pointed and pale chocolate brown. The
mandibles have a dense porcellanous plate ventrally in front (Plate LXIII, fig. 1 a),
with a black touch externally. The front edge is bevelled to a blunt point. The shafts
taper posteriorly, and are marked by longitudinal lines.

The first foot has a slightly conical short dorsal cirrus which does not extend so far
outward as the upper lobe, the base of which is broad and the tip acutely conical,
whilst three black spines pass into it, with a tuft of simple (dorsal) bristles which
project above the lobe. The ventral lobe is shorter and broadly lanceolate. In the
second foot the dorsal cirrus extends beyond the superior lobe, and another tuft of
bristles (jointed) projects from that inferiorly. In the anterior feet the dorsal cirrus
is large and vascular.

MAPPHYSA SANGUINEA. 445

In its progress backward the foot undergoes considerable change, the dorsal cirrus
increasing in bulk whilst the superior lobe increases in depth and thickness and develops
a posterior flap which is prominent superiorly. At the tenth foot, for example (Plate
LXXIV, fig. 8 a), a posterior view shows ja broadly conical superior lobe, the posterior
flap projecting superiorly and sloping inward inferiorly, the tuft of simple dorsal bristles
slanting upward and outward above the spines, whilst the inferior jointed bristles project
below it. The ventral lobe is conical with a long line (margin) inferiorly, corresponding
to the bulbous border.

About the twenty-first foot, sometimes a little in front, a clavate process appears at
the inner base of the dorsal cirrus, the first indication of the branchias, which thus
pertain to the dorsal cirrus. In an example in which these organs began on the twenty-
first foot, the twenty-second had a simple but longer process. On the twenty-third the
increasing process had a short spur on its inner side. Both processes had considerably
increased in length on the twenty-fourth foot. The twenty-fifth and twenty-sixth had
each two divisions springing from a common base, and so had the twenty- seventh on the
left, but the right branchia of the twenty-seventh had a stem with three divisions (Plate
LXXIV, fig. 8). Four divisions characterized the thirty-second foot, and the processes
were longer. Five divisions soon make their appearance, and about the fiftieth foot there
are six (in the large example taken as the type), and these form a pectinate series which
project upward from a more or less horizontal stem, the longest being to the exterior, the
shortest to the interior. This condition, however, may arise from the last formed process
budding from the base of that adjoining, and becoming continuous with the oblique stem.
The basal trunk from which all these arise is stout.

Behind the foregoing the maximum number of branches is found, viz. eight, and as
the processes are long and bright red, they form a conspicuous series along the dorsum.
After attaining the maximum the branchiae gradually diminish in number, and by-and-by
in size, until in front of the tail a short process of two divisions and then of one occurs,
the last twelve to fifteen segments or thereabout being devoid of them. Probably the
condition as regards reproduction of the tail has much to do with the arrangement in
individual specimens. In sickly and dying forms the branchiae become pale.

An urn-shaped Infusorian (somewhat like Biosceca) occurred in great numbers on the
branchiae.

The dorsal bristles, which leave the foot above the spines, have anteriorly translucent
slender straight shafts, the terminal region being marked from these by a slight
curve, beyond which the tip appears to be flattened, is a little dilated, and then tapers to a
fine point. In the middle of the body the curve alluded to is more gentle (Plate
LXXXIII, figs. 5 and 5 a, and also Plate LXXVII, fig. 16), and the dilatation of the tip more
distinct. The edge is also very finely serrated. Amongst the foregoing is a group of
brush-shaped bristles (Plate LXXXIII, fig. 5 a), the distal margin being of considerable
breadth and with a series of fine teeth, the edge at each side having a longer process.
A few of these appear on the tenth foot and continue to the posterior end, where (just in
front of the tail) they increase in size and some have considerably coarser fimbriae (Plate
LXXXIII, fig. 5 b) which are also oblique.

In the inferior group of bristles the shaft is curved toward the tip, dilated and

446 MARPHYSA SANGITINEA.

bevelled, whilst it is also marked by distinct striae, which slope obliquely from the
serrated anterior edge downward and backward (Plate LXXXIII, fig. 5 c). The distal
piece is articulated in the usual manner — broad at the base and tapered to a fine point.
A few fine striae slope obliquely downward and backward from the edge.

Three black spines occur in front, but at the fiftieth foot two are found. At the
ninetieth foot only a single black spine is present, and this remains to the posterior end.
At the fiftieth foot, however, a pale brown spine passes at an angle from the black spine,
and projects at the inferior edge of the setigerous process, a little above the ventral lobe.
At the ninetieth (Plate LXV, fig. 14) a brown spine accompanies the black, and two
project inferiorly, the tips of these having wings. In the terminal segments one or
two black spines are present, and the oblique brown bristle has the two terminal wings
and is bifid at the tip.

Habits. — These animals lurk in their long galleries in cracks and in fissures of rocks
at and near low water-mark, the crevices usually being filled up with dark mud or muddy
sand and gravel, in which their tunnels are formed. They also occur under large stones
lying on a sandy bottom, the posterior end of the tunnel penetrating deeply into the sand,
so that when the stone is raised they disappear into the tube and require a considerable
amount of digging to secure them, and even then only fragments may be obtained. The
gallery in fissures of rocks is sometimes in the form of a siphon, and is frequently more
than 18 ins. long. Thus the tail as well as the head of the annelid can readily be bathed
by the advancing tide. They do not appear to care for constant immersion, and their
galleries, though sheltered, are sometimes barely moist.

When disturbed by splitting the rocks it withdraws the end of the body exposed, and
clings tenaciously to its tunnel. If an attempt be made to draw it out by the anterior
end rupture readily ensues, and however skilful the collector may be the majority of the
specimens are more or less mutilated. When both ends of the tunnel are exposed it
withdraws itself to the most distant part. In several instances at St. Peter Port,
Guernsey, and also at Polperro, Cornwall, the little commensal, Harmothoe marphysx,
McL, accompanies Marphysa in its galleries. 1

On the whole it is somewhat sluggish in its tube, but when placed freely in a vessel
of sea-water it wriggles actively, making a series of screw -like coils after the manner of
Eteone.

This annelid is used as bait by the fishermen of St. Peter Port and the Channel
Islands generally under the term " Varme," being highly prized as a tempting bait for
the capture of whiting pout, wrasses, and other ordinary fishes as well as dog-fishes. The
men search for them with spades at low water, and they are plentiful on an oyster-bed
to the north of the pier at St. Peter Port. If the tube passes straight into the muddy
sand few entire examples are procured, but if a bed of rock lies beneath seven or eight
inches of sand they are often captured entire. Three or four inches of the anterior end
of the worms are placed in vessels amongst a little black or decaying " wrack," and they
can thus be kept about a fortnight. If the rest of the body were retained, the worms
would quickly break into fragments and putrefy.

A large example was conveyed from Herm to Norwich in August, but the great
1 ' Monogr. Brit. Aimel./ vol. i, p. 339.

MARPHYSA SANGUINE A. 447

heat and close quarters in that town proved fatal. Others, however, have been sent
alive from Jersey to St. Andrews, and survived a considerable time. Although fragments
of shells have been found near their galleries, it is probable these were due to other
animals. The species seems to be phytivorous.

This is one of the many additions made to the British Fauna by the acute and
indefatigable Colonel Montagu (1807), who searched the tidal region of the South
Devonshire coast with such perseverance and success, and whose figures by his niece,
Miss Dorville, are so useful to his successors. He observes that it is the largest
" Nereis " yet discovered on the British shores, extending sometimes to fifteen inches,
and frequenting rocky situations. The much larger Nereis virens, Sars, had not then
been described.

This does not appear to be the Eunice sanguined of Delle Chiaje, 1 either in regard
to structure or coloration, though tentacular cirri are also absent in it.

Milne Edwards, in one of his early papers, 2 gives a general account of the circulation
of this form, and compares it with the condition in Terebella.

A paper was published by H. Koch 3 on the so-called development (viviparous) of
Marphysa sanguinea, but the figures and descriptions show that the author had before
him a parasitic Lumbriconereid such as Oligobranchus, or perhaps, as Ehlers states,
Nematodes.

This form is prominently used by De Quatrefages (1865) in his description of the
anatomy of annelids — for he considered that its structure was both complicated and
superior to that of most of the Errant Annelids. Moreover, he credited it with a single
otocyst in the buccal segment, but Fauvel, 4 who has not actually found otocysts in any of
the Eunicidse, thinks a misapprehension had occurred. De Quatrefages separated the
American examples under the name Marphysa Leidii, but this appears to be unnecessary.
He also observed (1844) that an example appeared, from its coloration, to have
regenerated the head and adjoining segments.

Ehlers (1868) had only an incomplete spirit-specimen for description, but his
account is on the whole good, though he misinterpreted the condition of the segmental
organs. He pointed out Koch's error in supposing that this species was viviparous, the
appearances having been due to parasitic Nematodes.

Marion and Bobretzky (1875) observe that the fishermen of Marseilles employ this
form for bait under the name of Moureda, and they mention the occurrence of examples
with simple branchia3 throughout.

Webster found Marphysa sanguinea in great abundance on the Virginian coast and at
various stages from 2*5 mm. to 20 cm. in length. The youngest had eighteen to twenty-
two feet, no tentacles, no branchise, and no indication of frontal lobes. It had bidentate
setee in the anterior segments. There are five eye-specks. In the next stage (2*5 to 4 mm.)
one tentacle is present, bidentate seta3 have disappeared from two or three of the anterior
segments, and there is a slight depression of the anterior margin of the head. When

1 < Descrizione/ iii, p. 104, etc., Tav. 103, f. 9, 10, etc.

2 < Ann. Sc. nat./ 2 e ser., t. x, p. 204, 1838.

3 'Noveaux Mem. de la Soc. helvetique/ viii, p. 1, Taf. i and ii, 1845.

4 ' Ann. Sc. nat./ 9 e ser., t. vi, pp. 4—7, 1907.

448 MARPHYSA BELLI.

4 mm. long, simple branchiae appear on the tenth setigerous segment. In the larger
examples (10 to 20 mm.) the eyes are always two, and the branchiae commence on any
segment between the tenth and twenty-third (as in the adult), and they are usually
bifurcate. The dental apparatus of these, according to the author, agrees with that in
the adult. The brush-shaped bristles do not occur in the youngest forms, or until three
tentacles and branchiae are present. If the author is accurate in his diagnosis, the
younger forms resemble Nematonereis. The same author mentions that in those from
New Jersey the branchiae commence on the tenth foot — a considerable variation from the
usual condition in Britain.

Cosmovici l describes the segmental organs of this species as consisting of a
ciliated funnel in the dissepiment in front, and a tube leading externally at the base
of the foot.

Mr. Borradaile 2 gives an interesting account of the early stages in the development
of a Ceylonese Marphysa. The yellowish eggs are in an oval mass of mucus with a stalk,
and he found that the larvae, after escaping from this, afterwards secreted another mucus
investment in which they lived gregariously.

Benham states that it is the " rock-worm " of the Channel Islands.

Grravier 3 (1909) describes an instance of reproduction of head and anterior segments
and tail in an example of this species from St. Vaast-la-Hougue. The tapered anterior
end is conspicuous.

2. Marphysa Belia, Audouin and Edwards, 1833. Plate LV, fig. 5 ; Plate LV, fig. 6 — foot ;
Plate LX, fig. 12— head; Plate LXIII, figs. 2 and 2 a— teeth ; Plate LXV, figs.
11 and 11 a— teeth; Plate LXXXVI, figs. 3 and 3 d— bristles.

Specific Characters.— Head bluntly conical, with a median groove inferiorly, but
evenly rounded in front, only a slight notch appearing in spirit. Tentacles proportionally
long and slightly crenate. Body 6 — 7 ins. long, and attenuated for a Eunicid. It is
somewhat reddish anteriorly, fawn coloured posteriorly, with a dark purple central streak
from the blood-vessel. It is highly iridescent throughout. Peristomial segment twice
the breadth of the others, that following being about the average breadth. Proboscis,
armed with maxillae which rise little above the horizontal, are blackish-brown anteriorly
and greyish or pale chocolate at the base. Basal processes foot-shaped with a terminal
point. Great dental plates have seven teeth, the azygos plate on the left has seven or
eight teeth, the curved plate in front about six teeth. The right anterior curved plate
has seven to eight teeth. The mandibles have black tapering shafts, and the cutting
anterior edo*es are sinuous. Branchiae commence on the fourteenth foot and continue to
the thirty-fourth or a little further, and thus range from twenty to twenty-five in number.
The central stems have most filaments, the first having about ten, the twenty-eighth

1 ' Thesis/ p. 114, pi. xxvi, fig. 7.

3 < Proc. Zool. Soc./ 1901, vol. ii, p. 714, pi. xxxix.

3 ( Ann. Sc. nat./ 9 e ser., t. ix, p. 150, 1 text-fig.

MAPPHYSA BELLI. 449

nineteen divisions, and the last about thirteen. Foot with a short dorsal cirrus in front,
more elongated in the branchial region and posteriorly, a short bluntly conical setigerous
region with two or three blackish-brown spines in front, and bristles of three kinds, viz.,
simple tapering dorsal bristles, compound falcate bristles with narrow terminal pieces
which are bifid and winged, and thirdly, a few compound forms with tapering terminal
pieces as in Marphysa sanguinea. This type of foot extends to the end of the branchial
region. The posterior foot has also the three kinds of bristles — considerably diminished
posteriorly — but has, in addition to the black spine, one or two long, bifid winged hooks
(modifications of spines) which slightly slant downward.

Synonyms.

1833. Eunice Bellii, Audouin and Edwards. Ann. Sc. nat., l e ser., t. xxvii, pi. ii, f. 1 — 4, 8,9 ; t. xxviii,

p. 223.

1834. „ „ idem. Annel., p. 149, pi. iii, f. 1—4, 8, and 9.

1836 — 7. Eunice Belli, idem. Cuvier, Regne Anim:, 2 e eel., iii, p. 200 (note).

1851. Eunice Bellii, G-rube. Fam. Annel., pp. 44, 123.

1865. Marphysa Belli, De Quatrefages. AnneL, i, p. 333.

1869. „ „ G-rube. Mittli. St. Vaast-la-Hongue, p. 36.

1874. „ Bellii, Marenzeller. Adriat. Ann el., p. 59.

1885. „ ,, Cams. Fauna Medit., p. 213.

1887. „ „ Elders. Florida Annel., p. 95, Taf. xxviii, f. 1—8.

1902. „ „ Marenzeller. Potychset. des Grundes (Kaiserlichen Akad.), p. 17.

1903. „ „ Mcintosh. Ann. Nat. Hist., ser. 7, vol. xi, p. 560.

1904. „ „ Allen. Journ. M. B. A., N.s., vii, p. 225.

1905. „ „ Graeife. Arbeit. Zool. Stat. Triest, xv, p. 321.

Habitat. — Under a stone near low water-mark of ordinary tide near the boatman's
house, Herm (1868). Plymouth (Allen).
/ . Shores of France. Mediterranean. American shores (Ehlers).

Head (Plate LX, fig. 12) bluntly conical, evenly rounded in front, with scarcely a
trace of a notch dorsally, but with a well-marked ventral median groove— probably
indicating the palpi. The median and two lateral tentacles are proportionally long and
slightly crenate, the median exceeding the others in length. A small eye lies at the base
of the inner lateral tentacle — externally. The mouth presents a median frill at the
posterior end of the groove of the snout and a prominent and rounded posterior lip. The
peristomial segment is twice the breadth of the others, the footless segment succeeding it
being of the normal breadth. At its ventral edge it is somewhat coarsely crenate.

Body 6 — 7 ins. long — attenuated for a Bunicid, so that at first sight it resembles
Lumbriconereis ; slightly tapered towards the head, and ending in a slender tail. It is
somewhat reddish anteriorly and highly iridescent throughout. Behind the branchiae is a
dark purple central streak from the blood-vessel, the rest being iridescent fawn-colour.

The maxillae (Plate LXV, fig. 11) are greyish or pale chocolate at the base, the
curved fang anteriorly being blackish brown. A dark line separates them from the basal
processes, which are somewhat elongate, the outer border sloping inward, then forming an
elevation, and lastly curving to a slipper-shaped tip. The tips and inner edges are dark,

114 ■

450 MARPHYSA BELLI.

whilst a greyish area occurs anteriorly next the latter. A small process projects from
each posteriorly. The great dental plates have seven teeth ; the azygos plate on the left
has seven or eight teeth ; the curved plate in front about six teeth ; the right anterior
curved plate seven or eight teeth. The mandibles (Plate LXV, fig. 11 a) "have black
tapering shafts and the cutting anterior edges are sinuous.

The branchiae commence on the fourteenth foot as a process of ten divisions, all of
considerable length, and arising from the outer border of the main stem, the tip of which
makes the tenth filament. Sixteen divisions occur on the twentieth foot, nineteen on the
twenty-eighth, and thirteen on the last. The filaments are therefore much more numerous
than indicated by De Quatrefages, provided the forms are the same, for he gives six to
eight as the number of divisions. The total number of branchiae on each side varies from
twenty to twenty-five. When the body is stretched the beautiful red branchiae glide
alternately, each passing to the opposite side, the right being anterior. In contraction
they form a dense group.

The first foot has a short dorsal cirrus, shaped somewhat like an awl-handle that is,

constricted at the base, dilating, and again tapering towards the tip. The setigerous
region is short and bluntly conical, and is supported by a single blackish spine, or by
two or three. It carries four kinds of bristles, viz., dorsally simple tapering bristles,
with faint traces of spikes on the edge, and a few brush-shaped forms with nearly equal
lateral filaments (Plate LXXXVI, fig. 3*6), then a mixed group of jointed bristles, the
majority having slightly curved shafts, dilated and bevelled at the tip, to which a rather
narrow bifid terminal piece with wings is articulated, and lastly one or two of the type
of Marphysa sanguinea with similar shafts, but with long tapering terminal pieces ending
in a fine point. Behind the setigerous lobe is a slightly flattened conical flap, whilst
inferiorly is a somewhat thick and broadly conical ventral cirrus.

The changes which the foot undergoes to the thirteenth consist chiefly in the increase
of its various parts. Thus at the eighth foot (Plate LV, fig. 6) the dorsal cirrus is
more elongate, the setigerous and the posterior lobes are more prominent, the bristles
longer, and the ventral cirrus is longer. These features are still more pronounced at the
thirteenth foot, in which the increase in size is considerable. The dorsal cirrus is now
long and tapering, the setigerous region of greater depth, the posterior lobe broader
(deeper) at the base, and more tapered at the tip, whilst the ventral cirrus or lobe has the
shape of a broadly conical flap. The setigerous region is supported by three spines, and
bears the elongated slender simple bristles superiorly (Plate LXXXVI, fig. 3), along with
certain brush-shaped forms (Plate LXXXVI, fig. 3 b) which have ]ong lateral filaments.
Inferiorly are the compound falcate bristles (Plate LXXXVI, fig. 3 a), which are charac-
terized by the slight curvature of the shaft and the dilatation of the end, which is spinous
at the edges, and by the comparative narrowness of the terminal process, and lastly, those
having similar shafts, but with long tapering terminal processes (Plate LXXXVI, fig. 3 c)
after the manner of those of Marphysa sanguinea. The posterior lamella of the setigerous
region is somewhat lanceolate in front. The ventral cirrus is rather short, massive, and
conical.

The addition of the branchise is the only noteworthy change in the feet of the
branchial region, two or three spines being present in the setigerous process. The post-

MARPHYSA K1NBERGI. 451

branchial feet, again, differ in so far as the lower of the two spines separates from the
upper and is joined by a second, and their tips are differentiated into bifid winged hooks.
Thus at the fortieth foot the main spine is blackish brown, and both the tips and the
concavity are directed upward. A pale spine considerably less lies above it. The two
bifid hooks have their convexity directed upward, and their tips project a short distance
below the spine. The older' is usually dark brown, and the younger pale brown, the tips
in both being pale, bifid, and winged (Plate LXXXVI, fig. Bd). The four kinds of
bristles are still present, though their number appears to be fewer. This type of foot
continues to the posterior end, as a rule one strong blackish brown spine and long bifid
hook guarded by a wing being found in each setigerous process, with the three kinds of
bristles — all, however, diminished in size. Both the posterior lamella and the ventral
cirrus are more pointed posteriorly.

Habits. — In confinement in a vessel it keeps its body in screw coils amongst sand.

This is one of the many forms which the able authors 1 of the ' Annelids of the
Coast of France ' have added to the literature of the subject, and their description so far
as it goes is good.

De Quatrefages 2 considered that he had found another species allied to this form at
Chaussey, but the description does not appear to distinguish it from the earlier one.
He omits notice of this form, moreover, in the ' Anneles.' It may be, however, that
the author thought that the description of Audouin and Edwards did not apply to a
species with three kinds of bristles, besides the brush-shaped ones, in the anterior
feet. He afterwards, perhaps, considered that the differences were due to defective
descriptions.

Ehlers describes the branchias of the American forms as occurring on the fifteenth
foot, and being fourteen pairs in number, whereas Marenzeller notes in the Adriatic
forms that they commence on the twelfth, and that their number is from fifteen to
twenty-five. Considerable variation is thus present.

3. Marphysa Kineeegj, 3 ii. s. P]ate LXXIV, figs. 9 and 9 a — feet and branchise ;

Plate LXXXIII, figs. 6-6 b— bristles.

Specific Characters. — Head bluntly conical, without a trace of a notch dorsally in
spirit ; ventrally a well-marked median groove ; median and lateral tentacles of con-
siderable length, the former the longer ; no eyes visible in the preparations. Body
small, scarcely f in. in length, more massive in the preparation anteriorly than posteriorly.
It is somewhat flattened anteriorly, rounded posteriorly, but is incomplete. The maxillas
are chocolate brown, with darker blades, which do not rise much above the horizontal.
Posterior processes — to which they are articulated — are lobate. Great dental plates have
six teeth, the anterior being long and recurved, and a considerable portion posteriorly being
smooth. Azygos plate with seven teeth. Anterior curved plate on the right has six

1 Audouin and Edwards, ' Anne]./ p. 149, pi. hi, figs. 1-4 and 8—9, 1834.

3 ' Magasin de Zoologie/ An. 1843, p. 4, pi. ii, figs. 1, 2.

3 Named after the late Prof. Kinherg, of Stockholm, who did much to advance our koowledge of
the Marine Annelids.

452 MARPHYSA KINBERGt.

teeth. Mandible with the oblique anterior edge smooth, the outer angle, however, being
prominent. The branchise form an isolated tuft, commencing about the sixteenth
and continuing to the thirty-sixth foot. The anterior gills have from five to nine
filaments, those best developed twenty-four or twenty-five. The foot has a short subu-
late dorsal cirrus in front, but it becomes longer in its progress backward. The setigerous
region bears longer dorsal bristles than in Marphysa Belli, and brush-shaped forms, whilst
the inferior group is composed only of those with long tapering tips, as in Marphysa
sanguinea. Posteriorly the dorsal bristles are longer, and the long bifid brown ventral
hook projects inferiorly.

Habitat. — Dredged in the ' Porcupine ' Expedition of 1870 nine miles off Cape Finis -
terre in eighty-one fathoms on a hard bottom with sandy mud. Though not procured in
what are usually considered British waters, it has been thought proper to include this
species here.

Proboscis. — The maxillse are pale chocolate brown with darker blades which do not
rise much above the horizontal. They are articulated posteriorly to two rather long
lobate processes which are somewhat narrowed at the commencement. The great dental
plates have six teeth, the anterior being long and recurved, and a considerable posterior
region being smooth. The azygos plate on the left has about seven teeth. The anterior
plate on the right has six teeth ; that on the left has fewer. A dark horny patch also
occurs in front on each side. The mandibles have the oblique anterior edge smooth, the
outer angle, however, being prominent.

The branchias commence on the sixteenth foot, thus differing from Marphysa Belli
from the Channel Islands. The initial filament occurs only on one side (the right). The
seventeenth on. the right has a considerable gill of nine filaments ; that on the left five or
six. They are present in all on twenty segments, the last having fewer filaments. The
whole forms a somewhat isolated and dense tuft, some of the best developed (Plate
LXXIV, fig. 9 a) having twenty-four or twenty-five filaments which are longest in the
middle, and arise as usual from the side of the branchial stem.

At the tenth foot (Plate LXXIV, fig. 9) the dorsal cirrus is a proportionally large
subulate process — somewhat constricted at the base. The setigerous region has a bluntly
conical outline, with a conical lamella projecting beyond it. It is supported by three
black spines. Dorsally it bears a dense group of translucent simple bristles, the tips of
which have a very slight expansion and then taper to a fine point (Plate LXXXIII,
fig. 6). Ventrally are the even more numerous jointed bristles (Plate LXXXIII, fig.
6 a), which have translucent shafts with dilated distal ends showing fine striaa, and
serrations on the edge. The terminal piece forms a long tapering spear which diminishes
m length from the upper bristles downward. Moreover, when fracture occurs near the
attachment of the distal piece a portion remains at the socket, a condition not usually
seen in such forms, and indicating a differentiation of the point of attachment (Plate
LXXXIII, fig. 6 b). Each foot has a rete mirabile of branching blood-vessels.

The closeness in external resemblance between this species and M. Belli, Aud.
and Edw., is interesting — associated as it is with the divergent structure of the
bristles.

NEMATONEBEIS UNICORNIS. 453

Genus LXXIL— Nematonerims, Schmarda, 1861.

Head smoothly rounded, with a single median tentacle. Body elongated and
distinctly segmented ; two segments devoid of feet. Upper dental apparatus with an
uneven number of plates. The body- wall of Nematonereis unicornis is thick and muscular,
after the type of the Eunicicla3 and Lumbriconereidse. The nerve-area is pyriform with
the narrow end inferiorly — as a short pedicle. The area has a thick coating of cells which
stain deeply. A small neural canal exists in the centre superiorly. The oblique muscles
are attached to the upper and outer border, and some fibres pass down to the apex of the
nerve-area. The hypoderm forms a thicker layer in the centre inferiorly, and the fibres
of the circular coat curve inward along its inner border and apparently cross below the
nerve-area. On the inner aspect of the area in certain parts is a coarsely granular
brownish layer which does not stain well, and is probably fatty. Foot with dorsal and
ventral cirri; simple bristles, with a few brush-shaped forms, dorsally; jointed hooks
inferiorly.

Ehlers (1868), characterizes the genus as having a single tentacle ; two segments
devoid of feet. Foot with dorsal and ventral cirri, simple and compound bristles.
Upper jaws with an uneven number of jaw- plates.

1. Nematonereis unicornis, Grube, 1840. Plate L1V, fig. 3; Plate LXIII, figs. 5 and
ha— teeth; Plate LXXV, figs. 3 and 3 a— feet ; Plate LXXXIV, figs. 1-1 c—
bristles.

Specific Characters. — Head smoothly rounded in front ; two large black eyes at its
posterior border and between them a subulate median tentacle. Body five to six inches
long, pale pinkish and iridescent in front, posteriorly cream-coloured with a pale
orange streak from the intestine. The anterior of the first two segments is the broader.
The body terminates posteriorly in four (Ehlers) anal cirri. Maxillse short and
strongly hooked anteriorly, their posterior appendages of an elongated conical outline.
Anterior plates have a triangular brown patch in front. Great dental plates have four
or five large teeth. 1 Mandibles hatchet-like, the expanded anterior blade with parallel
lines. The vascular anterior feet {e.g., tenth) have a subulate dorsal cirrus. The
setigerous region has a single brown spine ; above it two groups of winged bristles with
attenuate tips. Below the spine are jointed hooks which have the dilated end of the
shaft minutely serrate. Ventral cirrus pear-shaped. Behind the foregoing the dorsal
cirrus by-and-by becomes more slender and has a distinct granular enlargement at its
base. A single group of winged bristles with attenuate tips and a few brush-shaped
forms occur above the spine, whilst the jointed bristles beneath are stronger. The whole
setigerous region is proportionally less, but it is more prominent. A single powerful
hook projects above the ventral cirrus from the twentieth foot backward.

Synonyms.
1840. Lumbriconereis unicornis, Grube. Actin. Ecliin. u. Wiirm., p. 80.

1843. „ pectinifera, De Quatrefages. Mag. de Zool., 2 e ser., y, p. 6, pi. ii, f: 3 — 8.

1861. Nematonereis unicornis, Schmarda. Neue Wirb. Thiere, ii, p. 119, pi. xxxii, f. 254.

1 Unfortunately the only example available for illustration was injured (Plate LXIII, fig. 5).

454 NEMATONEREIS UNICORNIS.

1864. Lumbriconereis unicornis, Claparede. Glanures Zoot., p. 112, pi. iv, f. 2.

1865. Nematonereis pectinifera, De Quatrefages. Hist. Nat. des AnneL, p. 372.
» j, Grubei, idem. Ibid., p. 373.

contorta, idem. Ibid., Explic. des Planches, p. 13, pi. viii, f. 24—27.

,, Lumbriconereis unicornis, idem. Ibid., ii, p. 652.
1868. „ oculata. Elders. Borstenw., p. 374, Taf. xvi, f. 19—22.

„ Nematonereis unicornis, Claparede. Annel. Nap., p. 151.
1875. ,, ; , Marion and Bobretzky. Ann. Sc. nat., 6 e ser., t. ii, p. 15.

1879. ,,■ v Langerhans. Zeitsclir. f. wiss. Zool., Bd. xxxiii, p. 295.

1885. „ „ Cams. Fauna Medit., p. 214.

1903. ., „ Mcintosh. Ann. Nat. Hist., ser. 7, vol. xi, p. 560.

1004. „ „ Allen. Journ. M. B. A., n.s., vii, p. 225.

1905. „ „ Willey. Rep. Ceylon Pearl F., iv, p. 284.

,, ,, oculata, Graeffe. Arbeit. Zool. Stat. Triest, xv, p. 321.

1906. , „ De St. Joseph. Ann. Sc. nat., 9 e ser., t. iii, p. 213.

Habitat, — Amongst muddy sand in fissures of gneiss between tide-marks, St. Peter
Port, Guernsey. Plymouth (Allen).

Ceylon (Willey). Off Pernambuco. Cape Verde Islands. Sea Point, Cape Town,
probably. Canaries and Madeira (Langerhans).

Head evenly rounded in front, forming a short blunt cone. At its posterior border
are two large black eyes, and between them a subulate median tentacle which is less than
the antero -posterior diameter of the head.

Body five to six inches or more long, pale pinkish iridescent anteriorly with a red line
from the dorsal vessel, and a central streak from the intestine, whilst this posteriorly is
pale orange from its glands, the rest being pale cream-coloured. The annelid is nearly
cylindrical from end to end.

Proboscis (Plate LXIII, fig. 5). — The maxillae are short and strongly hooked anteriorly
— like the upper half of the letter S. At the posterior end a dark brown band separates
them from the posterior appendages which have an elongated conical outline. The
anterior plates have a triangular brown patch in front. The great dental plates have a
series of comparatively large teeth four or five in number. The mandibles (Plate LXIII,
fig. 5 a) are hatchet-like, and the expanded anterior blade is marked by parallel lines.

The first two segments are devoid of feet, the anterior being somewhat broader than the
posterior. The first foot is small and has a short, subulate, almost conical dorsal cirrus, and
an ovate, massive ventral cirrus. The setigerous region is little developed, being less than
the ventral cirrus, and has a single dark brown spine. The dorsal bristles have tapering
winged tips. Below the spine is a group of jointed hooks, the distal process being winged
and bifid at the tip — the whole differing little from the same structures in the tenth foot.

At the tenth foot (Plate LXXV, fig. 3) the dorsal cirrus has considerably elongated
— forming a subulate organ. It is now separated by a considerable interval from the
deep, truncated setigerous region, bearing dorsally a group of four winged bristles
(Plate LXXXIV, fig. 1), which have the tips dilated above the shaft and then rather
abruptly narrowed to finely tapered and long processes. The wings are minutely
striated. Below there is a group of smaller winged bristles with finely attenuate tips.
Appearances would seem to indicate that the setigerous lobe has a flap.

NEMATONEREIS UNICORNIS. 455

The foregoing are above the single strong brown spine. Below the spine are the
jointed hooks which have the dilated end of the shaft minutely serrate (Plate L XXXIV,
fig. 1 b). The ventral cirrus is now pear-shaped from the outward growth of a conical
process. The vascularity of the foot is great.

At the twentieth foot (Plate LXXV, fig. 3 a) a single powerful brown spine supports
the setigerous region, which has similar bristles and hooks. A single large hook crosses
the spine obliquely from above and projects over the ventral cirrus. It has a broad
wing at the tip, and the main fang is large (Plate LXXXIV, fig. la). In the foot,
however, the secondary process above the main fang is not distinct. The dorsal cirrus
is more slender.

At the fiftieth foot the dorsal cirrus is even more slender than in front, and has a
distinct granular (glandular) enlargement at its ventral base. A single group of winged
bristles emerges above the spine, their wings are narrower, curves less marked, and tips
very attenuate. Below the spine the jointed bristles have increased in strength, the distal
ends of the shafts are striated, and a few serrations occur on the edge of the wings
guarding the terminal piece. The whole region is less and its outline more convex (less
flattened). The powerful hook projects above the ventral cirrus— which extends beyond
the massive glandular pad as a blunt cone. In this foot a brush-shaped bristle (or two)
occurs amongst the dorsal bristles (Plate LXXXIV, fig. I c).

Proceeding backward the dorsal cirrus becomes less, but the glandular swelling
remains at its ventral base. The setigerous region forms a broad cone with the tip of the
powerful brown spine at the apex. Dorsally is a small group of the two kinds of winged
bristles (larger uppermost, smaller below), and one or two brush-shaped. The jointed
hooks project further outward (are longer). The great hook has increased in strength,
and extends as far as the tip of the ventral cirrus, which is elongated and somewhat
lanceolate, whilst the glandular swelling has moved inward and is less.

Reproduction. — Claparede, in his ' Grlanures ' (1864), observes that in the females ova
were only developed behind the sixty-second segment. The young Nematonereis has four
eyes, but two disappear, and Grravier thinks that the maxillary apparatus of this and
Lysidice resemble each other. De St. Joseph found the minute young pelagic.

Habits. — It is fond of coiling itself in the vessel like a corkscrew, but this is probably
connected with its surroundings.

This was one of the fruits of Prof. Ed. Grube's researches on the fauna of the
Adriatic and Mediterranean, published in 1840.

Ehlers (1868) gives the species four eyes.

The Nematonereis Sclimavdse of the ' Challenger,' l from the south of Pernambuco,
is a closely allied form, but does not appear to be identical, and like the British form
lives in mud (red). The same form would appear to range to St. Vincent, Cape Verde
Islands, and to Cape Town, at both of which places fragments were obtained by the
' Challenger.' Schmarda found it in the Atlantic, and Willey in Ceylon.

It would seem that the three species mentioned by De Quatrefages (1865) may be
varieties of the same form.

1 ( Annelida/ p. 262, pi. xxxvii, figs. 6 — 8; pi. xviiiA, figs. 16 — 19.

456 LYSIDICE PUNCTATA.

Grube (1870), who examined the specimens in the Parisian Museum, showed that the
Blainvillea of De Quatrefages was a Nematonereis.

Nematonbebis sp. Plate LV, figs. 7 and 8— head and tail ; Plate LVI, fig. 6— body.

A form sent by Dr. Allen from Plymouth differs considerably in outline from the
foregoing, and is much less rigid, yet the general structure appears to agree. The loss
of the example by the artist after delineation prevents a satisfactory decision. This form
is considerably paler than that from the Channel Islands, has about seventy segments, and
the tail (Plate LV, fig. 8) has two shorter and two longer cirri. The head (Plate LV,
fig. 7) agrees with V. unicornis, and the dental armature is similar, but the feet are some-
what longer and more conspicuous and the spines are dark. The colour anteriorly varies
from pinkish to pale green, then the intestine gives a brownish hue to the central region
whilst the posterior region is brownish-yellow.

KinbergV third family of the Eunicea is the Lysidicea, distinguished by seven
maxillas, part Ii edentate; three tentacles, neither antennge nor palpi; buccal segment
double. Of the genera — Amphiro and Lysidice— the former is distinguished by the
presence of pectiniform branchiaa, whilst the latter has none.

Genus LXXIII. — Lysidice, Savigny, 1820.

Head free from the buccal segment, bearing three tentacles and two eyes ; two large
palpi.

Feet with cirri, bristles, and spines.

1. Lysidice punctata, Bisso. Plate LV, fig. 3; Plate LXIII, figs. 6 and 6 a — teeth; 2
Plate LXXV, figs. 4-4 b— feet; Plate LXXXIV, figs. 2-2 a\ and 9— bristles.

Specific Characters. — Head broad and flattened, with a median notch anteriorly,
while inferiorly are two thick prominent palpi. Median tentacle is short and somewhat
fusiform, tapering distally, and situated rather in front of the lateral. Eyes two,
black, of considerable size. Colour pale brownish, dappled with pale dots. Body 6 — 7 ins.
long, tapering a little anteriorly, but more distinctly so posteriorly, where it terminates in
four caudal processes. The first three segments are of a deeper brown hue and dappled
with pale spots. First and second devoid of feet. The fourth segment is pale or whitish
anteriorly, and in some the whole segment is white (Ehlers). The brownish colour with
pale dots continues to the twentieth or twenty-fifth segment, and then fades into the
paler brownish iridescence of the succeeding region. Segments in the middle and
posterior part of the body boldly marked.

Proboscis has strong, curved maxillas, with a shoulder posteriorly and long posterior
appendages with a notch in the middle. Great dental plates have four prominent teeth
on each side anteriorly — the anterolateral plates. The mandibles are proportionally

i ' Ofvers. af K. Vet.-Akad. Fork/ 1864, No. 10, p. 565.
2 As L. Ninetta,

LYSIDICE PUNCTATA. 457

large and deeply concave from side to side, so that about half the length of the upper
jaw-apparatus lies in the hollow curved anterior region. A longitudinal black bar on
the upper surface. Foot with a dorsal cirrus slightly constricted at the base, and,
in spirit, not extending beyond the tips of the bristles. Setigerous region a flattened
cone with two strong yellowish spines (tenth foot). The dorsal bristles are curved at
the end of the shaft, and the winged tip tapers to a fine point. Below the spines are
jointed bristles, dilated at the end of the shaft and serrated on the convex edge. Terminal
piece short, with a bifid hook and two wings. In the posterior feet a strong hook passes
obliquely downward and pierces the skin below the compound bristles.

Synonyms.
1826. Leodiee punctata, Risso. Hist. Nat. de FEurope, t. iv, p. 421.
,, „ triantennata , idem. Ibid., p. 422.

1833. Lysidice Ninetta, Audouin and Edwards. Ann. Sc. nat., t. xxvii, pi. xii, f. 1 — 8 ; t. xxviii, p. 235.

1834. „ „ idem. Hist. Annel., p. 161, pi. no, f. 1—8.

1836-7. „ „ M. Edwards. Cuvier's Regne Anim. Illust., pi. xi, f. 1 and 1 a.

1844. ,, torquata, De Quatrefages. Ann. Sc. nat., 3 e ser., t. xiv, p. 361.

1851. „ Ninetta, Grube. Fam. Ann el., pp. 45 and 124.

1853. ,, rufa, Gosse. Ann. Nat. Hist., ser. 2, vol. xii, p. 385.

1855. „ punctata, Grube. Arch. f. Naturges., Jalir. 21, p. 95.

1863. ,, Ninetta, Keferstein. Zeitschr. f. wiss. Zool., Bd. xii, p. 101, Taf. ix, f. 10—16.

1864. ,, punctata, Grube. Insel Lussin, p. 79.

„ „ Mahagoni, Claparede. Glanures Zoot., p. 116, pi. ii, f. 4.

1865. „ Ninetta, De Quatrefages. Annel., i, p. 375.

„ „ „ Johnston. Cat. Worms Brit. Mus., pp. 140 and 341.

„ ,, rufa, idem. Ibid., pp. 141 and 341.

„ „ torquata, De Quatrefages. Annel., i, p. 376, pi. ix, f. 19, 20.

„ Leodiee triantennata, idem. Ibid., p. 354.

1867. Lysidice Ninetta and rufa, Parfitt. Trans. Devon. Assoc, ii, p. 20 (sep. copy).

1868. „ „ Elders. Borstenw., ii, p. 366, Taf. xvi, f. 12—18.

„ „ punctata, Grube. Mitth. St. Vaast-la-Hougue, etc., p. 36 (126).

, idem. Arch. f. Naturges., Bd. xxxvi, p. 301.
Ninetta, Marion and Bobretzky. Ann. Sc. nat., 6 e ser., t. ii, p. 15.

Grube. Schies. Gesellsch., 20th June, 1877, p. 25 (sep. abdr.).

Langerhans. Zeitschr. f. wiss. Zool., Bd. xxxiii, p. 295.

idem. Canar. Annel. in Nova Acta Leop.-Car., xlii, p. 112.

Cams. Fauna Medit., p. 213.

Mcintosh. Ann. Nat. Hist., ser. 7, vol. xi, p. 560.

Allen. Journ. M. B. A., n.s., vii, p. 225.

Graeffe. Arbeit. Zool. Stat. Triest, xv, p. 321.

Bohn. Ann. Sc. nat., 9 e ser., t. iii, p. 101.

Habitat. — Abundant in fissures of gneiss in Guernsey in muddy sand. If the rock is
easily split, from the presence of an old crack, the greater is the probability of obtaining
large specimens. A fragment also occurred in the Grouliot Caves, Sark. At Herm small
specimens are frequently obtained along with Polydora, Dodecaceria, and other Annelids,
by breaking off the coating of Lithothamnion on stones between tide-marks. Whether it
merely occupies the holes made by other forms or whether it enlarges these is an open

115

1870.

„ torqv

1875.

„ Nine

1878.

)) ).

1879.

)) >

1881.

>> >

1885.

>> >.

1903.

)) );

1904.

)) )

1905.

>) }.

1906.

)) ).

458 LYSIDICB PUNCTATA.

question. It also occurs under the large coarse Ascidians, the test of which forms the
arch of the tunnel, the stone forming the base. It probably enlarges its tunnels. It also
was found in Gellepora dredged off St. Peter Port, and amongst the roots of Antenmilaria
and sponges (Col. Fraser and Capt. Powell) off the south coast of Sark. Isle of Wight,
Luccomb Chine (W. C. M.). Weymouth (P. H. Gosse). Plymouth (Baird and Allen).
It extends to Ireland, where it has been procured at Connemara (A. Gr. Moore) and Gal way
(Prof. B. P. Wright).

Shores of France. Keferstein found them in tubes of mucus and mud near low
water-mark at St. Vaast; Mediterranean; Canaries (Langerhans).

Head broad and flattened, with a median notch anteriorly, while inferiorly are two
thick prominent lobes (palpi). Eyes two, black, of considerable size. The median
tentacle is a short, somewhat fusiform process, constricted at the base and again
diminished toward the tip. It is placed somewhat in front of the two lateral tentacles,
which have a similar shape. The colour of the head is pale brownish dappled with pale
dots, the nuchal border being rather darker.

Body elongated, reaching 6 — 7 ins., slightly tapering anteriorly and more decidedly so
posteriorly, where it terminates in four caudal processes — two longer dorsal and two shorter
ventral cirri, the former being also wider at the tips. The dorsal arch is rounded, the
ventral surface flattened, with a median groove, and uniformly tinted.

The first three segments are tinted of a deeper brown hue than the rest of the body
and dappled with pale spots. The first is broader than the two succeeding. The first
and second are devoid of feet, as in Notocirrus. The fourth segment is pale anteriorly,
and thus makes a contrast with the preceding. In some the posterior border of this
segment is brownish, speckled with pale dots. In others the fourth segment is white, as
also is the half of the succeeding (Ehlers). The coloration indicated (brownish with pale
dots) stretches to the twentieth or twenty-fifth segment, and fades into the general brownish
iridescence of the body, with a slight shade of purple. A specimen trawled off the south
coast of Sark by Col. Fraser and Capt. Powell had only a white patch between the eyes,
instead of the usual white belt on the fifth segment.

The segments in the middle and posterior part of the body are boldly outlined, while
the feet are less differentiated, the bristles being borne on the prominent edge of the
segment, which is also marked by a transverse median riclge. Posteriorly the body
terminates in four cirri, which spring from below the vent, the upper being longer than
the inferior pair and their bases wider apart.

The proboscis (Plate LXIII, fig. 6) has a pair of strongly curved maxillas with a
shoulder posteriorly, and long, slightly tapered posterior appendages, which have a shallow
notch in the middle. The great maxillary plates have four prominent teeth on each side
anteriorly ; the curved anterior plate on the left has four rather blunt and rounded teeth,
that on the right about six, each has a blunt conical nodule in front, and there are
horny bars laterally. The mandibles (Plate LXIII, fig. 6 a) are proportionally large and
deeply curved from side to side, so that in many preparations the upper jaw apparatus
lies in the cavity thus formed. This curved anterior region is large, occupying about half
the total length of the teeth. It has on its upper surface a longitudinal black bar. The
dental limbs posteriorly terminate in two blunt processes.

LYSIDICE PUNCTATA. 459

The first foot lias a short dorsal cirrus somewhat tapered distally, a slight setigerous
lobe bearing both simple and compound bristles, and a short ovate ventral cirrus. In the
upper tuft of simple bristles are a few brush -shaped forms. The foot has a single spine.

At the tenth foot (Plate LXXV, fig. 4) the dorsal cirrus is longer, forming a tapering
process slightly constricted at the base. It does not extend in spirit beyond the tips of
the bristles. The setigerous process has greatly increased in size, forming a broad (or
deep) flattened cone with two strong yellowish spines. The simple winged dorsal bristles
(Plate LXXXIV, fig. 2) are curved at the commencement of the tip, which is somewhat
dilated, and then tapers rather quickly to a fine point. The edges of the wings are finely
serrate. These bristles issue above the spines.

Beneath the spines is a dense group of the jointed bristles (Plate LXXXIV, fig. 2 a),
the shafts being enlarged and curved toward the end, and serrated along the convex edge.
When the bristle is seen from the front only one of the convex edges is distinctly spinous,
the other being faintly so, but this may be due to position. The terminal piece is
comparatively short, has spinous wings, a double tip from a lateral hook, and a
terminal spur.

At the thirtieth foot (Plate LXXXIV, fig. 4 a) a single spine supports the setigerous
region, its point projecting beyond the edge. A strong hook, however, passes obliquely
downward and pierces the skin below the compound bristles, which are somewhat larger
than in the tenth foot. The hook (Plate LXXXIV, fig. 2 a) has a double wing at the tip,
which is bifid.

The structure of the simple and especially the compound bristles is more satisfactorily
made out in the fiftieth foot (Plate LXXV, fig. 4), the latter kind being considerably
larger than in the tenth foot. A single strong spine is present, and the powerful hook
(Plate LXXXIV, fig. 9) occupies the position indicated in the thirtieth foot.

In the posterior feet, viz., those in front of the tail, the dorsal cirrus becomes shorter,
projecting outward little beyond the tip of the setigerous region, which forms a broad
short cone, with the spine piercing the apex. The dorsal bristles are fewer and shorter,
but their serrations are very distinct. The compound bristles are also shorter and fewer,
and the serrations on the enlarged ends of the shafts are somewhat indistinct. The
ventral hook has the same oblique position, its brownish shaft crossing that of the spine.
The ventral cirrus is very short, bluntly conical, and does not reach a vertical line from
the apex of the setigerous region.

In some the spine towards the tip of the tail is dark brown. Occasionally two
occur, similarly coloured. A developing hook, again, is pale.

Reproduction. — In the beginning of April the females from Jersey carried ova — far
advanced. Lo Bianco 1 gives November as the month in which he found this species ripe
at Naples.

The young form has two pairs of eyes but only a median tentacle. By-and-by the
paired tentacles develop and the two anterior eyes disappear. 2

The Nereis cristatella of Montagu (MS. drawings, Linn. Soc, Plate XXXVI, fig. 3,
1808) appears to be this species.

1 ' Mitth. Zool. Stat. Neap./ xiii, p. 487.

2 Gravier, 1900.

460 GONIADIDJE.

The Lijsidice Mahagoni of Claparede (1864) may be included in this species, for the
only distinction is the absence of the slight median notch in the anterior border of the
snout.

De Quatrefages (1865) placed this form amongst his Lumbriconereids.

Bhlers (1868) rightly made considerable reformation in the synonymy of this abundant
southern form. Apparently a considerable change is also necessary in the synonymy of
foreign species.

The Lijsidice mavgaritacea of Claparede l from the Mediterranean seems to be closely
allied, though the spines are black.

The Lysidice collaris of Grube 2 from the Red Sea does not appear to differ essentially
from L. punctata. The comparatively slight differences existing in the teeth of such
species as this, and the close approach in the structure of the bristles and other features,
is suggestive. 3 This author 4 considered Risso's L. punctata as identical with L. Mnetta,
Aud. and Bdw.

The occurrence of a Marphysa at Marseilles, viz., M. fallax, Marion and Bobretzky, 5
mimicking this species so closely that they are readily confounded, is interesting, whether
the form is other than the young of the common Marphysa or not.

Family XII.— G-oniadid/K, Malmgren, 1865 (=G-oniadba, Kinberg, 1865).

Cephalic lobe elongate, tapering, with four equal terminal processes. Horny teeth
in two to five rows, terminal lateral, dorsal transverse, ventral transverse and radiate.
Feet biramous in the middle and posterior part of the body.

Grube's 6 seventh family, Grlycerea, included both Glycera, Sav., and Goniada, Aud.
and Edw.

Kinberg (1865) made a somewhat minute subdivision of the groups of teeth, which
he termed maxillae, ranging them into angular, terminal, lateral, transverse, transverse
dorsal, and transverse ventral maxillse. He describes the " branchiae " as terminal,
probably referring to the tips of the processes of the feet. He divides the genera
according to the presence or absence of the angular teeth into two divisions, and sub-
divides by the number of the transverse maxillae and terminal maxillae.

The view of Ehlers (1868) that Goniada and Glycinde should form one genus has
certain recommendations, but it should be remembered that the structure of the dorsal
division of the foot, both in soft parts and in bristles, shows a decided divergence,
and the same may be said of the teeth. 7 Moreover, Ehlers does not mention that
between the body-wall of each of the common forms (Goniada maculata and Glycinde

1 < Annel. diet. Nap./ p. 143, pi. viii, fig. 3, 1868.

2 ' Monatsb. Kg]. Akad. Wiss. Berlin/ June, 1869, p. 15 (sep. copy).

3 Vide, for instance, Marenzeller, ( Sudjap. Annel./ p. 136, Taf. v, fig. 2, 1879.

4 'Mitth. St. Vaast-la-Hougue/ p. 108.

5 ( Ann. Sc. nat./ 6 e ser., t. ii, p. 13, 1875.

6 < Fam. der Annel./ p. 59, 1851.

7 < "Challenger" Annel./ p. 342.

GONIADID^

461

Nordmanni) there is this difference, viz., that in the latter the dorsal longitudinal
muscles have a well-marked fold inferiorly, whilst in the former such is not present.

The Groniadidae formed the first family of the Euniciformia Grlycerina of Levinsen
(1883), and fall under Benham's Grlyceridae, the tenth family of the Nereidiformia. In
the former classification there is no doubtful association, whereas in the latter the
Sphserodoridge follow, and then the Ariciidas.

Goodrich observes that, as in Nephthys and Glycera, the segmental organ in Goniada
(Fig. 86) has no opening into the coelom, whilst a ciliated canal leads to the exterior —
through the septum, where it enlarges to form a lobed terminal organ — intermediate
between Nephthys and Glycera. The ciliated organ acts as a genital duct in Goniada
maculata. Fage, 1 again, sees in the closed nephridium, with its four rows of solenocytes,
and in the sudden development of the genital funnel at the moment of reproduction for

end o

r %-

Fig. 86. — Diagrammatic reconstruction of the nephridium and genital funnel of a ripe Goniada maculata (Goodrich).
end org. End organ, gf. Genital funnel, nd. Nephridial duct. np. Nephridispore. (Goodrich.)

the transmission of the sexual elements, a parallelism rather with the Phyllodocidas than
the Grlycericlas.

Ehlers (1868) groups his Glycerea polygnatha (Goniada) as follows : —

a. The terminal region of the proboscis without jaw-plates.

Two jaws with hooks and twenty paragnathi.
Ex. Eone Nordmanni, Malmgren.

b. The distal region of the proboscis with two rows of horny points (paragnathi).

1. Two chief jaws, several V-shaped jaws.

Ex. Goniada maculata, (Erst.

2. Two chief jaws and a ring of spiked horny points.

c. Surface of the proboscis with two longitudinal rows of horny points, with smaller
points on the ventral surface and angular plates on the side.

Ex. Glycinde multidens, F. Mailer.
It is probable that the British forms range to America, and that, perhaps, the
synonymy of the species requires readjustment.

1 ' Ann. So. nat./ 9 e set'.,

t. iii ; p. 311.

462

GONIADA MACULATA.

Genus LXXIV.— Gonjada, Audouin and M. Edwards.

Head slender, elongated and conical, with four terminal tentacles. Body rounded
anteriorly, flattened posteriorly ; segments with two rings. Proboscis minutely papillose,
with a row of V-shaped denticles on each side toward its base, and at its end two larger
pectinate jaws and an adjacent series of smaller denticles. Feet single in front, double
behind, the former having compound bristles, the latter both simple (dorsal) and compound.
The nerve-cords (Fig. 87) lie below the insertion of the oblique muscles, which pass from
below the bristle-bundles on each side with a slight inclination downward and inward,
and meet for insertion on each side of the hypodermic wedge above the nerves, which
lie in a somewhat triangular or pyriform area of the hypoderm, and each has a
small neural canal toward the upper and narrow part in section. Segmental organs
open ventrally, whilst internally they end in a tuft of short blind branches provided
with solenocytes. Genital funnel is trumpet-like and ciliated, fuses with and then opens
at maturity into segmental canal, and thus conveys genital products to the exterior
(Goodrich).

1. Goniada maculata, (Ersted, 1843. Plate LVI, figs. 2 and 3 ; Plate LX1V, figs. 6-6 d
—head, teeth; Plate LXXV, figs. 12-12 a— feet; Plate LXXXV, figs. 1-1 a'—
bristles.

Specific Characters. — Head an elongated tapering conical process, with four short
tentacles at the tip, and consisting of eight segments. Body 4— 6 ins. in length, tapered
anteriorly and posteriorly, rounded in front, flattened posteriorly, and terminating in two

Fig. 87. — Transverse section of the anterior third of Goniada maculata. Enlarged.

slender cirri. Proboscis has towards the base a row of about ten dark brown V-shaped
denticles on each side. They diminish in size distally, and also to some extent proximally,
in extrusion. Distally the organ has a circle of about twenty bluntly conical papillas, and
within this are two lateral denticles with a rounded base and a toothed distal region. A
series of ^-shaped intermediate small denticles, varying in size, forms a belt between the
larger pair. Feet simple in front, with three long papillas or lobes, but with two divisions at
the twentieth foot, the dorsal having a spine, a lanceolate lobe with a process at the base

GONIADA MACULATA. 463

dorsally, and a group of simple bristles. The ventral division has an upper, a middle,
and a ventral lobe, a strong spine, and two groups of compound bristles. The feet
increase in length posteriorly, especially the tips of the processes of the ventral division.
Arwidsson l found a reduced blood-system in Goniada maculata as well as in
Glycera, and the perivisceral corpuscles are similar in both, especially those containing
haemoglobin.

Synonyms.

1843. Goniada maculata, CE rated. Annul. Dame. Consp., p. 33, f. 16, 23, 91, 95, 97, 98.
,, ,, ,, idem. Regionis Marinis, p. 78.

1844. (?) ,, minusada, De Quatrefages. Ann. Sc. nat., 3 e ser. 14, p. 359.
1844 — 5. Glycera virldescens, Sars. Nyt Mag. f. Naturvid., vi, p. 381.
185-1. Goniada maculata, Grube. Fain. Aimel., pp. GO and 131.

1853. Glycera virldescens, Stimpson. Mar. Invert. Grand Manan, p. 33.

1865. Goniada maculata, Johnston. Cat. Worms Brit. Mus., p. 189.

„ „ „ Carrington. Annel. Southport, pp. 6 and 7.

„ „ ,, De Quatrefages. Annel., ii, p. 193.

1867. „ „ Malmgren. Annul. Poly oh., p. 182.

1868. „ „ Ehlers. Borstenw., ii, p. 704, Taf. xxiv, f. 36—48.

1873. „ „ Kupffer. Jaliresb. Coram, deutsch. Meere (' Pommerania '), p. 150.
„ „ „ Verrill. Americ. Journ. Sc. and Arts, vol. v, p. 103.

„ „ „ Sars. Bidrag Christ. Fauna, iii, p. 30.

1874. „ „ Smith and Harger. Trans. Conn. Acad., iii, pp. 13 — 16.
„ „ „ Malm. Goteb. vet. o. vitt. Handl., xiv, p. 86.

„ „ „ Mcintosh. Ann. Nat. Hist., ser. 4, vol. xiv, p. 200.

1875. „ „ Mobius. Jahresb. Comm. deutsch. Meere (' Pommerania^), p. 169.
„ „ „ Mcintosh. Invert, and Fishes St. Andrews, p. 123.

1879. „ „ Tauber. Annul. Danic, p. 103.

,, „ „ Hansen. Nyt Mag. f. Naturvid., xxiv, p. 268.

fi „ „ Langerhans. Zeitschr. f. wiss. Zool., xxxiii, p. 302, Taf. xvi, f. 37.

1886. ,, „ Harvey Gibson. Proc. Lit. Phil. Soc. Liverp., vol. xi, p. 155.

1893. „ „ Levinsen. Yidensk. Ud. " Hauchs," p. 332.

1897. „ „ Arwidsson. Bid. t. k. Svensk. Yet.-Akad. Handl., Bd. xxiii, aft. iv, No. 6,

p. 8, Taf. i, f. 3—10.

1898. „ „ idem. Bergens Mus. Aarb., No. xi, p. 36, pi. ii, f. 25—28, and pi. iv, f. 60

and 61.
1901. „ „ Whiteaves. Geol. Surv. Canada, No. 722, p. 79.

1905. „ „ Mcintosh. Ann. Nat. Hist., ser. 7, vol. xv, pp. 37 and 51.

Habitat. — The frequency with which this species was encountered in the stomachs of
the cod and haddock at St. Andrews by my mother was remarkable. It also frequents
the sand near low water-mark at St. Andrews, especially near the entrance to the harbour
at the East Sands. Dredged in muddy sand twenty-five miles west of Yalentia (J. Gr.
Jeffreys). S.W. Ireland, log 55, 1886; Ban try Bay, thirty-five fathoms, 1866, R.I.A.
Valentia Harbour (J. Gr. Jeffreys). S.W. Ireland, 110 fathoms off the Blasquet (J. Gr. J.).
In the Minch (off Hebrides) (J. Gr. J.), July, 1866. Seventy fathoms, Outer Haaf,
1 < Bih. k. Yet.-Akad. Handl./ Bd. xxiii, p. 8, Taf. 1, figs. 3—10.

464 GONIADA MACULATA.

Skerries, 1868 (J. G. J.). St. Magnus Bay, Shetland, eighty to one hundred fathoms
(J. G. J.). Dingle Bay, Ireland, thirty to forty fathoms, stones and sand, ' Porcupine,' 1 869.

Examples of the second type come from the Minch, off Bergen, Norway (A. M.
Norman). Denmark, Sweden, and Finmark. ' Porcupine,' 1870, at Station XVII a, in
795 fathoms, and in Bono Bay in twenty-five fathoms, and Station XXIX, 227 fathoms ;
bottom temperature, 53° P. North Sea (Ehlers). Madeira, where it is smaller than in
the North Sea (Langerhans). North America. Canada (Dr. Whiteaves).

Head (Plate LXIV, figs. 6 and 6 a) forming a tapered or elongated conical process,
with four short cirri anteriorly, and consisting of eight segments.

Body about 4 — 6 ins. long and of 194 segments (Ehlers), tapered at each end, but
much more distinctly so anteriorly, rounded in front, and becoming more flattened
towards the middle and posteriorly, and terminating in the vent, below which are two
cirri of moderate length. Female of a pinkish colour in front as far as the change in
the feet, then dusky brown, with a tinge of olive throughout the rest of its extent, the
dorsum being mottled with brown marked by pale lines and separated at each segment-

* w n

/

Fig. 88. — Teeth of Goniada maculata, from a Canadian example. Enlarged.

junction. Just in front of each junction a transverse line cuts off a narrow segment of
the brown pigment. The ventral surface is pale pink in front, greenish throughout the
rest of its extent from the large ova. These were extruded from the broken posterior
end of the animal, and had a very transparent capsule, with pale greenish yolk. The eggs
are comparatively large, though perhaps not quite ripe, and measure about *0105 mm.
No eyes. Six V-shaped denticles on one side and six or seven on the other in this
example.

Another example, also a female (16th June, 1904), was of a fine scarlet colour, paler
towards the snout — which was pale at the tip and pinkish behind, and dull yellowish
towards the tail. The red colour seemed to be generally diffused dorsally and ventrally
over those parts having it, and was bright at the bases of the feet, which showed some pale
patches, giving them a mottled aspect. Anteriorly the iridescent dorsum was divided into
a median and two lateral areas, the latter striated by longitudinal creases in the segments,
but at the change of segments this ceased, the dorsum then showing a similar disposition
to that in the foregoing female, though the pigment was red, the narrow bar in front of
the segment- junction being distinct. This remarkable coloration does not appear to be
due to the perivisceral fluid, which was pale, but to pigment in muscles and other parts,
as further examination proved, for this was likewise loaded with pale greenish ova with
large nuclei. The muscles of the proboscis and other parts appeared to be reddish.

GONIADA MACULATA. 465

The tail is dull greenish on the dorsum and on the feet, and terminates in a
moderately tapered region ending in the vent, below which are two cirri of average
length — also dull greenish.

The proboscis (Plate LXIV, fig. 6 a) is elongated and cylindrical, the first or basal
region (in extrusion) being minutely and densely papillose. The papillae become fewer
in the distal region, and disappear before reaching the ring of large papillae at the tip.
When firmly contracted the proboscis is closely striated. The basal region is armed by a
row of dark-brown V-shaped denticles on each side (Plate LXIV, figs. 6 a and 6 b), about
ten in number in well-developed examples. They diminish in size distally (in extrusion),
and also are somewhat less at the proximal end. Slight variation occurs in these teeth.
Thus Norwegian examples show the V-shaped teeth with longer limbs and more acute
apices, and they are also more closely arranged, though not more numerous. In some
British specimens a smaller number may exist on one side, and the last may be irregularly
formed or in two pieces. At the end of this region of the organ is a circle of twenty
papillae, of a bluntly conical form, and just within them two lateral denticles (Fig. 89) and
a few smaller intermediate ones. The larger denticles have a rounded or globular end,
connected by a neck with the somewhat palmate distal region which has two large curved
teeth (Plate LXIII, fig. 6d), the distal fang of great proportional size, the next smaller,
and the succeeding teeth gradually diminishing, the whole number being from six to
eight, the British examples presenting a much less globular basal end than that shown
by Ehlers. 1 The general outline in Norwegian examples agrees, though the number of
processes or teeth may vary. The intermediate small denticles (Plate LXIV, figs. 6 e
and 6 c') vary in aspect according to the position, as a rule having a basal end with two
flattened lobes or plates, a body terminated by two diverging main fangs, and a smaller
lateral fang on each side; but when viewed from the inner surface of the gullet four
projecting spikes meet the eye. Three of these occur in a row between the two larger
teeth, and on the opposite wall of the organ a pair of small V-shaped denticles are found,
and often another near each great denticle. The retractor of the proboscis is very long.
The buccal (peristomial) segment is about three times the antero-posterior diameter of
the preceding, and increases considerably in transverse diameter. The next segment
bears the first foot, which has two somewhat elongated lower lobes with bristles between
and above, and superiorly a flattened process continuous with the upper border of the
former, and representing the superior lobe or lamella.

At the tenth foot (Plate LXXV, fig. 12) a single median spine occurs in the conical
setigerous region, with a tuft of bristles superiorly and inferiorly, a long median papilla,
and a dorsal and a ventral elongated lamella (cirrus) tapered at the tip. The dorsal has
a flattened and slightly enlarged tip, a condition due to constriction or sinuosity of the
margin. The foot is thus characteristically trificl.

At the twentieth foot the three lobes and the bristles have a similar arrangement,
and so at the thirtieth foot, which, however, is somewhat larger. Between this and the
subsequent region a change in the complexity of the foot occurs, often about the forty-
first or forty-second.

1 Op. cit. ; pi. xxiv, fig. 43.

116

466 GONIADA BREMITA.

At the fiftieth foot (Plate LXXV, fig. 12 a) is a rather elongated slightly tapered
dorsal lobe, which appears to be homologous with a cirrus. Then the dorsal bristles with
their supporting spine occur above the large lanceolate lamella beneath. Inferiorly is the
rounded setigerous process of the region with its spine, and two groups of bristles, whilst
two long conical papilla project distally. At the ventral edge is a conical cirrus, which
does not project so far as the former.

The foot remains of similar structure till near the tip of the tail, the lobes somewhat
increasing in length, as also do the bristles. The bristles of the ventral lobe considerably
exceed the others, their very long terminal pieces projecting far outward. In the caudal
region the lower division shows superiorly three lobes, an upper and under and a smaller
median, with the ventral inferiorly.

The structure of the feet just detailed is observed in examples from the Outer Haaf,
Skerries, Shetland, from St. Andrews, Norway, and Canada. It is therefore the least
complex, both anteriorly and posteriorly. A tendency to the enlargement of the lobe
beneath the dorsal cirrus exists in females with large ova from Canada. Considerable
variety in regard to the length and breadth of the various lobes of the foot is observed.

The other type is that represented by examples from Cape Finisterre and Cape
Sagres, in which the anterior feet have five processes, viz., a dorsal and a ventral cirrus,
the former being near the base of the foot, a long and large posterior lobe, and two smaller
anterior. The dorsal lobe is long and narrow, and has a slight constriction at its base.
In this type the posterior feet have broadly lanceolate dorsal lamellae and three long
digit-like papillae inferiorly — above the elongate ventral cirrus, as exemplified in the
following species.

The dorsal bristles (Plate LXXXV, fig. 1) are translucent, with straight shafts, the
tip having a slight bend and dilatation, then being finely tapered distally. The ventral
bristles (Plate LXXXV, figs. 1 a and 1 a') have long shafts, slightly curved toward the
end. From the articulation the long, slender terminal piece tapers to a fine point. The
tips of the posterior bristles, especially the central series in the fan, attain a much greater
length.

Reproduction. — Some of those procured off the S.W. of Ireland had large ova in July.

Bhlers (1868) characterizes the proboscis as having small cordate papillae, with a
row of seven to nine arrow-shaped jaw-plates on each side, and with twenty terminal
papillae, two- to eight-toothed chief jaws, three ventral points, and four double points
(paragnathi).

The Goniada Alcoclciana of Dr. Carrington 1 appears to be a variety of this species,
though, at first sight, the shape of the body leans to Glycinde.

2. Goniada eeemita, Audouin and M. Edwards, 1834.

Specific Characters.— -Head like that of G. maculata, with four short tentacles at the
tip, and consisting of seven rings. Body elongated, four to seven inches in length and of
300 segments, rounded dorsally and slightly flattened ventrally, where a median groove

1 c Annel. of Southport/ 1865, p. 6.

GONIADA EEEMITA. 467

runs from end to end, and with the feet deeply cut. The first sixty-six feet are uniramous
three-lobed, with a lanceolate or sausage-like dorsal and a lanceolate ventral cirrus. The
feet behind have two divisions, the upper with a leaf-like dorsal cirrus, a lanceolate lobe,
and simple bristles ; the inferior division boldly trifid, with a lanceolate ventral cirrus and
compound bristles. Proboscis armed distally in extrusion with two large black teeth,
each with a flattened base, a short curved and blunt process, and three curved teeth,
whilst a belt of numerous spiked and complex denticles (somewhat ^-shaped) occurs
between them. Feet proportionally longer than in G. maculata, and from about the
tenth have five processes, viz., a dorsal cirrus, a three-lobed spinigerous region, and a
ventral cirrus. A change occurs at the sixty-ninth foot, the seventieth being biramous.

Synonyms.

1834. Goniada eremita, Audouin and Edwards. AnneL, p. 247, pi. 7a, f. 1 — 4.

1865. „ „ De Quatrefages. Annel., ii, p. 191.

1868. „ „ Bhlers. Borstenw., ii, p. 704, Taf. xxiv, f. 49—51.

1875. ,, „ Marion and Bobretzky. Ann. Sc. nat., 6 e ser., t. ii, p. 17.

Habitat — Specimens dredged in the ' Porcupine ' Expedition of 1870 in eighty-one
fathoms off Cape Finisterre, and in forty-five fathoms (eight miles N.W.) off Cape Sagres.

The head appears to be somewhat less attenuate than in the common form, but this
may have been partly due to the complete extrusion of the proboscis.

Body somewhat larger and longer than in Goniada maculata, of a firmer consistence,
more rounded dorsally, and grooved ventrally. First two segments without feet, the first
the broader. Proboscis (Fig. 88) with fifteen to eighteen V-shaped teeth on each side.
Rounded keel of the great fang less, and the adjoining flattened plate proportionally
longer ; terminal fang more massive, and the smaller teeth behind it fewer in number.
The row of spiked ^-shaped denticles is at least four times as numerous as in G. maculata.
Anteriorly the feet are proportionally longer than in G. maculata, and about the tenth
(Plate LXXXVII, fig. 20) have four lobes or papillse, viz., a large lanceolate dorsal lobe
(cirrus), which arises on the dorsal edge, nearer the base than the other processes, and
has a constricted base ; a larger and longer digit-like posterior lobe ; and two slightly
shorter anterior lobes which occur at the spinigerous region, whilst below is the long
lanceolate ventral lobe or cirrus. Only compound bristles are present. This therefore
differs from the condition described in Goniada maculata. At the thirteenth foot the
dorsal cirrus is a long sausage-shaped structure with a constricted base, and it has now
receded toward the body.

A considerable increase in length of the foot and its processes takes place at the
twentieth foot, the dorsal cirrus arising near the base, its tip reaching outward dorsally
to the middle of the foot. The long, pointed posterior lamella is prominent, and of the
two smaller anterior processes the inferior is the less, the lanceolate ventral lobe or cirrus
extending beyond the rest. At the thirty-first foot a single spine and only compound
bristles are found, as in front, and so at the fiftieth foot. Immediately behind the fiftieth,
however, a change occurs, two processes appearing instead of the single cirrus dorsally.
In the example this took place at the fifty-third foot, a lamella being suddenly developed

468 GONIADA EREMtTA.

beneatli and external to the cirrus. At the seventieth foot the dorsal cirrus is now
double (Plate LXXV, fig. 12 6), a long spathulate lobe inferiorly and a lanceolate lobe
superiorly. A dorsal spine is also present, but there are no dorsal bristles.

In the posterior feet the vertical diameter has considerably increased, and the dorsal
and ventral divisions are widely separated. The upper dorsal lobe is ovate and pointed
at the tip, the lower is somewhat lanceolate (Plate LXXV, fig. 12 c). The dorsal bristles
are simple and rather short, with a tapering tip slightly curved. A strong spine emerges
at the border of the lower lobe. The inferior division is the longer, has a setigerous
process with a strong spine, and a broad fan of compound bristles, the tips of which make
a gradational series, the longer tips being median, the shorter superior and inferior. It
has terminally a longer upper papilla or lobe (digit-like), and a pair of shorter lobes
inferiorly. The region is completed by the flattened, conical ventral cirrus, the tip of
which reaches the bases of terminal papillse above it.

Fig. 89. — Lateral tooth of Goniada eremita, Aud. and EdAv. Enlarged.

In their original description of this species from the Mediterranean, Audouin and
Edwards (1834) overlooked the toothed jaws and the denticles toward the tip of the
extruded proboscis, though they represented both in an allied species from New Holland.
The characters of the anterior as contrasted with the posterior feet also escaped them.

De Quatrefages (1865) also omitted to notice the distal armature of the proboscis
in extrusion. He, however, observed the distinctions between the anterior and the
posterior feet, though he overlooked the dorsal bristles, which probably had been

removed.

A careful account of this form was given by Ehlers (1868). In his examples the
change in the dorsal division occurred at the sixty-third foot, somewhat more posterior
than in the present form. The tail has two slender cirri. He describes the great length
of the proboscis with its sixteen terminal papilla) , its pair of three-toothed jaws, and the
band of smaller denticles, but his figure of the latter represents them as too minute.

GLYCINDE NORDMANNI. 469

Genus LXXV.— Glycinde, Fritz Mutter.

Head acutely conical, generally of ten segments, the terminal having four small
tentacles, whilst at the base clorsally are two small eyes. Body elongated, of two
divisions, the anterior region more smoothly rounded and having simple feet, the posterior
with two widely separate divisions of the foot, and stout dorsal bristles with a hooked tip
and a long tapering wing. Proboscis has a row of papillae slightly longer than in Goniada,
and behind these two larger denticles with four to five hook-like teeth, and a series of
smaller denticles, twenty-two to twenty-four in number, with several teeth (usually four).
The anterior feet are trifid, and the lobes long and pointed. Bristles arranged in a fan
and all compound, the longest tips occurring in the middle, and they are serrated. The
posterior feet have diminished upper lobes and stout bristles with slightly mucronate tips
and a tapering distal wing, a much enlarged lamella in front of the inferior setigerous
lobe, and a diminished ventral lobe (cirrus).

1. Glycinde Nordmanni, Malmgren, 1866. Plate LXIV, figs. 7-7 6— body, tail, teeth;
Plate LXXV, figs. 13-13 d— feet; Plate LXXXV, figs. 2-2 k— bristles.

Specific Characters. — Head of ten segments; two small eyes at the base clorsally.
Body resembling that of Goniada, feet commencing as simple and short, then bifid and
trifid organs as they increase in length, provided with short compound bristles. At the
tenth foot three long lanceolate lobes, a single spine, and a fan-like arrangement of
compound bristles in two groups. Before the fiftieth foot is reached the organ is bifid,
with two spines, and a prominent double lobe in each division, the upper and lower
representing the cirri. A change in the length of the foot occurs at the sixty-fifth foot.
The lanceolate lamella of the ventral lobe greatly increases in size posteriorly (e.g., one
hundredth foot), and the mucronate bristles project from the dorsal division.

Synonyms.

1865. Eone Nordmanni, Malmgren. Nord. Hafs-Armul., p. 409.

idem. Annul. Polych., p. 69, Tab. xi, f. 64.

Ehlers. Borstenw., ii, p. 703.

Mcintosh. Rep. Brit. Assoc. (1868), p. 837.

Malm. Goteb. vet. o. vifct. Hancll., xiv, p. 86.

Ehlers. Zeitschr. f. wiss. Zool., Bd. xxv, p. 56.

Tauber. Annul. Danic, p. 104.

When. Chset. c Yega '' Exped., p. 403.

Levinsen. Yidensk. Meddel. Foren. Kjobenh., p. 221.
, idem. Ibid., p. 221.

idem. Yidensk. Ud. "Hauchs," p. 332, Tab. i, f. 1—6.
1898. Glycinde Nordmanni, Arwidsson. Bergens Mus. Aarb.. No. xi, p. 50, pi, iii, f. 45—47 ; pi. iv,

f. 64 and 65.
1905. „ „ Mcintosh. Ann. Nat. Hist., ser. 7, vol. xv, p. 38.

Habitat. — In the stomach of the cod and flounder, St. Andrews Bay (E. M.). Not
uncommon in St. Magnus Bay, Shetland, where it was first dredged by Dr. Gwyn Jeffreys

1867.

>)

1868.

Goniada

1869.

>>

1874.

>)

1875.

?>

1879.

>y

1883.

))

))

))

>>

Eo

ne

1893.

(?)

}■>

470 GLYCINDE NORDMANNI.

in eighty-five to one hundred fathoms in 1867. He also procured it off North Unst in
ninety to ninety-five fathoms, and off Balta in fifty fathoms. In Bressay Sound it
occurs in eighteen fathoms. In the ' Porcupine ' Expedition of 1869 it was met with in
Dingle Bay, Ireland, in thirty to forty fathoms, and at Station XXXVI in 725 fathoms
in mud and sand. In the Expedition of 1870 it was dredged in 257 fathoms in the
Atlantic on a bottom of sand and at a temperature of 497°. In 110 fathoms off the
Blasquet (J. Gr. Jeffreys). Bay of Gralway (E. P. Wright).

Siberian Coast (Wiren). Foreign forms like that of the 'Challenger' appear to
come from comparatively shallow water.

Head (Plate LXIV, fig. 7) acutely conical, composed of ten segments, with four
short tentacles at the apex. At the base posteriorly are two eyes, which in some are
indistinct (spirit-preparations). Body about 3 — 4 ins. long, resembling that of Go?iiada,
tapered anteriorly and also posteriorly, where it is flattened and terminates in two
cirri (Plate LXIV, fig. 7 a). The surface has the same glistening aspect as in Goniada.
Of a pale pinkish or skin-colour in life.

Proboscis. — In this form none of the lateral V-shaped denticles are present. At the
termination of the first region of the proboscis (in extrusion) is a row of papilla? (Levinsen
shows eighteen), which are somewhat longer than those in Goniada macidata, and just
within these are two larger denticles placed near each other, which show four hook-
like teeth, two larger and two smaller. In the preparations, the arrangement of these
seemed in some to differ from Malmgren's figure (Plate XI, fig. 64 b), which shows a
decreasing series of four teeth in lateral view ; whereas in some this was not evident
(Plate LXIV, figs. 7 V and 7 b"). The smaller denticles appear to number from twenty-two
to twenty-four, and have two prominent teeth superiorly — most having thus a bifid aspect
■ — and a trifid plate inferiorly, an aperture on each side occurring at this point (Plate
LXIV, fig. 7 b). Levinsen (1893) gives a detailed account of the armature of the
proboscis with figures of the paragnathi, and these structures seem to differ from those
of the British form, which shows comparatively few paragnathi along the wall of the
organ.

The first feet are simple and short, bifid and then trifid as they increase in length,
and provided with short compound bristles. At the tenth foot three long lanceolate lobes
occur (Plate LXXV, fig. 13), an upper, a median, and a lower. The translucent com-
pound bristles stretch like a fan on each side of the tip of the median lobe, the longest
tips being in the centre of the group, and they diminish on each side (superiorly and
inferiorly) .

The thirtieth foot (Plate LXXV, fig. 13 a) has still only three divisions, but they
are more massive than at the tenth, and they are considerably flattened. The two groups
of compound bristles differ, since one (the upper) has the most conspicuously elongated
tips (Plate LXXXV, fig. 2b). The single spine is powerful. Between the thirtieth
and fiftieth feet the appendage attains greater complexity, the sixtieth (Plate LXXV,
fig. 13 b) presenting a dorsal division of two lobes, a spine, and a few bristles, but
these are often included in the tissues. This condition is evidently due to the growth
of a dorsal lobe on the upper process of the trifid foot, which the spine of the division
enters. The lower division of the foot consists of an upper long, bluntly conical

GLYCERINE. 471

setigerous region with two fan-like groups of compound bristles, which are arranged in
three divisions, an upper series of five or six with comparatively short tips, a second and
larger series with longer, finely tapered tips, and a ventral series with long tips dor sally,
shorter ventrally.

A great increase in the lamella at the inferior setigerous lobe occurs at the seventieth
foot, so that in outline the parts are broadly lanceolate. The bristles are longer, more
slender, and the terminal pieces finely tapered. The lobe also extends much further
beyond the ventral lobe than in the fiftieth foot. The characteristic bristles (Plate
LXXXV, figs. 2 and 2 a) now project from the lower superior lobe, it and the upper
having a broadly conical outline.

A considerable change occurs at the one hundredth foot, for the lower lobe now far
exceeds the superior in bulk, the broadly lanceolate outline being due to the approxima-
tion of the two flaps. The upper and lower bristles have shorter tips, the intermediate
have very attenuate tips. The ventral cirrus is proportionally smaller. In the superior
division, again, the setigerous lobe has assumed the form of a short and broad cone (Plate
LXXV, fig. 13 c), with the bristles projecting freely. The dorsal lobe is small. The
great length and slenderness of the tips of the ventral bristles posteriorly is a feature of
note (Plate LXXXV, fig. 2 c).

The foot retains, with modifications, the structure just mentioned to the posterior
end, the characteristic dorsal bristles projecting still more prominently, and with a
distinct curve below the hook at the tip, which has a delicate process (Plate LXXXV,
fig. 2 b).

A change in the length of the foot occurs about the sixty-fifth segment, for there-
after it becomes considerably longer, and remains so proportionally to the posterior end.

Reproduction. — In an example dredged in Bressay Sound in July, the body was filled
with advanced ova. In the same month another of small size (l\ in.) dredged in 110
fathoms off the Blasquet, S.W. Ireland, had well-developed ova. Large ova also were
present at the beginning of the same month in an example of average size from Balta
(fifty fathoms). Another at St. Andrews on 16th June had numerous large ova in the
coelomic space.

It is remarkable to find such slight differences between the Gli/cinde trifida of the
c Challenger ' l from Queen Charlotte Sound, near Long Island, New Zealand, and the
British form.

Verrill's Eone gracilis from Vinyard Sound 2 does not seem, so far as the description
goes, to differ materially from the foregoing.

Family XIII. — Glyceric.

Head long, in the form of a pointed ringed cone, with four small tentacles at the
tip, and having a central space communicating with the body-cavity. Several authors
state that " at the base of the snout are a pair of palps," and Ehlers observes that they
are retractile. Body elongated, rounded, slightly tapered towards the head, and more

1 Op. cit., p. 341.

3 ( U. S. Comm. F. and F./ 1873, p. 596, 1873.

472

GLTCBRID^J

distinctly diminishing toward the tail, which terminates in two short cirri. The segments
are narrow, and have two or more rings. Feet with two divisions — a dorsal and a ventral,
or with the latter only. Bristles of two kinds, viz., simple dorsal bristles with a narrow
wing and serrated edge, and compound bristles, with the tips of the shafts dilated and
split for the tapering serrated terminal piece.

Proboscis 1 powerfully muscular, with the first region easily protrusible, massive,
glandular, with four hooked teeth which have a bifid or simple basal spur, and the pairs
of which can cross each other vertically. Glands occur at the base. Alimentary canal
simple.

Blood-vessels present or absent. The coelomic corpuscles are large and tinted
reddish, and move with special activity at the ciliated bases of the feet. These corpuscles
contain haemoglobin. 3

The body-wall in the Grlyceridse (Fig. 90) is distinguished by its firmness and

Fig. 90. — Transverse section about the anterior third of Glycera lapidum. Enlarged.

muscularity. The cuticle is fairly thick, and is tough. Beneath is a hypoderm of
moderate depth, with numerous nuclei, which stain deeply, and areolae — lined
internally by a thin basement-tissue. The circular muscular coat is of exceptional
thickness. It diminishes (in transverse section) almost to a few fibres in the mid-dorsal
line (where the hypoderm is thicker), whilst ventrally, on each side of the nerve-area, it is
also thin, the space being considerably broader than that in the mid-dorsal line. A
narrow belt of circular fibres crosses this area ventrally to the other side, but
very few turn upward to clasp the inner ends of the ventral longitudinal muscles.
The dorsal longitudinal muscles are massive, separated by a gap in the middle line,
and with a well-marked lobe curved inward ventrally to a blunt point. They cover
a proportionally greater area than the ventral muscles, which form a long and slightly
fusiform mass on each side, and meet over the nerve-cords in the middle line as
they lie in a triangular area of their neuroglia — broad externally and pointed internally.
Each cord has a neural canal of moderate size at its upper third near the middle line.
It is doubtful, however, if in any respect such appears " to have generally the

1 Vide Jourdan, ' C. R. Ac. Sc./ t. cxii, p. 882.

2 Bay Lankester, f Proc. Roy. Soc./ No. 140, 1873, p. 3 (sep. copy).

QtLYCEBIDM

473

same relations as have the muscles and neural sheath, including the notochord, of a
vertebrate." : The alimentary canal has a powerful layer of longitudinal muscles
externally, and in some species (e.g., G. lapidum) this layer holds two large vessels (dorsal
and ventral).

Gills absent or present, sometimes retractile. The large nerve-cords (e.g., in G. lapi-

Flg. 91. — Diagrammatic reconstruction of the nephridial complex of Glycera siphonostoma, showing the nephridial
sac (empty), the ciliated organ, and the nephridium. co. Ciliated organ, ns. Nephridial sac. si. Soleno-
cytes. cc. Chambered caecum, nd. Nephridial duct. (Goodrich.)

dam, Fig. 90) are wedged in a hypodermic area between the large ventral muscles, which
touch at their dorsal edges so as to form an arch over the nerves. The external circular
muscular layer ceases before reaching the nerve-area, so that externally are only hypo-
derm and the specially thickened cuticle. The oblique muscles, attached above the
cords, are slightly developed. Segmental organs (Fig. 91) blind internally, opening by a

F IG . 92.— Diagrammatic reconstruction of a portion of the nephridium of Glycera siphonostoma, cut so as to show the
course of the canals. The inner surface would be applied to the wall of the nephridial sac. cb. Cell-body
(solenocyte). ml. Main branch. /. Flagellum, continued into the chamber, t. Tube with cilium in it.
nd. Nephridial duct. ch. Chamber, on, s. Outer surface, in. s. Inner surface. (Goodrich.)

pore on the ventral surface. Canal, which branches, passes inward and forward
through the next septum to the genital funnel, which is known to accumulate waste
products, and then spreads over the nephridial sac (or pouch of ccelomic epithelium).
Solenocytes arranged in small groups (Goodrich). Exit of genital products not known.
1 Ray Lankester, ' Ann. Nat. Hist./ ser. 4, vol. xi, p. 92.

117

474 GLYCERID.E.

GrrubeV seventh family, Grlycerea, held both Grlycera and Goniada, and he pointed
out the correspondence of the proboscides of these with those of the NephthydidaB and
Phyllodocidae.

Levinsen (1883) places the Grlyceridas as the second family of his Enniciformia
G-lycerma, the second division of the Euniciformia. Benham includes the family under
his Nereidiformia.

G-ravier 2 is of opinion that the rings of the prostomium in this group are superficial
and only affect the epidermis, and thus they correspond with the condition in other
annelids.

Goodrich 3 holds that Ehlers misinterpreted the exact structure of the segmental
organs (Figs. 91 and 92) in Glycera— which consist of a segmental organ, the ciliated
organ, and a segmental sac. The organ opens to the exterior by a minute pore outside
the large bundles of ventral longitudinal muscles. The pore leads to a slender canal
which passes through the body-wall to the septum, pierces it, and joins the segmental sac.
He thinks the ciliated organ and sac gather up through phagocytes the waste products
in the ccelom (with which there is no direct communication), whilst the purpose of the
segmental organ is to eliminate solid excretory matter from the ccelomic fluid. The
function of the solenocytes is possibly analogous to the Malpighian capsules of vertebrates,
viz., to excrete liquid. Fage 4 terms the ciliated organ the cilio-phagocyte organ, and it
is always attached to the nephridium. The nephridial mass in close contact with the
ciliated organ is divided into a multitude of minute canals, and the surface is hirsute with
a peculiar form of solenocyte. The organ, which is globular in shape, is remarkably
developed in Glycera svphonostoma, and in section it is elaborately reticulated. The
ciliated organ is a simple fold of the peritoneal epithelium which covers the dissepiment.
In Glycera alba, H. Ratlike, the nephridial mass forms three lobes, each with solenocytes.
It is held that the segmental organ does not transmit the genital products.

Dr. Thomas Williams (1852) pointed out the nature of what he called the " chyl-
aqueous fluid" in Glycera alba, and mentioned that in this form cilia occurred internally
at the base of the feet. He combated the opinion of De Quatrefages that the perivisceral
fluid was always circulated by vibratile cilia. He states that the ccelomic fluid is colour-
less, but that the corpuscles are blood-red, whereas " the true blood conforms with
the annelidan law of perfect fluidity : it bears no visible elements, and is light red in
colour."

Kinberg 5 termed the upper tentacles " antennas," the lower " palpi." He distin-
guished for purposes of classification the lateral pharyngeal papilla?- from the terminal
(between and in front of teeth), and he separated his "ovaria" ( = branchiae, And. and
Edw. ; " Kiemen," Grube) from his branchiae (=lingulse), the modified terminal regions
of the ventral cirri.

1 Tarn. Aimel./ p. 59, 1851.

3 < C. R. Ac. So./ t. cxxvi, p. 972, 1898.

3 ' Quart. Jo urn. Micr. Sc,/ vol. 41, pp. 439—457; and ' I ate mat. Congress Zool./ Cambridge,
1899, p. 196.

4 Ami. So. nat., 9 e ser., t. iii, p. 297, 1906.

5 'Of vers. af. k. Vet.-Akad. Fork/ No. 2, p. 244, 1865.

GLYCERINE. 475

De Quatrefages (1865) described a dorsal and a ventral-blood vessel accompanying
the intestine in the Glyceridae, though he also noted that the reddish fluid accumulated in
irregular places, and that in some the circulation was diffuse. He distinguished the
genera by the uniramous or biramous feet, and their nature.

Ehlers (1868) divides the Glyceridse into two great groups, according to the
structure of the feet. In the first the foot has two anterior but only a single posterior
lobe, and under this section fall two forms, viz., Glycera capitata and the British Glycera
lapidum, De Quatrefages. The other British forms are included under the second
division, viz., those in which two anterior and two posterior lobes are present. This
division has three sections. In the first the anterior lobes are long, whilst the posterior
are either truncated or rounded or very slightly pointed. Of those devoid of branchiae is
a British representative, viz., Glycera siphonostoma. In the second section in which the
two anterior lobes of the foot are long, whilst one of the posterior lobes is longer than the
other and with simple branchia?, is the well-known Glycera alba, H. Bathke. In the third
section, comprising forms in which the anterior lobes of the foot are long and the two
posterior somewhat long, and with simple dorsal branchia3, is G. albican*, De Quatre-
fages.

Ehlers unites Goniada with Glycera, and there is much to be said in favour of this
view, and comparatively little against it. He groups the Glyceridaa thus :

A. Proboscis with four hook-like jaws issuing from glandular sacs. Foot in all the
segments well formed (uniform) : — Glycera tetragnatha.

1. Foot simple (of one division) with only a single spine and a bundle of jointed

bristles.
Hemipodus, De Quatrefages.

2. Foot with two divisions, two groups of bristles, and two spines.

Glycera (Sav.).

B. Proboscis with several irregularly formed teeth, without glandular sacs. Anterior
and posterior feet unlike (ungleich) : — Glycera polygnatha.

Goniada and Bone.

Claparede (1868) observes that the perivisceral cavity in Glycera is divided into two
by muscular floors. In the majority of the Annelida Errantia it is divided into three
longitudinal chambers. He was of opinion that this family (Glyceriens, Grube) may best
be divided into two genera, the one Glycera, like Savigny's original species G. unicornis,
being devoid of teeth, the other, Ehynchobolus, having them.

The blood- system is reduced, but Wiren found blood-vessels in Glycera alba, and
Arwidsson 1 in this, in G. Goesi, G. siphonostoma, and G. capitata.

Ray Lankester again met with hasmoglobin in the perivisceral corpuscles of Glycera
(species not stated) at Naples. 2 This subject has been investigated by Cuenot/ Goodrich, 4
and Kollmann. 5 Goodrich found leucocytes and granular cells besides the corpuscles

1 ' Bih. k. Sv. Vet.-Akad. Handl./ Bd. xxiii, 1897.

2 ' Proc. Roy. Soc./ No. 140, 1873, p. 3 (sep. copy).

3 < Arch. Zool. exper./ 2 e ser., t. v, Suppl. 2, 1873.

4 ' Quart. Journ. Micr. Sc./ vol. 43.

5 'Ann. Sc. nat./ 9 e ser., t. viii, p. 147.

476 GLYCERIDJE.

containing hemoglobin, which were regular in form in Glycera sirjlwno stoma. Similar
results were attained by Kollmann, who further found the corpuscles containing
hemoglobin dividing karyokinetically in Glycera convoluta.
Grube, in 1870, thus groups the Glyceridae :

1. Glyceridae with branchiae.

A. Branchiae of the upper edge of the foot situated near the tip — a simple or clavate

process. The two anterior lamellae of the foot elongated.

(a) The upper posterior lobe resembling the anterior, the inferior shorter.
Branchiae simple.
G. alba, H. Rathke ; G. convoluta, Kef.

B. Both hind lobes short and blunt; only a small papilla present.

Foreign species.
G. Branchiae situated on the anterior border of the foot.

(a) Branchiae simple.

G.fallax, De Quatref.

(b) Somewhat forked, simple; lobes triangular, the anterior longer.

G. unicornis.

(c) Branchiae branched.

Foreign sp., e.g., G. dibranchiata (Massachusetts Bay).

2. Glyceridae without branchiae.

A. Foot with two anterior lobes, and only one broad posterior lobe.

G. capitata.

G. gigantea ( = G. siphonostoma, D. Ch.), De Quatref. Anterior segments
longer than the rest, the middle third as broad as long. Body 1 foot long,
300 segments. Anterior lip of foot with blunt tip ; posterior upper lip
short, triangular, the inferior less, shorter, bluntly rounded. Papillae of
proboscis digit-like, blunt.

G. decor uta, De Quatref. Foot similar, anterior lip pointed, hind lip short and
high, anterior segments shorter than in G. gigantea. Papillae of proboscis
fringe-like, oval, or clubbed.

Grube (1878) characterizes the family Glycerea (Gr.) as having a conical head-lobe
of many annuli. Tentacles and subtentacles two, short; sometimes none, often two
obtuse short tentacular organs posteriorly; no eyes. Body long or short, vermiform,
rounded, with numerous biannulate segments ; two anal cirri. No blood-vessels.

Buccal segments several ; the first long, naked, the rest bear feet. Proboscis clavate
or sub-cylindrical, with minute papillae, often with a circle of larger ones at the orifice.
Jaws two or four, sometimes with intermediate paragnathi, anterior and basilar. Branchiae
fixed to walls of pinnae, often retractile, simple sacs or bifurcate, rarely ramose, or none.
Feet uni- or biramous, with dorsal and ventral cirri, the former minute, papilliform,
inserted above the base of the foot. Bristles simple and compound.

A careful account of the proboscis of Glycera convoluta, Kef., was given by C. Gravier

GLYCERA LAPIDUM. 477

in 1898. He divides the structure into three regions — the anterior or pharyngeal, the
middle or proboscis (trompe pharyngienne), and the posterior region or ventricula
(stomach). From the structure of the teeth this would appear to be G. capitata. The
great development of the nervous system of the proboscis is emphasized.

This author further adds that Glycera has transverse bands of cilia on the segments
of body and head. He figures and describes a nuchal organ on each side of the basal
cephalic segment. A longitudinal groove occurs on the prostomium ventrally. Gravier
concludes that the head of a Polychaate differs fundamentally from a normal metamere.

Viguier 1 again considers all in front of the mouth as the prostomium.

In regard to distribution, none were found beyond 470 fathoms in the ' Challenger,'
but Ehlers mentions Glycera capitata at 664 fathoms in the ' Porcupine.' Hemipodus has
not hitherto occurred in British waters; as a rule it is southern and abounds in the
Strait of Magellan ('Challenger'). It is probable that most of the European forms
stretch to the American shores, and that further study will simplify the present list of
species.

Genus LXXVI. — Glycera, Savigny, 1820.

Head acutely conical with eight or more rings. Buccal (peristomial) segment large.
Eyes absent or present ; two retractile palpi (Ehlers). Segments with two or three rings.
Feet bifid ; a dorsal division with a stumpy dorsal cirrus often above the foot and simple
bristles with very narrow serrated wings, and a ventral with compound bristles and a
flattened ventral cirrus. Ventral spine considerably longer than the dorsal. Branchiae
present or absent. A modified blood-vascular system. Perivisceral corpuscles reddish.
Proboscis large, powerful, with four black teeth, the tips of which are strongly hooked
and emerge from a follicle with large glands. Muscular layers, nerve-cords, and segmental
organs as in the type.

Kinberg (1865), in his characters of the genus, placed reliance on the structure of
the teeth (his maxilla), which he described as having a long and a short lateral process,
whilst the " branchiae " were terminal, and stated that with the ventral cirrus there were
generally five lobes on the feet. He did not enter into the minute characters of the
bristles. Glycera and Hemipodia were his two genera.

Group I.
No branchiae.

1. Glycera lapidum, De Quatrefages, 1843. Plate LV, fig. 4; Plate LXIV, figs. 9 2 and
9 a ; Plate LXXVI, figs. 1-1 b— feet, etc. ; Plate LXXXV, figs. 3-3 b —bristles.

Specific Characters.— Read a moderately elongated cone of eight segments. Body
elongated (3 — 4 ins.), and of 150 — 170 segments, which are three-ringed, the median in
spirit-preparations being more elevated than the others. It terminates posteriorly in two

1 c Annel. Infer. Bay of Algiers/

3 This is termed G. capitata in the explanation.

478 GLYCERA LAPIDUM.

small cirri. Colour yellowish-white or in some slightly pinkish. Proboscis clavate, with
a rich covering of filiform papillae with crenated edges, four blackish-brown teeth hooked
at the tip and also thinned off at the other end, and with a bifid spur at one side. The
foot has a single spine in front, but about the thirty-fifth the dorsal lobe, instead of being
folded backward, projects straight outward, and dorsal bristles are present, though the
second spine only appears about the forty-fifth foot. The typical foot presents three
pointed lobes and a small dorsal cirrus above it, the third or inferior lobe, which Ehlers
says is larger, being the conical ventral cirrus, whilst the hind lip is broad and rounded.
The ventral bristles (compound) are in two groups and fan-like in arrangement ; more-
over, the extreme dorsal and ventral bristles are stronger than the central in the posterior
region of the body.

In the epitocous form the body is longer, the feet are longer and more prominent,
and the spines and bristles longer and more attenuate.

Synonyms.

1845. Glycera alba, Johnston. Ann. Nat. Hist., xv, p. 147, pi. ix, f- 1 — 10-

1851. „ „ Williams. Eep. Brit. Assoc, 1851, p. 172, pi. ii, f. 5, and p. 235, pi. xi, f. 61.

1853. Nereis teres, Dalyell. Pow. Creat., ii, p. 144, pi. xx, f. 1 and 2.

„ „ dorese, Leach. In Brit. Mus. Coll. {fide Johnston).

1862. (?) Glycera capitata, Keferstein. Zeitschr. f. wiss. Zool., Bd. xii, p. 105.

1864. Glycera alba, Grnbe. Insel Lnssin, p. 83.

1865. „ ,, Carrington. Ann el. Southport, p. 7.

„ „ capitata (olim alba), Johnston. Cat. Worms Brit. Mus., p. 186, pi. xv b, f. 1 — 10.

„ „ Baird. Ibid., p. 342,

,, ., lapidum, De Qnatrefages. Annel., ii, p. 187.

1867. ,, alba, Parfitt. Trans. Devon. Assoc, ii, p. 26 (sep. copy).

1868. „ lapidum, Ehlers. Borstenw., ii, p. 652.

1874. ,, capitata, Mcintosh. Ann. Nat. Hist., ser. 4, vol. xiv, p. 200.
}} ., „ Malm. Gloteb. vet. o. vitt. HandL, xiv, p. 86.

1875. „ „ Mcintosh. Invert, and Fishes St. Andrews, p. 124.

1878. „ ,, idem. Trans. Linn. Soc, Zool., i, p. 503.

1879. ,.. ., Hansen. Nyt Mag. f. Naturvid., xxiv, p. 268.

1883. „ „ Levinsen. Vidensk. Meddel. Foren. Kjobenh., p. 219.

1885. „ „ Mcintosh. ' Challenger ' Annel., p. 343.

v ., Cams. Fauna Medit., p. 224.

1891. ,, ,, Horn el. Trans. Biol. Soc. Liverp., vol. v, p. 244.

1894. „ „ De St. Joseph. Ann. Sc nat., 7 e ser., t. xvii, p. 31, pi. ii, f. 39 — 42.

1896. ., lapidum, Michaelsen. Polych. dentsch. Meere, p. 27.

1898. , „ Arwidsson. Bergens Mus. Aarb., xi, p. 15, pi. i, f. 7 and 8; pi. iv, f. 55.

1905 > }) y} Mcintosh. Ann. Nat. Hist,, ser. 7, vol. xv, pp. 39 and 50.

Forma epitoca.

1865. Glycera setosa, Johnston. Cat. Worms Brit. Mus., p. 342,

1876. ,, „ Mcintosh. Proc. Roy. Soc, xxv, p. 217.
1879. , „ Hansen. Nyt Mag. f. Naturvid., xxiv, p. 269.

1883. , „ Levinsen. Vidensk. Meddel. Foren. Kjobenh., p. 219.

GLYCERA LAPIDUM. 479

Habitat. — St. Andrews Bay, three to four fathoms and between tide-marks in fissures
of rocks amidst sandy mud ; nine miles off Balta, fifty to sixty fathoms, J. G. Jeffreys,
1868; four to eight fathoms, Bressay Sound, Shetland; dredged south of Sark (epitocous
male). Polperro, on gravel (Dr. Baird). Lamlash Bay (Dr. Howden). Coast of Corn-
wall, twenty to twenty-three fathoms. In fifty fathoms Dingle Bay ; off Bundoran,
Donegal Bay, twenty to thirty-five fathoms, ' Porcupine,' 1869. In muddy sand at Station
III, < Porcupine,' 1870, Lat. 48° 31' N., Long. 11° 3' W., in 690 and 500 fathoms.
Stomachs of cod and haddock, St. Andrews (E. M.). Scottish Fishery Board, Station
LXVI. Trawled off Sark by Col. Fraser and Capt. Powell; thirty to fifty miles west of
Valentia, Ireland, in 90 — 125 fathoms (J. Gr. J.) ; fifty-three fathoms, ' Knight Errant,'
3rd and 4th August, 1880, Station III; 664 fathoms in the 'Porcupine' (Ehlers).

Off Bergen, Norway (Canon Norman). In clayey mud in the Norwegian Fjords,
where O. F. Miiller early procured it. As G. setosa, surface of sea in Waigat Strait,
' Valorous ' Expedition. Station III, ' Valorous,' twenty fathoms, shell-sand, 1875.
' Valorous,' 1100 fathoms, 1875, Station VI. Shores of the North Atlantic. West coast
of North America (Ehlers). Off Fayal, Azores. Off Setubal. Mediterranean in
6 Porcupine ' Expedition of 1870, at Station XVII a in 795 fathoms, and thirty miles W.
of Cape Mondays, 100 fathoms; and at Station 50, Lat. 36° 14' N., Long. 17° 30' E., seven
to fifty-one fathoms. Mud and muddy sand, Bono Bay, twenty-five fathoms. On a sandy
bottom at Station XXX, in 386 fathoms, and with a bottom temperature of 53°, and in
forty-five fathoms eight miles N.W. of Cape Sagres, ' Porcupine,' 1870.

Head with a tapering series of about eight segments, the terminal having four slender
tentacles. No sign of retractile palpi.

Body 3 — 4 ins. long, 150—170 segments (Ehlers), slightly tapered anteriorly, and
more distinctly so posteriorly, rounded on the dorsal and somewhat flattened on the ventral
surface, diminishing to a slender tail, and terminating posteriorly in two small cirri.
Each segment has three distinct rings, the median being more elevated than the anterior
and posterior. In section, besides the neural canals at the upper border of each

nerve several smaller (branches) exist near the middle line below it. Groups of nuclei
occur in the sheath, especially externally. The cords lie at the outer border of the area.
The apex of the triangular nerve-area is occupied between the ganglia by a fibrillar
neurilemma with nuclei externally.

Colour of a small example translucent yellowish-white, opaque along the gut.
Ventrally the double nerve- cord is visible through the skin, and is pinkish anteriorly,
where it is dilated, and a pink patch also appears both dorsally and ventrally in front
of the ganglia. The nerve-cord is pale posteriorly. The four black jaws are seen
through the transparent skin. The intestinal canal is straight—pale behind the jaws and
dilated for some distance, then brownish to the tail. The lines of the segments have a
clear patch on each side. Some are slightly pinkish.

Internally active ciliary motion at the base of the feet, probably in connection with
the segmental organ, and it did not extend into the foot. The peritoneal corpuscles
have projecting processes like a bread-fruit. In section the gut has a thick layer of
longitudinal muscular fibres under the peritoneal or coelomic investment, and this

480 GLYCERA LAPIDUM.

coat clasps two vessels at opposite poles, the contents of these being coagnlable and
staining deeply. The circular nmscnlar coat follows, and then the dense epithelial coat
intern all v.

The proboscis in extrusion forms a long clavate organ, richly papillose, the individual
papilla being filiform (Plate LXXVI, fig. 1 a) with crenated edges. At the termination
of the first region are four blackish-brown teeth, each consisting of an elongated portion
hooked at the sharp tip, and also thinned off at the other end, where its muscles of
attachment are. On one side is a spur which becomes bifid after a short distance, and
to it other muscles are fixed. In the ordinary position in extrusion the four teeth are at
the angles of a square, and the long spur is vertical, the shorter passing inward obliquely
from each angle. The axis of the hooked tooth is nearly transverse. The aperture of
the proboscis is also transverse and in the centre. A symmetrical arrangement of the
muscular fibres further gives a character to the truncated terminal region at the
teeth. When the organ by a rupture of the body-wall is thrust out in its unrolled
position, the first region is marked by a series of longitudinal strise with the villi
internally. At the end is a ring of long processes like ruptured muscular fasciculi,
beyond which the four thick muscular masses for the teeth occur, followed by a somewhat
clavate part with ridges internally, and containing food. Such probably represents a
gizzard. A constriction separates this from the head-region, which at its commencement
has a series of strong muscular bands for retraction.

So far as can be observed the first foot has three anterior lobes and a large rounded
posterior lobe. The dorsal is the most slender, the median next, and the ventral is much
broader, its outline being lanceolate. The posterior lobe is bluntly rounded. The dorsal
bristles are comparatively short and finely tapered. The ventral spread out like a fan
above and below the spine. The differences between the outer and central bristles of the
fan are slight.

At the tenth foot (Plate LXXVI, fig. 1) the bristles are longer, but only one spine
occurs. The median lobe or papilla is longer. The change in the condition of the foot
occurs about the thirty-fifth, and is often recognized by the fact that the dorsal lobe,
instead of being folded backward, stands straight outward. The dorsal bristles are now
present (Plate LXXXV, fig. 3), but only a single spine, the second appearing about the
forty-fifth, though variation probably exists. The form remains similar at the fiftieth
foot, but the upper and lower compound bristles (Plate LXXXV, fig. 3 b) are thicker or
stronger than the median (Plate LXXXV, fig. 3 a), a feature still more marked at the
ninetieth foot (Plate LXXVI, fig. 1 b). The pointed condition of the three lobes is a
marked feature, and all are more or less anterior, though the ventral lobe is generally
sloped backward. The only posterior lobe is the bluntly rounded one behind the bristles.
A single spine supports the foot, the simple dorsal bristles passing from the spine toward
the dorsal papilla or lobe. At the base of the foot superiorly is the short papilla on the
side of the body. The differentiation of the ventral bristles into a diminishing series from
each edge is noteworthy. The foot appears to remain constant in its parts from the
anterior to the posterior end.

Reproduction. — Specimens procured in Bressay Sound in July were laden with
nearly ripe ova. Dalyell's example spawned in July.

GLYCERA EHLERSI. 481

Habits. — This species frequents sandy gravel and mud (Shetland). Dr. Johnston
found his under stones and buried in gravel or sand, not far below the surface, and, as
mentioned, it sometimes occurs between tide-marks in fissures of sandstone.

In the epitocous form (G. setosa) the body becomes more elongate and attenuate and
the feet are more prominent and longer — with attenuate bristles — which with the foot
give a character to the form. Two specimens procured in the 'Valorous' were males.
In these both spines and bristles become greatly elongated, but the latter retain their
essential structure. The example from eight fathoms in Bressay Sound seemed also to
be a male. It was much smaller than the foreofoine.

This would appear to be the species referred to by 0. F. Muller (1788) as Nereis alba,
judging from his description and figures.

Malmgren (1867) included under G. capitata both the northern type of CErsted and
the southern G. lapidum, making its sexual form, G. setosa, a separate species.

Ehlers (1868) showed that (Ersted's Glycera setosa, and Johnston's G. alba, as well as
the G. lapidum and G. Mulleri of De Quatrefages, all pertained to this species.

In Verrill's Check List for the U.S. Commission (1879) Rhynchobolus albus, capitatus,
and dibra?ichiatus appear.

2. Gtlycera Ehlerst, Arwidsson, 1898. Plate LXXXVI, figs. 4, 4', and 4 a— bristles ;

Plate LXXXVII, fig. (>— foot.

Specific Characters. — Head very long, attenuate, of more than twenty rings, muscles
separated by a median streak. Body elongated for a Glycera, segments two-ringed;
dorsal surface rounded and marked by a slight median groove. The ventral surface has
a median ridge flanked by a groove at each side. Proboscis with rather long digit-like
papillge 1 slightly tapered towards the tip. Jaws with a powerful curved main fang having
a broad base with two long spurs. Foot has the dorsal cirrus unusually large, two long
papilla? or lobes close to each other and almost parallel anteriorly, and a single flattened
lobe with a rounded edge posteriorly. Ventral cirrus short and conical, its tip nearly
in a line with the edge of the posterior lobe.

Synonyms.

1868. Glycera lapidum, Elders. Borstenw., ii, p. 652.

1898. „ Ehlersii, Arwidsson. Bergens Mus. Aarb., No. xi, p. 19.

1905. „ „ Mcintosh. Ann. Nat. Hist., ser. 1, vol. xv ; p. 41.

Habitat. — Eighty fathoms, St. Magnus Bay, 1867. Nine miles off Balta, fifty to
sixty fathoms, 1868 (Dr. J. Gr. Jeffreys). Fifty-three fathoms, Station III, ' Knight Errant/
3rd and 4th August, 1880.

Shores of France (St. Malo).

1 Grube describes them as short and leaf-like. He apparently thinks this like G. capitata, but
with the anterior lobes of the foot longer, and the segments two-ringed.

118

482 GLYCERA SIPHONOSTOMA.

Head long, attenuate, composed of numerous (apparently twenty-two) rings, with the
four minute tentacles at the tip, and two longer dorsal and two shorter a little below
(palpi of Ehlers). In the preparations the muscles of the snout are separated by a
median longitudinal streak.

Body rather small, about three inches in length, tapering a little anteriorly, and much
more gradually posteriorly. The dorsal surface is rounded and marked by a slight
median groove. The ventral has a median ridge flanked by a groove at each side. The
segments are two ringed. The proboscis has rather long simple digit-like papillae slightly
tapered towards the tip. 1 The black jaws have a powerful, curved fang, and a broad
base which has two spurs, so that it is triradiate, but it is difficult to remove them
uninjured from the muscles.

The typical foot (Plate LXXXVII, fig. 6) has a large dorsal cirrus at the base
superiorly, two well-marked pointed lobes near to and almost parallel with each other
anteriorly, and only a single short and rather deep lobe posteriorly. The ventral cirrus
forms a short conical lobe which does not project so far outward as the anterior lobes, the
tip, indeed, being in a line with the edge of the posterior lobe. Two spines are present.

The dorsal bristles (Plate LXXXVI, fig. 4) are simple and long, with a slight
forward and then a backward curvature as the tip flattens before tapering to a delicate
tip. The ventral bristles (Plate LXXXVI, fig. 4 a) have a similar curvature of the shaft
which slightly enlarges, then is narrow below the expanded distal end, which is split for
articulating with the flattened and finely tapered terminal blade. One bristle near the
dorsal edge of the ventral fascicle is somewhat larger than those near it.

3. G-LYOERA siphonostoma, Delle Chiaje, 1825. Plate LXIV, figs. 8, 8 a — proboscis and
tail; Plate LXXVI, figs. 2-2 c— feet; Plate LXXXV, figs. 4-4 b— bristles.

Specific Characters. — Head a conical process of nine segments. 2 Body rounded,
upward of a foot long and half an inch in diameter, slightly tapered anteriorly, more
distinctly so posteriorly, where it terminates in two cirri. . Segments two-ringed. Colour
pinkish or pale skin colour, with a reddish patch from the brain, and a pale brownish gut
posteriorly. Proboscis massive — with four powerful black hook-like teeth. Papillae
lanceolate and streaked like a leaf. 3 Foot comparatively short, with an ovoid dorsal
cirrus at its base ; two long anterior broadly lanceolate papillge or lobes about equal in
length ; and two short blunt posterior lobes besides the ventral cirrus, which is of con-
siderable length and has the tip pointed, but it does not extend so far outward as the
anterior lobes. The nerve-cords are surrounded by a thick sheath of the neuroglia, to
the investment of which the muscular fibres are attached. Numerous nuclei and cells
more immediately surrounding the cords in the interganglionic areas stain deeply. The
sections would seem to show that the cords are circular in the interganglionic and

1 Ehlers says they are leaf-like.

3 Grube states that each ring bears minute processes (Harchen), probably palpocils.

3 Grube describes them as stiff and thread-like.

GLYCERA SIPHONOSTOMA.

483

elongate-ovoid in the ganglionic area, where their vertical diameter is considerably
increased. Segmental organs as described (p. 473).

1825.

1827.
1828.
1841.

j y
1853.
1862.
1863.

1864.
1865.

1867.
1868.

))
1870.
1874.

1875.

}}
1879.

1881.
1885.

))
1891.
1893.
1894.
1897.
1901.
1904.

)}
1905.
1906.

Synonyms.
Lumhricus siphonostoma, Delle Chiaje. Mem., ii, pp. 413—428, Tav. xxviii (2), f. 20—24.
Nereis (Glycera) dulia, De Blainville. Diet. Se. Nat., xxxiv, p. 451.

alba, Brno-mere. Encycl. Meth., Vers, i, p. 134, Tab. lvi, £. 21, 22.
Glycera dubia, Blainville. Diet. Sc. Nat., lvii, p. 483, Tab. xix, f. 1.

„ syphonostoma, Delle Chiaje. Descriz. e not., iii, p. 84, v, p. 98, Tav. ci, f. 21—24.
Lumhricus siphonostoma, idem. Ibid., Tav. ci, f. 21 — 24 (descrip. Plate).
(?) Nereis (Nephthys) hirsuta, Dalyell. Pow. Creator, ii, p. 145, pi. xxi, f. 1—3.
Glycera convoluta, Keferstein. Zeitschr. f. wiss. Zool. (partim),^e Claparede.
„ tessellata, Grube. Arch. f. Naturges., Jahrg. xxix, p. 41, Taf. iv, f. 4.
» „ idem. Insel Lussin, p. 83.

„ dubia, De Quatrefages. Annel., ii, p. 179.
„ gigantea, idem. Ibid., p. 183.
„ decorata, De Quatrefages. Annel., ii, p. 181.
,, tessellata, idem. Ibid., p. 190.
„ siphonostoma, idem. Ibid., p. 198.

„ dubia, Johnston. Cat. Worms Brit. Mus., pp. 185 and 342.
» >> Parfitt. Trans. Devon. Assoc, ii, p. 26 (sep. copy).

tessellata, Elders. Borstenw., ii, p. 654, Taf. xxiv, f. 2, 33, and 34.
Bhynchobolus siphonostoma, Claparede. Annel. Nap., p. 182, pi. xvi, f. 2, et pi. xxxi, f. 5.
Glycera gigantea, Grube. Jahr. Schles. Gesell, 1869, p. 56.

dubia, Mcintosh. Ann. Nat. Hist., ser. 4, vol. xiv, p. 200.

„ idem. Invert, and Fishes St. Andrews, p. 124.
tessellata, Marion and Bobretzky. Ann. Sc. nat., 6 e ser., t. ii, p. 17.
dubia, Langerhans. Zeitschr. f. wiss. Zool., xxxiii, p. 301, Taf. xvi, f. 36.
tessellata, idem. Canar. Annel. in Nova Acta Leop.-Car., xlii, p. 113.
Mcintosh. < Challenger ' Annel., p. 343, pi. xlii, f . 5.
„ Carus. Fauna Medit., p. 224.

nigripes and dubia, Hornel. Trans. Biol. Soc. Liverp., vol. v, p. 244.
tessellata, Marenzeller. Zool. Ergebn., ii, p. 10.

gigantea, De St. Joseph. Ann. Sc. nat., 7 e ser., 1 t. xvii, p. 22, pi. ii, f. 20 — 29.
siphonostoma, Arwidsson. Bih. Sv. Vet.-Akad. Handl., Bd. xxiii, p. 21.
tesselata, Treadwell. U.S. Comm. F. and F., p. 201.

„ Moore. Proc. Acad. Nat. Sc. Philad., vol. x, p. 464.

gigantea, Allen. Journ. M. B. A., n.s., vol. vii, p. 226.

siphonostoma, Mcintosh. Ann. Nat. Hist., ser. 7, vol. xv, pp. 41, 50, 51, and 56.
tesselata, De St. Joseph. Ann. Sc. nat., 9 e ser., t. iii, p. 229.
siphonostoma, Eisig. Fauna u. Fl. Neap., xxviii, p. 236.

Habitat. — Tossed on the West Sands of St. Andrews after storms, and occasionally
found in the sand at extreme low water (E. M.). Under stones on sand, between tide-marks,
Guernsey and Herm. Dredged off the Hebrides, and in the Minch ; in ninety-five fathoms
off Balta; in eighty fathoms, St. Magnus Bay (J. G. Jeffreys, 1866). In muddy sand,
four to eight fathoms, Lochmaddy, 1865 and 1872, as well as between tide-marks, in Loch
1 By a misprint in the volume this has occasionally been entered as 8 e ser.

484 GLYCERA SIPHONOSTOMA.

Portan, near Lochmaddy, and at Paible. Scottish Fishery Board, Station LXVI, off
Cromarty? Plymouth (Allen). Coimemara (A. G. Moore). One hundred and ten
fathoms, thirty miles west of the Blasquet, south-west Ireland, ' Porcupine,' 1870. East
coast of England. Loch Goil (E. W. Hoyle, 1878).

' Porcupine,' 1870, off Cape Guardia, Shores of France. Shores of the Adriatic,
Lussin Piccolo (Grube). Quarnero (Ehlers). Canada. Japan (Moore and self). Off
Fayal in the Azores (' Challenger '). Madeira and Canaries (Langerhans). Off the Cape
of Good Hope (' Challenger'). Porto Rico (Treadwell).

Head forming a conical process of thirteen segments.

Body in life upwards of a foot long and nearly half an inch in diameter, a little
narrower anteriorly, and tapering more distinctly posteriorly. Each segment has two
rings, that bearing the foot being somewhat broader as a rule. In good examples the
surface of the body is thrown into a series of papillge, especially laterally on the ridges of
the segments, so that a tessellated aspect is given to the general surface, hence probably
the origin of the name given by Grube.

Proboscis. — In the preparations this is a short, massive, clavate organ, having the
four powerful black hook-like teeth at the tip. The inner surface of the organ is
minutely papillose (Plate LXIV, fig. 8), each papilla being somewhat lanceolate in outline,
and many are obliquely streaked. The teeth are only of use for seizure after the
extrusion of the proboscis, which the animal ejects with much force. Claparede (1868)
attributes the coloration of this organ to reddish granular pigment between the muscular
fibres. He also observes that the perivisceral cavity is divided into two chambers by the
union of the dorsal muscles below the intestine. The body terminates posteriorly in two
short tapering cirri (Plate LXIV, fig. 8 a) .

The nerve-cords occupy a similar position to those of G. lajpidum, and the area has
the same shape, and, as in that species, there are alternations in position according as the
section passes through the ganglia or between them. Two upper and several subsidiary
neural canals also are present.

The dorsum has a pinkish blush, marked in front by the whitish opacity of the
retracted proboscis, and acquiring a very faint pale purplish hue posteriorly in contrac-
tion. Others have a uniform pale flesh-colour with a pale snout and a reddish patch
(ganglia) at its base. Some are mottled pinkish white, with the black jaws visible
through the integument, and a few are of a reddish flesh colour. A large example at
Herm had a general purplish skin colour. Posteriorly the gut in the pale forms tints the
central line brownish.

The first foot has a large conical and a smaller lobe, a single spine, and two tufts of
bristles.

At the tenth foot (Plate LXXVI, fig. 2) three main divisions appear in lateral view,
viz., a large digit-like dorsal, a smaller median, and a shorter ventral with a broad base,
though posteriorly the setigerous process also shows two small elevations of its edge.
The dorsal bristles are simple, translucent, tapered, with just a trace of a wing. The
ventral are compound. Two spines are present. An ovoid papilla (dorsal cirrus) occurs
on the body at the base of the foot superiorly, and is especially conspicuous anteriorly in
this form.

GLYCEBA SIPHONOSTOMA. 485

The twentieth foot has two long conical papillae anteriorly, and two flattened
truncated lobes posteriorly, besides the dorsal cirrus and the shorter ventral cirrus..

The dorsal bristles (Plate LXXXV, fig. 4) are translucent, simple, with a slight
forward curvature of the shaft below the tip which inclines a little backward and then
forward at the finely tapered extremity. The edges of the tip present traces of wings.

The ventral bristles (Plate LXXXV, figs. 4 a and 4 b) have somewhat stouter shafts,
also with a slight bend forward and then backward toward the dilated end, below which
a slight constriction occurs. The dilated end of the shaft is split for articulation with the
thickened base of the terminal blade, which then is flattened and gradually tapered to a
fine point.

Both dorsal and ventral bristles extend beyond the anterior conical lobes.

The two anterior lobes have increased in size at the thirtieth foot (Plate LXXVI,
fig. 2 a), and the bristles are proportionally longer. The ventral lobe is somewhat less.
Little change occurs from the foregoing to the ninetieth foot, and indeed to the posterior
end of the fragmentary examples. The bristles are arranged in two bundles in each
division of the foot. The foot is therefore characterized by the two pointed anterior and
the two blunt posterior lobes.

A typical foot, e.g., the thirtieth, is comparatively short, and the dorsal cirrus is
more closely connected with it than in some forms, so that in profile it is distinctly a part
of the outline of the foot. The two anterior lobes are broadly lanceolate with pointed
tips and are about the same length. The ventral lobe does not extend so far outward,
but, as it arises near the base of the foot, it is of considerable length, and the tip is also
pointed. The two posterior lobes form short blunt flaps. The characters of the foot
remain more or less distinct to the posterior end, as, for instance, in the twentieth from
the tip of the tail (Plate LXXVI, fig. 2 b).

In some forms, e.g., that dredged by the ' Porcupine ' off Cape Guardia, in 1870, the
two anterior lobes are considerably elongated (Plate LXXVI, fig. 2 c).

The posterior portion of the body is readily reproduced, a provision which renders
the frequent ruptures during its violent muscular exertions less serious (Plate LXIV,
fig. 8 a).

Reproduction. — At Naples Lo Bianco 1 found this form mature from December to
April.

Habits. — They are active animals, progressing through the water with a wriggling
screw-motion — probably as they do through wet sand. When irritated the proboscis is
ejected, and the organ is used to perforate the sand. They apply the pointed snout to
the surface, and then by a vigorous thrust of the proboscis a considerable space is
penetrated. They seem to be liable to wounds and ruptures, and are irritable.

In the cavity (stomach) behind the proboscis in one from the south west of Ireland
was a considerable portion of a Gapitella, so that like many other Annelids the species is
carnivorous.

The early description of Delle Chiaje (1825) with the accompanying figures is

recognizable, especially by aid of the subsequent observations of Claparede and a specimen

kindly sent by E. Grrube. Delle Chiaje records the actions of the proboscis on immersing

the animal in spirit.

1 ' Mitth. Zool. Stat. Neap./ xiii, p. 491.

4 «6 GLYCERA ALBA.

The Nereis alba, Bruguiere, in the ' Encyclop. Methodique ' (1827), in so far as the
structure of the foot is given, approaches the southern variety of this species.

The Glycera Rouxii of Audouin and Milne Edwards (1834) agrees with this form very
closely, though some are doubtful on this head.

Leidy's Glycera americana 1 would appear to resemble this species in essential points.

The Glycera gigantea of De Quatrefages 2 is apparently this or a closely allied form.

In all probability the Glycera mitts 3 and G. clubia of Dr. Johnston pertain to this
species, and the same may be said of Glycera nigripes, a variety.

Grube 4 distinguishes G. Bouxii, And. & Edw., from this species by the fact that both
hind lobes are long, yet a little shorter than the anterior. All the lobes are thus some-
what long, pointed or a little rounded. Ventral cirrus somewhat pointed, about as long as
the hind lobe. Papillas of proboscis digit-like, blunt.

Roule (1896) describes a Glycera convoluta, Keferstein, from the ' Caudan,' but there
is nothing to indicate its special differentiation.

De St. Joseph, while entering Glycera tesselata, Grube, from St. Marguerite, also
states that he found Glycera tridactyla, Schmarda, at St. Raphael, a species which
Marenzeller (and apparently he also) identifies with Keferstein's Glycera convoluta — here
regarded as a synonym of the British form.

Eisig 5 considered that this species exhibited mastigo-helicoidal locomotion, and
cryptoid when boring in sand. Those deprived of the head did not survive.

Group II.
With branchiae.

4. G-lycera alba, De Blainville, 1828. Plate LXV, fig. 1 — proboscis; Plate LXXVI, figs.
3-3 c — feet; Plate LXXXV, figs. 5 and ha — bristles.

Specific Characters. — Head acutely tapered, with eight segments and four short
tentacles at the tip. Body of considerable length (6 to 8 ins.), the segments (100, Ehlers)
two-ringed, rounded dorsally and somewhat flattened ventrally, each surface with a
median line. The proboscis is short and clavate, and its papillae are small and clavate.
It is marked by longitudinal bands, and is transversely wrinkled distally in extrusion.
The dorsal cirrus is short with a constriction at the base. The foot is somewhat short
anteriorly, longer posteriorly, and in front soon presents two spines and a branchial
process, the wall of which differs in structure from the lobes. The branchiae occur on all
the feet, except the first and last (about twelve in each case), and arise from the dorsal
edge of the foot near the tip. In front of the foot are two flattened lanceolate lobes, and

i c Fauna Rhode Isl. and N. Jersey/ p. 147, pi. xi, figs. 49 and 50, 1855. See also Yerrill for
same form (his Bhynchobolus americanus).
^ <Annel./ii,p. 183, 1865.

3 ' Cat. Worms Brit. Mus./ p. 185.

4 < Jahresber. Schles. Gesellsch./ 15th December, 1889, p. 56 (1870).

5 ' Fauna u. Fl. Neap./ xxviii, p. 263, 1906.

GLYCERA ALBA. 487

behind are a conical lobe shorter than the anterior, and a bhint lobe inferiorly. The ventral
cirrus is bluntly lanceolate and reaches only to the tip of the blunt lobe just mentioned.

The nerve-cords in section are proportionally smaller than in either Glycera
lapidum or G. siphonostoma, but they have the same relations to the ventral muscles and
surface. Large neural canals sub-divided by bridles occur superiorly.

Synonyms.

1791. (?) Nereis alba, Linn. Syst. Nat. Gmelin, ed. 13, i, pt. 6, p. 3119.
1806. (?) „ „ Turton's LinnaBus, iv, p. 89.
1825. „ „ De Blainville. Diet, Sc. Nat., xxxiv, p. 452.

1828. Glycera alba, idem. Ibid., lvii, p. 484.

1843. „ „ Rathke. Beitr. Fauna Norweg., in Nova Acta Acad. Nat. Leop.-Car., t. xx, 1, p.

173, Tab. ix, f. 9.
„ GErsted. Annul. Danic. Consp., p. 33, f. 24, 103, 105, 110.

1844. (?) „ albicans, De Quatrefages. Ann. Sc. nat., 3 e ser. 14, p. 358, pi. ix, f. 6.
1844-5. „ alba, GErsted. Kroyer's Nat. Tids. (Drobak), p. 411.

„ „ „ idem. Regionis Marinis, p. 78.

1851. „ „ Grube. Fam. Annel., pp. 60 and 131.

„ „ „ Sars. Nyt Mag. f. Naturvid., vi, p. 208.

„ „ setosa, idem. Ibid., p. 208.

1857. „ aZ6a,-Danielssen. Reise, p. 61.

1858. „ „ idem. Ibid., p. 117.

1862. „ convoluta, Keferstein. Zeitschr. f. wiss. Zool., Bd. xii, p. 106, Taf. ix, f. 28 and 29.

1864. „ „ G-rube. Insel Lussin, p. 83.

„ „ „ Kolliker. Kurzer Ber. Schott,., p. 9.

1865. ,, alba, De Quatrefages. Annel., ii, p. 186.
„ ,, Danica, idem. Ibid., p. 187.

„ „ alba, Baird. Johnston's Cat. Worms Brit. Mus., p. 342.

„ „ setosa (partim), Johnston. Ibid., p. 342.

„ (?) „ fallax, De Quatrefages. Annel., ii, p. 184, pi. ii, f. 2, and pi. ix, f. 18.

„ (?) „ albicans, idem. Ibid., ii, p. 186.

1867. „ alba, Parfitt. Trans. Devon. Assoc, ii, p. 26 (sep. copy).

„ Bhynchobolus convolutus, Malmgren. Annul. Polych., p. 69, tab. xiv, f. 82.

1868. „ „ Claparede. Annel. Nap., p. 186, pi. xvii, f. 3.
„ Glycera convoluta, Ehlers. Borstenw., ii, p. 663, Taf. xxiv, f. 29 and 30.
„ ,, alba, idem. Ibid., ii, p. 660.

1873. „ „ Kupifer. Jahresb. Comm. deutsch. Meere (' Pommerania '), p. 150.

}> „ „ Willemoes-Suhm. Zeitschr. f. wiss. Zool., Bd. xxiii, p. 348, Taf. xviii, f. 1 — 3.

1874. „ Goesi, Mcintosh. Ann. Nat. Hist., ser. 4, vol. xiv, p. 200.
„ „ alba, Malm. Goteb. vet. o. vitt. Handl., xiv, p. 86.

1875. „ „ Mcintosh. Marine Fauna St. Andrews, p. 124.

99 „ „ Mobius. Jahresb. Comm. deutsch. Meere, p. 169.

1879. „ „ Hansen. Nyt Mag. f. Naturvid., xxiv, p. 268.

1883. „ „ Levinsen. Vidensk. Meddel. Foren. Kjobenh., p. 219.

1885. „ „ Cams. Fauna Medit., p. 223.

„ „ convoluta, idem. Ibid., p. 224.

1889. „ „ Wiren. Biol. Foren. Forh. Verhandl. in Biol. Vereins Stockh., Bd. ii, p. 32.

1893. „ „ Levinsen. Yidensk. Ud. " Hauchs/' p. 332.

48 8 GLYCERA ALBA.

1894. Glycera convoluta, De St. Joseph. Ann. Sc. nat., 7 e ser., t. xvii, p. 27, pi. ii, f. 30—38.

1897. „ alba, Arwidsson. Bih. Vet.-Akad. Handl., Bd. xxiii, p. 14, Taf. i, f. 11—13.

1898. „ „ De St. Joseph. Ann. Sc. hat., 8 e ser., t. v, p. 344.

convoluta, Gravier. Compt. Rend. Ac. Sc, t. 126, p. 973.
1901. (?) „ Goesl, Hornel. Trans. Biol. Soc. Liverp., vol. v, p. 244.

1904. „ convoluta, Allen. Journ. M. B. A., n.s., vol. vii, p. 226.

1905. „ alba, Mcintosh. Ann. Nat. Hist., ser. 7, vol. xv, pp. 42, 50, and 56.
» » G-raeffe. Arbeit. Zool. Stat. Triest, xv, p. 323.

convoluta ( = G. retractilis, De Qnatref.), idem. Ibid., p. 323.
1906 - „ „ Bohn. Ann. Sc. nat., 9 e ser., t. iii, p. 109.

Habitat. — Coast of Wales and Cornwall (Baird and Laughrin) ; Polperro ; Plymouth.
Stomach of flounder and haddock, St. Andrews and West Sands after storm (E. M.), and
dredged off B. Rocks. Outer Haaf, Skerries, Shetland, in seventy fathoms, 1868 ; nine
miles off Balta, fifty to sixty fathoms, 1868 ; eighty fathoms, St. Magnus Bay, Shetland,
1867 (J. G. J.). Bressay Sound, four to eight fathoms. CumbraB (Kolliker). In 165
fathoms in muddy sand off Ireland, < Porcupine,' 1870. Thirty to forty fathoms, Dingle
Bay, Ireland, ' Porcupine,' 1870. Off Bundoran, Donegal Bay, ' Porcupine,' 1869. Ken-
mare River, Roy. Irish Acad., 18th August, 1890. Station III, ' Knight Errant,' 3rd and
4th August, 1880, fifty-three fathoms. Dredged in twenty-four fathoms off the Bass
Rock, Firth of Forth (Mobius). Southport Sands, near low water-mark in company with
Sulcator arenarius and other forms.

Forty-five fathoms off Cape Sagres, and off Cape Guardia, 'Porcupine,' 1870. North
Sea. Faroe (Willemoes, Suhm). Japan Sea (Moore, etc.). N. America? (Levinsen).

Head typical, with eight segments, the tapered tip having four short tentacles.

Body of considerable length, viz., 6 — 8 ins. or more, slightly tapered in front, more
distinctly attenuated posteriorly. It is rounded dorsally, somewhat flattened ventrally,
and marked on each surface by a median line. The segments are well defined, and
consist of two rings, that from which the foot comes off being slightly the broader.
Posteriorly the body is flattened, and ends in a pair of slender cirri. In section the
main and subsidiary neural canals are distinct, the latter being somewhat larger than
usual.

Proboscis. — In extrusion the organ is short and clavate, with numerous longitudinal
strise, besides transverse wrinkles behind the teeth at the tip, a definite fold marking off
this region from that bearing the teeth. The papillae are small and clavate (Plate LXV,
fig. 1), the tip generally presenting a pale ovoid area from pressure. They seem to be
somewhat smaller on the transversely wrinkled distal region. The teeth occupy elevated
bosses symmetrically arranged in the extruded organ, the mouth being in the centre of
a four-lobed area, each lobe passing between the teeth.

In section the body -wall shows more massive dorsal longitudinal muscles, which have
only a slight incurvation at the lower edge, and thus they are in contrast with those of
G. lapidum. The ventral longitudinal muscles, however, are similarly developed, and
meet over the small nerve-area inferiorly to a greater extent than in either of the other
forms mentioned, only a narrow pedicle passing upward at the ganglia. The circular
muscles, as in the others, are powerfully developed. A conspicuous transverse muscle

GLYCERA ALBA. 489

passes from side to side between the edges of the dorsal and ventral longitudinal muscles
under the proboscis. This seems to take the place of the oblique muscles in this region.

The first foot has a single group of two spines, three small lobes, the superior being
the shorter, and it bears both dorsal and ventral bristles which extend beyond the lobes.
The dorsal basal papilla appears on the first foot and it is conspicuous on the second.
The ventral lobe is the shortest in this foot, the two upper being about equal in length.
The spines diverge distally, each going to its bunch of bristles (dorsal and ventral), so
that in the succeeding feet each spine is connected only with its own group of bristles.
The foot thus early shows two spinigerous lobes, two long lobes superiorly extending
considerably beyond these, a shorter lanceolate lobe interiorly, and the basal papilla
dor sally.

In its passage backward the dorsal lobe or process, which first appears as a papilla
in the anterior region, assumes a more or less erect position, until in the typical foot it
arches upward and. outward. Moreover, its structure differs from that of the other lobes
which are hypodermic, for it shows a close series of spiral or crossed fibres, evidently
muscular, and which are probably connected with the respiratory functions of the parts.

Beneath the foregoing are two flattened lanceolate lamellse — between which the dorsal
bristles emerge — supported at the base by their spine. These bristles are translucent
and faintly yellowish (Plate LXXXV, fig. 5), with a curvature at the tip and a very
slight enlargement, after which the bristle tapers to a fine point.

The ventral bristles again emerge between a short blunt posterior lobe and a
lanceolate flap in front. The bristles (Plate LXXXVI, fig. 5 a) form two groups which
slope gently from the spine. The ventral lobe is bluntly lanceolate, and reaches only as
far as the tip of the truncate lobe above. The dorsal papilla is short, with a constriction
at the base. The ventral spine is considerably longer than the dorsal.

At the tenth foot (Plate LXXVI, fig. 3) the dorsal lobe is long and with an
enlargement at the base dorsally. The other anterior lobe is narrower and somewhat
longer, whilst the ventral is shorter and somewhat conical. The upper posterior lobe is
papilliform and considerably shorter than the two anterior lobes. The lower posterior lobe
is also somewhat pointed or papilliform in this region, but afterwards becomes less acute.
The basal papilla (the dorsal cirrus of Ehlers) forms a rounded knob above the base of
the foot dorsally.

A considerable change occurs at the twentieth foot, for the slight elevation at the
base of the dorsal papilla of the foot has now assumed the form of a short branchia
(which is first visible about the twenty-fourth or twenty-fifth foot) extending more than
half the length of the upper papilla. The other lobes are similar to those in front except
that the lower posterior lobe is now shorter and more truncated. The changes noted are
still more evident at the thirtieth, the fiftieth, and the seventieth foot, the last (Plate
LXXVI, fig. 3 a) presenting a typical condition. The dorsal papilla or cirrus is now on
the dorsal base of the foot; the branchia extends considerably beyond the tip of the
papillse. The surface of the organ is corrugated, and a series of crossed fibres (Plate
LXXVI, fig. 3 b) which indicate the contractile condition of the parts is evident. The
various lobes are typical and stand out prominently.

Towards the tip of the tail the branchia diminishes to a small process at the base of

119

490 GLYCERA ALBA.

the dorsal lobe, and the dorsal process or cirrus moves inward on the foot. All the long
lobes of the foot are now more pointed, this condition being especially marked in the
ventral lobe. The lower posterior lobe remains short and rounded.

The branchiae seem to be readily thrown off in the preparations, and probably even
m life, as many in certain examples appear to be in process of reproduction. About ten
or twelve segments anteriorly and posteriorly are devoid of them.

In life the dorsum of the foot appears to be specially branchial in function, the
interior being boldly ciliated, and the rich crimson perivisceral corpuscles rush to and fro
and revolve rapidly. In the sickly animal the corpuscles tint the dorsum of the foot as
well as the superior process. Long motionless processes resembling cilia occur on the
slight convexity within the ventral process.

In a form from the south-west of Ireland (1885— log 29, 40 fms.) the lower
posterior papilla or lobe is longer than usual, and considerable variation exists in the
length of the processes. Thus all the lobes as well as the ventral cirrus are long and
pointed in an example dredged by the ' Porcupine ' off Cape Sagres (Plate LXXVI,
fig. 3 e).

In all probability the Olycera unicornis of Savigny 1 is this species. Its locality was
unknown. He did not distinguish the branchia from the other processes of the feet.

Keferstein 3 described a new species from St. Vaast-la-Hougue which he termed
Glycera convoluta. So far as can be determined by a perusal of the literature of the
subject and of specimens having this name, it would appear to be only a variety of the
widely distributed Glycera alba, H. Rathke, and in this Bidenkap 3 and De St. Joseph 4
now agree. Such also was the opinion of Grube, though he indicated certain differences
in the feet and in the papillae of the proboscis.

It is doubtful if the Rhynchobolus convolutus of Claparede 5 (Glycera convoluta, Kef.)
is other than a variety of G. alba, though he gives fourteen spurious segments to the
prostomium, and he places the branchia further from the tip of the foot than Keferstein.

Willemoes-Suhm (1873) gives a brief account of the sense-organs (nuchal organs) of
this species from Faroe. It is a protrusible structure on the dorsum of the segment on
each side in front of the first bristles.

Levinsen (1883) would be inclined to place Malmgren's Glycera Goesi under this
form.

Wiren 6 ' refers to the composition of the perivisceral fluid of this species and the
presence of haemoglobin.

This may be the species referred to by Miss Newbigging from the Clyde at Millport. 7

1 ' Syst. des Ann el./ p. 37.

3 ' Zeitschr. f . wiss. Zool./ xii, p. 106, Taf. ix, figs. 28 and 29.

3 < Christ. Yidensk. Selsk. Forhandl./ 1894, p. 77.

4 ' Ann. Sc. nat./ 8 e ser., t. v, p. 345.

5 ' Annel. Nap./ p. 186, pi. xvi, fig. 3, 1868.

6 "Oni blodet och blodomloppet hos Glycera alba, H.R./' f Biol. Foren. Forhandl./ Stockholm,
Bd. ii, p. 31, 1889.

Communic. Millport Mar. Lab./ i, 1900, p. 4.

7 t

GLYCERA GOESI. 491

5. Glyceka goesi, Malmgren, 1867. Plate LXV, fig. 2— head; Plate LXXVI, figs.
4-4 6— feet; Plate LXXXV, figs. 6 and 6 a— bristles.

Specific Characters. 1 — Head a short cone of eight or nine segments with the nsual
terminal tentacles. Body about 5 or 6 ins. in length or more, and segments two-ringed.
The surface is firm and somewhat tessellated at the sides, and the anterior end appears to
be less tapered. Brauchise begin on the thirty-second foot and continue nearly to the
posterior end (incomplete). Proboscis massive, clavate, brownish. The lateral spur of
each tooth is of moderate length and has a triangular anterior crest or process. Papillas
conical with interspersed globular ones. Typical foot of considerable depth, with the
dorsal cirrus at the base of the foot. Branchia simple, on the anterior surface of the foot
and directed forward and slightly upward. It appears about the thirty-second foot and
continues to the posterior region of the body. Two long lobes occur anteriorly at the tip
of the foot, and two shorter flattened lobes — the lower by-and-by being the shorter—
occur posteriorly, and a short ventral cirrus with an oblique and rather blunt tip.

Synonyms.

1845. (?) Glycera Bouxii, CErsted. (Drobak.) Nat. Tids., n.r., Bd. i, p. 411, f. 1.

1865. (?) „ hranchialis, De Quatrefages. Annel., ii, p. 182.

1867. Glycera Goesi, Malmgren. Annul. Polych., p. 71, Tah. xiv, f. 81.

1874. „ „ Malm. Goteb. vet. o. vitt. HandL, xiv, p. 86.

1875. ., Goesi, Ehlers. Zeitschr. f. wiss. 1 Zool., Bd. xxv, p. 56.

1897. „ Bouxii, Michaelsen. Polych. Deutsch. Meere, in Komm. deutsch. Meere., n.f., Bd. ii,

p. 27, Taf. i, f. 9 and 10.
„ Goesi, Arwidsson. Bih. t. K. Sv. Vet.-Akad., Bd. xxiii, pp. 4 and 19, Taf. 1,1 14 — 16;

Taf. 2, f. 1—5.

1898. „ Mesnili, De St. Joseph. Ann. Sc. nat., 8 e ser., t. v, p. 339.
1903. „ Goesi, Moore. Proc. Acad. Nat. Sc. Philad., vol. v, p. 464.
1905. „ Goesi, Mcintosh. Ann. Nat. Hist., ser. 7, vol. xv, p. 43.

Habitat. — 110 fathoms, thirty miles west of the Blasquet, S.W. Ireland. Bay of
Gal way, fifteen to twenty fathoms. Dredged in 160 fathoms fifty-five miles west of
Valentia (J. G. Jeffreys, 1870). In the Minch, at eighty fathoms, as well as at Station
XL V in 426 fathoms in the 'Porcupine' Expedition of 1869. Plymouth (Allen, as O.
Mesnili) .

Extends to Sweden. Japan (J. P. Moore). In forty-five fathoms off Cape Sagres,
i Porcupine,' 1870.

Head (Plate LXV, fig. 2) a short cone of eight or nine segments with the terminal
tentacles as usual.

Body of considerable size, probably 5 or 6 ins. in length, though incomplete
posteriorly, and the segments are two-ringed. The surface is firm and somewhat

1 Grube characterizes this form (G. unicornis) as having the hind lobes of the foot shorter than
the anterior. Papilla of the proboscis small, leaf-like; others knob-like.

492 GLYCERA GOESI.

tesselated at the sides, and the anterior end appears to be less tapered than in the
ordinary forms, being rounded at each side of the base of the snout. In life the body is
pale reddish.

Proboscis (Plate LXV, fig. 2 a) massive and clavate, of a pale brown colour in the
preparations, with the four teeth in the normal position. The lateral spur of the tooth is
of moderate length, and has a somewhat triangular anterior crest or process, that is, it
projects toward the central region of the organ in extrusion (under cover of the muscles).
The papillae are conical, and amongst them are a few more or less globular.

In section of the body-wall the nerve-area is wider transversely, and thus has a more
extensive muscular covering than in its allies. The gut has a thin muscular coat, and the
dorsal vessel projects freely above it, only the lower border being fixed closely to the
intestine. The peritoneal layer surrounds both.

The foot is of moderate length and considerable depth, having anteriorly two
elongated papillae and posteriorly two slightly shorter and also pointed papillae. Dorsally
is a single branchial stem, whilst ventrally is the cirrus, which is somewhat lanceolate.
The dorsal cirrus occurs at the base of the foot.

The typical foot (Plate LXXVI, fig. 4) of this species differs from that of Glycera
alba, H. Rathke, to which it is somewhat allied by the position and size of the branchia,
and by the fact that both posterior lobes are somewhat elongated and conical.

The tenth foot (Plate LXXVI, fig. 4 c) has a comparatively large and slightly clavate
dorsal cirrus, two long anterior lobes, the upper occasionally being flattened and bifid at
the tip, two flattened conical posterior lobes about equal in length and considerably shorter
than the anterior, and a short ventral cirrus with an oblique and rather blunt tip.

The chief difference at the twentieth foot is the slight increase of the ventral cirrus,
the tip of which is more pointed.

About the thirty-second foot a papilla appears on the anterior surface, and midway
between the base and the tip — the first evidence of the branchia ; and in the subsequent
feet this develops into a conical process, of comparatively small size, which is directed
forward and slightly upward (Plate LXXVI, fig. 4' representing the thirty-eighth foot),
and it continues to the end of the body, which, however, is not quite complete. This
and the long dorsal cirrus give a character to the foot. As an exceptional condition one
branchia, the sixth after the commencement on the right side, was large (almost as long
as the foot), and consequently it impinged against the foot in front, and curved upward.
The position of the branchiae makes it difficult to see them when the feet are placed
vertically on a slide.

Both dorsal (Plate LXXXV, fig. 6— dorsal) and ventral bristles (Plate LXXXV,
fig. 6 a — ventral) are comparatively slender and long, but there is little in their minute
structure to differentiate them from allied forms.

The foot alters little in its progress backward (Plate LXXV, fig. 4b), and the
prominent and pointed posterior lobes are easily observed under a lens, for they are
usually directed backward whilst the anterior point forward. Occasionally, however, the
lower lobe is blunt, thus simulating the condition in Glycera alba, H. Rathke, but the
normal arrangement is as mentioned. In one also, whilst the lobes were of equal length,
the lower was the broader.

GLYCERA GOESI. 493

(Ersted's Glycera Bouxii (1844 — 5) is a species requiring further investigation, for his
figure of the foot more resembles that of G. siphono stoma than that of G. Goesi.

The Glycera fallax of De Quatrefages 1 is probably this species or G. alba, in which
the branchiae (ovaria, Kinberg) have assumed a globular form either from softening or
otherwise. The same explanation suffices for Mr. Parfitt's Glycera vesiculosa, n. s., in
which the author places the branchiae on the anterior aspect of the foot, as in the present
species.

The Glycera branchialis of De Quatrefages 2 seems to be an allied form, the branchiae
springing from the anterior and upper part of the foot. His G. albicans 3 is a similar
form, but he describes the branchiae as arising from the upper and posterior part of the
foot.

Ehlers (1868) states that the branchiae in G. unicornis spring from the anterior border
of the foot — just as occurs in that from St. Magnus Bay, only two processes characterize
the branchiae of that species. Moreover, he describes the branchiae as present on all the
feet between the twenty-third and last four, and mentions that some are bifid. He
observes that two kinds of papillae occur on the proboscis.

Such, therefore, would agree with the Glycera Meckelii, And. & Edw., 4 in regard to the
branchiae.

The Glycera decipiens of Marenzeller, 5 from the Bay of Miija in Japan, appears to
resemble this species very closely, both in regard to the position of the branchiae and the
structure of the feet.

There is much in the structure of Glycera sagittarise procured off the Tetuaroa
Islands by the 'Challenger' 6 that agrees with this species, which may yet be found to
have a very wide distribution. It is true the branchiae commence only after the thirtieth
foot, but variation may occur in this respect.

Marenzeller (1902) gives a careful criticism of the synonymy of this form, which he
considers to be the Glycera Bouxii of Audouin and Edwards. He shows that the lower
posterior lip is shorter than the upper, and that the branchiae (which are simple) are
retractile and on the anterior border of the foot, and that while the latter is also the con-
dition in G. unicornis, Savigny, the branchiae in that form are forked.

Arwidsson (1897) describes the branchiae as occurring on segments 29 to 190, and
gives a sketch of the teeth of this form and of Glycera alba, the teeth of the latter having
a much broader base.

This is not the Rhynchobolus siphonostoma of Delle Chiaje (1842) from Naples, a
species having a great development of the branchiae, which are bifid and conspicuous, but
it may be Malmgren's G. Bouxii, QErsted.

1 ' Ann. Sc. nat.,' 3 e ser., t. xiv, p. 358 ; and ' Annel./ ii, p. 184, pi. ii, fig. 2.

2 Op. cit., ii, p. 182.

3 Op. cit., ii, p. 186 ; and ' Ann. Sc. nat./ 3 e ser., t. xiv, p. 358, 1850.

4 ' Regne Anim., Annel./ pi. xiv, fig. 2.

5 « Sudjap. Annel./ op. cit., p. 140, Taf. vi, fig. 3, 1879.

6 < Annelids/ p. 346, pi. xlii, fig. 8, pi. xxii a, fig. 10 ; 1885.

494

ARICIIDiE.

Family XIV. — Arioiim], Audouin and Edwards, 1834.

Head (prostomium) a pointed cone, with or without eyes, and devoid of processes.
Peristomial segment without appendages. Body tapered anteriorly and posteriorly,
flattened dorsally, rounded ventrally. Segments narrow. Proboscis a frilled rosette.
Tail with two long cirri. Feet dorsal in position, the ventral division being dorso-lateral
anteriorly and dorsal posteriorly. Branchiae ligulate, dorsal. Rows of papillae behind the
anterior feet.

Bristles of several kinds — elongated and tapering bristles peculiarly serrated so as to
appear camerated, simple tapering bristles, or the bases of the stronger ventral serrated
forms modified by friction.

The nerve-cords (Fig. 93) anteriorly pass from the ganglia to the ventral surface,
where they are at first widely separated, a broad band of transverse fibres intervening.

Fig. 93. — Transverse section about the anterior third of Aricia Latreillii, Aud. & Edw.

They lie outside the circular layer, and beneath and external to the oblique bands at the
inner edge of the ventral longitudinal muscles. Then the median space contracts, the
nerves approach each other at the inner edge of a somewhat triangular area of hypoderm,
the circular coat bending upward along the inner edge of the ventral longitudinal muscles
to the raphe on each side at the summit, to which the oblique muscles are attached. The
hypodermic area in this region is larger than it is posteriorly, and the surface is raised,
not grooved. By-and-by, however, a broad groove appears and then the narrow one
posteriorly. Toward the middle of the body the trunks have been pushed further
inward and are closely enveloped laterally by the ventral muscles, the fibres being bound
by a thin band of the circular muscle, which is intimately applied to the sheath of the area.
The latter is now somewhat clavate in outline (Fig. 94), springing by a slightly wider
base from the line of the ventral median groove, and marked by transverse fibres at the
base (externally) ; indeed, this narrower region below the cords is transversely streaked
throughout and has minute (stained) nuclei here and there. The cords occur at the
upper or inner dilatation, and are ovoid in outline, with a large neural canal in the centre
superiorly. Occasionally two smaller lateral canals are visible on the same level. Groups

ARICIID^E.

495

of deeply stained nuclei occur below the cords. The oblique muscles are attached to the
upper arch, to the middle of which the gut is fixed by strong converging muscular fibres,
so different from the usual lax band attached first to the ventral vessel and then to the
arch of the cords.

Audouin and Milne Edwards regarded the genus Arid a, Savigny, as intermediate
between the Errant Annelids and the Tubicola, and also possessing certain modifications
of structure found in the Terricola.

Their bodies are long, flattened dorsally and rounded ventrally, with numerous
segments. Head small, conical, of one segment, devoid of appendages. The proboscis
is unarmed. From the second segment backward are feet, the first twenty or thirty
having two divisions which differ from each other, whilst the subsequent feet have
divisions resembling each other and dorsal in position.

(Ersted 1 (1844), grouping the Annelids under three great divisions (his orders), viz.,
Maricolae, Tubicolee, aud Terricolse, further subdivided the Maricolaa into the Chsetopoda,

(S^ffiMffii

m

Fig. 94. — Highly magnified view of the nerve-area in Aricia Latreillii towards the middle of the body. The hypoderm and

cuticle are below.

which he split into the G. trematodina, G. vera, and G. terricolina, the family Ariciae
constituting the second of the last section. He characterized them as having a somewhat
rounded body of a definite number of segments, a simple alimentary canal without maxillge,
and a head with rudimentary appendages. But his family contained diverse forms now
included under several families, and those dealt with here fell under his Aricias verse,
in which the pinnae and branchiae are dorsal, and the tentacular appendages absent or
rudimentary.

Grube 2 chiefly followed the arrangement of (Ersted in his tenth family Ariciea, Aud.
& Edw., grouping the Spionidas, Sphaerodoridas, and Cirratulidas as well as the Ariciidas
under this heading, but he separated the Opheliidde.

Dr. Thomas Williams 3 observes that the segmental organ is very prominent in this
group, though limited to the two posterior thirds of the body. The ova and sperm-

1 ' Arch. f. Naturges./ Bd. x, p. 102.

2 < Fam. der Anriel./ p. 64, 1851.

3 < Philos. Trans./ 1858, p. 125, pi. vii, fig. 17.

496 ABICIULE.

bearing masses exhibit the same relation to the tube as in Nereis, and they occupy the
floor of each segment. The areolated tissue in which the ova are contained is distinguishable,
he states, into two parts, one of which is densely and intimately connected with the blood-
vessels, the other forming a mere utricular receptacle for the generative elements.

The second division of Levinsen's x Syllidiformia contains six families, viz., Spionidse,
ChaetopteridaB, Cirratulidse, Ariciidaa, Chlorgeniidge (?), and Ophelidse (?) They are termed
the Syllidiformia spionina. This arrangement might be improved, as also might Benham's
view that the family lies between the Sphserodoridas and Typhloscolecidse of his sub-order
Nereidiformia.

Kinberg describes the Ariciea as devoid of jaws, pharyngeal papillae near the margin
of the mouth small or obsolete, tentacles, antennae, and palpi absent or minute, no eyes,
cephalic lobe conical, terminal ; buccal segment devoid of feet ; tentacular cirri two or
none ; a change in the structure of the segments ; branchiae cirrose, dorsal, absent from
the anterior segments ; setaa numerous, capillary and annulate. He considered the
dorsal cirrus a second branchia, whilst a third was at the extremity of the dorsal division
of the foot, and a fourth a papilla near the ventral division. He arranged the genera
according to the presence of cephalic appendages — Arioia having four ; by the tentacular
cirri, the position of the branchiae, and the condition of the bristles. His uncertainty on
the subject is further shown by his separating Anthostoma of Schmarda as a distinct
family. 2

De Quatrefages (1865), like others, separated the Ariciidae from Auclouin and Milne
Edwards' group, containing Aonia, Ophelia, and Cirratulus, as a separate family, and
arranged them in two great divisions according to the nature of the proboscis, sub-
dividing the first series by the structure of the feet, the bristles, tentacles, and caruncle.

Cunningham 3 says: " In the Ariciidae I need only confirm the account given by
Mcintosh, that in the middle of the body the nerve-cords are thrust inwards by the great
ventral longitudinal muscles, which contain between them a narrow lamina preserving the
connection between the nerve-cords and the epidermis. A single median neural canal
runs above the nerve-cords as in Nerine."

In regard to organs of sense, pairs of otoliths were described by Bobretzky 4 in Arlcia
capsulifera of the Black Sea in certain segments behind the third, and in Aricia (Erstedi,
Claparede, 5 by Marion and Bobretzky 6 at Marseilles, a form with six pairs of otocysts.
Langerhans 7 further mentions in A. acustica similar organs situated dorsally in segments
eight to eleven, and, as Fauvel points out, dorsal to the feet, whereas the ciliated organs
are ventral to the dorsal division. The latter author 8 has recently found in Scoloplos
armiger an otocyst containing grains of quartz in connection with the external ciliated

1 ' Vidensk. Meddel. Kjobenliavn/ p. 180, etc., 1883.

2 ' Ofvers. af K. Yet.-Akad. Fork/ No. 9, 1866, p. 337.

3 'Quart. Journ. Micros. Sc./ vol. 28, p. 273, 1888.

4 'Black Sea Annel/ (Not seen.)

5 < Glanures Zoot./ p. 42.

6 'Ann. Sc. nat./ 6 e ser., t. ii, p. 68.

7 'Zeitschr. f. wiss. Zool./ Bel. xxxiv, p. 88, Taf. iv, figs. 1—9.

8 ' Ann. Sc. nat./ 9 e ser., t. vi, pp. 17—25, text-figs, iv and v, pi. iii, figs. 19 and 20, 1907.

ARICIA CUVIEEL 497

organ in a small nnmber of the anterior segments only (4 — 5 to 13 — 15), whilst the lateral
ciliated organs occnr in every segment. The otocyst is situated at the base of the dorsal
cirrns and opposite the ciliated organ in the form of a diverticulum of considerable depth.
Its walls have thick cuticle without cilia, and the surrounding cells are finely granular
and stain deeply, and their extremities are fibrillar. Thus two sense-organs occur in the
Ariciidae, viz., the ciliated organs and the otocysts.

Hacker (1896) includes the Ariciidge amongst those with non-pelagic larvas, and he
alludes to a six days' old larva of Aricia faetida, described by Salensky, 1 with three ciliated
rings in addition to the cilia of the prototroch.

Kostanecki 2 subjected the eggs of Aricia to an acid solution (acetic and nitric), and
then to a hypertone solution, with the result that certain ova segmented into three or
four blastomeres, and a few reached the six- to eight-celled stage.

An interesting memoir lately published by Ceruti 3 places the genera Aricidea and
Paraonis on a proper footing, and demonstrates that they pertain rather to the group of
the Spionidse than to the Ariciidse. This group will subsequently be dealt with.

The distribution of the Ariciidge is still in need of elucidation, but certain European
forms would appear to stretch to the eastern American shores, where, under new names,
it is not always easy to identify them. Some are littoral, whilst others frequent water -of
considerable depth.

Genus LXXVII. — Aricia, Sayigny, 1820.

Head pointed, without tentacles ; mouth with a foliaceous proboscis. Body more or
less elongated, flattened dorsally and rounded (convex) ventrally, little tapered anteriorly,
but slender posteriorly, and ending in two slender caudal cirri. Regions of the body
two, the anterior with rudimentary feet and peculiar spines, the posterior with dorsal and
ventral divisions better developed. Branchiae commence about the fifth foot, and
continue throughout. Bristles club-shaped serrated forms and peculiar serrated (came-
rated) forms with tapering tips. Body-wall as in the family.

1. Aricia Cuvieri, Audouin and Edwards, 1833. Plate LXXVI, figs. 5-5 c — feet; Plate
LXXXY, figs. 7-7 b— bristles; Plate LXXXVII, fig. 18— body.

Specific Characters. — Head a short pointed cone without trace of rings or eyes.
A transverse line separates it from the peristomial segment ventrally, but dorsally
a crescent is cut from the latter at the base. Body slightly diminished in front, gradually
tapering posteriorly and reaching eight or more inches in length. It is flattened dorsally
and rounded on the ventral surface anteriorly. Segments narrow. In transverse section
dorsally it is concave anteriorly and flattened posteriorly ; convex ventrally. Peristomial
segment narrow dorsally, but broad ventrally. Proboscis forms in extrusion a frilled

1 ' fitud. Devel. Annel./ i, taf . vii, fig. 8, a, v.

2 e Journ. R. Micros. Soc./ 1909, p. 575 (from 'Bull. Intemat. Acad. Sc. Cracovie,' 1909, pp.
238—253, 16 figs.).

3 c Mitth. Zool. Stat. Xeapel./ Bd. xix, pp. 459—512, Taf. xviii and xix, 1909.

120

498 ARICIA CUVIERL

rosette. Feet dorsal in position, the ventral cirrus being dorso-lateral. Posteriorly body
tapers to a blunt point with the anus at the tip dorsally, and with two very long and very
slender cirri arising at the anterior part of it.

The first region of the body comprises twenty-one feet, but these vary considerably
amongst themselves, commencing with a rudimentary foot anteriorly and ending in
modified feet toward the twenty-first. From the sixth foot backward the posterior of
the three rows of ventral bristles is darker, and is composed of spines increasing in
strength, so that they form a conspicuous row of brown dots. The foot becomes more
free and the posterior row of papillae more conspicuous, extending ventrally beyond the
bristles at the seventeenth foot. From the nineteenth to the twenty-fourth these papillse
go to the mid-ventral line. The rows of ventral bristles attain their maximum about the
fifteenth or sixteenth foot. The dorsal cirrus increases in size in its progress backward
and has an enlargement at the base. The branchiae commence at the fifth foot, and are
of a rich red colour in life. In the succeeding region of the body (at and after the
twenty- second foot) the dorsal division has in front a long tuft of serrated and slightly
camerated bristles, and behind is a large dorsal cirrus with blood-vessels. Below, after an
interval, is a smaller cirrus, also with a distinct vessel. The ventral division consists of a
setigerous process with a long papilla (ventral cirrus ?) and a few very slender bristles of
the same type as the dorsal. The camerated or peculiarly serrated bristles disappear
posteriorly, long simple tapering bristles taking their places in both divisions of the foot.

Synonyms.

1833. Aricia Cuvieri, Audouin and Edwards. Ann. Sc. nat., t. xxix, pi. xv, f. 5 — 13.

1834. „ „' idem. Annel. (Faun. Lit. Fr. ii), p. 258, pi. vii, f. 5—13.
1837. „ „ M. Edwards. Regne Anim. Illust., pi. xvii, f. 1.

1840. ,, „ Grube. Actin. Echinod., etc., p. 69.

1851. ,, „ idem. Fam. Annel., pp. 68 and 135.

1865. „ ,, De Quatrefages. Annel., ii, p. 283.

1867. „ „ Malmgren. Annul. Polycli., p. 71.

1869. „ „ Mcintosh. Rep. Brit. Assoc. (1868), p. 339.

1873. „ ., Sars. Bidrag Christ. Fauna, iii, p. 31, Tab. xviii, f. 17—23.
„ „ ,, Willemoes-Suhm. Zeitschr. f. wiss. Zool., Bd. xxiii, p. 348.

1874. „ „ Mcintosh. Ann. Nat. Hist., ser. 4, vol. xiv, p. 200.
9} „ ,, Malm. Goteb. vet. o. vitt. Handl., xiv, p. 91.

1875. „ „ Mcintosh. Invert, and Fishes St. Andrews, p. 124.
1879. „ „ Tauber. Annul. Danic, p. 106.

1893. „ „ Levinsen. Vidensk. Ud. "Hauchs," p. 336.

1904. „ „ Allen. Journ. M. B. A., n.s., vii, p. 227.

1905. ,. „ Mcintosh. Ann. Nat. Hist., ser. 7, vol. xv, pp. 44 and 50.

1906. „ „ De St. Joseph. Ann. Sc. nat., 9 e ser., t. iii, p. 167.

Habitat. — Dredged in twelve fathoms west of Peel, Isle of Man ; ' Porcupine '
Expedition of 1869 in 164 fathoms, on a bottom of greyish sand, stones and coral; and
in same Expedition at a depth of 422 fathoms; ' Porcupine ' Expedition, 1870, Station
VI, Lat. 48° 26' N., Long. 9° 44' W., in 358 fathoms on sand, bottom temperature 50°.
At Station VIII, in 257 fathoms ; Royal Irish Academy's Expedition to S.W. Ireland,
No. 122.

AEICIA CTJVIEKL 499

Bergen, Norway (Sars, Norman); Greenland; 'Porcupine,' 1870, Station XXV, in
374 fathoms, and bottom temperature 53 "5°.

Head a short pointed and very mobile cone without trace of rings or eyes. A
transverse line separates it ventrally from the peristomial segment, but dorsally a
crescent is cut from the latter at the base. It is dull salmon-yellow in colour like the
rest of the body, with an opaque brownish patch in the centre.

Body (Plate LXXXVII, fig. 18) of considerable length, slightly diminished in front
and gradually tapering posteriorly. It is somewhat flattened on the dorsal surface and
rounded on the ventral surface anteriorly. The anterior region has twenty-one segments.

Posteriorly the body diminishes to a blunt point, with the anus at the tip dorsally,
with two lateral flaps, a ventral process and papilla, and with two very long and very
slender cirri passing oif from the anterior part of it as in A. Lcdrelllil (Plate LVI, fig. la).
The dorsal processes (branchiae and feet) towards the tip of the tail gradually leave the
middle bare and pass to the sides in a diminished condition, even ceasing as distinct lateral
papillae before reaching the line of the vent.

The peristomial segment is narrow dorsally, but broad ventrally, the mouth forming
a puckered orifice in the centre of the ventral surface towards its posterior border. Two
crescentic lips occur laterally, while the posterior margin of the mouth has a series of
symmetrical longitudinal furrows passing forward from the segments behind.

The proboscis forms a deeply frilled organ, which projects from the mouth as a button-
like process. There are about ten frills in the form of a rosette.

The first bristled segment of the body has dorsally a low papilla, from which a tuft
of tapering and finely camerated pale-yellow bristles emerges, and a minute cirrus is
present. The bristles spread over the dorsum in a fan-like manner, almost approaching
the middle line. The ventral division is formed dorsally of similar bristles at its upper
and lower edges and here and there along the row, but the main part of the row consists
of the broken basal regions of the bristles, which often retain the camerated marks on one
side and which have a distinct curve toward the tip, the convexity being on the smooth
edge. The tips of these fragmentary bristles, especially inferiorly, have been worn
apparently from use, so that they are smoothly rounded. The second foot has a longer
dorsal cirrus and a similar tuft of bristles in the division. The ventral row of bristles is
longer, and it has posteriorly a flap with several (four) papilla. The bristles have the
same character as in the first foot, and the area forms a file-like surface from the broken
ends of the bristles. The third foot is similar, though longer from above downward, and it
has a posterior fringe of six papillae. The same may be said of the fourth foot, only the
dorsal cirrus is longer. At the fifth foot a branchia appears on each side dorsally and it
approaches the middle line more nearly than to the dorsal cirrus. The fimbriae or papillae
behind the foot are also more numerous, and the four rows of bristles show signs of wear.

The feet have the same structure (Plate LXXVI, fig. 5, which represents the tenth),
to the nineteenth, though the following changes are noted. From the sixth foot back-
ward the posterior of the three rows of ventral bristles is darker and composed of spines
increasing in strength so that they form a conspicuous row of brown dots. The foot also
becomes more free and the posterior row of papillae more conspicuous, whilst they extend
ventrally beyond the bristles at the seventeenth or eighteenth foot, though in others with

500 AEIOIA CUVIERI.

thirty-one segments anteriorly it is the twentieth, about five free papillae, that is, beyond
the foot, occurring at the eighteenth foot. At the nineteenth, twentieth, twenty-first,
twenty-second, twenty-third, and twenty-fourth the papillae extend to the mid-ventral line
along the anterior edge of the segment. The ventral rows of bristles attain their maxi-
mum development about the fifteenth or sixteenth foot, where there are twenty-one, and
diminish subsequently, the last occurring on the twenty-first foot as a small patch
considerably less than the first. The dorsal cirrus increases in size in its backward
progress, and shows an enlargement at the base.

The dorsal bristles (Plate LXXXV, figs. 7, 7', 7") are finely tapered and marked by
minute transverse rows of spikes, which give a camerated aspect to the structure. They
curve inward towards the branchiae anteriorly, but after the alteration of the feet they
project over them. The strong, brown, curved spines of the ventral rows (Plate LXXXV,
figs. 7 a and 7 6) do not, in those best developed, show any trace of the camerated con-
dition, but so many intermediate forms occur that it is not improbable that originally the
majority sprang from such at first. In glancing along the ventral divisions of the feet,
the strong brown spines of the posterior row become distinct about the sixth foot, and
increase in size in the following feet. The rows of papillae behind the spines and bristles
of the anterior feet evidently perform important functions in the operations of the region.
They all slope backward.

The twenty-second foot (Plate LXXVI, hg. 5 a) inaugurates a change in the structure,
for, whilst the branchiae continue as before in the mid-dorsal line, the ventral division has
no rows of strong bristles. The dorsal division has in front a long tuft of camerated
bristles tapering to fine points, and behind the large dorsal cirrus. Below, after an
interval, is another smaller cirrus, then a setigerous process with a long papilla, and a
few very slender bristles of the same type as the others (camerated). Lastly, the papillae
form a row to the mid-ventral line. The function of these ventral papillae may be con-
nected either with tube or tunnel in sand. This type of foot (Plate LXXVI, fig. 5 b) is
continued to the thirtieth, and then the camerated bristles disappear, simple long tapering
bristles taking their places in both divisions of the foot.

The posterior feet show to the exterior of the large flattened branchial process a long
tuft of bristles wi th a knife-shaped cirrus behind it, the cirrus having a narrow pedicle
with a broad blade tapering to a point beyond it (Plate LXXVI, fig. 5 c); then comes a
cirrus, followed by the ventral division in the form of a bifid process with a long tuft of
tapering bri sties, and a short conical ventral cirrus externally. The foot is dorsal in
position, even the ventral cirrus being dorso-lateral.

The branchiae, as Claparede so clearly showed, 1 have two vessels, with a numerous
series of transverse or connecting trunks.

Only twenty ventral rows occur in an example from S.W. Ireland (122), but the
papillae are larger. Another small example, dredged at 422 fathoms during the * Porcu-
pine ' Expedition of 1869, had only nineteen of the bristled rows of the ventral division.
Moreover, whilst the strong upper brown spines showed no trace of transverse markings
(camerae) the inferior did so. The great thickness of the upper spines made it probable
that either a new growth had taken place in the bristle sac, or that an increment had

1 ' Recherches sur la Struct. Annel. Sedent./ pp. 106, 109, and 192-3, pi. xiv, figs. 3 and 4, etc.

ARICIA EDWAKDSI. 501

ensued on the base of the old bristle. Another small example had only eighteen of these
ventral rows of bristles. Thus the number of the ventral rows seems to increase with
age.

In a small specimen from 164 fathoms in the ' Porcupine ' Expedition of 1869 all the
bristles were pale yellow, and a few of the stout curved stumps occurred in the twenty-
second foot (Plate LXXVI, fig. 5 a) amongst the slender bristles, so that they may
occasionally be overlooked.

Posteriorly towards the tail the blood in the various dorsal and dorso-lateral pro-
cesses, which in this region are proportionally long, gives a deep red colour to the parts.

Most specimens consist of the anterior region only, so that they probably frequent
the sand or gravel.

Reproduction. — Lo Bianco 1 found A. foeticla, Clap., ripe from January to June at
Naples. The eggs are of a greenish colour, and deposited in a cylindrical vermiform
mass of mucus.

This was the type of Audouin and Edwards (1834) which Cuvier had procured on
the French shores, and their description and figures are for the most part good.

De St. Joseph (1906) distinguishes between this species and A. Latreillii, Aud. and
Eclw., by the shorter anterior region in the latter, the smaller number of papillose
segments, and by the diminution in the number of papillee in each segment.

2. Aricia Edwardsi, an. var. Plate LXXX VII, fig. 8 — foot.

An injured specimen dredged in Busta Voe in forty fathoms by Dr. Gwyn Jeffreys
in 1868 offers certain distinctions which would appear to be specific. Thus, whilst in the
outline of the head and in the general aspect of the body, and in the commencement of
the branchiae on the fifth bristled segment, it agrees with A. Cuvieri, it differs in the
darker brown spinigerous rows anteriorly, in the length and slenderness of the papillae
behind the feet, in the conspicuous size and shape of the spine-like bristles of the ventral
row, and in the paucity and minuteness of the papillae which extend ventrally on the
segments. As in the majority of these forms, only the anterior and a portion (about
three quarters of an inch) of the succeeding region had been secured by the dredge.
The head is a short acute cone, and the frilled proboscis projects ventrally from the
peristomial segment. The anterior region consists of twenty-one segments, the ventral
rows being dark brown, and the last four or five very small. At the tenth foot (Plate
LXXXVII, fig. 8) the subulate dorsal cirrus is rather more elongate than in either A.
Cuvieri or A. Latreillii, and the bristles consist of a longer inner (i.e., next the median
line) and a shorter outer series of tapering spinous forms. After a considerable interval
a long papilla somewhat enlarged at the base occurs to the exterior. It is accompanied by
a very powerful and nearly straight spine with a sharp tip, followed by the dark-brown
series, some of which are curved at the tip as in the previous forms but far stronger, and
moreover they show that the tips are not serrated. The smallest forms are at the extreme

1 < Mitth. Zool. Stat, Neap./ xiii, p. 484.

502 ARICIA LATREILLII.

ventral edge, but all along the row much stouter spine-like bristles occur, and they are
either nearly straight or slightly bent at the tip. Short tapering serrated bristles are
present throughout. In the ventral division of the succeeding feet all the curved spine-
like bristles are stronger than in A. Guvieri and the papilla have the same character as
above. At the twenty-first foot the ventral division has a small group considerably more
elongated than in front, but with the terminal curvature, yet no trace of serrations occurs.
Even in this, the last of the series, the bristle-like spines are remarkably strong. They
are accompanied by the short, tapering, serrated (camerated) bristles. The papillaa
remain elongated. The condition of the preparation does not admit of minute description,
since the parts are much altered, but the change occurs at the twenty-first foot, which
has an elongated ventral cirrus and a tuft of the usual bristles serrated at the tip. At
the thirty-first the branchia is fairly large, the dorsal cirrus slightly enlarged in the
middle, and a tuft of somewhat stiff bristles serrated at the tip. Between it and the
ventral division is a cirrus, whilst the ventral setigerous lobe is bifid and bears a few
slender elongated bristles spinous at the tip. The ventral cirrus is subulate.

The ventral surface is comparatively smooth anteriorly, for none of the rows of
papilla) meet in the middle line, those most nearly approaching it being the eighteenth,
nineteenth, twentieth, and twenty-first, though some occur laterally from the seventeenth
to the thirtieth foot.

3. Aricia Latreillii, Audouin and Edwards, 1834. Plate LVI, figs. 1, la.

Specific Characters. — Head as in A. Guvieri. Body 6 — 8 ins. long, pinkish anteriorly,
yellowish-white posteriorly, distinguished by having about thirty-one segments in the
anterior division ; rows of papillaB on the ventral surface from the twenty-fourth back-
ward more numerous than in A. Guvieri.

Synonyms.

1834. Aricia Latreillii, Audouin and Edwards. Annel., p. 259.
1882. „ „ Mcintosh. Trans. Linn. Soc, Zool., i, p. 504.

Habitat. — Sandy shores of the Bay of St. Andrews at and beyond low water-mark,
and in Laminarian roots tossed on the West Sands after storms. Firth of Forth (J. T.
Cunningham) .

Head acutely pointed, forming a cone as characteristic as in A. Guvieri.

Body 6 — 8 ins. long, very little tapered in front, and more distinctly so posteriorly,
where it terminates in two long slender cirri. In general appearance it resembles
A. Guvieri, but the anterior region is composed of thirty or thirty-one segments with the
brownish ventral bristles, is somewhat more firm, and the rings bearing papillaB on the
ventral surface from the twenty-fourth backward are more numerous. The body is
somewhat flattened dorsally or even hollowed in front, and rounded ventrally, and the
segments are very numerous — 300 to 400. The proboscis is sometimes ejected as a frilled
lobate organ. In life it is of a yello wish- white colour, slightly pinkish anteriorly.

The body- wall anteriorly is characterized by the great development of the circular

ARICIA LATREILLII. 503

muscular bands and the powerful muscles of the feet, but the dorsal and ventral
longitudinal muscles are less massive in proportion. Behind the middle of the body,
again, the great increase of the ventral longitudinal muscles is a feature of note, the
dorsal being comparatively small.

The first region of the body comprises from thirty to thirty-three bristled segments,
the first and last being more or less modified. The strong ventral bristles of the region
as a rule are somewhat paler than in A. Guvieri, the stout ventral bristles of which are
sometimes even dark brown.

The first segment is larger than the buccal, but less than those which follow. Like
the rest the foot is dorso-lateral in position, the superior division indeed being wholly
dorsal, and consisting of a tuft of transversely serrated (camerated) and finely tapered
bristles with a cirrus behind and rather towards the outer border. The ventral division
is dorso-lateral, and it has a cirrus and a long papilla with a group of the long tapering
camerated bristles, and a series of strong and short curved forms with blunt tips generally
smoothly rounded, and having spinous rows (abraded) on the shaft below. Many of these
are arranged at an angle to the longer forms, and the shorter forms are ventral. The
second foot has longer dorsal bristles and a longer and larger cirrus, whilst fewer long
serrated bristles occur in the ventral division, which has three papillae. The third foot
has a few tapering serrated bristles superiorly and a thinly scattered series along the rest
of the line. All are shorter than those in front. The stout abraded forms with the
rounded ends and serrations on the convex edge of the curve (as on the tapering bristles)
extend from the upper to the lower edge of the division. The upper papilla is longest
(and may represent the cirrus), and seven shorter follow. The fourth foot has still fewer
and shorter tapering serrated bristles, but the curved stout forms are more conspicuous.
The one increases in strength as the other wanes.

The tenth foot has a much longer dorsal cirrus and a series of strong dorsal bristles.
The ventral division has very short tapered forms here and there along the line, the
whole length of which bristles with the stiff curved and serrated forms, the tips of which,
moreover, are more or less acute. About fifteen papillae occur along the line. The
twentieth foot has a longer and denser tuft of dorsal bristles, and the cirrus has increased
in size. The short tapering serrated bristles are still present along the line, whilst the
curved spine-like forms are fully developed and thicker than in front. Many of the points
are sharp, while others are rounded and blunt. The papillae are more numerous, and the
line is longer.

At the twenty-ninth foot the dorsal cirrus has assumed the form of a bellied knife,
that is, whilst its inner edge (that next the branchia) is more or less straight, the outer
springs from a broad base, curves outward, and then backward to the subulate tip. The
row of the large curved yellow spines is now short, but traces of serrations are still visible,
and the short serrated bristles accompany them. The papillae are conspicuous, the
longest being* that next the dorsal division, and another a little to its exterior.

The knife-like shape of the dorsal cirrus is conspicuous at the thirtieth foot, and so
is the great tuft of serrated (camerated) tapering bristles. Externally after an interval
is a long cirrus-like papilla, and a somewhat shorter one to the exterior. Yentralwards
little is seen except the row of papillae but the tips of a few tapering serrated bristles at

504 ARICIA LATEEILLII.

and below the second or shorter papilla just mentioned. No strong spine-like forms can
be observed. The dorsal division of the thirty-first foot is like the preceding, but the
cirrus-like papilla to its exterior is larger, whilst its neighbour, also above noticed, has
now assumed a conical outline, is much larger, and has a broad base, whilst a small tuft
of long, slender, and finely spiked bristles projects from a furrow between a slight elevation
and its base. The papillae are present along the edge of the foot beneath, but all trace of
the curved spine-like forms and short slender bristles has vanished.

The papillae behind the feet pass down to form a continuous row from side to side
for seven segments in front of the thirty-first, besides several imperfect rows before these,
and for five segments behind the thirty-first.

The type of foot just described is modified only in a slight degree in its progress
backward to the tail. Thus at the fortieth foot the cirrus next the dorsal division is
longer, and the ventral setigerous process is bifid, whilst a ventral cirrus appears at its
lower border. The dorsal bristles have forked bristles with probe-points and spinous
inner edges amongst the others. The length of the ventral bristles has rather increased
and the spikes on the tips are directed distally.

At the ninetieth foot the branchia has increased in size, whilst the dorsal cirrus is
narrower and smaller.

In the feet some distance in front of the tail the branchia has notably increased in
size, and the dorsal cirrus is still knife-shaped, the intermediate cirrus being filiform, as
also is the tip of the long horn of the setigerous region and the ventral cirrus. The long
and strong dorsal bristles are only faintly serrated with spikes distally, and the forked
bristles continue. The tips of the ventral bristles are similar.

No noteworthy change occurs till the diminution of the parts at the tip of the tail.

In an example of this species procured in sand at low water, 15th June, 1904, there
were respectively thirty-one and thirty-three of the anterior segments with the deep-
brownish bristles. The eighth on the right, from injury, was out of line, and the seventeenth
was double, also apparently the result of an injury. The rows of ventral papillae began on
the twentieth and continued to the thirty-fifth, at which the specimen abruptly terminated.

Reproduction. — In a female procured on 15th June the body ventrally and laterally
was of a dull yellow or buff colour from the masses of dull yellowish eggs, which occurred
in each segment. The eggs formed masses at the bases of the feet and were visible through
the dorsal wall. Such a condition seems to be common at this time. In section the ova
occupied not only the coelomic space on each side of the alimentary canal, but passed on
each side of the middle line between the dorsal longitudinal layer and the circular coat of
the region.

Habits.— This species is a borer in sand, a life for which its mobile, pointed snout
and the conformation of its elongated body adapt it. It is not easy, however, to secure
perfect examples, especially if attempts are made to remove them from the hard sand.
When lying free in the water they sometimes keep up an undulatory movement of the
body as if for respiratory purposes.

1 • Mitth. Zool. Stat. Nap./ xiii, p. 484.

ARICIA GRTJBBI. 505

4. Arioia Guubei, n. s. Plate LXXXV, figs. 8-8 b— bristles ; Plate LXXXVI, fig. 1

— foot.

Specific Characters.— Read typical. Anterior region of fifteen bristled segments.
Each foot has a dense tuft of camerated or spinose bristles finely tapered, with a stout
dorsal cirrus posteriorly. An interval separates it from the ventral division, which has
a series of similar but shorter bristles, many of which have shafts with rounded tips.
The last three feet of the region have dark brownish spines with long hastate tips. The
rows of papillae posteriorly are separated by an interval from the bristles. These papillae
occur on the ventral surface of bristled segments thirteen, fourteen, fifteen, and sixteen,
and it may be in one or two succeeding segments. The twenty-third foot has the broadly
lanceolate branchia, then the elongated dorsal cirrus shaped like a pointed shoe, with a
tuft of long serrated (camerated) bristles, amongst which are some simple forms. There
are four spines. In the interval between this and the ventral division is a short and
rather broad conical papilla or cirrus. The ventral division has a somewhat clavate
setigerous region supported by two spines, and a pointed lanceolate lobe attached to it
distally and projecting in front of it. The ventral cirrus forms a broad process, some-
times with a pointed tip to the exterior of its base at the twenty-third foot. The dense
bristles of the dorsal division anteriorly are transversely spinous or camerated forms;
those in the upper series of the ventral division are shorter, but of similar structure
mixed with broken shafts with rounded tips like stout spines. The last three anterior
feet have four dark-brownish hastate spines. At and after the twenty-third foot slender
tapering serrate (camerated) bristles, a few simple bristles, and four spines are found
dorsally, whilst the ventral division has a few very slender bristles with minutely spinose
tips, but none were perfect.

Habitat. — Dredged in the 'Porcupine' Expedition of 1869 in 422 fathoms on a
bottom of sand, stones, and coral. ' Porcupine ' Expedition of 1870 at Station VI in 358
fathoms ; and at Station VIII in 257 fathoms.

The proboscis is extruded as a foliaceous button.

A comparatively small form, with the typical arrangement of snout anteriorly, and
about fifteen of the largely developed anterior feet with vertical rows of ventral bristles.
The anterior feet, e.g., the tenth (Plate LXXXVI, fig. 1), have a dense tuft of camerated
bristles finely tapered (Plate LXXXV, fig. 8) and a stout dorsal cirrus, then an interval
devoid of appendages. The inferior division has a more or less complete series of similar
though shorter bristles, and also of broken shafts with rounded tips, more or less modified
so as to perform the functions of stout spines (Plate LXXXV, fig. 8 a), whilst the majority
of the anterior feet have pale or slightly yellowish bristles in the inferior division; the
last three have about four dark brownish hastate spines with long tips (Plate LXXXV,
fig. 8 b). The row of papillae behind these feet is separated by a considerable interval
from the bristles, so that the foot has a character of its own. Rows of papillae occur on
the ventral surface of bristled segments thirteen, fourteen, fifteen, and sixteen, and it may
be partially on one or two others behind.

The change in the form of the foot takes place about the seventeenth, and at the

121

506 ARICIA NORVEGICA.

twenty-third the foot has dorsally on each side of the median line the broadly lanceolate
branchia, then the elongated dorsal lobe, which is shaped somewhat like a pointed shoe,
with a tuft of slender, tapering, serrate (camerated) bristles, amongst which are some
simple forms, and with four spines. In the interval between this and the ventral division
is a short and rather broad conical papilla. Then the bifid ventral lobe follows with a
somewhat clavate setigerous region supported by two spines, and a pointed lanceolate
lobe in front of it. The bristles were almost wholly abraded, but such as remained
conformed to the elongated slender forms with finely tapered tips and minute serrations
directed distally.

At the base of the ventral division is a bluntly pointed lamella apparently
homologous with the ventral cirrus.

No forked bristles were observed, but the condition of the examples was not
favourable.

This form, therefore, differs from the Aricia Kupfferi of Bhlers in the occurrence of a
short cirrus between the dorsal and ventral lobes, though it also has hastate spines in the
last three feet of the anterior region. It likewise differs from A. norvegica in having
the intermediate cirrus, and in the shape of the ventral cirrus.

5. Aricia norvegica, If. Sars, 1873. Plate LXV, fig. 3— head; Plate LXXXV, figs.
9-9 b— bristles ; Plate LXXXVII, figs. 2 and 2 a— feet.

Specific Characters. — Head a short cone as in the type. Anterior region of the body
has fifteen bristled segments. Branchiae commence on the fifth. Dorsal cirrus flattened,
slightly enlarged towards the base, and tapered distally. Anterior rows of ventral
bristles pale, the last three or four feet with dark brown hastate spines. Rows of papillse
behind the foot more acutely pointed than in the previous forms, and eight to twelve in
number (Sars). Form and colour as in A. Cwvieri (Sars). At the twenty-third foot the
dorsal division has four or five spines and a long tuft of slender bristles with finely
tapered serrated tips. The cirrus is narrow and has a tapered extremity. The ventral
division has one spine and a small group of equally slender bristles, the tips of which are
serrated. The blunt setigerous lobe has a conical lobe to its outer side, whilst at the
base externally is a conical ventral cirrus nearly equal in size to the last mentioned.

Synonyms.
1871. Aricia norvegica, M. Sars. Yidensk.-Selsk. Forhandl. for 1871, p. 5 (sep. copy).

1873. „ „ idem. Bidrag Christ. Fauna, iii, p. 36, Tab. xvi, f . 1 — 8.

1874. „ „ Malm. Goteb. vet. o. vitt. HandL, xiv, p. 91.

1875. „ „ Mobius. Jahresb. Comm. deutsch. Meere, p. 160.

1877. „ groenlandica, Mcintosh. Trans. Linn. Soc, Zool., i, p. 504, pi. lxv, f. 5—9.

1879. „ norvegica, Tauber. Annul. Danic, p. 106.

„ „ Hansen. Nyt Mag. f. Naturvid., xxiv, p. 269.

1893. „ „ ' Levinsen. Vidensk. Ud. " Hauchs," p. 336.

1905. „ „ Mcintosh. Ann. Nat. Hist., ser. 7, vol. xv, p. 44.

Habitat— Hi. Magnus Bay, Shetland (J. G. Jeffreys).

ARICIA NORVEGICA. 507

Drobak, Norway (Sars). Station XL VII, off the coast of New York (' Challenger'),
at the great depth of 1340 fathoms in bine mnd. Also in 1200 fathoms off Rio, San
Francisco, in mnd. Jan Mayn (Hansen). Variety from the ' Valorous ' Expedition, 1875.
Bergen, Norway (Canon Norman).

Head (Plate LXV, fig. 3) a short cone as in other species.

Body typical in outline. The anterior region consists of fifteen bristled segments
(Sars says fifteen to sixteen). The branchiae commence on the fifth bristled segment.
The general aspect of this (anterior) region differs from that of the allied species (1) in the
flattened condition of the dorsal cirrus, which occasionally has a filiform process at its
outer edge ; (2) by the pallor of the anterior rows of ventral bristles and the large dark
brown spines of the last three or four ; (3) by the more acutely pointed papillae in the
row behind the foot (ventral division) and the tendency to show a larger area of bristles
and flap. Along the dorsum is a series of brownish dots at the segment- junctions, but
these were not observed in the spirit-preparations.

The proboscis forms a frilled button like a rosette, the folds passing to the central
aperture.

The first foot has two papillae, each with a short tuft of tapering camerated bristles.
The foot is thus bifid from the commencement, a condition not always seen in allied
families. At the tenth foot (Plate LXXXVII, fig. 2) the dorsal cirrus has a broadly
lanceolate outline with a long tip, and the shafts of the long bristles are smooth, the
tapering tips being camerated (Plate LXXXV, fig. 9). The bristles of the inferior
division form a dense series, and are short, tapering, and camerated. The posterior
papillae have tapered tips. The twelfth, thirteenth, fourteenth, and fifteenth feet show
numerous dark brown hastate spines (Plate LXXXV, fig. 9 a) ; the posterior papillae are
sometimes bifid. One of the spines (the upper) usually projects freely.

Amongst the dorsal bristles are some bifurcate spinous forms (Plate LXXXV, fig.

9 b).

The anterior branchiae form broad, tapering, flat processes, nearer each other than
behind, where they are considerably longer and narrower, and leave much of the dorsum
bare. Posteriorly they greatly increase in length and have filiform tips.

At the twenty-fifth foot, which shows the average condition after the change,
a considerable interval occurs between the branchia and the dorsal division, which has
four or five spines and a long tuft of slender bristles with very finely tapered smooth tips.
The cirrus is narrower and has a long tapered extremity. The ventral division has one
spine and a small group of equally slender bristles, the tips of which, however, are serrate.
The blunt setigerous lobe has a conical lobe to its outer side, whilst at its base externally
is a ventral cirrus, also of a conical outline and approaching the former in size.

Tauber 1 (1879) appears to think that the Aricia Kujpfferi of Ehlers is a variety of
this form which certainly agrees with it in the absence of the cirrus between the dorsal
and the ventral divisions of the foot. Both have a similar ventral cirrus, and the general
characters agree except that the ventral surface of A. norvegica has no rows of papillae,
whereas that of A. Kupfferi has. The absence of the intermediate cirrus just referred

1 ' Armulat. Danic./ p. 106.

508 ARICIA ARMANDI.

to is characteristic of A. foetida of the Mediterranean, as well as of a fragmentary form
from Malahide.

The Aricia gramlandica 1 of the cruise of the ' Valorous ' appears to be a variety of this
species.

6. Aricia Armandi, 2 n. s. Plate LXXXVII, figs. 19 and 20— feet.

Specific Characters.— Snout somewhat truncated (?). Body, so far as known, of the
typical outline; anterior region of nineteen segments, and the ventral spinous areas are
pale yellow. Branchiae arise on the fifth foot. Foot differs from A. Guvieri in the
absence of the cirrus between the dorsal and ventral divisions, and in the great length
of the papilla in the ventral rows. From A. Kupfferi and A. norvegica the species
diverges in the number of the anterior segments, and in the form and arrangement of
the papilla behind the ventral division of the foot, and on the ventral surface of certain
segments.

Habitat. — Dredged in St. Magnus Bay, Shetland, in eighty fathoms, by Dr. Gwyn
Jeffreys in 1868.

The specimen is in a softened and fragmentary condition, so that the condition of
the head is unsatisfactory. The snout is truncated, the tip of the usual cone being
absent, but whether this is natural or due to injury is uncertain.

The anterior region of the body, which is of comparatively small size, consists of
nineteen segments, though the softened state of the parts gives insecurity as to a segment
in front. The ventral spinous areas are of a pale yellow hue. The general outline is of
the normal type, but only the anterior region and a portion of the succeeding about the
same length are present. The branchiae appear to arise on the fifth foot. The long
papillae are continued a short distance downward at the seventeenth foot and pass nearer
the middle line on the eighteenth, nineteenth, and twentieth feet, diminishing again on
the succeeding two or three feet, the region thus having a remarkably hirsute aspect from
their comparatively great length.

The first foot is injured dorsally, only a tuft of long, slender, serrated bristles
remaining. The ventral division has a short row of spine-like stumps of the moderately
strong serrated bristles with the tips smoothly rounded, whilst the edge of the bristle
beneath is faintly serrated. A few tapering serrated bristles of moderate length are
likewise present along the row. At the ninth foot the dorsal cirrus seems to be of
moderate length, and the dorsal bristles are long, serrated (camerated), and tapered.
The upper process of the ventral division is longer than the others and cirrus-like,
a considerable interval occurring between it and the dorsal division. A dense series of
strong spine-like bristles occurs along the edge, the upper being stouter than the lower.
Each is a stout process, curved at the tip, which is smooth and blunt. No trace of the
serrated edge beneath is now seen. By transmitted light they are dull yellow. A few
serrated tapering bristles are in the row bounded posteriorly by the papillae, which are

1 ' Trans. Linn. Soc, Zool./ vol. i, p. 504, pi. lxv, figs. 5—9, 1877.

2 Named after M. Armand De Quatrefages, who for many years was the leading authority on
the Annelids in France.

SCOLOPLOS. 509

largo and tapered, and project considerably beyond the spines. Little change occurs at
the sixteenth foot except that the bent tips of the spines towards the middle and lower
part of the row are slightly clavate, that is, bluntly rounded and somewhat enlarged.
The posterior papillae are still tapered and rather long. The seventeenth and eighteenth
feet are similar, but have fewer bristles, thus making a shorter row. A few short,
tapering, serrated forms are also present ; moreover, there are a few bifid forms with
the limbs well separated and with a mucro at the tip, their inner edges being spinous.
They radiate from a narrow base, as the long bristles do, only their tips are curved and
modified for special service. The dorsal cirrus is elongate-lanceolate, the tips of the
bristles being serrated (camerated), whilst the long shaft is smooth. The papillae are very
conspicuous, and at the nineteenth foot they have the margin of the division to them-
selves, for the stout and modified spine-like bristles are few and project little, whilst near
the upper edge of the ventral division a few very slender and apparently smooth bristles
with finely tapered tips project. Amongst these is a short (broken ?) slightly bent shaft
with a rounded tip. These bristles would appear to be modified by their surroundings,
as well as by the adaptability of the tissues which form them. The dorsal cirrus is
a conspicuous flattened process, and the dorsal bristles are longer, the whole foot
initiating the change, though the projection of the spinigerous lobe is scarcely marked,
At the twenty-third foot the alteration is complete, the large branchia being internal,
then follows the somewhat subulate dorsal cirrus, with a slight enlargement on its external
edge, and the long dorsal bristles with slight serrations on their finely tapered tips,
accompanied by a few bifid forms. The ventral lobe is clavate, bifid at the tip, the
shorter process being rounded and containing the spine, whilst the other (ventral) forms
a considerable subulate appendage. The bristles are few, slender and tapering, scarcely
a trace of serrations being visible. The ventral cirrus is a subulate process resembling
the appendage above it.

Whilst approaching Aricia Cuvieri this form differs in the number of the anterior
segments, the absence of the cirrus between the dorsal and ventral divisions, and in the
great length of the papillae in the ventral rows. The number of the anterior segments
likewise differentiates it from A. Kujpfferi and A. norvegica, as also do the form and
arrangement of the papillae behind the ventral division and on the ventral surface
of certain segments.

Genus LXXVIII. — Scoloplos (Blainville), CErsted.

Head pointed, peristomial segment a blunt cone. Proboscis a foliate rosette. Body
widened anteriorly, flattened, diminishing posteriorly, and ending in a vent with four
papillae and two cirri. Regions of the body two, anterior and posterior ; the former with
the feet lateral, the latter with the feet dorsal. Branchiae dorsal, ligulate, decreasing in
size posteriorly and disappearing. Inferior division of the foot in the anterior region
with a small papilla. Long, tapering serrated dorsal bristles, and stout, serrated spine-
like bristles ventrally. In the posterior region the bristles are all slender and serrated.
The structure of this type generally approaches that of Aricia, though the rows of papillae
on the feet are absent.

510 SCOLOPLOS ARMIGER.

Scoloplos armiger, 0. F. MUler, 1 788. Plate LVI, fig. 7 ; Plate LXV, figs. 4 and 4 a—
head and tail; Plate LXXXVI, figs. 1 and 1'— bristles; Plate LXXXVII fi>s 4-
4 5— feet. ' 5 •

Specific Characters.— -Head an acutely pointed cone set on the larger truncated cone
of the peristomial segment. Often with deep-seated dark specks at the base. Body
flattened and widened anteriorly, then rounded ventrally, and flattened dorsally in its
progress to the tail, which ends at the vent surrounded by four papilla?, whilst on each side
is a long slender cirrus. Anterior region of about eighteen segments with lateral feet,
having shorter, stouter, serrated bristles. The fifteenth foot is composed of a conical
dorsal cirrus or lobe, with, in front of it, a tuft of rather long bristles, the shafts of which
are smooth. The tapering tips are peculiarly spiked and end in smooth, hair-like points.
The ventral division has two short conical papillaa, one of which is below the bristle-
tuft and probably represents the ventral cirrus, whilst the other is behind the middle of
the dense, deep row of bristles, which are shorter than the dorsal but of the same type.
Feet posteriorly are dorsal, the branchia being nearest the middle line, then follows the
dorsal cirrus of the shape of a bellied knife and about half the length of the branchia, and
lastly the dorsal fascicle of serrated bristles. The ventral division is a little shorter and
boldly bifid, the upper process being the longer. The small group of slender bristles
emerges between them.

Branchias commence in the form of a small papilla on or about the seventeenth
bristled foot, attain considerable size on the twentieth foot, and almost reach the tip of
the tail. Below the foot from the eighteenth bristled segment to the thirtieth (fewer in
some) a papilla occurs on the side of the body.

Synonyms.

1788. Lumbricus armiger, 0. F. Miiller. Zool. Danica, i, p. 22, Tab. xxii.
1828. Scoloplos „ De Blainville. Diet. Sc. Nat., lvii, p. 493.

1842—3. „ „ (Ersted. Kroyer's Nat. Tids., p. 125.

1843. Aricia MueUeri, H. Ratlike. Nova Acta Leop.-Car., xx, i, p. 176, Tab. viii, f. 9 — 15.
„ Scoloplos armiger, (Ersted. Annul. Danic. Consp., p. 37, f. 8.

idem. Gronl. Annul. Dorsibr., pp. 106, 107, 109, 201, f. 113, 117, 118.

1844. „ ,, idem. Arch. f. Naturges., x, i, p. 104.
„ „ „ idem. Regionis Marinis, p. 78.

1844—5. ,., „ idem. KroyerVNat. Tids. (Drobak), p. 412.

1851. Aricia armigera, Grube. Fam. Annel., pp. 68 and 135.
1853. „ „ Sars. Nyt Mag. f. Naturvid., vii, p. 381.

1856. ,, „ Koren. Ibid., ix, p. 95.

1858. ,, armiger, Danielssen. Reise, 1857, p. 53.

1859. „ „ idem. Ibid., 1858, p. 129.

1865. 3 , armigera, De Quatrefages. Annel., ii, p. 286.
1867. Scoloplos armiger, Malmgren. Annul. Polych., p. 72.

1869. „ „ Mcintosh. Rep. Brit. Assoc. (1868), p. 338.

1870, „ „ Grube. Arch. f. Naturges., xxxvi, p. 316.

SCOLOPLOS ARMIGEP. 511

1871. Scoloplos armiger, Mobius. Bericht. Exped. ( Pommerania/ p. 197.
„ „ ,, Ehlers. Sitz. Phys.-Med. Soc, Erlangen., iii, p. 79.

1873. „ „ Sars. Nyt Mag. f. Naturvid., xix, p. 240.

„ „ „ Mobius. Jahresb. Comm. deutsch. Meere, p. 107.

1874. „ „ idem. Die Zweite deutsch. Nordpol., p. 255.
„ „ ,, Malm. Groteb. vet. o. vitt. Hand]., xiy, p. 91.

„ ,, „ Mcintosh. Ann. Nat. Hist., ser. 4, vol. xiv, p. 200.

1875. „ „ idem. Invert, and Fishes St. Andrews, p. 124.

„ „ armiger, Mobius. Jahresb. Comm. deutsch. Meere, 1874, p. 160.

1877. „ „ Hansen. Nyt Mag. f. Naturvid., xxiv, p. 6.

1878. „ „ Lenz. Jahresb. Comm. deutsch. Meere, 1874 — 5, p. 11.
„ ,, „ Mcintosh. Trans. Linn. Soc, ZooL, i, p. 504.

1879. „ ,, Hansen. Nyt Mag. f. Naturvid., xxiv, p. 6.

„ „ „ Theel. K. sv. vet. Akad. Handl., Bd. xvi, No. 3, p. 45.

„ „ „ Tauber. Annul. Danic, p. 106.

1881. „ „ (acoustic capsules), Mau. Zeitschr. f. wiss. ZooL, Bd. xxxvi, p. 389, pi.

xx vi and xxvii.

1882. Aricia arctica, Hansen. Nord. Nordhav. Exped., vii, p. 34, Tab. v, f. 20 — 26.

1883. Scoloplos arctica, Wiren. Chast. ' Vega ' Exped., p. 405.

1888. „ armiger, Cunningham and Ramage. Trans. Roy. Soc. Edin., p. 200, vol. xxxiii, p.

642, pi. xxxviii, f. 7.

1889. Aricia armigera, Marenzeller. Arch. f. Naturges., lv, p. 132.

1891. „ „ Hornel. Trans. Biol. Soc. Liverp., vol. v, pp. 249 and 252. c

1894. Scoloplos armiger, De St. Joseph. Ann. Sc. nat., 7 e ser., t. xvii, p. 94, pi. v, f. 119 — 120.

1897. Aricia armigera, Birula. Ann. Mus. Zool. FAcad. Imp. St. Petersb., p. 21.

1898. „ Millleri, De St. Joseph. Ann. Sc. nat., 8 e ser., t. v, p. 356, pi. xx, f. 167.
1901. Scoloplos canadensis, Whiteaves. Geol. Surv. Canada, No. 722, p. 79.

1904. „ armiger, Allen. Journ. M. B. A., N.s.,. vii, p. 227.

1905. „ Millleri, Mcintosh. Ann. Nat. Hist., ser. 7, vol. xv, p. 45.
„ „ armiger, idem. Ibid., p. 54.

1909. „ „ Moore. Proc. U.S. Nation. Mus., vol. xxvii, p. 139.

Habitat. — Abundant between tide-marks at Lochmaddy, Nortb Uist, and also dredged
on a sandy bottom in the Laminarian region. On a sandy bottom at Paible, North Uist,
in four to five fathoms. In the sandy mud of the bathing pond at the Step-Rock, St.
Andrews. Dredged in four to five fathoms and also in fifty fathoms off Balta, July, 1867
(J. Gr. Jeffreys). Common in Herm, in sand between tide-marks, August, 1868, and in a
similar region in Cobo Bay, Guernsey. Off Symbister Harbour, Whalsey, Shetland,
in three to five fathoms, July, 1871. Fragment from Malahide, Co. Dublin (A. C.
Haddon).

Extends to Finmark (A. M. N.); Baltic (Mau); Nova Zembla (Theel); and to Canada
(W. C. M.); Labrador (Moore); Greenland (CErsted) ; and 1100 fathoms in 'Valorous,'
1875. Siberian Coast (Wiren). Northern seas generally (Malmgren).

Head (Plate LXV, fig. 4) forms an acutely pointed cone without rings or subdivisions,
though when actively thrust amongst the surrounding structures it frequently presents
crenations at the edges. In some, four deep-seated dark specks occur in front of the
mouth arranged in a crescent, the posterior pair being nearest each other. In others there
is less definition, the dark specks (often two) being irregular. It is doubtful if these are eyes.

512 SCOLOPLOS ARMIGER.

The peristomial segment follows as a larger truncated cone. According to the degree of
extrusion of the proboscis a rounded or a somewhat cylindrical process projects from the
mouth, or a foliate rosette of six or seven lobes, like those of Anthostoma, lie over (i.e.
beneath) the mouth.

Body 3 to 6 or more inches in length, flattened and widened in the anterior region
before tapering to the pointed snout. Behind this anterior region the body is rounded
ventrally and flattened dorsally, and it continues so to the tail. The segments are distinctly
marked. Colour of a deep but not a bright red, inclining to orange posteriorly. Others
are of a general orange colour. The dorsum has a bluish-green iridescence anteriorly. In
ripe males the posterior region is cream-coloured. The body diminishes considerably
posteriorly and terminates at the vent, the rim of which apparently is divided into four
papilla3, whilst on each side is a long slender cirrus. In an example from Symbister
Harbour, Whalsey, Shetland, the tail (Plate LXV, fig. 4 a) has in addition to the papillae
(four) two other processes besides the cirrus, but as this tail had been reproduced the
exact nature of these additional processes is uncertain, and they may have been abnormal.
In another example from the same site the tail corresponded with the type, viz., had four
small lobes or papillae and two long cirri. These cirri show peculiar hypoderm, the
granules and cells towards the tip often being isolated. In all the slender posterior region
has proportionally much longer branchiae and foot-processes, so that it is hispid or fringed
in a characteristic manner.

The muscularity of the body- wall anteriorly is great, the vertical and transverse
muscles being apparently strong. As soon as the nerve-cords reach the ventral surface
they lie under a broad band of muscle continuous in the middle line and apparently
homologous with the oblique muscles. When the body-wall is completely formed the dis-
proportion between the dorsal and ventral longitudinal muscles is great. The dorsal are
comparatively thin and narrow, whereas the ventral form nearly half the area of the
body in section. Each is a massive muscle stretching almost to the dorsal edge of the
body, and increasing in diameter to the mid- ventral line, where it is deepest. The nerve-
cords are carried inward, and are connected with the hypoderm only by a very narrow
pedicle. The area is ovoid, with a large neural canal superiorly. The inner border of
each muscle often separates along with the nerve-area in section, a condition which,
perhaps, occurs more easily from the fan-like series of fasciculi, which traverse the muscle
from the great vertical band on each side of the gut. The oblique muscles, which are
fairly developed, are fixed to the outer edge of the upper arch of the area.

Very fine examples from Lochmaddy, North Uist, have the median ridges of segments
running from the feet ventrally well developed, especially posteriorly. In a specimen
from Balta (four to five fathoms) the lateral fillets below the ventral division of the foot
are tinted brownish in the spirit-preparation, as if from pigmentary deposit, from the
twenty-first foot to the fifty-first, the end of the fragment.

The anterior region consists of about eighteen segments, having shorter, stouter,
serrated bristles. The feet are little developed, though from the first segment backward
two bristle-bundles, which soon assume distinct characters, are present. Thus at the
fifteenth foot the dorsal division has a conical dorsal cirrus of considerable size, whilst in
front of it, and arising from a rounded elevation, is a tuft of rather long bristles, the shafts

SCOLOPLOS APMIGER. 513

of which are smooth (Plate LXXXVI, figs. 1 and 1 '),but the tapering tips are peculiarly
spiked, and they terminate in smooth hair-like points. Four somewhat short spines
support the bristles. The ventral division has two short conical papillae, one of which is
below the bristle-tuft, and is probably the representative of the ventral cirrus, whilst the
other is a little below the middle of the bristles, which form a dense and deep row, shorter
than the dorsal but of the same type. At the ninth foot, however, the papillae are
smaller, and the inferior division has numerous short strong bristles with somewhat blunt
tips, which are strongly curved and serrated on the convex side. They are modified
forms of the longer series, and correspond with the stronger series in Aricia. The same
is seen in the tenth foot (Plate LXXXVII, fig. 4).

The branchiae commence in the form of small papillae from the fifteenth to the
seventeenth foot, attain considerable size on the twentieth foot, rather increase in pro-
portional length in their progress backward, and almost reach the tip of the tail.

Below the foot from the eighteenth bristled segment to about the thirtieth (fewer in
some) a papilla occurs on the side of the body. It varies in prominence, it may be in
some cases from preparation, and is very well marked in Zetlandic examples from
Symbister. This feature is variable. Thus, in an example from Lochmaddy the papilla
appears on the seventeenth bristled segment. At the twenty-fourth and twenty-fifth it
has become scale-like, and adheres to the front edge of the ridge from the foot.

The eighteenth bristled foot shows considerable alteration. Externally, that is,
ventrally, is a small papilla, then a bifid process with the bristles. The dorsal division
consists of a longer conical papilla with a fan-shaped tuft of bristles. The ridge of the
segment below the feet also becomes more prominent in this region.

At the twentieth foot (Plate LXXXVII, fig. 4 a) the ventral division presents a
somewhat lanceolate outline, and the bristles are slender and almost smooth. The dorsal
papilla is constricted at the base, then dilated, and slightly tapered to the tip. It bears a
fan-shaped tuft of long bristles, with smooth shafts and tapering serrated tips. It
adjoins the branchia.

The thirtieth foot (Plate LXXXVII, fig. 4 b) has ventrally a bifid setigerous region,
with a rounded enlargement at the base ventrally. The bristles are few, long, slender,
and very finely serrated — quite different in aspect from the dorsal. The dorsal lobe is
constricted at the base, with a lanceolate terminal region tapering to a point. The
bristles have smooth shafts and boldly serrated tips. The branchia is a richly ciliated,
flattened lanceolate organ, with central vessels which apparently have pinnate lateral
branches.

Cunningham and Ramage figure l a bifid bristle as occurring in this species, where, it
is not stated. A mistake may have happened, especially as the inner edge of the fork is
smooth.

On the whole, while there are certain differences in the proportional length of the

1 'Trans. R. Soc. Edin./ vol. xxxiii, pi. xxxviii, fig. 7 e. This recalls the case of a foreign
observer, who showed this kind of bristle in another species in a proof of the plate. The absence of
these spikes was pointed out, the author asserting that the limbs of the fork were smooth. When the
paper and its plates subsequently appeared the spikes had taken their proper places.

122

514 SCOLOPLOS ARMIGER.

dorsal cirri and the papilla anteriorly, as well as variations in the length of the bristles,
specific separation of the northern and southern representatives does not seem to be
necessary or expedient. The following remarks will indicate the leading features relating
to the varieties : —

The variety Mulleri (the Aricia Mulleri of De St. Joseph) is characterized by the
frequency of shorter bristles in the anterior feet — such bristles as are represented by
De St. Joseph in his figure 167, Plate XX. The structure thus figured and described,
however, might be explained by the occurrence of injury to the tapering bristles and the
rounding of the broken tips. As formerly observed, 1 they are modifications of the longer
series, not structures sui generis, and are apparently due to influence of habitat, e.g.,
constant friction, and they vary much amongst themselves. De St. Joseph has the credit
of drawing special attention to them, and to the fact that in the first six segments of the
posterior region of the body the ventral division has two protuberances below it, with a
minute elevation near them, and a conical papilla on the body below. It is not explained
that a similar papilla occurs in the typical Scoloplos armiger from Greenland and the
northern waters. The two forms very closely resemble each other, the main features
which differentiate them being the somewhat longer rows of bristles in the southern form,
and the better developed fleshy ridge behind them with its flattened conical elevation in the
ventral division, and the two papillse just below it ; whilst the ventral row of bristles has
a variable number of short and somewhat truncate forms, which must not, however, be
regarded as specific. They are only modified forms of the ordinary tapering serrate kind
belonging to the division, and, as stated, are probably due to circumstances of habitat, and
are perhaps more abundantly present in those from the Channel Islands than in the more
northern examples. In this form a papilla appears in the middle of the fifth foot, and a
conspicuous one from the tenth to the thirteenth. At the seventeenth foot two papilla
occur, by the addition of one below the median, whilst the eighteenth has three, the
bristles being above (dorsal to) the upper. At the nineteenth foot the lateral ridge ends
dorsally in three conical processes, with a thick papilla beneath. In very fine examples
from Lochmaddy the papilla commences on the fifteenth, and continues to the twenty-
second foot.

In the typical Scoloplos armiger from Greenland, the first fifteen anterior feet have
somewhat shorter (i.e., from above downward) ventral rows of bristles. The dorsal cirrus
is a well-marked conical process, which gradually increases in length posteriorly. The
fillet behind the ventral row of bristles forms a convex flap, highest in the middle, and
without any evident differentiation till the fifteenth foot, where a papilla projects from its
median convexity, one or two of the preceding feet showing a slight thickening at the
same part. The row of ventral bristles is shorter. Though the bristles are longer, the
row is still shorter at the sixteenth foot, and a papilla occurs on each side behind it. The
seventeenth foot has three papilla, whilst the eighteenth has the enlarged lateral fillet
with an isolated papilla below it, and this for three or more segments. A survey of a
series of specimens from Greenland shows considerable variations in the second division
of the body in the shape of the dorsal cirrus, which may be fusiform or lanceolate and
stalked, and in the length of the bifid ventral lobe. Yet these and other features are

1 'Ann. Nat. Hist./ 7th ser., vol. xv, p. 45.

SCOLOPLOS ARMIGER. 515

found in those from St. Andrews, Lochmaddy, the Channel Islands, and Shetland. These
minute differences, and especially those mentioned by De St. Joseph, indicate the slender
grounds on which specific distinctions may be founded.

Reproduction. — Hornel (1891) on 2nd March procured, on a sandy beach at Egre-
mont, small gelatinous pear-shaped brownish egg-masses, the annelid being in close
proximity to them. The brownish cocoons are smaller than the green ones of Arenicola,
and are anchored amongst the bare ripple-furrows of the sand, the cylindrical stalk
descending two or three inches into the sand. He watched their development, and
corroborates Cunningham and Ramage, 1 who found the gelatinous transparent cocoons
with opaque white eggs and embryos in the estuary of the Forth at low water in
February. The larvae had three transverse bands of cilia, a well-developed prostomium
with two eyes, and by-and-by a pair of anal cirri as well as dorsal cirri appear. Similar
cocoons and eggs were found by Leschke 2 in Kiel Bay in May. Late larvae are found
from July to September at St. Andrews (1890). 8

Garstang, 4 again, mentions February as the breeding season at Plymouth.

The males are distinguished in January, when the body is distended with ripe sperms,
by the cream-coloured posterior region.

In January the females are recognized by their buff-coloured bodies filled with
similarly tinted ova, which, though far advanced, vary considerably in size — probably
because they are in different stages.

Habits. — When removed from their haunts in the sandy mud they push their finely
pointed snouts ceaselessly around it, and often coil their bodies into a spiral. If a little
sand is in the vessel it is soon transformed by the aid of the mucus into a central mass
permeated by the bodies of the examples, while the heads are pushed in various directions.
Their food seems to be the sandy mud in which they dwell.

0. F. Midler gave the anterior region seventeen segments, and he found the species
in mud. To what group the Lumbricus squamatus 5 of Abildgaard belongs is doubtful.
It may be one of the Spionidae.

Auclouin and Edwards 6 describe an Aricia sertulata which is probably synonymous
with 0. F. Midler's Lumbricus armiger.

(Ersted 7 (1843) gives the anterior region fifteen segments with dark bristles. In
the succeeding segments the ventral pinna is bifid, the superior acuminate, and the
ligulate branchiae thrice as long, the ciliated margin regularly decreasing and finally
vanishing towards each end of the body. Subulate bristles in all the segments. Tail
truncate without cirri. In this description there is not much at variance with the
common British examples, for considerable variation occurs in the number of the anterior

1 < Trans. E. Soc. Edin./ vol. xxxiii, p. 651, pi. xl, figs. 14a— 14f, 1888.

2 ' Wiss. Meeresuiitersuch./ vii, p. 126, 1903.

3 'Aim. Nat. Hist./ 6th ser., vol. vi.

4 < Journ. M. B. A./ vol. iii, p. 225.

5 ' Zool. Danica/ iv, p. 39, Tab. civ, figs. 1—4.

G ' Aim. Sc. nat./ l e ser., t. xxix, p. 399. See also De Blainville, 'Diet. Sc. Nat./ xxxiv, p. 450,
and lvii, p. 483.

7 < Annul. Dan. Consp./ p. 37, figs. 8, 106, 107, 109.

516 NAINEREIS.

segments, and it only needs the effect of friction to alter the form of the anterior ventral
bristles. Too rigid adherence to (Ersted's 1 description would give neither papillae nor
cirri to the tail.

Mau (1881) furnishes a detailed account of the body-wall, bristles, circulatory,
nervous, muscular, digestive, and reproductive systems, and enables a comparison to be
made with southern examples. Certain corrections are necessary in regard to the nerve-
trunks, but he pointed out the position of the segmental organs which are at the base of
the ventral division of the feet.

There is little in the description of Hansen's 2 Aricia arctica from Jan Meyen to
distinguish it from this species.

Genus LXXIX. — Nainereis, JDe Blainville, 1828.

Head button-shaped. Body of two regions, slender and elongated, terminated by
two rounded dorsal papillae and two short ventral cirri. Branchiae commence on the
sixth foot and continue nearly to the posterior end. The structure of the foot in the
second region is less complex than in Aricia.

The generic description of 0. Fabricius was : — Nais, furnished with bifid lateral
processes, abdominal cirri, and a quadrifid tail. By subsequent authors, such as
Malmgren, the title was altered to Naidonereis, but there seems to be no special reason
why the original name of De Blainville should not be retained.

The arrangement of the nerve-cords anteriorly seems to be similar to that in Aricia
and Scoloplos, and they take position under the raphe, where the circular bands and
oblique muscles meet. The muscles of the body-wall do not show the disproportion so
characteristic of the species just mentioned, for the ventral are only a little larger than
the dorsal. On the other hand the oblique and vertical are very strong, both uniting in
the median raphe over the nerve-cords, which are at the inner end of a short, broad area
passing inward from the hypoderm. In the middle of the body this area has externally
the cuticle, then a somewhat thick hypodermic coat, which in the middle line presents a
symmetrical arrangement of its cells and areolae. Across the inner border of this coat
pass fibres — probably from the circular coat (unless the basement tissue, which is not
visibly differentiated, has them). Other fibres from the circular coat clasp the inner
borders of the longitudinal layer. Deeply stained cells occur next, and then the nerve-
area with its rounded cords and a neural canal superiorly in the middle line.

A new genus (Theodisca) was founded by Fritz Muller for Ariciidae, with strap-like
branchiae, and biramous feet with the ventral division bilobed. The proboscis is extruded
as a five-lobed foliaceous button. So far as can be observed all the known forms may
readily be placed under Nainereis.

An allied group is the Levinseniens of Mesnil and Caullery. The prostomium has a
median tentacle, as in Aricidea, and this is always absent in the Ariciidae proper, which
have a palpode. In the Ariciidae the first body-segment is achetous, the branchiae com-

1 ( Annul. Dorsibranch.,' p. 201, figs. 115, 117, 118.

2 ' Norske Nord.-Exped./ 1876—8, vii, p. 34, Tab. v, figs. 20—26.

NAINEREIS QUADBICUSPIDA. 517

mence on one of the twenty anterior segments, but are rudimentary anteriorly, and
gradually increase. In the Levinsenians they begin on the fourth setigerous segment and
only occur on a certain number of anterior segments, whereas in the Ariciidae they extend
to the posterior end, and their insertion is more dorsal than in the Levinsenians, which,
moreover, have more simple feet, with a dorsal and a ventral cirrus, though the latter is
often absent. The bristles are not annulated in the Levinsenians. They have three
anal cirri, whereas the Ariciidge have two or four. The proboscis is less complex than in
the Ariciiclas and there is no oesophageal caecum. The ventral nervous system is intra-
epidermic, whereas in the Ariciidae it is intra-coelomic.

Mesnil and Caullery 1 place Theodisea under the sub-genus Nainereis, Blv., of the
genus Aricia, And. and Edw. They enter T. anserina, Clap., Port Vendres; T. liriostoma,
Clap., Naples; T. mamillata, Cun. and Ramage (Claparede ?). In this group the
prostomium is rounded. Lo Bianco puts Nainereis as a synonym of Theodisea.

It is understood that Dr. Eisig is at present engaged with the Ariciidae, and is of
opinion that the genus Theodisea of Fritz Miiller is unnecessary, and that the old genus
Nainereis {Naidonereis of others) including N. quadricuspidata, seems to be sufficient for
the known forms.

1. Nainereis quadricuspida, Fabricius, 1780. Plate LXV, figs. 5 and 5 a — head; Plate
LXXXVI, figs. 2-2 b— bristles ; Plate LXXXVII, figs. 5-5 b— feet.

Specific Characters. — Head forming a rounded button-shaped process set on the
peristomial segment, which is notched at each side. Body about 3 ins. in length, slightly
tapered in front and considerably diminished towards the tail, which ends in two rounded
dorsal papillae and two short ventral cirri. Anterior region of thirteen bristled segments.
Branchiae commence on the sixth foot and continue of comparatively large size till the
eighth or tenth segment from the tip of the tail. At the tenth foot the branchia is
flattened and acutely lanceolate, and it is separated by a space from the dorsal division
of the foot, which bears a fan-like tuft of tapering bristles with smooth shafts and spinose
tips. The dorsal cirrus behind has an enlarged base and gently tapered tip. After an
interval the slightly convex and long inferior division occurs, with a prominent blunt lobe
or papilla rather below its middle posteriorly, and directed inward and backward. A
dense series of the strong and slightly brownish, curved (club-shaped) bristles occupies
the whole length of the division, with a few tapering serrated forms amongst them.
Most of the short, strong forms show distinct serrations towards the curve. The ventral
division ends in a notch ventrally.

After the change in the structure of the foot occurs, as at the twenty -third, the
slightly tapered dorsal cirrus has in front of it a group of long tapering bristles with
smooth shafts and serrated tips, and a few bifid forms— all supported by four spines. A
low rounded papilla closely adjoining the foregoing represents the inferior division with
two spines and a few slender serrated bristles. Posteriorly a ventral cirrus appears
beneath the division.

1 ' Bullet. Scientif. de la France et de Belgique/ Tom. xxxi, 4 e ser., ii, 1898, p. 143.

518 NAINBRBIS QUADRICUSPIDA.

Synonyms.

1780. Nais quadricuspida, Fabricius. Fauna Groenl., p. 315, n. 296.

1828. Nainereis „ De Blainville. Diet. Sc. Nat., lvii, p. 491.

1843. Scoloplos „ CErsted. Gronl. Annul. Dorsibr., p. 200, f. 106—110.

1849. Aricia „ Leuckart. Arch. f. Naturges., xv, p. 198, Tab. iii, f. 11.

1851. ., quadricuspis, Grube. Fam. Ann el., pp. 68 and 135.1

18-53. „ ,, Stimpson. Synops. Mar. Invert. Gr. Manan, p. 33.

1863. Scoloplos quadricuspis, idem. Ann. Groenl. in Proc. Acad. Nat. Sc. Philad., p. 140.

1865. ,, quadricuspida, De Quatrefages. Anne]., ii, p. 287.

1867. Naidonereis ,, Malmgren. Annul. Polych., p. 73.

1869. „ „ Mcintosh. Rep. Brit. Assoc, (1868), p. 338.

1875. „ „ Ehlers. Zeitschr. f. wiss. Zool., Bd. xxv, p. 59, Taf. iv, f. 28.

1878. „ „ Mcintosh. Trans. Linn. Soc, 2 ser., Zool. i, p. 504.

1897. Aricia „ Birula. Zool. Mus. Kaiser. Akad. Wiss. St. Petersb., p. 20.

1901. „ „ Whiteaves. Geol. Surv. Canada, No. 722, p. 79.

1905. Naidonereis ,, Mcintosh. Ann. Nat. Hist., ser. 7, vol. xv, p. 48.

Habitat. — Procured in numbers between tide-marks, Lochmaddy, August, 1865,,
generally under stones amidst sandy mud ; a young example under Lithot 1 1 amnion
between tide-marks, Lerwick, Shetland.

Greenland (O. Fabricius).

Head (Plate LXV, figs. 5 and 5 a) forming a rounded and somewhat button-shaped
process set on the peristomial segment, which forms a short cone and is two-ringed
(being notched on each side).

Body about 3 ins. in length, tapering very slightly in front, but considerably
diminishing towards the tail, which ends somewhat abruptly in two rounded dorsal
papillae, and two short ventral cirri.

The anterior region consists of thirteen segments with the specially modified ventral
division of the feet. The branchiae in the sole example commenced on the sixth foot,
but it may be that those on the dorsum of the body at the fifth foot had been lost.
They remain of comparatively large size to within eight or ten segments from the tip of
the tail. When fully formed, as at the tenth foot (Plate LXXXVII, fig. 5), the branchia
is a flattened and somewhat acutely lanceolate process. A convex border of some extent
intervenes between it and the slight elevation of the dorsal division of the foot bearing
a fan-like tuft of tapering bristles (Plate LXXXVI, fig. 2), which have smooth shafts
but spinose tips (with a marked bend) simulating cameration. The dorsal cirrus behind
has a somewhat enlarged base and gently tapered tip (which is not acute). A smooth
margin intervenes between this division and the slightly convex and long inferior
division, which has a prominent blunt lobe or papilla rather below its middle posteriorly,
directed downward and backward. A dense series of the strong and slightly brownish
curved (club-shaped) bristles (Plate LXXXVI, fig. 2 a) occupies the whole length of the
division, with here and there a long serrato-camerated tapering bristle or two projecting
beyond them. Most of these short and strong bristles show distinct serrations towards
the curve, as if they were developed from broken forriis. The tips of many are rounded,

NAINEREIS MAMMILLA.TA. 519

but others are cracked. It may be that enlargement of these occurs after fracture,
though some of similar form in a developing condition have been seen in the tissues.
The prominent ridge of the foot ends in a notch ventrally, and the surface trends to the
mid-ventral line.

A change occurs at the fourteenth foot, and is well marked at the twenty-third
(Plate LXXXVII, fig. 5 a), the slightly tapered dorsal cirrus having in front of it a group
of long tapering bristles with smooth shafts and serrated tips (Plate LXXXVI, figs. 2 b
and 2//). These are supported by about four spines. Moreover bifid bristles appear
in the dorsal division about the twentieth foot, and continue to the posterior region. The
shafts in the twentieth foot are rather strong, cylindrical, and translucent, and further
show, by the play of light, indications of serrations, though none are actually present.
The limits of the fork are " finished " distally and slightly enlarged, and the inner edges
are spinous. A low rounded papilla closely adjoining the foregoing represents the
inferior division of the foot and has two spines which pierce the tip, and a few slender
serrated bristles. The edge below is boldly convex.

The chief changes in connection with the posterior feet are (1) the diminution in the
convexity of the body-wall below the foot, (2) the diminution in the size of the dorsal
cirrus, (3) the increase in the size of the ventral division of the foot, and (4) the appear-
ance of a ventral cirrus beneath it (Plate LXXXVII, fig. 5 b) a little in front of the tail.
The bifid bristles of the dorsal division are continued posteriorly, and they have stout
shafts, the sides of which present serrations, the bifid tip being short but stout, the inner
edges being spinous, and the terminal spine on each side projecting beyond the truncated
ends of the bifid region.

CErsted (1843) added little to the account of 0. Fabricius, but he gave recognizable
figures and grouped the form under the Ariciaa, a family of his Cha?topoda Terricolina.

Grrube (1851) included Nainereis in his last family of the tribe Papacia of the
Appendiculate Polycheeta.

A young form found in Lithothamnion between tide-marks, Shetland, July, 1871,
belongs to this species. The snout is bluntly rounded, and is followed by two double-
ringed segments. The bristled segments are forty-two or forty-three, and the caudal
cirri are comparatively large. It is about 6 mm. in length.

The original account of 0. Fabricius (1780) is fairly complete, without touching on
the microscopic characters of the bristles, and shows how acutely this zoologist had
observed Nature. Even the habits and food of this species did not escape him.

It is probable that the Scoloplos minor of CErsted 1 is this species. The author
describes the head as globose, and the tail is furnished with four filiform appendages.

The head of Theel's Aricia Tullbergi, from Nova Zembla, agrees in outline with this
form, but in other respects it differs, notably in the absence of dorsal bristles, yet the
latter may have been accidental, for such is a very exceptional condition in the group.

2. Nainereis mammtllata, Glaparede.

Cunningham and Pamage found an example which they referred to the above species

1 ' Kroyer's Nat. Tids./ 1842—3, p. 125.

520 NAINEREIS.

in Laminarian roots at the Birnie Rocks near Grranton Quarry. They describe it l as
having an obtuse rounded head, and a pair of eyes; branchise commencing on the fifth
segment ; feet with a dorsal division of a single cylindrical process with a fascicle of long
capillary bristles in front, a ventral division consisting of a mammiform projection with a
nipple-like process at the end, on the anterior surface exteriorly a row of capillary setae,
proximally a large number of short thick setse, bifid at the apex (the latter being present
only in the first ten to fifteen segments). Amongst the dorsal bristles in the middle of the
body are some two-pronged.

As the preparations made by the authors have disappeared, it is not possible to
re-investigate the subject at present. The form, described as Theodisca Mamittata,
appears to be a Nainereis.

Nainereis, sp. 1. Plate LXXXV, figs. 10 and 10 a— bristles.

A form which appears to be related to the above genus has more than once occurred
at St. Andrews, but the condition of the examples is such that no complete account
can at present be given, and since the bristles were drawn the preparation has been
mislaid.

Habitat. — Tossed on shore at the West Sands after storms both in summer and
spring. It would appear, therefore, to reside in the sandy ground in the bay.

Head somewhat flattened, bluntly conical. Body of considerable size, probably 6
to 8 ins. long, somewhat flattened dorsally and ventrally, rounded at the sides. It is
slightly narrowed anteriorly and more distinctly tapered posteriorly, where it terminates
in four short cirri. A median ridge occurs at the ventral surface. Claparede's form had
longest dorsal cirri at vent. A folded proboscis seems to have issued from the anterior
end, but it is injured. The segments are both narrow and numerous. Dorsally each
segment anteriorly has a long narrow bar with pointed ends, and behind it a ridge con-
necting the dorsal papillse of opposite feet. Ventrally each segment has three transverse
furrows or creases.

Nainereis, sp. 2. Plate LXXXV, figs. 10 and 10 a— bristles; Plate LXXXVI, fig. 19—

bristle.

Specific Characters. — Head somewhat flattened, bluntly conical (from above). Eyes
unknown. Body rather large, flattened, slightly narrowed anteriorly. Ventral surface
with a median ridge. Segments narrow ; dorsally each has a narrow bar with pointed ends
in front, and behind it is a ridge connecting the dorsal papillge of opposite feet. Ventrally
each segment has three transverse furrows. Tenth foot has a prominent dorsal papilla
for the bristles, which are simple, golden, stout, and slightly curved. After an interval
a long elevated ridge represents the inferior division, and by-and-by this forms a flap
sloping backward, the dorsal papilla fusing with it superiorly. Ventral bristles have a
sharp hook at one side of the blunt tip, with a wing. At the ventral edge are long, simple
bristles with finely tapered tips.

1 'Trans. Roy. Soc. Edin./ vol. xxxiii, pp. 642—643, 1888.

NAINEREIS. 521

The feet are very small in front, but they conform to the type behind. About the
tenth foot is a somewhat prominent dorsal papilla for the bristles, which are simple,
golden, stout, and slightly curved. They have longitudinal striae and also a few transverse
ones (Plate LXXXVI, fig. 19). The tip seems to be slightly hooked and to have a wing
in some. The difficulty is their absence, as few feet show any. After an interval a long
elevated ridge represents the inferior division, and by-and-by this forms a flap sloping
backward, the dorsal papilla fusing with it superiorly, so that both are included in the
free flap. The ventral bristles have a sharp hook at one side of the blunt tip, with a
wing. At the ventral edge are long simple bristles with finely tapered tips, to which
mucus adheres.

Both dorsal and ventral bristles seem to be extremely brittle.

This form, whilst agreeing in certain respects with Nainereis, leans also to the
Spionidse. It was tossed on the West Sands, St. Andrews, after a storm.

123

INDEX TO VOL. II

PAGE

AlciopidgB .....
Amblyosyllis ....

. 112
. 224

Eteone picta
pusilla

PAGE

. 100
. 106

lineata ....
Anaitis .....

. 225
. 70

spetzbergensis .
Eulalia

. 104
. 50

Jeffreysii

. 73

bilineata

. 50

Kosteriensis

. 72

imbricata .

. 64

rosea .....

. 71

macroceros

. 60

Arabella .....

. 394

nebulosa

. 53

iricolor '.

. 395

tripunctata

. 63

Aricia .....

. 497

viridis

55

Armandi ....

. 508

Eumida

66

Cuvieri ,

Edwardsi an. var. Cuvieri .

. 497
. 501

sanguinea .
Eunice

66
425

Grubei ....

. 505

fasciata

425

Latreillii ....

. 502

floridana

439

norvegica ....

. 506

norvegica .

434

Ariciidee .....

. 494

vittata

431

Autolytea .....

. 209

Eunicidaa .

343

Autolytus .....

. 209

Eurysyllis . . . .

241

aurantiaca ....

. 214

paradoxa .

241

ehbiensis ....

. 243

Eusyllidea

160

? female bud

. 223

Eusyllis .

173

inermis ....

. 247

Blomstrandi

176

longiferiens

. 245

tubifex

173

pictus . . . . .

. 211

Exogone . . . .

149

prolifer . . . . .

. 215

gemmifera .

var. S charm

151
154

Castalia ......

. 120

naidina

150

arctica . . . .

. 125

Exogonea . . . .

149

fusca ......

. 127

punctata .

. 121

Grenetyllis .
citrina

94
96

Dalhousiella .

. 134

hibernica .

97

Carpenteri . . . .

. 134

lutea . . . .

95

Drilonereis .

. 392

Grlycera .

477

longa .

. 393

alba .
Ehlersi

486
481

Eteone ......

. 98

Groesi .

491

arctica .

. 102

lapidum

477

var. Robertiana .

. 103

siphonostoma

482

Glyceridas .
Glycinde .

Nordmanni
Goniada

eremita

maculata
Goniadidae
Grubea

clavata

Hesionidee
Hyalincecia

sicula

tubicola

Leocrates

atlantica
Leptonereis

Vaillanti
Lumbriconereis .

acutifrons .

fragilis

G. — fragment

gracilis

hibernica .

impatiens .

Latreillii

(Zygolobus) laurentianus
Lysidice ....

punctata

Magalia

perarmata
Marphysa .

Belli .

Kinbergi

sanguinea
Micronereis

variegata
Microsyllis

Marenzellei
Myrianida

pinnigera
Mysta

barbata
Mystides .

Lizziae

INDEX.

PAGE

471

Nainereis .

469

mammillata

469

quadricuspida

462

sp. 1 .

466

sp. 2 .

462

Nematonereis

460

sp. .

234

unicornis

235

Nephythydidge
Nephthys .

114

417

caeca .
ciliata

417
419

cirrosa
Grubei
Hombergii

130
130

hystricis

incisa

Johnstoni

268

264
371

388
372

387

longisetosa

pansa
Nereicola ovata
Nereidse .
Nereis

cultrifera

385
383
379
376
390
456
456

diversicolor

Dumerilii

fucata

longissima

Marionii

pelagica

Schmardaai

virens

136

Nerilla

136

antennata

442

Notocirrus

448

scoticus

451

sp. ? .

442

Notophyllum

260

foliosum

261

159

Odontosyllis

160

ctenostoma

228

fulgurans

229

gibba

108

Onuphididee

108

Onuphis .

109

brevibracbiata

110

britannica

523

PAGE

516

519
517
520

520

453

456

453

1

6

8

23

36

33

17

27

38

34

29

40

286

249

267

280

312

302

336

325

295

267

291

330

147

147

400

400

403

46

46

178
182
178
183
403
404
407
404

524

Onupbis conchylega

fragosa

quadricuspis
Ophiodromus

flexuosus .
Ophryotrocha .

puerilis

Phyllodoce

groenlandica

lamelligera

maculata .

Paretti

rubiginosa

vittata
Phyllodocidee
Pionosyllis

altera osetosa

divaricata .

hyalina

lamelligera

prolifera

Scoloplos .

armiger
Sphaerosyllis

hystrix
Staurocephalidaa
Stauroceplialus .

B

ciliatus var. Robertianee

INDEX.

PAGE

PAGE

410

Stauroceplialus Kefersteini . . 358

415

pallidus

. 363

413

rubrovittatus

. 353

116

Syllidaa .

137, 233

117

Syllidea (proper)

. 187

364

Sylline .

. 186

364

rubropunctata .
Syllis

. 186
. 188

74

abyssicola

. 207

86

armillaris .

. 188

76

brevicirrata

. 196

89

brevicollis .

. 208

82

Buskii

. 206

92

cornuta

. 200

93

var. Collingsii

. 202

41

cucullata .

. 191

161

Cunninghami

. 195

238

gracilis

. 203

164

Krohnii

. 192

166

macroceras

. 194

236

spongicola

. 197

161

Trypanosyllis .

. 169

509

cseliaca

. 240

510

zebra ....

. 169

156

156

Xenosyllis

. 155

350

Kinbergi .

. 155

352

362

Zygolobus (Lumbriconerei

8) .

. 390

359

laurentianus

. 390

ADLABD AND SON, IMPR., LONDON AND DORKING.

PLATE LI

Fig.

1. Trypanosyllis zebra, Grube, from Plymouth. Enlarged.

2. Pionosyllis (Typosyllis) hyalina, Grube, from Plymouth. Enlarged.

3. Syllis gracilis, Grube, from Plymouth. Enlarged.

4. Syllis spongicola, Grube, from Plymouth. Enlarged.

5. Head of Odontosyllis fulgurans, Aud. and Edw.

6. Onuphis britannica, McL, from Plymouth, removed from its tube and seen from the

dorsal surface. Enlarged.
6 a. Head and anterior region of the foregoing, still further enlarged.

Plate LI.

\f I

'

PLATE LIL

Fig.

1. Nereis jjelagica, L. Adult female (atokous) of a russet-brown hue in September,

St. Andrews. Enlarged.

2. An epitokous male on 17th February, St. Andrews. The tail is in process of

reproduction. Enlarged.

3. Nereis cultrifera, Grube, from East rocks, St. Andrews, in the atokous condition. The

tail is developing.

4. Nereis diversicolor, 0. F. Muller, from St. Andrews. Enlarged.

4 a. Example with the proboscis extruded, from St. Andrews. Slightly enlarged.

5. Nereis Dumerilii, Aud. and Edw., from a sketch at Lochmaddy. Slightly enlarged.

6. Nereilepas fucata, Say., from the Moray Firth. In a shell of Buccinum undatum with

a hermit-crab.

7. Nereis Schmardsei, De Quatref., from the dorsum. Enlarged.
7a. Tail of the same. Still more enlarged.

Plate LII.

;

.

PLATE LIII.

Fig.
1. Nereis virens, Sars, from an example tossed on shore on the West Sands, St.
Andrews, and carried alive to Murthly, where it was drawn. Natural size.

Plate LIU.

PLATE LIV.

Fig.

1. Eunice fasciata, Risso, from St. Peter Port, Guernsey. Enlarged.

2. Marphysa sanguined, Mont., from St. Peter Port, Guernsey. The branchise in this

figure are less characteristic than usual. Their structure is seen in Plate LXXIV,
fig. 8. Enlarged.

3. Nematonereis unicornis, Grube, from St. Peter Port, Guernsey. Enlarged.

4. Arabella iricolor, Mont., from St. Peter Port, Guernsey. Enlarged.

5. Eyalinoecia tubicola, 0. F. Miiller, from Plymouth, removed from its tube and viewed

from the dorsum. Enlarged.
5 a. Head and anterior region of the foregoing. Enlarged.
5 b. Caudal region of the same. Enlarged.

6. Lumbriconereis Latreillii. Aud. and Edw. Enlarged.

Plate LIV

PLATE LV.

Fig.

1 . Staurocephalus rubrovittatus, Grube, from Herm. Enlarged.

2. Staurocephalus Kefersteini, McL, from North. Uist and Channel Islands. Enlarged.

3. Lysidice punctata, Pisso, from Jersey. Enlarged.

4. Glycera lapidum, De Quatref., from St. Andrews. Enlarged.

5. Marphysa Belli, And. and Edw., from Plymouth. Enlarged.

6. Eighth foot of the foregoing, x 25.

7. Head and anterior region of Nematonereis, from Plymouth. Enlarged.

8. Tip of the tail of the same. Enlarged.

9. Head and anterior region of Ophryotrocha puerilis, Clap, and Meczn., from Plymouth,

Enlarged.

R.M.Figs

- - \ , - T -i , 1 1

PLATE LV1.

Fig.

1. Aricia Latreillii, Aud. and Edw., from St. Andrews. Enlarged.
1 a. View of tip of tail with vent and cirri. Enlarged.

2. Goniada Q?iaculata, (Ersted, from the sand, near low water-mark, St. Andrews.

Enlarged,
o. Mid-dorsal segments of the same. Enlarged.

4. Ophryotrocha puerilis, Clap, and Meczn. Enlarged.

5. Tip of tail of the foregoing. Enlarged.

6. Nematonereis from Plymouth, 7th September, 1905, from the dorsum. Enlarged.

7. Scoloplos armiger, 0. F. Miiller. Dorsum shown anteriorly ; posterior region seen

laterally.

PLATE LXXI.

Fig.

1. Foot of Amblyosyllis lineata, A. Costa. The process above the setigerous region is

often more pointed, and the cirrus has its basal region directed dorsally. X 55.

2. Foot of Autolytus pictus, Ehlers. X about 55.

3. Foot of Autolytus aurantiacus, Claparede, viewed obliquely from above. X about 55.

4. Foot of female bud of Autolytus prolifer, 0. F. Miiller. X about 55.

5. Foot of nurse-stock of Myrianida pinnigera, Mont. X about 55.

6. Foot of female bud of the same with developing swimming bristles. X about 55.

7. First foot of Nereis pelagica, L. X 15.
7 a. Tenth right foot of the same. X 15.

7 b. Thirty-seventh right foot of the foregoing. X 15.

7 c. Fifty-seventh foot (right). X 15.

7 d. First foot of a large epitokous male from the West Sands, St. Andrews. X

about 12.
7 /. Thirty-seventh foot of epitokous form. X 12.
7 g. Fifty-seventh foot of the foregoing. X 12.
7 h. Eightieth foot of the same. X 12.

7 i. Twenty-eighth foot of a young example of Nereis pelagica from St. Andrews with

clavate parasitic growth. Viewed from the dorsum. X 90.

8. First foot of Nereis cultrifera, Grube. X 15.

8 a. Tenth foot of the same. X 15.

8 b. Thirty- seventh foot of this species. X 15.

8 c. Fifth foot from tip of tail. X 15.

8 d. Nineteenth foot of epitokous male. X 15.

pi.iixxr.

f-p.

VtMTrap ampi

A U.W.& W. CM.de

PLATE LXXIL

Fig.

1. Twentieth foot of the epitokous male Nereis cultrifera, Grube. X 15.
1 a. Thirty-seventh foot of the foregoing. X 12.

1 b. Fifty-seventh foot of the same male. X 10.

1 c. Foot of this form near tip of tail. X 15.

2. First foot of Nereis Schmardsei, Be Quatref. (irorata, Malmgren). x 15.

2 a. Tenth left foot of the same, from the front. X 15.

2b. Sixty-fifth foot of a ripe epitokous female from St. Vaast (Prof. Grnbe's). X 15.

3. First foot of Nereis Marioni, Aucl. and Edw. X about 14.

3 a. Tenth foot of the same. X 15.
3 b. Thirty-seventh foot. X 15.

3 c. Fifty- seventh foot, x 15.
3d. Sixty-eighth foot. X 15.

4. First foot of Nereis Dumerilii, Aud. and Edw. X 15.

4 a. Tenth foot. X abont 15.

4 6. Thirty-seventh foot. X 21.
4 c. Fifty-seventh foot. X 13.

4 d. First foot of epitokous form (male) of the same species from the surface in Castle
Bay, Barra, in May. X 21.

4 e. Sixth left foot of an epitokous male. X 21.
4/. Thirty-seventh foot of the same male. X 18.

5. First foot of Nereis diversicolor, 0. F. Muller, from St. Andrews. X about 21.

5 a. Tenth foot of the same from Herm. x 15.
5 b. Thirty-seventh foot. X 15.

PI. I.XX11

% AH.W.del

f a PAY. M Trap impr.

PLATE LXXIII.

Fig.

1. First foot of Eunereis longissima, Johns t. X 15.
la. Tenth foot. X 15.

1 b. Sixty-sixth foot of an example (var.) from the Bay of Tunis. Enlarged.
1 c. Sixty-seventh foot of an epitokous female. X 18.

1 d. Foot of the foregoing near the tip of the tail. X 15.

2. First foot of Alitta virens, Sars. X 13.

2 a. Tenth foot (right). X 12.

2 b. Thirty-seventh foot. X about 12.

3. First foot of Nereilepas fucata, Sav. X about 15.

3 a. Tenth foot. X about 15.

3 b. Twenty-seventh foot of epitokous female. X 12.

3 c. Fifty-seventh foot of the foregoing. X 12.

4. Foot of Staurocephalus rubrovittatus, Grube, from the middle of the body. X 60.

4 a, Foot of a young pelagic form of the same, caught in tow-net off Sark by Mrs.

Collings. X 60.

5. Typical foot of Staurocephalus Kefersteini, Mel. x Zeiss oc. 2, obj. D.

6. Twentieth foot of Staurocephalus ciliatus, Kef., var Robertianse, from St. Peter Port,

Guernsey. X 65.

7. Foot of Ophryotrocha puerilis, Clap, and Mecz. X 100.

8. First foot of Lumbriconereis fragilis, 0. F. Mull. X 25.
8 a. Tenth foot of variety of the foregoing. X 25.

8 b. Fiftieth foot from a specimen dredged off Sark by Col. Fraser and Capt. Powell.
X 25.

8 c. Foot (sixtieth) from the posterior region. X 25.

9. Tenth foot of Lumbriconereis Latreillii, Aud. and Edw., from Bantry Bay. X 55.

9 a. Foot (about fifteenth) of the same. X 55.

10. Tenth foot ? of Lumbriconereis impatiens, Clap., from Plymouth. The long hooks

have been omitted. X 55.
10 a. One hundredth foot of the foregoing form. The long hooks have been omitted.

x 55.
10 b. Fifty-seventh foot of a specimen from the ' Knight Errant.' x 55.

PL. LXX1LX.

^

,

1

A-

5

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•,A;A

8 <P

f ^ PWM-Trap impr.

\Y.C.:M.&A'.HW.del.

PLATE LXXIV.

Fig.

1. Tenth foot of Lumbriconereis hibemica, Mel. X about 60.
1 a. Thirtieth foot of the foregoing. X about 60.

2. Foot of Lumbriconereis G. X Zeiss oc. 2, obj. A, full draw-tube.

3. Foot of Lumbriconereis (Zygolobus) laurentianus, Grube (from a somewhat softened

preparation from the stomach of cod, St. Andrews) ; middle third of the body.
X 35.

4 Sixth foot of Drilonereis Elisabeths, McL, turned round so as to show the dorsal

process. X about 35.

5. First foot of Arabella tricolor, Mont., from St. Peter Port, Guernsey. X 50.

5 a. Tenth foot. X 50.

5 b. Foot from the middle of the body. X 50.

5 c. Foot from the posterior region. X 50.

6. Typical foot of Notocirrus scoticus, McL, from Lochmaddy; a, branchial lobule

(cirrus) ; b, spine ; c, bristles. X 350.

6 a. Foot showing two papillse from example procured off Peel, Isle of Man. X 50.

7. Tenth foot of Eunice fasciata, Risso. X 35.

7 a. Ninetieth foot. X 21.

7 b. Foot near tip of tail. X 35.

8. Twenty-seventh foot of Marphysa sanguinea, Mont. X about 30.

8 a. Tenth foot. X about 35.

9. Tenth foot of Marphysa Kinbergi, McL X about 35.

9 a. Thirtieth foot also from the ' Porcupine ' example, off Cape Finisterre. X

about 35.

10. Tenth foot of Eunice vittata, D. Chiaje. X about 55.
10 a. Twentieth foot. X about 55.

10 b. Fortieth foot, from Guernsey. X about 55.

11. Tenth foot of Eunice norvegica, L., from Norway. X 21.

11 a. Twentieth foot of the same. X 21.

12. Head and anterior region of Lumbriconereis Latreillii, Aud. and Edw. Enlarged.

Pl.IiXXlV.

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PLATE LXXV.

Fig.

1. Tenth foot of Eunice norvegica var. B. A., from the ' Porcupine' Expedition of 1870.

x 21.

1 a. About the fortieth foot. X 21.

2. Anterior foot of Eunice floridana, Pourt. X 21.

2 a. Anterior foot of young example. X 21.

3. Tenth foot of Nematonereis unicornis, Grube, from St. Peter Port, Guernsey. X 90.

3 a. Twentieth foot. X 90.

4. Tenth foot of Lysidice punctata, Risso. X 21.

4 a. Thirtieth foot. X 21.
4 6. Fiftieth foot. X 21.

5. Twentieth foot of Onuphis britannica, McL,from Valentia, ' Porcupine.' X 21.

6. Tenth foot of Onuphis brevibrachiata, Ehlers. Enlarged.

6 a. Thirtieth foot of the same with an unusual number of branches to branchige.
Enlarged.

7. Tenth foot of Onuphis conchylega, Sars. X 21.

8. Ninth foot of Onuphis quadricuspis, Sars, from Cape Rosier, Canada. X 30.

9. First foot of Onuphis frag osa, Ehlers. X 55.
9 a. Third foot from Bergen. X ?.

9 b. Twentieth foob of the same. X P.

10. Twentieth foot of Hyalinoecia sicula, De Quatref. X 21.

11. Tenth foot of Hyalinoecia tnbicola. X 30.

12. Tenth foot of Goniada maculata. x 30.
12 a. Fiftieth foot of the same. X 30.

12 b. Seventieth foot of Goniada eremita, Aud. and Edw., from Cape Finisterre,
'Porcupine,' 1870. X 15.

12 c. Posterior foot (hundredth) of the foregoing. X 15.

13. Tenth foot of Glycinde Nordmanni, Mgrn. X 50.

13 a. Thirtieth foot, from the < Porcupine,' 1870. X 40.
13 b. Sixtieth foot, X 40.

19 c. Hundredth foot. X 40.

13 d. Unknown foot (seventieth?). X Zeiss oc. 2, obj. D.

Pl.LXXV.

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PLATE LXXVI.

Fig.

1. Tenth foot of Glycera lapidum, De Quatref. X 21.
1 a. Papillse of the proboscis. X Zeiss oc. 2, obj. D.

1 b. Ninetieth foot of the foregoing.

2. Tenth foot of Glycera sipliono stoma, D. Chiaje, from Herm. x 21.

2 a. Thirtieth foot of a specimen dredged off Cape Gruardia, ' Porcupine,' 1870. X 21.
2 b. Twentieth foot from the tip of the tail. X 21.

2 c. Anterior foot of example from Cape Guardia, ' Porcupine,' 1870. X 21.

3. Tenth foot of Glycera alba, H. Eathke. X 21.

3'. Sixteenth foot of example from Donegal Bay, ' Porcupine,' 1869. X 21.

3 a. Seventieth foot from a specimen dredged by the ' Porcupine,' 1870, off Cape

Sagres. X 21.
3 b. Branchia showing crossed fibres. X Zeiss oc. 2, obj. A, with full draw-tube.

3 c. Sixtieth foot of an example with longer processes dredged by the ' Porcupine ' off

Cape Sagres, 1870. X 21.

4. Typical foot (thirty-eighth) of Glycera Goesi, Mgrn., from St. Magnus Bay. X Zeiss

oc. 2, obj. A, with draw-tube.

4 a. Tenth foot. X Zeiss oc. 2, obj. A, with draw-tube.

4'. Thirty-eighth foot. X Zeiss oc. 2, obj. A, with draw-tube.

4 b. Seventieth foot. X Zeiss oc. 2, obj. A, with draw-tube.

5. Tenth foot of Aricla Guvieri, Aud. and Edw. X about 21.
ba. The twenty-second foot. X about 21.

5 b. Twenty-third foot. X about 21.
5 c. Posterior foot. X about 21.

PI. LXXVI

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PLATE LXXVIL

Fig.

1. Upper bristle of Nephthys coeca, 0. F. Mull. X Zeiss oc. 2, obj. D. 1 a, base of the

foregoing indicating the arrangement of the spikes on the edge ; 1 b, barred
bristles ; 1 c and 1 c', the same showing the rows of spikes ; 1 d, lateral view of a
barred bristle ; X 250 1 e, long capillary bristle from the variety of Nephthys coeca.

2. Capillary bristle of Nephthys Hombergii, Lamk. X Zeiss oc. 2, obj. D.

3. Barred bristle of the same. X Zeiss oc. 2, obj. D.

4. Capillary bristle of N. Hombergii, variety Jeer sivalen sis, from Lochmaddy. X Zeiss

oc. 2, obj. D.

5. Barred bristle from the same. X Zeiss oc. 2, obj. D.

6. Capillary bristles from Nephthys ciliata, 0. F. Mull. X Zeiss oc. 2, obj. D.
6 a. Outline of another bristle showing the curvature, x Zeiss oc. 2, obj. D.

7. Barred bristles from the same. X Zeiss oc. 2, obj. D.

8. Capillary bristle of Nephthys longisetosa, (Erst., from St. Magnus Bay, Shetland.

X Zeiss oc. 2, obj. D.
8 a. Barred bristles of the same. X Zeiss oc. 2, obj. D.

8 h. Edge of the capillary bristle showing the spikes. X more highly.

9. Capillary bristles of Nephthys Grubei, Mel. X Zeiss oc. 2, obj. D.

9 a. Barred bristles of the foregoing. X Zeiss oc. 2, obj. D.

10. Capillary bristle of Nephthys JoJmstoni, Ehlers. This figure, like some others, has

been reduced proportionally so as to get it into the plate. X Zeiss oc. 2, obj. D.

11. Barred bristle of the foregoing. X Zeiss oc. 2, obj. D.

12. Capillary bristle of Nephthys cirrosa, Ehlers. X Zeiss oc. 2, obj. D.

12 a. Outline showing the proportional size of this long bristle which has been reduced
as before.

13. Barred bristle of the foregoing. X Zeiss oc. 2, obj. D.

14. Capillary bristle of Nephthys incisa, Mgrn. X Zeiss oc. 2, obj. D.

15. Barred bristle of the same. X Zeiss oc. 2, obj. D.

16. Capillary bristle of Marphysa sanguined, Mont. Zeiss oc. 2, obj. D, with draw-tube.

Pl.LXXVII

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PLATE LXXVIII.

Fig.

1. Bristle of Eulalia nebulosa, Mont. X Zeiss oc. 2, obj. D.

2. Bristle of Eulalia viridis, 0. F. Mull. X Zeiss oc. 2, obj. D.
2 a. Bristle from anterior foot. X Zeiss oc. 4, obj. F.

3. Bristle of Eulalia macroceros, Grube. X Zeiss oc. 4, obj. F.

4. Bristle of Eulalia tripunctata, Mel. X Zeiss oc. 4, obj. F, with 2-in. draw-tube.

5. Bristle of Eumida sanguinea, (Erst. X Zeiss oc. 2, obj. D.

6. Bristle of Phyllodoce lanielligera, Gmel., from St. Andrews. X Zeiss oc. 2, obj. F.

7. Bristle of Phyllodoce grwnlandica, (Erst., from St. Andrews. X Zeiss oc. 2, obj. F.

8. Bristle of Phyllodoce rubiginosa, De St. Jos. X Zeiss oc. 2, obj. F.
8 a. Another bristle. X Zeiss oc. 2, obj. F.

9. Bristle of Phyllodoce vittata, Ehlers, from St. Magnus Bay. X Zeiss oc. 2, obj. F.

10. Bristle of Anaitis rosea, Mel. X Zeiss oc. 2, obj. F.

11. Bristle of Anaitis Kosteriensis, Mgrn. X Zeiss oc. 2, obj. F.

12. Bristle of Anaitis Jeffreysii, Mel. X Zeiss oc. 2, obj. F.

13. Bristle of Genetyllis lutea, Mgrn. X Zeiss oc. 2, obj. F.

14. Bristle of Genetyllis citrina, (Erst., from the Minch. X Zeiss oc. 2, obj. F.

15. 15 a, and 15 b. Bristles of Genetyllis hibemica, Mel. X Zeiss oc. 2, obj. F.

16. Bristle of Eteone spetsbergensis, Mgrn., from St. Andrews. X Zeiss oc. 2, obj. F.
16 a. Bristle of the same from Gaspe Bay, Canada. X Zeiss oc. 2, obj. F.

16 b. Bristle of the foregoing from Lochmaddy, North TJist. X Zeiss oc. 2, obj. F.

17. Bristle of Eteone pusilla, (Erst., from Shetland. X Zeiss oc. 2, obj. D.

18. Bristle of Eteone arctica, Mgrn., var. Eobertiana. X Zeiss oc. 2, obj. D.

19. Bristle of Mysta barbata, Mgrn., from the twenty-ninth foot, West Sands, St.

Andrews. X Zeiss oc. 4, obj. F.
19 a. Bristle of the same from a posterior foot. X Zeiss oc. 4, obj. F.

20. Bristle of Mystides Lizzim, Mel. X Zeiss oc. 2, obj. D.

21. Bristles of Notophyllum foliosum, Sars. a, lateral view, X 420; b, end of shaft in

profile, X 700 ; and c, the same viewed laterally, X 700.

22. Bristle of Eteone picta, De Quatref . X about 700.

23. 23 a, and 23 b. Bristles of Phyllodoce maculata, L. X Zeiss oc. 2, obj. F., and

draw-tube, full.

24. Long swimming bristle (spiral) of male Phyllodoce maculata, L., in July. X Zeiss

oc. 4, obj. D.

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PLATE LXXIX.

Fig.

1. Portion of shaft of bristle from the dorsal lobe of OpModromus flexiwsus, D. Chiaje.

X 750.
1 a. Bristle with longest terminal process from the same. X Zeiss oc. 2, obj. F.

1 b. Bristle with shorter terminal piece. X 700.

2. Dorsal bristle of Gastalia punctata, 0. F. Mull. X Zeiss oc. 2, obj. F.

2 a. Ventral bristle of the foregoing. X Zeiss oc. 2, obj. F.

2 b. Bristle with shorter terminal region from the same. X 700.

3. Ventral bristle of Gastalia arctica, Mgrn., from south-west Ireland. X Zeiss oc. 2,

obj. D.

3 a. Tip of bristle, more highly magnified.

4. Tip of shaft and terminal piece of the bristle of Gastalia fusca, Johnst. X 700.

5. Bristle of Leocrates atlantica, Roule, with short tip. X about 100.
5 a. Tip of a bristle (terminal piece). X about 400.

6. Tip of terminal piece of similar bristle in Leocrates Glaparedii, for comparison. X

about 400.

7. Tip of bristle of Dalhousiella Garpenteri, Mel. X about 400.

8. Bristle of Mag alia per armata, De St. Jos. X Zeiss oc. 2, obj. D.

8 a. Another of the same with longer tip. X Zeiss oc. 2, obj. D.

9. Bristle of Pionosyllis (Syllis) hyalina, Grube. X Zeiss oc. 4, obj. F.

9 a. Simple bristle in the posterior segments of the same. X Zeiss oc. 4, obj. C.
10. Bristle of Xenosyllis Kinbergi, Mel. X Zeiss oc. 2, obj. D., with draw-tube.

11 a and b. Compound bristles of Sphserosyllis hystrix, Clap. X Zeiss oc. 4, obj. F.

12. Simple bristle of the foregoing. X Zeiss oc. 2, obj. D, and draw-tube.

13. Long swimming bristles of the same. X Zeiss oc. 2, obj. F.

14. Bristle of Pionosyllis prolifera, Krohn, from Scalloway, with long tip. X Zeiss oc. 2,

obj. D.
14 a. Another example of the foregoing with slightly shorter tip. X Zeiss oc. 2, obj. D.
14 b. Bristle from a bud of the same species dredged at Scalloway. X Zeiss oc. 2, obj. D.
14 c and 14 c' . Bristles of Pionosyllis prolifera var. Malmgreni. X 700.

15. Bristle of the same form with shorter (average) tip from Guernsey. X Zeiss oc. 4,

obj. F.

16. Bristle of the foregoing from Luccomb Chine, with less distinctly bifid tip. X Zeiss

oc. 4, obj. F.

17. 17 a and 17 b. Bristles of Pionosyllis divaricata, Keferst. X Zeiss oc. 4, obj. F.

18. Bristle of Trypanosyllis zebra, Grube. X Zeiss oc. 2, obj. F, draw-tube, full.

PI. LXXIX.

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PLATE LXXX.

Fig.

1. Bristle of JEusyllis tubifex, Gosse. X Zeiss oc. 2, obj. D.

2. Bristle of Eusyllis Blomstrandi, Mgrn. X Zeiss oc. 2, obj. D.

3. Bristle of Autolytus A (referred to on p. 222 of vol. ii, Part I), x Zeiss oc. 2, obj. D.

4. Bristle of Odontosyllis fulgurans, Clap, x Zeiss oc. 2, obj. D.

5. Peculiar tip of spine from the posterior segments of Odontosyllis ctenostoma, Clap.,

from Plymouth. X Zeiss oc. 2, obj. D.

6. Bristle of Odontosyllis ctenostoma. X Zeiss oc. 2, obj. F., with 3-in. draw-tube.

7. Bristle of Odontosyllis gibba, Clap., var. Bobertiana, dredged off St. Peter Port,

Guernsey. X Zeiss oc. 2, obj. D.

8. Bristles of Syllis armillaris, 0. F. M. x 280.

8 a. 1 Posterior bristle of the foregoing, x 350.

9 and 9 a. Simple and compound bristles of Ioida macrophthalma, Johnst. X 350.

10. Bristle of Syllis cornuta, H. Rathke, var. Colling sii, off Sark. The bifid tip is not

very distinct in this figure. X Zeiss oc. 2, obj. D.

11. Bristles (two) of Syllis cucullata, McL, from Luccomb Chine. X Zeiss oc. 2, obj. D.

12. Compound bristle of Syllis Krohnii, Ehlers, from Lochmaddy. X 700.

13. Compound bristle of Syllis macroceras, Grube, Isle of Wight, x Zeiss oc. 2, obj. D.

13 a. The same from an example from the Gouliot Caves, Sark. X Zeiss oc. 2, obj. D.

14. Compound bristle of Syllis brevioirrata, McL, with longer tip. x Zeiss oc. 2, obj. D.

14 a. A form of the same with shorter tip. X Zeiss oc. 2, obj. D.

15. Bristle (hamate) of Syllis spongicola, Grube. X Zeiss oc. 2, obj. D.

16 and 16'. Bristles of Syllis cornuta, H. Rathke, St. Magnus Bay, Shetland, x as above.
16 a. Bristles of Syllis cornuta, H. Rathke, off the Hebrides. X Zeiss oc. 2, obj. D.

16 b and 16 c. Bristles of the same procured off Donegal Bay by the 'Porcupine.'

X Zeiss oc. 2, obj. D.
17. Upper anterior bristle of Syllis gracilis, Grube. X Zeiss oc. 2, obj. D.

17 a. Lower bristle from the same region, x as above.

17 b. Strong bristle from the posterior region of the foregoing, from Herm. X as above.

17 c. Similar bristle from an example from St. Andrews. X as above.

18 and 18 a. Bristles of Syllis Buskii, McL, from Guernsey, x Zeiss oc. 2, obj. D.

19. Bristle of Syllis brevicirrata var. plana, from Herm. x Zeiss oc. 2, obj. D.

20. Bristle of Amblyosyllis lineata, Grube. X Zeiss oc. 2, obj. D.

21. Bristles of Autolytus pictus, Ehlers. X 700.

22. Bristle of Autolytus aurantiacus, Clap. X about 700.

22 a. Another example seen from the front, the terminal piece being twisted, x about 700.

23. Compound bristle of Autolytus prolifer, O. F. M., seen obliquely. X Zeiss oc. 2, obj. D.

23 a. Swimming bristle of a bud. x Zeiss oc. 2, obj. D.

24. Falcate bristle of Myrianida pinnigera, Mont, x Zeiss oc. 2, obj. D.

24 a. Swimming bristle of a bud of the foregoing, x Zeiss oc. 2, obj. D.

25. Falcate bristles of the tenth foot of Nereis pelagica, L. x Zeiss oc. 2, obj. D.

25 a. Dorsal bristle of the same. X Zeiss oc. 2, obj. D.

25 b. Sabre-shaped bristle of the epitokous form, x Zeiss oc. 2, obj. D.

26. Bristle of Sylline rubropunctata, Grube. x Zeiss oc. 2, obj. C.

27. Bristle of Bhyllodoce Baretti, De Blv. X Zeiss oc. 2, obj. C.

1 The small bifid bristle also accidentally labelled 8 a in the middle of this plate pertains to Syllis gracilis.

Pl.LXXX

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PLATE LXXXI.

Fig.

1. Bristle with shorter tip in inferior division of the tenth foot of Nereis cultrifera,

Grube. X Zeiss oc. 2, obj. C.

1 a. Dorsal bristle of the foregoing. X Zeiss oc. 2, obj. C.

2. Bristle from the dorsal division (with slender tapering tip) of Nereis Schmarclsei, De

Quatref. (irorata, Mgrn.). X Zeiss oc. 2, obj. D.

2 a. Bristle with short tip from same division. X Zeiss oc. 2, obj. D.

2 b. Bristle with longer tip from ventral division. x Zeiss oc. 2, obj. D.

2 c. Swimming bristle of the same. X Zeiss oc. 2, obj. D.

3. Dorsal bristle of Nereis Marionii, And. and Edw. X Zeiss oc. 2, obj. D.

3 a. Stouter bristle from the lower group of the inferior lobe of the foregoing. X

Zeiss oc. 2, obj. D.

4. Dorsal bristle of Nereis Dumerilii, Aud. and Edw. X Zeiss oc. 4, obj. F.

4 a. Ventral bristle of the same. X Zeiss oc. 4, obj. F.

4 b. Swimming bristle of epitokous form. X Zeiss oc. 2, obj. C.

4 c. Ventral bristle (with short tip) of the same epitokons example. X Zeiss oc. 2,

obj. C.

5. Dorsal bristle of Nereis diversicolor, 0. F. Mull. X Zeiss oc. 2, obj. C.

5 a. Upper bristle of the lower ventral tuft of the same. X Zeiss oc. 2, obj. C.

5 b. Lower bristle of the lower ventral tuft. X Zeiss oc. 4, obj. C.

6. Dorsal bristle of Eunereis longissima, Johnst., var., from Bay of Tunis. X Zeiss

oc. 2, obj. D.

6 a. Falcate bristle from the ventral lobe. X Zeiss oc. 2, obj. D.

7. Bristles of the dorsal lobe of Nereis (Alitta) virens, Sars; a, entire tip; 7b, articula-

tion viewed from behind ; 7 c, portion of shaft. X 350.

8. Superior bristle of the anterior third of Nereis (Nereilepas) fucata, Savigny. X Zeiss

oc. 2, obj. D.
8 a. Upper bristle of the inferior series with short tip. X Zeiss oc. 2, obj. D

8 b. Falcate bristle of the ventral series of the foregoing, x Zeiss oc. 2, obj. D.

9. Dorsal bristle of Staurocejphalas rabrovittatus, Grube, from Herm. X Zeiss oc. 2,

obj. D.

9 a and 9 b. Ventral bristles with bifid terminal pieces. X Zeiss oc. 2, obj. D.

10. Bifid dorsal bristle of Staurocephalus Kefersteini, McL, from Lochmaddy. X Zeiss

oc. 2, obj. F.

10 a. Simple tapering serrated bristle below the foregoing. X Zeiss oc. 2, obj. F.

10 b. Falcate compound form with bifid tip (one of medium length from the middle of

the group). X Zeiss oc. 2, obj. D.

11. Simple tapering dorsal bristle of Slaurocephalus ciliatus, Keferst., from St. Peter

Port, Guernsey. X Zeiss oc. 2, obj. F.

11 a. Bifid bristle (stout). X Zeiss oc. 4, obj. F.

116. and lib'. Compound bristles from the ventral series, with a tip of medium length.
u X Zeiss oc. 4, obj. F.

Tl. iXXXI

EX.fiiAHW.dLel.

f a EWMIrap :mpr.

PLATE LXXXIL

Fig.

1. Dorsal bristles of Ophryotrocha puerilis. X Zeiss oc. 2, obj. D.

1 a. Ventral bristle. X Zeiss oc. 2, obj. D.

2. Dorsal bristle of Lumbriconereis fragilis, 0. F. Mull. X Zeiss oc. 2, obj. D.

2 a. Winged hook of the fiftieth segment. X Zeiss oc. 2, obj. D., reduced.

2 b. Hook from the same foot with terminal prolongation of the guard. X Zeiss oc. 2,

obj. D, reduced.

3. Dorsal winged bristle of Lumbriconereis Latreillii, Aud. and Edw. X Zeiss oc. 2,

obj. D.

3 a. Articulated and winged hook from foot in front of tenth. X Zeiss oc. 2, obj. D,

reduced.
3 b. Articulated and winged anterior hook with shorter terminal piece (behind tenth
foot). X Zeiss oc. 2, obj. D., reduced.

3 c. Posterior hook. x Zeiss oc. 2, obj. D.

4. Long winged hook from the first foot of Lumbriconereis ivrpatiens, Clap. X Zeiss

oc. 2, obj. D.

4 a. Another example from the same foot which shows a tendency to form an articulation.

X Zeiss oc. 2, obj. D.
4b. Stronger dorsal bristle from fifty-seventh foot. X Zeiss oc. 2, obj. D.

4 c. More slender form from the same foot. X Zeiss oc. 2, obj. D.
4d. Hook from the middle of the body. X Zeiss oc. 2, obj. D.

5. Winged bristle of Lumbriconereis hibernica, Mel. X Zeiss oc. 2, obj. D, draw-tube.
ba. Long, winged tapering hook from the tenth foot. X Zeiss oc. 2, obj. F.

5 b. The same viewed from the front so as to show both wings at the tip. X Zeiss oc.

2, obj. D.
be. Long hook from the thirtieth foot. X Zeiss oc. 2, obj. D.

5 d. Long hook considerably behind the foregoing. X Zeiss oc. 2, obj. D.

6. Tenth foot of Lumbriconereis gracilis, Grube, from Lochmaddy. X Zeiss oc. 2, obj. D.

6 a. Bristles from thirtieth foot of the foregoing. X Zeiss oc. 2, obj. D.

7. Tenth foot of Lumbriconereis Gr. X 200.

7 a. Elongated hook from the middle of the body. X Zeiss oc. 2, obj. D.

8. Dorsal bristle (above spine) of Lumbriconereis (Zygolobus) laurentianus, Grube. X

Zeiss oc. 2, obj. D.

8 a. Bristle with bow-like curve below the spine. X Zeiss oc. 2, obj. D.

8 b. Long winged hook of the twentieth foot seen from the front. x Zeiss oc. 1, obj. D.
8 c. Profile of the same. X Zeiss oc. 1, obj. D.

PI. LXXXIt

A.HW. del

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PLATE LXXXIII.

Fig.

1. Spine of Drilonereis longa, Webst., var. Elisabeths, Mel. X 100.

1 a. Tip of foot with bristles. X Zeiss oc. 2, obj. D.

2. Dorsal bristle of Arabella tricolor, Mont, x Zeiss oc. 2, obj. D.

2 a. Bristles below the former with bold serration. x Zeiss oc. 2, obj. D.

3. Bristle of Notocirrus scoticus, Mel. X Zeiss oc. 2, obj. D.

4. Dorsal bristle of Eunice fasciata, Bisso. X Zeiss oc. 2, obj. D.

4 a. Falcate bristle, x Zeiss oc. 2, obj. D.

5. Anterior dorsal bristle of Marphysa sanguinea, Mont. x Zeiss oc. 2, obj. D.

5'. Another showing slightly broader tip. X Zeiss oc. 4, obj. D, with 2-in. draw-tube.

5 a. Brush-like bristle. X Zeiss oc. 4, obj. F.

5 b. Variety of the latter near the tip of the tail, slightly twisted. X Zeiss oc. 4, obj. F.

5 c. Ventral bristle. X Zeiss oc. 4, obj. F.

6. Dorsal bristle of Marphysa Kinbergi, McL, from the ' Porcupine,' 1870. X Zeiss oc. 2,

obj. D.

6 a. Jointed bristle. X Zeiss oc. 4, obj. D.

6 b. Tip of shaft of the foregoing turned so as to exhibit its structure. X Zeiss oc. 4,

obj. D.

7. Dorsal bristle of Eunice vittata, D. Chiaje. X Zeiss oc. 4, obj. D.

7 a. Falcate bristle (ventral). X Zeiss oc. 4, obj. F.

7 b. Great hook with a bifid crown, from Adventure Bank, ' Porcupine,' 1870. X Zeiss

oc. 4, obj. F.

8. Falcate bristle of Eunice norvegica, L., from Norway. x Zeiss oc. 2, obj. D.

8 a. Posterior hook, x Zeiss oc. 2, obj. D.

9. Dorsal bristle of Eunice norvegica, var. B.A. 'Porcupine.' x Zeiss oc. 2, obj. D.

9 a. Brush-like bristle. X Zeiss oc. 2, obj. D.

9 b. Falcate ventral bristle. X Zeiss oc. 2, obj. F.

9 c. Posterior hook, x Zeiss oc. 2, obj. D.
10. Brush-like bristles of dorsal series of Eunice floridana, Pourt. X Zeiss oc. 2, obj. D.
10 a. Falcate bristle. X Zeiss oc. 4, obj. D.
10 b. Posterior hook. x Zeiss oc. 4, obj. D.
10 c. Dorsal bristle of the same, x Zeiss oc. 4, obj. D.

pi. lxxxiit

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A.H.W. del

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PLATE LXXXIV.

Fig.

1. Dorsal bristle of Nematonereis unicornis, Grrube, from St. Peter Port, Guernsey.

X Zeiss oc. 2, obj. D, full draw-tube.
1 a. Posterior hook, x 350.
1 b. Falcate bristle. X Zeiss oc. 4, obj. F.

1 c. Brush-shaped bristle from a posterior foot. Zeiss oc. 2, obj. D.

2. Dorsal bristle of Lysidice punctata, Risso. X Zeiss oc. 2, obj. D. with 3-in. draw-

tube.

2 a. Falcate bristle. X Zeiss oc. 2, obj. D, with 3-in. draw-tube.

2 a/. Falcate bristle of the tenth foot. X Zeiss oc. 2, obj. D, with 3-in. draw-tube.

3. Falcate bristle of the first foot of Onuphis britannica, Mel. The folds of the two

wings make a complex tip. X 200.

3 a. Simple winged bristle from the twentieth foot. X 200.

3 b. Brush-like bristle from the twentieth foot. X Zeiss oc. 2, obj. F.

3 c. Powerful hook from the same segment. X 200.

4. Dorsal bristle of Onuphis brevibrachiata, Ehlers. X Zeiss oc. 2, obj. D.

4 a. Ventral bristle. X Zeiss oc. 2, obj. D.

4b. Powerful winged hook from the twentieth foot. X Zeiss oc. 2, obj. D.
4c. Brush-like bristle (anterior). X Zeiss oc. 4, obj. D.

5. Falcate bristle of first foot of Onuphis conchylega, Sars. Wings have been abraded.

X Zeiss oc. 2, obj. D.

5 a. Developing bristle (in the tissues), showing wings. X Zeiss oc. 2, obj. D.
5 b. Dorsal bristle. X Zeiss oc. 2, obj. D.

5 c. Ventral hook. X Zeiss oc. 2, obj. D.

6. Bristle of the first foot of Onupjhis (Diopatra) fragosa, Ehlers. X Zeiss oc. 2, obj. D.

6 a. Dorsal bristles of the twentieth foot. X Zeiss oc. 2, obj. D.

6 b. Ventral hook, x Zeiss oc. 2, obj. D.

7. Falcate bristle from the anterior region of Onuphis sicula, De Quatref. X 700.

7 a. Another example of the foregoing. X Zeiss oc. 2, obj. D.

7 b and 7 c. Dorsal bristles, the former with just a trace of a wing, x Zeiss oc. 2, obj. D.,
with draw-tube (full).

7 d. Posterior hook. X 700.

8. Falcate bristle of the first foot of Hyalinoecia tubicola, O. F. Mull, (small example).

X 200.

8 a. Dorsal bristle (anterior region). X Zeiss oc. 2, obj. D.

8 b. Brush-like bristle of the same region. X Zeiss oc. 2, obj. D.
8 c. Powerful winged hook from the eightieth foot. X Zeiss oc. 2, obj. D.
Sd. Lower bristle of the dorsal series with broad wings, from the eighth foot. X Zeiss
oc. 2, obj. D.

9. Hook of Lysidice punctata, Risso. X Zeiss oc. 2, obj. D. with draw-tube.

PL. JLXXXIV.

A.H.W. u W.C.KA&l

f a PAY.M-Tra-pimpr.

PLATE LXXXV.

Fig.

1. Dorsal bristle of Goniada metadata, (Erst. X Zeiss oc. 2, obj. D.

1 a, 1 a. Ventral bristles. X Zeiss oc. 2, obj. D.

2. Ventral bristle of Glycinde Nordmanni, Mgrn., in lateral view. X Zeiss oc. 3, obj. D,

with draw-tube.

2 a. A form with longer tip seen from the front. X Zeiss oc. 2, obj. D, with draw-tube.

2 /;. Dorsal bristle with process at tip. X Zeiss oc. 2, obj. D, with draw-tube.

3. Dorsal bristle of Ghjcera lapidum, De Quatref., from the ' Porcupine ' (muddy sand),

1870. X Zeiss oc. 2, obj. D.

3 a. Ventral bristle of the ordinary kind. X Zeiss oc. 2, obj. D.

3 b. Larger (upper and lower) ventral bristle from the ninetieth foot. X Zeiss oc. 2,

obj. D.

4. Dorsal bristles of Ghjcera siphono stoma , D. Chiaje. X 280.

4 a and 4 b. Ventral bristles. X 280.

5. Dorsal bristle of Ghjcera alba, H. Rathke, from Cape Gruardia, ' Porcupine,' 1870.

X Zeiss oc. 2, obj. D.

5 a. Ventral bristle. X Zeiss oc. 2, obj. D.

6. Dorsal bristle of Ghjcera Goesi, Mgrn., from St. Magnus Bay. X Zeiss oc. 2, obj. D,

with full draw tube.

6 a. Ventral bristle. X Zeiss oc. 2, obj. D, with full draw -tube.

7. 7', 7". Dorsal (camerated) bristles of Aricia Cavieri, Aud. and Edw., the latter bristle

in profile. X Zeiss oc. 2, obj. D.

7 a. Strong brownish ventral bristle. X Zeiss oc. 2, obj. D.

7 b. Larger example. X Zeiss oc. 2, obj. D.

8. Dorsal bristle of Aricia Grubei, Ehlers, from the 'Porcupine,' 1870. X Zeiss oc. 2,

obj. D.

8 a. Stout spine-like bristle of the inferior division of the foot. X Zeiss oc. 2, obj. D.
8b. Hastate spine. X Zeiss oc. 2, obj. D.

9. Lateral view of dorsal bristle (camerated) of Aricia norvegica, Sars. X Zeiss oc. 2,

obj. D.

9 a. Hastate spine. X Zeiss oc. 2, obj. D.

9 b. Bifurcate bristle with spinous forks. X Zeiss oc. 2, obj. D.
10. Dorsal bristle of Nainereu sp., from St. Andrews. X Zeiss oc. 2, obj. A.
10 a. Tip of the same from the tenth foot. X Zeiss oc. 2, obj. D.

pi. lxxxv:

A.H.W. del

f a F.W.MTrap impr.

PLATE LXXXV1.

Fig.

1 and 1 a. Dorsal bristles of Scoloplos armiger, 0. F. Mull. X Zeiss oc. 2, obj. D.

2. Dorsal bristle of the tenth foot of Nainereis quadricuspida, Fab., Lochmaddy. X

Zeiss oc. 2, obj. D.

2 a. Somewhat clavate strong bristles (like spines) in the ventral division of the tenth

foot. X Zeiss oc. 2, obj. D, with full draw-tube.

2 b and 2 V '. Dorsal bristles from the middle of the body. X Zeiss oc. 2, obj. D, with

full draw -tube.

3. Simple dorsal bristle of Marphysa Belli, Aud. and Edw. X Zeiss oc. 2. obj. F.

3 a. Falcate compound bristle of Marphysa Belli, Aud. and Edw. X Zeiss oc. 2, obj. F.
3 b. Brush-shaped bristle of Marphysa Belli, Aud. and Edw. X Zeiss oc. 2, obj. F.

3 c. Compound bristle of Marphysa Belli with long tapering tip. It has been a little
compressed at the articulation. X Zeiss oc. 2, obj. F, with 1-in. draw-tube.

3 d. Long ventral hook, with bifid tip, of Marphysa Belli, Aud. and Edw. X Zeiss oc. 2,

obj. F, with 3-in. draw-tube.

4 and 4'. Dorsal bristles of Glycera Ehlersii, Arwid. X Zeiss oc. 4, obj. D.
4 a. Ventral bristles of Glycera Ehlersii, Arwid. x Zeiss oc. 4, obj. D.

5. Bristle of Eulalia G. (var. of E. biliueata ?) from deep water off St. Andrews Bay.

x Zeiss oc. 4, obj. C.

6. Bristle of Syllis liamaia, var. Gunning hami. x Zeiss oc. 4, obj. C.

7. Dorsal bristle from the anterior region of Lumbriconereis acutifrons, Mel. X Zeiss

oc. 2, obj. D, with 3-in. draw-tube.
7 a. One of the slender series from the same region as the foregoing. X Zeiss oc. 2,

obj. D, with 3-in. draw-tube.
7 b. Shorter winged hooks from the middle of the body of the same. X Zeiss oc. 2, obj.

D, with 3-in. draw-tube.

8. Typical bristle of Micronereis variegata, Clap. X 700.

9. Dorsal bristle of Leptonereis Vaillanti, De St. Jos. X 700.

9 a. Ventral (heterogomph) bristle of Leptonereis VaillaMi, De St. Jos. X 700.

10. Long dorsal bristle of Stauroceplialus pallidas, Langerh. X Zeiss oc. 2, obj. D.
10 a. Bifid dorsal bristle of Stauroceplialus pallidas, Langerh. x Zeiss oc. 2, obj. D.
10 6. Ventral bristle of Stauroceplialus palUdus, Langerh. x Zeiss oc. 2, obj. D.

11. Bristle of Grubea clavata, Clap. X Zeiss oc. 2, obj. D.

12. Bristle of Pionosyllis lamelligera, De St. Jos. X Zeiss oc. 2, obj. D.

13. Bristle of Pionosyllis alternosetosa, De St. Jos. x Zeiss oc. 2, obj. D.

14. Bristle of Trypanosyllis coeliaca, De St. Jos. X Zeiss oc. 2, obj. D.

15. Bristle of Pur y syllis par adoxa, Clap. X Zeiss oc. 2, obj. D.

16. Bristle of Autolytus elibiensis, De St. Jos. X Zeiss oc. 2, obj. D.

17. Bristle of Autolytus longiferiens, De St. Jos. X Zeiss oc. 2, obj. D.

18. Bristle of Autolytus inermis, De St. Jos. X Zeiss oc. 2, obj. D.

19. Bristle of Nainereis, from St. Andrews (West Sands). X Zeiss oc. 2, obj. D.

PLILXXXVI.

ARM. del.

Tir-rn.. RTOLTrap irrvpr.

PLATE LXXXVII.

Fig.

1. Tenth foot of Aricia Grubei, Mel. x Zeiss oc. 2, obj. A.

2. Tenth foot of Aricia norvegica, Sars, from Bergen, x Zeiss oc. 2, obj. A.
2 a. Twenty-fifth foot, x Zeiss oc. 2 5 obj. A.

3. Head and anterior region of Eurysyllis paradoxa, Clap, (after Ehlers). Enlarged.

4. Tenth foot of Scoloplos armiger, 0. F. Mull. Enlarged.
4 a. Twentieth foot. Enlarged.

4 b. Thirtieth foot, from Symbister Harbour. Enlarged.

4 c. Head of the foregoing. Enlarged.

5. Tenth foot of Nainereis quadricuspida, Fabr., from Lochmaddy, North Uist. x 2]

5 a. Twenty-third foot (about), x 21.
5 b. Posterior foot. X 21.

6. Typical foot of Glycera Ehlersii, Arwid., from behind, x 21.

7. Dorsal view of Liimbriconereis acuiifrons, Mel. Enlarged.

8. Tenth foot of Aricia Edwardsi an. var. A. Ouvieri. Enlarged.

9. Head of Staurocephalus pallidas, Langerh. Enlarged.
9 a. Foot of the foregoing. x 30.

10. Foot of Grubea clavata, Clap. x 30.

11. Foot of Pionosyllis lamelligera, De St. Jos. x 30.

12. Foot of Pionosyllis alternosetosa, De St. Jos. x 30.

13. Foot of Trypanosyllis coeliaca, De St. Jos. X 30.

14. Foot of Eurysyllis par adoxa, Clap. X 30.

15. Foot of Autolytus eMiensis, De St. Jos. x 30.

16. Foot of Autolytus longiferiens, De St. Jos. X 30.

17. Foot of Autolytus inennis, De St. Jos. X 30.

18. Lateral view of Aricia Ouvieri, And. and Edw. Enlarged.

19. Mnth foot of Aricia Armandi, Mel. X 35.

20. Twenty-third foot of the same. X 35.

21. Tenth foot of Goniada eremita, Aud. and Edw. X 35.

Pl.LXXXVJI.

INDEX TO VOL. I

(Not including the Introduction.)

PAGE

PAGE

Acanthicolepis .

. 311

Enipo . .395

asperrima .

. 311

Kinbergi .

. 395

Acholoe

. 396

Enopla

. 155

astericola .

. 397

Eunoa

. 291

Acoetidee .

. 399

nodosa

. 291

Amphinomidas .

. 219

tritoni

. 296

Amphinomina .

. 219

Euphrosyne

. 233

Amphiporidse .

. 155

armadillo .

. 238

Amphiporinse .

. 155

foliosa

. 234

Amphiporus

. 155

Robertson i .

. 240

bioculatns .

. 163

Euphrosynina .

. 226

hastatus

. 162

Eurythoe" .

. 224

lactifloreus

. 156

borealis

. 224

pulcher

158

Eusthenelais

. 425

spectabilis .

160

hibernica .

. 425

Anopla

181

Evarne

. 353

Antinoe

365

atlantica

. 363

finmarchica

368

Hubrechti .

. 360

mollis

369

impar

. 353

Sarsi .

365

Johnstoni .

. 358

Aphrodita.

241

aculeata

247

Gattyana .

. 285

Aphroditidse

240

cirrosa

. 285

Borlasia

193

Halosydna

. 384

Elizabeths

193

gelatinosa .
Harmothoe

. 384
. 313

Carinella .

203

antilopis

. 334

annulata

203

areolata

. 349

linearis

206

Fraser-Thomsoni

. 337

Carinellida3

203

Haliaeti

. 336

CephalotliricidaB

208

imbricata .

. 314

Cephalothrix

208

ljungmani .

. 332

linearis

208

lunulata

. 342

Cerebratulus

194

marphysas .

. 339

angulatus .

195

setosissima

. 345

444

INDEX.

Harmathoe spinifera

zetlandica .
Hermione .

hystrix

Laetmatonice

filicornis

producta var. britannica
Lagisca

Elizabethan

extenuata .

floccosa

Jeffreysii .
Leanira

hystricis
Lepidonotus

clava .

squamatus .
Lineidse .
Lineus

bilineatus .

gesserensis

lacteus

marinus

sanguineus

Malmgrenia

andreapolis

castanea
Meckelia .

asulcata
Micrnra

aurantiaca .

fasciolata .

fusca .

purpurea .

Nemerteans
Nemertes .

carcinophila

gracilis

Neesii
Nemertinas

PAGE

327

Panthalis .

330

GErstedi

264

Paramphinome .

264

pulchella .

Pholoe

258

minuta

258

Phyllantinoe

263

mollis

298

Polychaata

303

Poly no e

307

scolopendrina

298

Polynoidae

305

Prosorhochmus .

434

Claparedii .

434

273

Scalisetosus

280

assimilis

274

communis .

181

Si galion .

181

Buskii

191

Mathildas .

185

Sigalionidas

190

Spinther .

181

miniaceus .

188

oniscoides .

Stlienelais

379

atlantica

382

boa .

379

JefTreysii .

202

limicola

202

? sp. .

196

zetlandica .

201

197

Tetrastemma

196

Candida

200

dorsalis

flavida

155

melanocepliala

176

PobertianaB

180

vermicula .

176

178

Yalencinia

176

li nef or mis .

PAGE
400

400
221
222
436
437
370
371
217
389

270
174
174

372

377
372
427
431
427
406
227
232
230
407
415
408
421
417
424
414

164
167
172
170
165
166
169

207
207

ADLABD AND SON, IMPR., LONDON AND DORKING.

BAY SOCIETY.

(INSTITUTED 1844.)

OFFICERS AND COUNCIL,
1910-11.

president :

The Eight Hon. LOED AVEBUEY, P.O., D.C.L, LL.D, F.E.S, Pres.S.A, F.L.S., F.G.S, F.Z.S.

DiceHPreeibente :

Sir CHAELES ELIOT, M.A., D.C.L, LL.D., K.C.M.G, C.B., F.Z.S., F.E.G.S.
B. DAYDON JACKSON, Ph.D., F.L.S.
S. F. HAEMEE, M.A., F.E.S., V.P.Z.S.

Council :

Eobert Adkin, F.E.S.
G. T. Bethune-Bakkr, F.L.S. , F.E.S.
Eev. C. E. N. Burrows, F.E.S.
T. A. Chapman, M.D., F.Z.S.
Eev. Alfred Fuller, M.A., F.E.S.
A. E. Gibbs, F.L.S., F.E.S., F.E.H.S,
Albert H. Jones, Treas.E.S.
Henry Layer, M.B.C.S.

G. B. Longstape, M.D., F.E.S.

Edward Meyrick, B.A., F.E.S., F.Z.S., F.E.S.

A. D. Michael, F.L.S., F.Z.S., F.E.M.S.,

F.E.H.S.
Charles Oldham, F.Z.S., M.B.O.U.
Eobert F. Scharfe, Ph.D., B.Sc, F.L.S., F.Z.S.
Charles D. Soar, F.E.M.S.

{Treasurer :

F. DuCANE GODMAN, D.C.L, F.E.S., F.L.S., F.G.S., F.Z.S, F.E.S.

Secretary :

JOHN HOPKINSON, F.L.S, F.G.S, F.Z.S, F.E.M.S, F.E.Met.Soc, Assoc.Inst.C.E,

Weetwood, Watford.

{Trustees :

LOED AVEBUEY; JOHN HOPKINSON; ALBEET D. MICHAEL.

PRINTED BY ADLARD AND SON, LONDON AND DORKING.

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