AY SOCIETY

A MONOGRAPH

OF THE

BBITISH ANNELIDS.

PART II.

POLTCHJETA.

AMPHINOMIDiE to SIGALIONIM2.

Pages 215—442; Plates XXIV— XXVI, XXVI a, XXVII— XLII.

w. o. Mcintosh, m.d., ll.d , f.r.s., etc.

1900.

Issued to the Subscribe 'c for the Year 1898.

CARDED AUG 14 m

DIVISION MARlNJi
IMVSBTilBRAXHB

lrS~~~?LY^

%^Ofc^-^^

% %^*^t * /f

Ray Society

A Monograph of British Annelids, by W. C. Mcintosh,

Vol. I, Pt. II. 1900

Family I Amphinomidae p # 219

" II Aphroditidae ♦ 240

11 III Polynoidae . . . . . 270

" IV Acoetidae . 599

V Sigalionidae... ....... .......... 406

THE

RAY SOCIETY

INSTITUTED MDCCCXLIV.

This Volume is issued to the Subscribers to the Ray Society for the Tear 1898.

LONDON:

MDCCCC.

ML

v./

MONOGEAPH

OF THE

BRITISH ANNELIDS.

V.l. PART II.

POLTCH^TA.
AMPHINOMUbE TO SIGAUONIM.

Pages 215—442 ; Plates XXIV— XXVI, XXVIa, XXVII— XLII

BY

WILLIAM CARMICHAEL McINTOSH, M.D.Edin.,

L.R.C.S.E., LL.D., F.R.S., F.R.S.E., F.L.S., C.M.Z.S., etc.;

PROFESSOR OF NATURAL HISTORY IN THE UNIVERSITY OF ST. ANDREWS; DIRECTOR OF THE UNIVERSITY

MUSEUM AND OF THE GATTY MARINE LABORATOBY.

LONDON :
PRINTED FOR THE RAY SOCIETY.

PRINTED BY ADLARD AND SON, BARTHOLOMEW CLOSE, E.G.

°IS.1

DEDICATED

TO

CHARLES HENRY GATTY, Esq., M.A.,

LL.D., F.K.S.E., E.L.S., F.Z.S., F.G.S., ETC.,

OF LELBK1DGE PLACE, EAST GRINSTEAD,

THE MUNIFICENT FOUNDER OF THE GATTY MARINE LABORATORY,

ST. ANDREWS.

TEMPOEAEY PBEFACE

When the author, as already explained in the Introduction to Part I, undertook the
task of placing the British Marine Annelids on a more satisfactory footing, he did not
foresee either the difficulties of isolated work in the country— apart from libraries and the
sea-shore — or the increasing strain of very responsible official duties. Again, when he
had at length secured the advantages pertaining to the sea and a university library, the
constant succession of fresh labours — connected respectively with the volume on the
Annelida of the ' Challenger,' the engrossing work accompanying the comparatively recent
development of the scientific aspects of the Fisheries, and the foundation and super-
intendence of the first Marine Laboratory in Britain — were inimical to rapid progress with
the subject. But this was not all; the revival of zoological science and zoological
teaching in a university so admirably situated for both, the reorganisation and develop-
ment of the University Museum, and the numerous official duties, requiring a large
amount of time, which devolved on him might have seemed to prove almost fatal to the
undertaking. But during this period he had unique opportunities of extending the
knowledge of the Annelids and their distribution, so that not even the absorption of time
in upholding science and medicine in the oldest Scottish university— at, perhaps, the most
critical period of its existence— for a moment caused doubt as to the ultimate progress of
the work.

Such a brief explanation is necessary, for it is no less than a quarter of a century
since the first part was published.

The great extent of the field necessitates but a brief reference to anatomy and
embryology, were it only for the cost of plates and text. These and the historical parts
can follow either in this or another channel. Publication has become the more clamant
since in this country no book of reference on the subject is up to date, and hence the
group has not received that care from zoologists which is desirable — for instance, in
extending the species and accurately defining their distribution.

Amongst those who have largely aided the author by the collection of specimens the
late Dr. Gwyn Jeffreys holds a foremost place, since he took with him for several years
specially prepared bottles for the preservation of the Annelids, and thus forwarded them
in excellent condition, I desire to record my grateful remembrance of his constant care
in this respect. Exceeding even the foregoing in number and variety of specimens come the
collections made by several relatives in St. Andrews, one of whom for more than twenty
years enriched the museum of the author in a noteworthy manner. Few took deeper

viii TEMPORARY PREFACE.

interest in the progress of the work than the late Prof. George Busk, whose unselfish
devotion to zoology is another memorable example of the brotherhood of medicine and
natural history. The late Mr. Spence Bate, of Plymouth, Dr. Baird, of the British
Museum, Dr. W. B. Carpenter, Sir Wyville Thomson, Dr. Bowerbank, Dr. David
Robertson, of Cumbrae, Prof. Grube, of Breslau, Edouard Claparede, of Geneva, Dr.
Howden, of Montrose, Dr. Carrington, of Eccles, Mr. A. G. Moore, of Dublin, James
Duncan Matthews, of Aberdeen, and Profs. Dickie, Ogilvie, and Dyce, of the same place,
were of signal service in forwarding specimens. I have also to acknowledge similar and
valued aid from the late Prof. Allman, Canon Norman, Prof. Ray Lankester, Mr. Whiteaves,
of Canada, Ludwig von Graff, of Gratz, Mr. Brooking Rowe, Plymouth, Miss Florence
Buchanan, Prof. G. S. Brady, Prof. Percival Wright, Prof. Hoavcs, Sir John Murray,
Prof. Haddon, Dr. Scharff, Dublin, Mrs. Collings, late of Sark, Dr. E. P, Ramsay and Prof.
Haswell, of Sydney, Dr. John Anderson, late of Calcutta, Prof. Herdman, Mr. Beddard,
Dr. Sydney Harmer, Prof. Jeffrey Bell, Mr. J. T. Cunningham, Prof. Ewart, Mr. Shipley,
Mr. Parfitt, Exeter, Dr. Fulton and Mr. H. C. Williamson, of the staff of the Fishery
Board, Mr. Arnold Watson, Sheffield, Mr. Shrubsole, Sheerness-on-Sea, Mr. Sibert
Saunders, Whitstable, and Mr. Hornell, of Jersey.

For assistance in other respects I have gratefully to remember Prof. Giarcl, of Paris,
Prof. Alex. Agassiz, A. E. Verrill, John Murdoch, J. Walter Fewkes, Prof. H. E. Webster,
Dr. H. P. Johnson, Miss M. Lewis, J. E. Benedict, Dr. T. H. Montgomery, of the United
States, Dr. J. P. Hill, of Sydney, Prof. A. G. Bourne, Madras, Prof. W. B. Benham, Otago,
Prof. Harvey Gibson, Liverpool, J. Jennings Hind, Mr. Garstang, Plymouth, Prof. Gilson,
of Louvain, the late Prof. Loven, Stockholm, the late Dr. A. J. Malmgren, Helsingfors,
Dr. Emil Marenzeller, Vienna, Prof. A. F. Marion, Marseilles, MM. Pruvot and Racovitza,
Prof. G. 0. Sars, Dr. Fridiof JSTansen, Christiania, Prof. Hjalmar Theel, Stockholm, Dr. A.
Wiren, Upsala, Prof. Hubrecht, Utrecht, Prof. Haeckel, Jena, Dr. W. Michaelsen, Hamburg,
Prof. C. 0. Minot, Boston, Prof. Playfair McMurrich, Michigan. Dr. Langerhans, Madeira,
Louis Roule and Baron Jules de Guerne, Paris, Baron de St. -Joseph, Prof, Paul Hallez,
Prof. Barrois and M. Malaquin, Lille, F. Mesnil, Paris, M. Caullery, Lyons, Pierre Fauvel,
Caen, Julian Fraipont, Prof. Julin, Liege, Jules Bonnier, E. Brumpt, Dr. Kiikenthal,
Jena, Prof. Ehlers, Gottingen, Prof. B. Hatschek, Dr. R. Greef, Dr. F. Yejdovsky, Praag,
Prof. Hensen, Kiel, Prof. K. Brandt, Dr. J. Reibisch, Th. Kuhlgatz, Dr. V. Hacker,
Freiburg, P. Tauber and G. M. R. Levinsen, Copenhagen, Dr. R, Horst, Leiden, Dr. G. A.
Hansen, Dr. C. Apstein, Kiel, Dr. Apellof, Bergen, Ivar Ardwidsson, Dr. C. Aurivillius,
Dr. D. Carazzi, Florence, Dr. S. Orlandi, Di\ C. J. Cori, A. Birula, St. Petersburg, Prof.
Mobius, Berlin, Prof. Kishinouye, U. Takura, S. Goto, S. Hatai, and A. Iizuka, Tokyo,
the late Prof. Moseley, Dr. Murie, Capt. Fielden, Dr. Marcus Gunn, the Duke of Argyll,
and the late Sir Walter Elliot, of Wolflee. My colleague, Prof. Lawson, kindly looked
over certain proof-sheets when they were delayed by an illness; Dr. A. T. Masterman
also gave me his valued aid at this time. I am also indebted to Prof. Wiltshire for his
careful revision and suwestions.

Various Fishery officers in Scotland, especially Mr. Bain of Peterhead, Mr. Duthie of
Lerwick, Mr. Mair of Montrose, and Mr. Cooper, formerly of Aberdeen, have been of
much service in forwarding specimens used as bait or otherwise interesting

TEMPORARY PREFACE. ix

For the best of the coloured drawings from life I am indebted to my late sister, Mrs.
Giinther, the loss of whose delicate touch and experience has been greatly felt in the
completion of the work. I have also to thank Dr. A. T. Masterman, Rev. J. M. Anderson,
my artist— Miss Ada Walker, and Miss Burnet of St. Andrews, for aid in this respect.
The present part contains comparatively plain examples of the group, which is remarkable
for beauty of coloration in such as the Phyllodocidge, Nereidae, and many others.

It has been found impossible to represent all the parts required for identification from
the living form, because many have not been seen alive, or if so, under circumstances
inimical to anything else than preservation. Yet this is not altogether a drawback, since
many workers have only spirit-specimens to deal with, and the parts so preserved retain
certain useful characteristics. On this head, indeed, diverse views are held, one of the
most recent being that of Dr. H. P. Johnson, of California, who asserts that descriptions
from life are only advantageous in respect to colour.

W. C. M.

G-atty Marine Laboratory,
St. Andrews ;
July,

215

DESCRIPTION

OF THE

GENERA AND SPECIES

OP THE

BRITISH MAEINE ANNELIDS

(ANNELIDA POLYCIDETA).

29

217

Order POLYCH.ETA.

Marine worms usually with a definite head (prostomium), which dorsally bears eyes
and tentacles, and ventrally palps, besides possessing other sense-organs.

From the buccal region (peristomium), which frequently carries cirri or other organs,
the fore-gut often sends out a protrusible proboscis — armed or unarmed. (Esophagus may
have a pair of diverticula. Alimentary canal simple or branched, and a median dorsal
and median ventral mesentery may be present.

Body of one or more regions, generally elongate and cylindrical, sometimes flattened
and compressed, of numerous segments, which internally are marked rather by the
oblique muscles than by diaphragms. The last segment in many has long ventral (anal)
cirri. It is invested by a thin, tough cuticle covering a granular layer, 1 bounded
internally by basement-lining, then a more or less complete layer of circular muscular
fibres, while within are four longitudinal muscular bands, — as a rule, two dorsal and
two ventral. The oblique muscles converge from the dorso-lateral region to the vicinity
of the nerve-cords.

The segments in the free examples bear lateral processes of the body- wall (feet or
parapodia), and each foot is frequently divided into a dorsal and a ventral lobe, with a
spine (aciculum) to which motor muscles are attached, and a group of specially
differentiated bristles, besides a dorsal and a ventral cirrus.

Respiratory organs either free branchiae or the general surface.

Circulatory organs generally closed, contractile or non-contractile vessels. Blood
red, pink, green, or pale, corpusculated or non-corpusculated (?) ; occasionally absent.

Perivisceral (coelomic) fluid abundant, highly organised and corpusculated. In
certain forms the perivisceral corpuscles are coloured, and the whole may act as a
substitute for the circulatory fluid.

Nervous system consists of cephalic and subcesophageal ganglia with commissures,
separate or closely applied longitudinal ganglionated trunks, with or without neural
canals ; placed in or internal to the granular layer under the cuticle. A stomato-
gastric system of nerves also occurs.

Reproductive organs are developments of the perivisceral (coelomic) epithelium in
connection with the ventral blood-vessels. The products escape by the segmental
organs (nephridia), or by rupture of the body- wall. Sexes, as a rule, are separate, but
some are hermaphrodite. Polymorphism and asexual reproduction also occur.

The segmental organs may act as genital ducts, and are arranged in pairs in few or
many segments.

Larva a trochophore. Metamorphosis during development.

Free or sedentary (errant or tubicolous).

1 Formerly named hypoderm, a term, however, which has given rise to misapprehensions.

219

FAMILIES, GENERA, AND SPECIES, OE THE BRITISH MARINE

ANNELIDS.

(ANNELIDA POLYCELETA.)

Order— POLYCELETA.

Family I. — Amphinomim:.

Cephalic lobe rounded or compressed. A median and two lateral tentacles, though
the latter may be absent, an elongated dorsal caruncle, and four eyes. Body elongate,
oblong or ovate-oblong, feet with the dorsal and ventral divisions widely separated and
furnished with cirri. Mouth removed from the tip of the snout ventrally, with
modified segments laterally; protrusible proboscis devoid of jaws. Feet peculiarly
modified, the dorsal lobe being extended and merged into the dorsum, but with bristles,
branchiae, and cirri. Bristles brittle, calcareous, and tubular, with gelatinous contents ;
rarely hook-like spines. Buccal apparatus and proboscis large and complex; alimentary
canal often with a caacum in front. Anus dorsal. Two posterior appendages. Nerve-
cords either lie within the body-wall, the decussation of the oblique muscular bands
being beneath them, or the latter are attached at the outer border of each trunk.
Oviparous.

Sub-family — Amphinomina.

Cephalic lobe rounded, furnished with a median and two lateral tentacles; body
elongate; branchiae on the dorsum of the feet, which have their dorsal and ventral
divisions widely separated. Bristles brittle, calcareous, tubular, with gelatinous
contents.

Nerve-cords somewhat small and flattened, occupying an area bounded internally
by a transverse band of fibres, and externally by the circular muscular layer and the
granular layer beneath the cuticle. The oblique muscles are attached at the outer
border of each trunk. Intestine simple.

Savigny in his ' Systeme ' (1820) made the Amphinomse the fourth Family of his
first Order Nereids, characterising them as follows :— Branchiae large, complex,

220 AMPHINOMINA.

situated along the superior border of all the dorsal divisions of the feet, the first three
or four excepted, or behind the border, and extending to the ventral branch, resembling a
pinnatifid leaf (feuilles), with tufts or arbuscles, which generally divide at their origin
into several trunks, now coalescing, and again separating more or less distantly the one
from the other. Mouth with a short proboscis, opening longitudinally at the extremity,
without prominent folds or tentacles, and devoid of jaws. Eyes two or four ; antennae of
moderate length and generally complete. Sometimes the middle and exterior are
absent; the unpaired is always present and inserted at the front of the caruncle
superiorly, the latter extending to the third or fourth segment. Feet with large
separate divisions, each furnished with a single bundle of bristles and without a spine.
Cirri well marked, subulate, enlarged at the base, or in the form of two articulations, of
which the one, large and short, serves as a support for the other, which is completely
retractile. They are inserted within the orifice of the sheath, behind the bundle of
bristles. The anterior feet do not differ materially from the posterior. The feet of the
first and second segments exist in all the genera.

Lamarck 1 (1818) followed Savigny in his classification of the group.

OErsted 2 placed the family Amphinomacese in his division Maricolse, characterised
by having bristled pinnge, a depressed body, segments numerous and defined, and with
the alimentary canal often branched. They were finally distinguished as Chaotopoda
Trematodina, with branchiae completely ramose. He does not include any repre-
sentative, however, in his list.

Kinberg 3 describes the Amphinomea as having the mouth inferior, formed by the
anterior segments ; pharynx protrusible, devoid of papillae and jaws ; cephalic lobe with
four eyes and a caruncle. The cirri and branchiae do not alternate. He divided the
group into two families — the Amphinomacea and the Euphrosynea. His first family,
the Amphinomacea, included Chloeia, Sars, Notopygos, Grube, Lirione, n., Amphinome,
Bruguiere, Hermodice, n., and Murythoe, n. The characters were — cephalic lobe rounded ;
no tentacle; two antennae; two antenniform palpi; branchiae on few segments ; dorsal and
ventral divisions of the foot distinct. The representatives of these genera are more
characteristic of the warmer seas, as shown by the earlier writers, as well as by Kinberg's
paper and the voyage of the " Challenger." Even the expedition of the " Porcupine," in
1870, brought fine specimens of Chloeia from the Mediterranean. The labours of Prof.
M. Sars, however, introduced in 1861 a northern example of the genus Eurythoe ; 4 and his
distinguished son, Prof. Gr. 0. Sars, added another representative of the family in
Par amphinome? which he had found amongst the unpublished manuscripts of his
father.

In the catalogue of the Poyal College of Surgeons 6 an account of the circulation
and the branchial plexuses of Amphinome capillata is given.

1 ' An. sans Vert./ vol. v, p. 327.

2 ' Ann. Danic. Consp./ p. 4, 1843.

3 •' Ofversigt af Kongl. Vet. Akad. Forhandl./ Aug. 14, No. 1, p. 11, 1857.

4 ' Christ. Vidensk. Selsk. Forhandl./ 1861.

5 l Some .Remarkable Forms, &c, off the Norwegian Coast/ Christ., 1872, p. 45.

6 Vol. vi, p. 14.

PARAMPHINOME. 221

Ehlers (1864) placed the Amphinomea as the first family of his order Nereides,
characterising them as having the mouth on the under surface, surrounded by several
segments. The head is not clearly denned, and bears a caruncle which extends over
several segments. Under this family he includes Ohloeia, Notopygos, Lirione, Amphinome,
Hermodice, Eurythoe, Euphrosyne, Spinther, Aristenia, Hipponoe, Lophonota, Zothea, and
Didymobr'anchus.

Grube, in his ' Annulata Semperiana' (1878), comprehended the Euphrosynidse in his
family Amphinomea. He describes them as having an oval or elongate body, more or
less depressed, frequently tetragonal : and with one or two anal appendages. The
cephalic lobe is united with the buccal segment beneath, generally with a caruncle and
a bifid pre-buccal process. Tentacles one or three, sub-tentacles (palpi) two or none.
Eyes two pairs. Mouth inferior. Buccal segments several, rarely one. Dorsal cirri
single or double ; ventral single, cirri rarely absent. Fascicles of bristles double on each
side. Spines none. Bristles simple, now capillary, now unequally bifurcate; rarely com-
pound, hooked. Branchiae dorsal or marginal ; pinnate, ramose or simple, rarely absent.
Pharynx (proboscis) somewhat suboval, having neither jaws nor papilhe. This is a more
detailed diagnosis than he gives in his ' Eamilien der Anneliden ' (1851).

Claus, in his ' Grundzuge ' (1880), made the Amphinominas the first sub-family of the
Amphinomidse.

Cams describes the Amphinomea ('Prod. Faun. Medit.,' 1884) as having the mouth
on the ventral surface, surrounded symmetrically by several segments ; cephalic lobe little
differentiated, or represented by a caruncle on the dorsal surface covering several segments.
The genus Amphinome he characterises thus : — Head with three antenna?, four eyes, and a
distinct caruncle ; tentacular cirri two ; caruncle covering two to three segments, more or
less plicate; feet biramous, divisions distinct, a hiatus between the bristles; branchise
arborescent, branches and ramuscles numerous.

Genus I. — Paramphinome, Sars, 1867.

The genus Paramphinome was characterised by Sars as having a moderately
elongate vermiform body, segments few. Cephalic lobe small, produced posteriorly ; no
caruncle; no eyes. Five short tentacles, cylindrical and subequal— a median in the
posterior part of the head, two anterior, and two lateral. Dorsal and ventral cirri present
in the first segment of the body, elongate, similar to the cephalic tentacles ; rudimentary in
the other segments. Mouth a longitudinal fissure surrounded by four fleshy lips. Anus
terminal. Feet biramous, the dorsal and the ventral divisions, which are small, widely sepa-
rated. The dorsal bristles are of two kinds — a shorter simple serrated series, and a longer
wxd much more slender kind, with a spur at the base of the terminal region. Amongst these
are also some long, slender, and tapering bristles— faintly and sparsely serrated towards
the tip. The inferior bristles have in some the distal end of the shaft dilated and bifur-
cate, one of the divisions being short and simple, the other elongate and serrate; others
are long and slender, with fine serrations towards the tip, while a few are shorter and
more boldly serrate. Two strong and somewhat S-shaped hooks are in front of the

222

PARAMPHINOME PULCHELLA.

insertion of the dorsal bristles of the first segment. Branchias occur only on the anterior
segments, and are large and dichotomously divided, extending across the dorsum.

paramphinome pulchella.
Fig. 15.

Anterior end of Paramphinome pulchella, Sars. (After G. O. Sars.)
Enlarged.

Specific Characters. — Body nearly cylindrical, a little broader than high, slightly
tapered anteriorly and posteriorly. Setigerous segments twenty-four to thirty-three.
Cephalic lobe narrower than the first segment, produced into a conical process
posteriorly. Branchise commencing on the fourth setigerous segment, and from four to
six in number, dichotomously divided ; terminal processes cylindrical. Colour pale.
Length 9 mm.

Synonyms.

1868. Hipponoe Jeffrey sii, Mcintosh. Ann. Nat. Hist., 4 ser., vol. ii, p. 250.

1869. „ „ Idem. Trans. R. S. E., vol. xxv, p. 406, pi. xv, f. 1.

„ Paramphinome pulchella, M. Sars. Yidensk. Selsk. Christian. (Aar, 1868), 1869, p. 2o4.

1872.
1873.

1875.
1876.

1879.

Gr. 0. Sars. Remark. Forms An. Life, vol. i, p. 45, pi. iv, f. 19 — 35.

Sars. Bid. Christ. Fanna, p. 15.

Ehlers. Zeitsch. f. w. Zool., vol. xxv, pp. 17 and 31.

Hansen. Annel. Norsk. Nordhavs-Exped., 1876, Nyt. Mag. £. Nat.,

vol. xxiv, pi. i, p. 4.
Tauber. Ann. Danic, 78.

Habitat.— Dredged in St. Magnus Bay, Shetland, by Dr. Gwyn Jeffreys in 1867,
at a depth of 100 fathoms. G. O. Sars found it occasionally at Lofoten in water
ranging to 300 fathoms ; in the Christiania Fjord in 100 to 230 fathoms ; and also more
frequently at Aalsund on a muddy bottom in 20 to 30 fathoms near the shore. It has
even come from a depth of 600 fathoms. The species extends to Norway and the Arctic
seas (Hansen).

PABAMPHINOME PULOHBLLA. 223

In 1872 G. 0. Sars gave an excellent description with figures of this form under the
name of Paramphinome pulchella, M. Sars, mentioning that it is widely distributed
round the Norwegian shores. As he had an opportunity of observing living specimens
his description has been, as far as possible, added to the original account.

The head is rounded oval, convex dorsally, somewhat conical in front and even
more so behind, since it projects over the first segment almost to its posterior border.
It is furnished with five tentacles — two shorter in front and two longer on the projecting
median region. The unpaired is behind the latter on the tongue-shaped posterior part
of the head. No eyes are present.

The body is nearly cylindrical, a little wider than high, and tapering at both ends.
The bristle-bearing segments range from twenty-three to thirty-three, are about twice as
wide as long, and distinctly divided from each other. Each has a dorsal and a ventral
division of the foot, and a dorsal and a ventral cirrus, the latter organs being short —
except in the first and three last segments. The mouth has four fleshy lobes or lips,
and the proboscis is often everted in spirit-preparations. The anus is terminal and
crescentic.

The branchiae are confined to the anterior part of the body, and appear to be
normally five pairs, six pairs being less common, while four occur in younger specimens.
They increase in size from before backwards, the first being on the fourth bristled
segment. They are dichotomously divided, broad, fan-shaped organs, which even over-
lap on the dorsum. They have two primary divisions, each of which is more or less
subdivided. Two lateral vessels are visible winding along the body and connected by
branches with the gills, besides a median ventral vessel which anastomoses with the
former.

The dorsal and ventral divisions of the foot are widely separated. The bristles
of the superior lobe (Plate XXXV, figs. 19 b, c) are for the most part shorter and
stouter than the inferior, the former (b) being one of the largest, viewed somewhat
obliquely so as to show the faint serrations. When the bristle is turned round (as in c) 9
the latter are scarcely observable. Besides, there are some long, slender, and tapering
bristles faintly and sparsely serrated towards the tip. The inferior lobe of the foot has
likewise two kinds of bristles, a few (Plate XXXV, fig. 19 a) with the tip of the shaft
dilated and bifurcate, one of the distal portions being short and simple, the other
elongated and serrate. The rest of bristles in this lobe are either long slender forms
with fine serrations towards the tip, or short and more boldly serrate.

In the first body-segment, in front of the insertion of the dorsal bristles, are on eac
side two strong and somewhat S-shaped hooks. A

The want of eyes, and of a distinctly marked caruncle, as Sars says, distinguishes it
from other known representatives of the family. The form of the bristles resembles
that of other members of the group, but the remarkable hooks on the dorsum of the
first segment are unique.

Colour. — Pale and rather transparent, so that the dark intestine is readily visible.

The animal is somewhat sluggish, rolling itself up in a circle when irritated, so as to
make the long bristles project outward like those of a Nerine-l&rva,. When unmolested
it uncoils and moves hither and thither with a sluggish snake-like motion.

30

224 EURYTHOE BOREALIS

Genus II. — Eukythoe, Kinberg, 1857.

Body elongated, with flattened rectangular segments. Cephalic lobe large, rounded
or pentagonal ; eyes four, dorsal; a median, and two lateral tentacles ; caruncle trilobed.
Palpi forming two adnate lobes interiorly in front of the mouth, each with a tentacular
process or sty lode (Eacovitza).

Dorsal cirrus single ; bristles of the dorsal lobe linear, subarticulate, others subbifid,
with a serrate limb and a short process. Ventral bristles bifid. Branchige from the third
segment backward.

Eurythoe borealis, Sars, 1861. Plate XXVII, fig. 16.

Specific Characters. — Head somewhat horse-shoe shaped, with two short awl-shaped
tentacles (lateral) in front, and two longer a little behind. A curved line separates the
head into an anterior and a posterior region, the latter more elevated and furnished with
four reddish eyes, the anterior being about twice the size of the posterior. A little
behind the anterior pair is a filiform tentacle, while behind the eyes the caruncle
extends to the anterior border of the third bristled segment. Body elongate, straw -
yellow or pale vermilion, twenty-three to sixty- seven segments, and in transverse
section nearly square, though the ventral surface is most flattened. Branchiae commence
on the second bristled segment as a tuft of four papilliform processes, and generally arise
, behind and somewhat below the dorsal fascicle of bristles, and consist of a tuft of three

or four pale finger-like processes. They are continued almost to the tip of the tail
(antepenultimate segment), which terminates in a rounded papilla with a minutely
crenate margin. The dorsal division of the foot has a jointed cirrus, and bristles of two
kinds — bifid with a serrated limb of greater or less length, and somewhat stronger simple
bristles with serrate tips. The ventral division has also a jointed cirrus, and many bifid
bristles somewhat stouter and with shorter tips (about four serratures). Length one to
one and a half inches.

>-f *- ♦;

\k Synonyms.

1861. Eurythoe borealis, Sars. Vid. Selsk. Forh., 1861, p. 56 (p. 9 sep. copy).
1869. Amphinome vagans? Mcintosh. Trans. R. S. E., vol. xxv, p. 406, pi. xv, £. 1.
1876. Eurythoe borealis, Mcintosh. Trans. Z. S., vol. ix, p. 373.
18&6. ,, „ Eacovitza. Arch. Zool. Exp., 3e ser., iv, p. 179, pi. i, f. 1 — 6.

Mabitat.—JJnder a stone in a tide-pool in Herm, August, 1868. Sars found a
smaller example at Manger, in the neighbourhood of Bergen, in shell-sand. 1

1 It is doubtful if Dr. Benham's view that specimens six inches long occur all round the British
area refers ta this species, which appears to be rare in our seas.

EURYTHOE BOBEALIS. 225

The head is pale, somewhat horse-shoe shaped, with two short, conical, awl-shaped
tentacles in front, and two longer ones — stylodes (Racovitza) — a little behind, opposite the
wide part of the snout. A curved line separates the anterior from the posterior part of
the head (Plate XXVII, fig. 16), the former being flattened, the latter more elevated, and
furnished with four reddish eyes, the anterior being about twice the size of the posterior
pair. A little behind the anterior pair a filiform tentacle projects upward in the middle
line, and, immediately behind, the caruncle extends as a wrinkled ridge to the anterior
border of the third bristled segment, and is of the yellowish colour of the rest of the
dorsum.

The body is elongated, nearly cylindrical throughout the greater part of its length,
only a little tapered at the extremities. The number of segments is sixty-seven in the
larger, forty-one in the smaller specimens. In contraction the body is somewhat square
in transverse section, the inferior surface especially beiDg flattened. The great shorten-
ing which takes place causes the prominent rows of bristles to project very boldly. The
sulci between the first three bristled segments are somewhat less marked, and the slope
of the bristles more oblique, but the rest are very distinctly separated, indeed the body is
occasionally more or less moniliform. The first bristled segment is small and has no
branchiae. The second has a tuft of about four papilliform branchial processes. Each
segment is furnished with a dorsal and ventral division of the foot — bearing bristles,
besides the branchial tuft which springs from a point behind and rather below the dorsal
fascicle of bristles, and consists of about four finger-like processes arising from a common
origin. They continue almost to the tip of the tail, being apparent on the third last
segment. All are pale in colour.

The dorsal division of the foot (Plate XXXVI, fig. 16) has a jointed cirrus, and a
group of bristles consisting — of bifid forms with elongate finely serrate tips (Plate
XXXV, figs. 20 and 21), of those with shorter tips with four or five serratures, and of
somewhat stronger tapering simple bristles with serrate tips (Plate XXXV, fig. 22).

The inferior division has also a jointed cirrus, and many bifid bristles somewhat
stouter than in the superior division, and with shorter tips having about four serratures
(Plate XXXV, fig. 23).

The tail terminates in a rounded papilla with a minutely crenate margin.
The proboscis commences at the posterior third of the fourth bristled segment (as
indicated by two brownish specks), and occupies the fifth and sixth segments. It is
somewhat fusiform, truncated anteriorly and posteriorly.

The specimens were of a general straw-yellow hue, somewhat paler anteriorly, and
pinkish here and there from the dorsal blood-vessel.

The animal is sluggish, crawling slowly about, and contracting: into a bristled
mass when irritated. Occasionally the posterior extremity is twisted like a screw, and
it also coils its body into one or two convolutions, the condition it assumes in spirit;
hence the examination is rendered difficult, and most of the fragile bristles are broken
during the efforts to decipher their structure.

This form seems to be the same as that found by Prof. M. Sars in 1861, though the
brevity of his description leaves some doubt. His sole specimen was a small one of 10
mm. in length, and occurred at a depth of sixty to seventy fathoms. As in other types

226 EUPHROSYNINA.

of Annelids, however, forms which in the north frequent the offshore are found between
tide-marks in the south. The number of the branchige differs slightly in. the two forms,
Sars giving five as the usual number. Further inquiry is thus necessary before the
absolute identity can be satisfactorily ascertained.

Sub-family — E uphrosynina.

Cephalic lobe compressed, bending downwards anteriorly to the ventral surface.
Dorsally a median tentacle, two eyes, and a trilobed caruncle, ventrally two eyes, two
short lateral tentacles, and a pair of adnate palpi in front of the mouth.

Body oblong or ovate-oblong, segments few, feet crest-like, the dorsal and ventral
divisions being indistinct. Bristles of the dorsal region brittle and hollow, with simple
or bifid tips or hooks with jointed stems, accompanied by simple bristles. Branchias on
almost all the segments. Buccal apparatus large and complex; alimentary canal simple,
with only a trace of an anterior cascum. Anus dorsal. Two posterior appendages. The
nerve-cords are separate and comparatively large, and lie quite within the body- wall, —
the oblique muscles, which generally bound the longitudinal ventral muscles, in this
case decussating beneath them.

When Savigny established the genus Euphrosyne in 1820, he placed it, as his
nineteenth genus, under his fourth family, the Amphinomse of Bruguiere ; and several
subsequent authors, such as Milne Edwards, Kinberg, Ehlers, and Be Quatrefages, have
adopted the same arrangement.

Kinberg (1857) made the Euphrosynea the second family of his second group, Amphi-
nomea. A single genus and species only are mentioned. His description is — cephalic
lobe compressed ; neither antennas nor palpi; branchiae on many segments ; feet crest-
like and transverse. He subsequently (1867) made three groups of the Amphinomea, viz.
the Chloeia group, the Notopygos group, and the Amphinome group, and gave a brief
resume of the literature of each. He does not, in this paper, mention Euphrosyne or
Spinther.

Grube included the genus Euphrosyne under the family Amphinomidse both in his
earlier and later publications (1851 — 1878), having followed Savigny and the preceding
authors in this respect. It is placed under his primary division (tribe) Eapacia.

Ehlers 1 likewise adopted Savigny's classification, placing all those with the dorsal
caruncle under this group (Amphinomea). He 3 gives an account of a fossil form
(Meringosoma curtum) from the lithographic slate of Solenhofen which approaches
Euphrosyne in character. In his recent publication ('Florida Anneliden/ 1887) be
adheres to this arrangement. Amongst other features he noticed the hollow nature of
the bristles.

The Euphrosynidas in the classification of De Quatrefages 3 were grouped under the

1 'Borstenwurmer/ 1864.

2 ' Cassel/ 1869, p. 161, pi. xxxvi, f. 3.

3 'Armeies/ 1865.

SPINTHER. 227

Amphinomiens, and separated from the Aphroditidae by the Palmyridae, Eunicidae and
Lumbrinereidae. The Amphinomidae he characterised by the similar or subsimilar seg-
ments, by the absence of buccal armature or its simple nature, and by the presence of
arborescent branchiae on the segments. The separation of the Euphrosynidae and their
allies from the proximity of the Aphroditidae does not seem to be warranted on anatomi-
cal grounds.

Claus in his ' Grundzuge ' (1880) placed the Euphrosynina3 as a sub-family of the
Amphinornidae.

Carus l makes Euphrosine one of the genera of his Amphinomea, and describes it as
having the head with one or several antennas, two eyes, and a caruncle ; body ovate ;
feet biramous, rami confluent ; branchiae springing from numerous trunks more or less
branched.

In Benham's 2 (1896) classification they form part of his eighth family Amphinomidae,
placed between the Nephthydidae and Eunicidae, a position which, as already indicated,
cannot be supported on anatomical or other grounds of value.

Genus — Spinthek, Johnston, 1845.

Oniscosoma, M. Sars, 1850.
Cryptonota, Stimpson, 1854.

Body ovoid ; dorsum more or less convex ; segments few. Cephalic region incised
as in the other segments, and bearing dorsal bristles, lamellae and uncinate setae. Eyes
four, at the base of the dorsal tentacle. Buccal aperture inferior — set in the midst of the
neighbouring segments. Proboscis exsertile, short and semi-tubular. Intestine pinnate.
Anus posterior. Nerve-cords widely separated. Segmental organs absent. Neither
cirri nor branchiae.

Since few opportunities have occurred in this country of seeing living examples of
the genus, it will most conduce to brevity and clearness if the recent work of von Graff
be summarised.

Polychaeta with elliptical and dorsally convex bodies gently rounded anteriorly and
posteriorly, and having distinctly marked segments. The flanks have a series of short
parapodia with a dorsal lamella in each segment. In the centre anteriorly is a single
tentacle, while posteriorly two wart-like anal cirri occur. At the base of the dorsal
tentacle are four small eyes. On the ventral surface a little behind the snout is the
mouth. Posteriorly is the anus, with the reproductive aperture in front. The free dorsal
lamellae have a double row of chitinous bristles with simple or bifid tips. The parapodia
have a prominent hooked and jointed bristle, and from one to four of the same kind
undergoing development, besides from eight to thirteen simple bristles. The brain lies
under the dorsal tentacle. The sincipital region of Racovitza is the only part of the head
remaining, and consequently the middle region of the brain is alone present — both the

1 Op. cit., f Prod. F. Medit.'

2 < Camb. Nat. Hist./ p. 318.

228 SPINTHER.

anterior and the posterior being absent — the result, according to Racovitza, of parasitism.
The two ventral nerve-cords are widely separated, with a ganglion in each segment, and
a transverse commissure. The alimentary canal has a muscular pharynx without dental
armature, with paired diverticula of the middle and hind gut, and a dorsal blind gut. A
vascular system is present, but both branchiae and nephridia are absent. The sexes are
separate.

The genus Spinther was established by Dr. George Johnston in 1845 for an Annelid
(S. oniscoides) half an inch in length, and fully a quarter of an inch in breadth, which he had
received from W. Thompson of Belfast, who had dredged it in six to ten fathoms in the
neighbouring bay. It is remarkable that no undoubted example of this species has been
found in British waters since that date, though another species has once been procured in
the Minch. Dr. Johnston mentions its cream-yellow colour, the absence of distinct " head,
tentacula, and tentacular cirri," and gives the number of the dorsal lamellae at thirty.
He correctly noticed the general form of the " feet," the presence of the cirrus and other
features. He also distinguished the hooks and the various kinds of bristles, though his
figures were not drawn with that scientific accuracy — probably from deficient microscopic
power — which modern requirements demand. He grouped the genus under the Amphi-
nomidae. Unfortunately the type-specimen 1 is not in the British Museum, where Dr.
Johnston's collection of Annelids is. The ciliated pits on the anterior portion of the
cephalic region are probably sensory (Racovitza).

Michael Sars five years later (1850) described his Oniscosoma arcticum, n. g. et sp.,
which he had dredged in Komagfjord, in thirty to forty fathoms, on a sponge. This form
had twenty segments, a tentacle and four eyes in the third segment. He gave an
account of the dorsal lamellae, the marginal expansions with the bifurcate bristles,
and the ventral division with the hooks. He linked it with Euphrosyne.

Edonard Grube next year (1851), in his ' Familien der Anneliden,' considered that
Spinther leaned to the Siphonostomae or to the Amphinomea rather than to the Aphro-
ditidae.

A few years later (1854) Dr. Stimpson formed the genus Gryptonota for a similar
Annelid, giving most of the characters already known, and stating further that the
branchiae resembled those of Euphrosyne, though he could not satisfactorily make them
out.

Grube (1860), in describing Spinther miniaceus, a new species, placed it near
Amphinome.

In A. de Quatrefages' ' Histoire cles Anneles ' Johnston's species is given at the end
of the Ohloremiens, under the genera and species of uncertain position, it being noted
that while Johnston considered it near the Aphroditidae, Grube thought it approached the
Amphinomaceae and Siphonostomae, and that he (the author) was entirely of the latter
opinion, which he based on the structure of the feet, the presence of " albuminous "
matter in them, and the nature of the hooks.

Claparede in a note 3 states that the genera Spinther {Oniscosoma) and Gryptonota are
identical.

1 I am indebted to Prof. Jeffrey Bell for making a search.

2 { Arch. sc. Phys. et Nat./ t. xxii; 'Bibl. Univ. et Rev. Scientif./ Apr., 1865.

SPINTHEB. 229

Malmgren in his ' Aimulata Polychseta ' (1867) included Sars's species under the
genus Spinther — as 8. arcticus, Sars.

In 1865 I found in the Minch, off North Uist, a form which was placed under 8.
oniscoides, Johnst., but which is clearly S. miniaceus, Grube, or more correctly 8. arcticus,
Sars (non Wiren).

Hansen next (1882) describes a form which he identified with 8. arcticus, Sars, from
the Norske JSTordhavs-Expedition. The description and figures, however, as von Graff
says, agree better with S. oniscoides, though the somewhat indifferent drawings of the
bristles leave some doubt.

In the account of the Annelids of the Yega Expedition (1883) Wiren described a
species which he calls 8. arcticus, but which materially differed from Sars's form in size
and in structure.

The same year Levinsen, in his systematic account of the geographical distribution
of the northern Annelids, confused Sars's species with Wiren's, and made Hansen's
(8. oniscoides) a new species.

Drasche 1 further in 1885 gave an excellent anatomical description of S. miniaceus,
Grube. He found this species chiefly on the surface of the sponge Tedania anhelans,
Lieberk., at Trieste. He pointed out the double nature of the dorsal membranes — of
the structure of which he gives good figures. Underneath the cuticle is the hypoderm,
which is thickened in certain regions. The muscular investment which occurs next is
more or less continuous as an outer longitudinal and an inner circular layer, but in his
figure (pi. ii, fig. 7) of the ventral region the reverse arrangement is shown. His
description and figures of the cephalic ganglia (brain) are good, as are also his remarks on
the eyes. He gives a clear account of the alimentary canal and of the dorsal blind gut,
which joins the intestine in front of the rectum. His account of the circulatory system is
brief, since he had only examined the system in section — a dorsal vessel, a ventral trunk,
and transverse vessels in each segment being the chief features alluded to. He concludes
his paper with remarks on the body-cavity and its septa. Between each pair of septa are
the ganglia of the ventral cords and their commissure, the diverticula of the gut and the
transverse blood-vessels. The rest of the cavity, in his specimens, was filled with the
reproductive products. In the male the region of the dorsal blood-vessel is specially
connected with the development of the sperms, as Haswell and Selenka showed in the
Aphroditacea. He found no segmental organ (nephridium).

The most complete account of the genus, however, is that of L. von Graff (1887).
This author had an extensive series of specimens, and brought to bear on their structure
modern methods of investigation. After an historical introduction he gives the characters
of the genus and the three species known, viz. 8. oniscoides, Johnston, 8. miniaceus, Grube,
and 8. arcticus, Wiren. The adoption of the specific term arcticus for the latter, however,
seems to be open to some objections, since Sars's name arcticus has some claim to priority
over Grube's term miniaceus, and confusion may result from the application of the same
name to a different species, northern though it be. He shows that 8. miniaceus is the
smallest of the series, 8. oniscoides considerably larger (26 mm.), while 8. arcticus

1 Drasche, R. von, ' Beitrage z. feniereu Anat. der Polychasta/ Wien, 1885, 14 pp., 2 pis.

230 SP1NTHER ONISCOIDES.

reaches the length of 50 ram. A general structural description follows, and he points out
the essential differences of the dorsal lamella) in the three species. Bifid dorsal bristles
only are present in S. miniaceus, while in the other two simple bristles accompany the
former. The mouth opens on the ventral surface anteriorly, and four small brown eyes
lie at the base of the tentacle, each with a short thick nerve from the cephalic ganglion.
The nerve-cords are separate except for the transverse commissures. The alimentary
canal consists of fore, middle, and hind gut, besides caecal appendages. The pharynx is
muscular. The mid-gut possesses generally a pair of diverticula in each segment, and
terminates in an anus which has a pair of short cirri. Excretory organs are absent, but
a blood-system is present. There are no differentiated branchia3. The reproductive
elements are separated from the body-wall, and lie in the body-cavity. All are parasitic
on sponges.

Recently Racovitza 1 (1896) has carefully investigated the structure and homologies
of the cephalic region, and he is of opinion that only the sincipital region of the head
remains with the tentacle and the four eyes, and the much reduced brain internally ; the
latter (brain) consisting of the middle region only, whereas in Eurythoe' and Euphrosyne
both the anterior and posterior regions of the brain are present. This reduction of the
cephalic region he attributes to parasitism on Esperella and other sponges.

The species show a certain amount of variation, and von Graff thinks that the some-
what radiate arrangement of the parts anteriorly and posteriorly approaches the condition
in Myzostoma, which occurs on starfishes, while Spinther frequents sponges.

In the pinnate arrangement of the alimentary caeca Sjpinther differs from the typical
Euphrosynina, and approaches the Aphroditidse.

1. Spinther oniscoides, Johnston, 1845.

Specific Characters. — A dorsal cirrus at the end of the parapodium. The dorsal
lamella project at the free margins and have a series of strong, simple or bifid bristles.
The hooks of the parapodia are powerful, pectinate at the projecting margin, and the tip
is strongly curved. Ventral surface furnished with small papillae (warts) on the ridges
which flank a median furrow. Posterior end split to the anus.

Synonyms.

1845. Spinther oniscoides, Johnston. Ann. Nat. Hist., vol. xvi, p. 8, pi. ii, f. 7 — 14.

1854 u Gryptonota citrina, Stimpson. Marine Invest. Grand Marian, p. 35, pi. ii, f. 27.

1865. Spinther oniscoides, Johnston. Cat. Brit. Mus. ; p. 127, pi. xiv, f. 7 — 14.

1882. „ arcticus, Hansen. Annel. Norsk. Nordhavs-Exped., p. 44, pi. i, f. 1 — 5.

1883. „ major, Levinsen. Syst. Oversigt. nordiske Annulata, &c, Yidensk. Meddelels, p. 125.
1887. „ oniscoides, Graff. Zeitsch. f. w. Zool., vol. xlvi, p. 90, pi. vi — vih, and woodcuts.

Habitat.— Dredged off Castle Chichester, Belfast Bay, by Mr. Hyndman in 1844,
and off Anglesey in seventeen fathoms (Hornell) ? (fide Dr. A. Merle Norman). Abroad
1 < Arch. Zool. Exper./ 3 ser., vol. iv, p. 197, &c, 1896.

SPINTHER ONISCOIDES. 231

it has been met with in the Bay of Fundy, New Brunswick, by Stimpson and Leidy, and
at Station 275 (Ost Havet) den Norske Nordhavs-Expedition 1876-8 (Hansen). Verrill
includes it in his list of marine forms from Cape Cod to the St. Lawrence.

Body. — Length 11 to 26 mm., the British forms seen by Johnston approaching the
first-mentioned figure, ovate or somewhat oblong, convex dorsally, smoothly rounded at
each extremity. The more convex dorsum has a series of symmetrical double lamellae,
strengthened by simple or bifid bristles, the tips of which slightly project beyond the
skin, these lamellae being continuous with the feet, which in Von Graff's example amounted
to forty-eight, Johnston's specimens, apparently having fewer. The ventral surface ha s
a series of low ridges studded with minute warts, the anterior sloping forward and the
posterior backward, the middle line being marked by a furrow, likewise warty. In the
largest example (26 mm.) the thickness in the centre of the body was 2*5 mm., thinning
off at the sides, and the breadth about 11 mm. ; the height of the dorsal lamellae in the
same being 1*5 mm. The anus terminates some distance within the posterior border, the
adjoining pair of ventral ridges presenting no special differentiation.

Colour. — Johnston's examples were of a uniform cream-yellow colour, while Von
Graff's had a yellowish-ochre hue. Stimpson observes that his specimen was of a
beautiful lemon-yellow colour, resembling very much that of some sponges occurring
with it on a gravelly and muddy bottom in thirty-five fathoms in Hake Bay, Grand
Man an.

Read. — This region is merged into the general contour of the body, but bears
superiorly over the brain the ridge-like dorsal tentacle with four eyes at the base. The
mouth opens on the ventral surface some distance behind the anterior border, often as a
radiate button — from the protrusion of the proboscis.

Feet (parapodia). — These are distinguished by the possession of a ringed base, and
the presence of a short dorsal cirrus of about four or five segments, the lamellar ridge of
the dorsum curving smoothly away from it on the one hand, while on the other the
golden hooks emerge near its base ventrally. The hooks, which are excellently figured
by Von Graff, 1 have strongly curved and sharp points, while the terminal process of the
shaft is boldly serrated. One or two of these project freely, while internally one or two
are in course of development, the point appearing first, according to Von Graff, in a
cellular matrix.

It is remarkable that, notwithstanding the great increase of zoological explorations
of the British seas, no example of this species has been procured since Dr. Johnston
described it, with fair accuracy — both with regard to external form and the characters of
the bristles and hooks — from two examples forwarded by Mr. Wm. Thompson of Belfast.
Moreover, Prof. Jeffrey Bell informs me the type specimens cannot be found in the
British Museum. Under these circumstances the only alternative was to utilise what
had been so ably written by Von Graff and others, and place materials for ready
identification in the hands of future observers.

The species follows the habit of certain marine forms, adhering to sponges and
feeding on them, so that the alimentary canal, including the dorsal blind sac, is packed
with sponge-debris and spicules.

1 Op. cit., pi. viii, figs. 4 and 5.

31

232 SPINTHER MINIACEUS.

2. Spintheii miniaceus, Grube, 1860. Plate XXIV, figs. 1, 2.

Specific Characters. — Parapodia without cirri ; the dorsal lamellae are prominent
processes with leaf-like external borders. The supporting bristles are bifid and more
slender than in the previous species, and arranged in a radiate manner in the leaf-like
lateral lobes. The distal end of the shaft of the hook is smooth. The ventral surface
appears to be smooth, but it has rows of minute warts.

Synonyms.

1850. Oniscosoma arcticum, Sars. JSIyt. Mag. f. Naturvid., vol. vi, p. 240 (210 Sars' paper).

1860. Spinther miniaceus, Grube. Arcniv f. Naturges, Jahrg. 26, Bd. i, p. 74, Taf. 3, figs. 3 — 3 b.

1861. „ „ Grube. Ein Ausflug nacli Triest, pp. 31 and 140, Taf. 3, figs. 3—3 b.
„ „ arcticus, Sars. Sasrsk. aft. Vidensk. Forfchandl., p. 3.

1862. „ „ Sars. Christiania Yidensk. Selsk. Forhandl., p. 52.
1867. „ „ Malmgren. Annul. Polychaet., p. 1.

1876. „ oniscoides, Mcintosh. Traus. Z. S., vol. ix, p. 373, pi. lxvii, figs. 1 — 3.

1879. „ arcticus, Tanber. Annul. Danic, 77.

1883. ,, ,, Levinsen. Yidensk. Meddelelser; Kjobenhavn, p. 125.

1884. ,, miniaceus, J. Y. Carus. Prodr. Faunas Medit., p. 208.

1885. ,j „ Drasche. Beitrage zur Anat. Polycliast., Heft. 1, Taf. ii, Wien.
1887. „ „ Graff. Zeitsch. f. w. Zool., vol. xlvi, p. 93, Taf. i — v, and woodcuts.

1890. „ „ Mcintosh. Ann. Nat. Hist., Aug., 1890, p. 182.

1891. ; , oniscoides, Hornell. Trans. Liverp. B. S., p. 239.

1896. ,, miniaceus, Racovitza. Arch. Zool. Exp., p. 197, pi. hi, figs. 22 — 26.

Habitat. — The Minch, near North Uist, on a yellowish sponge (Mcintosh) ; on
Antennularia ramosa and yellow Halichondria in seventeen fathoms off Holy Island,
Anglesey (Hornell). Finmark and west coast of Norway ; Bay of Muggia at Trieste on
reddish sponges (Tedania) ; north coast of Denmark (Lutken). Stimpson (1853) found
the American example of the genus also on a yellowish sponge.

Colour. — The Hebridean specimen was of a straw-yellow hue, the lamellar processes
at the side being pale and marked with opaque-white spots. The Mediterranean examples
are of a cinnabar tint, or dull reddish, brownish, or violet-brown, according to the colour
of the sponges which they frequent.

Head. — The caruncle presents no feature of interest. The exserted pharynx is
smooth and trumpet-like, thus diverging from the condition in S. arcticus.

Body. — Ranging from "9 to 8*5 mm., the Hebridean example being about 5 mm., with
a thickness in the centre of 1 mm. in the largest, ovate or somewhat oblong, with the
sides flanked by the translucent lamellse, which dorsally form a series of symmetrical
processes provided only with bifid bristles, and in the largest examples 0*4 mm, in
height. The lamellae on the two sides of the body are separated by a median furrow.
Ventrally the surface is more or less smooth, only minute warts being present, and there
is no well-defined median furrow. This surface is covered with areola (polygonal spaces)

ETTPHROSYNE. 233

in the Mediterranean examples. The segments range from twelve to twenty-four. The
body terminates posteriorly in two somewhat globular anal cirri.

The bristles of the dorsal lamella are all bifid (Plate XXXV, figs. 5 a and 5 b), the some-
what delicate process resembling a web stiffened by the spiculge. They are arranged with
considerable regularity, and are all characteristically curved. A sensitive papilla also
occurs at the tip of each lamella laterally.

Feet. — These are more or less cylindrical. One conspicuous hook (Plate XXXV,
fig. 5 c) in the British example projected beyond each foot, generally another of similar
form (but shorter) within the tip, and the developing extremities of two in the tissues.
A group of simple bristles (eight to twelve) with tapering tips also accompany the
central hook.

Von Graff found considerable variation in the condition of the alimentary canal, so
that he made two series, viz. the Stenocoela and the Euryccela. He mentions also, as
Grube had previously done, that the ova wander into the bases of the feet, and probably
are the " opaque white spots " seen in the Hebridean example. The ripe ova have short
protoplasmic processes in the body, which elongate subsequently into structures like
pseudopodia, somewhat symmetrically arranged as in a Radiolarian. The spermatozoa
present a head with an acute tip, two granules at the base, and a long filament.

Grube's original description was carefully drawn up, and he gave the chief features
in regard to form and coloration, though his examples were small, having only twenty-
two parapodia. He, however, placed too much reliance on the reddish-orange colour as
distinctive from Johnston's species. Von Graff w r as at first inclined to place the
Hebridean example under 8. arcticus, Wiren, but further investigation showed that it
was S. miniaceus. 1

It is probable that the Sjpinther arcticus of Sars (1861) is this species, and there is
nothing in his description to invalidate this view. As already mentioned, the adoption of
his name would lead to considerable confusion.

Genus IV. — Etjphrosyne, Savigny, 1820.

Body generally short, oblong, equally narrowed at either end, segments not
numerous ; two thick styles posteriorly. Cephalic lobe narrow, frontal part narrowest,
a band passing downward to the inferior ridge. Eyes both on the dorsal and ventral
surfaces. One median tentacle situated in front of eyes. Lateral tentacles two, very
short, springing in front of the inferior eyes. Fascicles of bristles arranged on each side
of the segments. Superior bristles forming a transverse row, no barbs ; inferior grouped
in a broad pencil. Capillary bristles unequally bifurcate. Dorsal cirri two (or three —
Ehlers) on each side, the one at the inner border of the fascicle, the other at the outer
margin. Ventral cirri single. Branchiae dorsal, in row r s, and more or less ramose.
Palpi forming fixed lobes on each side of the mouth. Mouth opening on the ventral sur-
face and extending over several segments. Buccal apparatus complex, alimentary canal
simple.

1 Vide ' Ann. Nat. Hist./ August, 1890, p. 182.

234 EUPHROSYNE FOLIOSA.

The genus Euphrosyne of the family AmphinomaB was established by Savigny in his
' Systeme des Annelides ' (1820) for two species (E. laureata and E. myrtosa) from the
Red Sea. Briefly his description is as follows : — Mouth with a simple unarmed proboscis.
Eyes distinct, two in number, separated in front by the caruncle. Antennae incomplete,
the middle and exterior absent, the unpaired subulate. Feet slightly separated, but each
division provided with bristles having a minute denticle near the tip. Cirri nearly equal ;
an additional one resembling the others inserted at the upper extremity of each dorsal
division. The last pair of feet forming two small globular cirri. Branchiae situated
immediately behind the feet, extend from the dorsal to the ventral division, and
consist of seven separate arborescent tufts arranged transversely. Head very narrow
and much produced (tres rejete) posteriorly, split into two lobes in conformity with the
feet ventrally, and provided dorsally with a depressed caruncle which is prolonged to the
fourth or fifth segment. The body is oblong or ovate-oblong, with comparatively few
segments.

Lamarck adopted the foregoing views in the classification of the group.

Audouin and Milne Edwards followed Savigny in regard to the systematic position
and description of the genus, and gave an account of a new species (E. foliosa) from the
shores of France.

Grube, Ehlers, and subsequent authors made little change in the characters attributed
to the Euphrosynidae.

The Euphrosynidae frequent both the littoral region and comparatively deep water,
a distribution that characterises both foreign and British species. Thus Euphrosyne
foliosa is diagnostic of the tidal rocks of the Channel Islands, while the other species
have been procured only by the dredge. They extend to the American shores.

1. Euphrosyne foliosa, Audouin and Edwards, 1834. Plate XXIV, fig. 3.

Specific Characters. — Caruncle extending to the anterior part of the fifth segment,
with a filiform tentacle in front between the dorsal eyes. Branchiae much branched,
with ovate expansions at the tip. The middle cirrus between the fourth and fifth
branchial stems. Bristles longer than the branchiae, which are from eight to nine in
number. The serrated bristles have the longer fork peculiarly curved towards the tip.
Segments thirty-one to thirty-five.

Synonyms.

1833. Euphrosina foliosa, Audouin and M. Edwards. Ann. d. Sc. Nat., vol. xxviii, p. 201, pi. ix,

figs. 1 — 15.

18 34. u „ Ibid. Litt. de la France, vol. ii, p. 126, pi. ii, b, figs. 1—4.
1836-7. „ „ Cuv. Reg. An., Crochard's Edit., tab. viii, fig. 2.

1840. Euphrosyne foliosa, Grube. Actin., Echinod., &c., p. 84.

I 841 - » ,, Belle Chiaje. Descrizione, Tav. 62, Tab. 139, figs. 6—8.

„ Lophonota Audouinii, Costa. Ann. d. Sc. Nat. (2), vol. xvi, p. 270, pi. xiii, fig. 1.
1849. Euphrosyne foliosa, W. Thompson. Ann Nat. Hist., ser. 2, vol. .hi, p. 355.

EUPHROSYNE FOLIOSA. 235

J 851. Euphrosyne foliosa, Grube. Fam. der Annel., pp. 41 and 122.

1853. „ „ P. H. G-osse. Ann. Nat. Hist., 2 ser, vol. xii, p. 384.

1863. „ mediterranea, Grube. Archiv f. Naturges., 1863, i, p. 38, pi. iv, fig. 2.

1864. „ foliosa, Ehlers. Die Borstenw., p. 65.

„ „ racemosa, Ehlers. Die Borstenw., p. 67, Taf. i, figs. 1 — 11.

1865. „ foliosa, Johnston. Cat., p. 126.

33 33 39 De Quatrefages. Hist. Nat. Anneles, p. 408.

1868. „ „ Baird. Proceed. Linn. Soc, vol. x, p. 237 (1870).

1870. „ „ Grube. Archiv f. Naturges., 1870, p. 304.

1875. ,, Audouini, Marion and Bobretzky. Ann. Sc. Nat., 1875, p. 10.

1876. „ foliosa, Mcintosh. Trans. Z. S., vol. ix, p. 273.

1884. „ Audouinii (Claparede), V. Cams. Fauna? Medit., p. 207.

1888. „ foliosa, St. Joseph. Ann. d. Sc. Nat. (7), vol. v, p. 190.

1896. „ Audouini, Kacovitza. Arch. Zool. Exp., p. 186, pis. i et xi, figs. 7 — 13.

Habitat. — Belfast Bay, Hyndman; Weymouth, Grosse. Plentiful under stones near
the border of low water mark at Herm, Lankester, Mcintosh, Hornell. Off the
Hebrides, Gwyn Jeffreys. Birterbury Bay, Connemara, fifteen fathoms, Gwyn Jeffreys,
Haddon. Arran Isles and Bay of Galway, E. P. Wright. Off Marsden, co. Durham,
twenty to thirty fathoms, A. M. Norman.

It was discovered in European waters by Audouin and Milne Edwards, viz., on a
bank of oysters and Anoinira between Granville and Chausey in fifteen fathoms, and on
a small bank of the same kind at St. Malo. Ranges to the Mediterranean, where it
occurs at considerable depths.

Colour. — Some of those from Herm are of a very fine deep reddish orange on the
dorsum, the branchial processes being of this colour. Ehlers describes his E. racemosa
as of an intense red (" zwischen orange und ziunoberfarben "). Others, again, were quite
greyish, or of an earthy hue. The under surface is either pinkish or pale flesh-colour.
They tinge spirit brick-red. Gosse's example from Weymouth was of a bright cinnamon-
red, with the median ventral line purplish. Milne Edwards' example seems to have
been vividly tinted of a fine cinnabar colour, very marked on the branchiae, and
mingled with yellow and green on the dorsum. The cirri were yellow, with a band of
red in the middle; while the caruncle was of a vivid red. The pinkish ventral surface
had a median band of vivid red.

The head is wedged between the anterior segments, and is distinguished mainly by
the caruncle and eyes. It is more or less linear, with the elevated region somewhat
ovoid. The two dorsal eyes, which are black and of considerable size, lie at the anterior
border, and have the tentacle, which does not seem to taper much, rising between them.
The fillet, which runs forward from the caruncle, carries a pair of short tentacles, with
tactile hair-like processes, and bending over the tip of the snout, bears the two elongated
ventral eyes, which are smaller than the dorsal. 1 The dorsal part of the fillet is stated by
Ehlers in E. racemosa to be covered with cilia, and it is probable that they also occur on
the ventral surface of the band.

The body of this species forms a somewhat elongate oval, the dorsal surface being
elevated, the ventral flattened. Dorsally a broad groove, smooth but for the seo-ment-
1 J. Y. Cams, in his 'Prod. Faun. Medit.' (1884), gives the genus only one pair of eyes.

236 EUPHROSYNE FOLIOSA.

lines and the characteristic furrows of the cuticle, occupies the centre, the sides beino*
flanked by the cirri and the dense rows of bristles and branchiae. Bach row has a
palisade of bristles towards the front of the segment, the cirrus (Plate XXXV, fig. 1)
standing freely a little behind, while the branchiae form a series of arbuscles— after an
interval — extending outwards from the latter. The second cirrus occurs in the position
formerly noted, and is generally longer than the other appendages. The anterior part
of the median groove is occupied by the caruncle, which stretches from the front to the
anterior part of the fifth segment, the tentacle springing between the dorsal pair of
eyes in the second segment. 1 J. V. Cams says his E. Audouinii (= E. laureata, D. Oh.,
and E. racemosa, Ehlers) has a pair of tubercle-like antennae, seven branchial trunks, and
both bifurcate and smooth dorsal bristles. The ventral surface is marked in the British
examples by the presence of a slight median groove, and by the regularly arranged
segments. The first bristle-bearing segment lies on each side of the median fillet, and
ventrally forms a pad on each side of the middle line in front of the mouth; the second
and third slope obliquely inwards towards the first, whilst the halves of the fourth
are continuous behind the mouth, the outer ends being directed forward ; and a similar
inclination characterises the eight or nine which follow. The posterior segments, on
the other hand, have the outer regions sloped backwards. Two globular cirri terminate
the body posteriorly, the anus lying above them.

The segments are much more distinctly marked than in Spinther, being readily
recognised dorsally by the palisades of bristles flanked by rows of branchiae, and ventrally
by the sulci between each.

The feet are also fairly defined for a group in which the dorsal division forms
part of or becomes confluent with the dorsal arch of the body, and it is this structure
which, amongst other features, gives grounds for the classification followed in the
"Challenger" Annelida and in the present treatise. In each segment the innermost
structure is the cirrus, which is attached rather behind the middle of the segment, and
is a tapering organ of considerable length, with a filiform tip richly covered with cilia.
About the middle of the segment the palisade of bristles runs from the inner to the outer
border. The ordinary bristles have a well-marked spur with a slightly curved tip (Plate
XXXV, figs. 15 and 16). The serrated forms have a deeply cleft tip, both limbs being
curved, the longer doubly so when viewed laterally (Plate XXXV, f\g. 17). Milne
Edwards' figure of this bristle is quite diagnostic, though not absolutely correct. The
figure of the same kind of bristle given by Ehlers for E. myrtosa differs only in its
artistic treatment. Antero-posteriorly, again, this kind of bristle is straight (Plate
XXXV, fig. 18), The palisade of bristles just mentioned terminates at the dense tuft
of bifid bristles of the first-mentioned type (figs. 15 and 16) which projects from the foot
on each side. This tuft includes stout forms with comparatively short bifid tips, as well
as numerous slender forms with the tips almost in a line with the shaft and having
serrations on the edge of the tip (Plate XXXV, fig. 17); the short spur, moreover, has
a membranous guard from the tip to the side of the larger limb. The bristles in the
Hebridean example were remarkably developed all over. Schmarda found a golden

1 Ehlers describes the caruncle of his E. racemosa as having an ovoid basal and a compressed
upper region.

EUPHROSYNE FOLIOSA. 237

fluid in the axis of the bristles of E. polybranchia, and more careful investigation of
the British forms in life might lead to further information on this point.

In section about four rows of bristles are observed in the tufts, the more slender
being guarded by outer (i. e. anterior and posterior) rows of large bristles. The median
cirrus is situated opposite (in front of) the third branchia, and behind the palisade of
bristles ; De Saint-Joseph says between the second and third branchise.

The ventral cirrus lies between the parapodia, and is thus hidden in the ordinary
position of the parts.

The branch ia3 form a series of arbuscles with foliate tips ranged transversely behiud
the palisade of bristles. They vary in number from seven to eight. Each consists of
a short main stem, which rapidly and somewhat dichotomously divides into branches
terminating in the expanded ovate processes. The latter (Plate XXXV, fig. 3) have a
well-marked cuticle with the subjacent granular layer (hypoderm) filling up the central
region. The cells under the cuticular investment are larger and somewhat regularly
arranged. So far as the preparations show, the long cilia occur on the sheltered parts of
the larger branches.

De Saint-Joseph states that he has seen the red blood penetrating the branchige,
contrary to the opinion of Claparede, but without reaching the terminal enlargement of
the organ, which is shut off by a septum. This has not been observed in our examples.

Very considerable variation in the form of the branchise occurs during development,
young examples having few branches, and the tips more or less cylindrical or only slightly
tapered ; then they become broadly lanceolate. "What relation the condition of the
terminal processes of the branchiae have to injuries and regeneration is at present
unknown, but small forms do not always show elongated tips, some measuring about a
quarter of an inch presenting short branchial arbuscles with broadly ovate tips.

Reproduction. — Ripe ova occurred in the specimens from the Channel Islands in July.
The axial blood-vessels of the ovaries in the species from the Cape were mentioned by
Schmarda.

In describing this species for the first time Audouin and Milne Edwards observe
that it differs little from Eujphrosyne myrtosa, found by Savigny on the shores of the Red
Sea. In the latter species, however, he finds but seven branchiae, whereas in his species
eight occur ; and they are shorter than in E. lauredia, and more densely tufted than in
E. myrtosa, while the tips are large and ovoid. The caruncle, moreover, is narrow,
almost linear, rather elevated, instead of being ovoid, very large, and depressed.

The same form was mentioned as British in 1844 by Mr. W. Thompson, who
dredged it in Belfast Bay on shelly ground in six to ten fathoms, the discrimination of the
species having been made by Prof. Allman. Mr. Glosse, again, procured it at Weymouth
in 1853, and describes the minute tentacle at the tip of the caruncle as flattened and
truncate instead of subulate, and the general colour bright cinnamon-red rather than
cinnabar, while the median ventral line is purplish.

The E. mediterranean of Grube x appears to be closely allied if not identical with this
species. It comes from Lussin Piccolo, Villa Franca, and other places on the southern
shores, and the same remarks apply to Victor Car us' s E. Audouinii (z=:LopJwnata Audouinii,

1 ' Arch. f. Xaturges./ 1863, p. 38.

238 EUPHROSYNE ARMADILLO.

Costa (?), E. laureata, D.Ch., E. mediterranea, Grube, and E. racemosa, Ehlers). So far
as the description of Ewphrosyne racemosa of Ehlers goes there is little to distinguish
it from E. foliosa, for it has jet to be proved that the variation in the number of the
branchiae (eight to nine in E. foliosa and five and six in E. racemosa) is not due to age, and
that the position of the middle cirrus is not due to the same cause.

Grube, who examined the species in the Parisian Museum, found that the median dorsal
cirrus did not extend between the fourth and fifth branchial process, as in the figure of
Audouin and Edwards, but between the second and third, as is also the case in Ehlers'
E. racemosa. Both from the structure of the branchiee and the structure of the bristles
Grube concludes that the E. racemosa of Ehlers is synonymous. Grube was likewise of
opinion that the E. laureata and E. myrtosa of Savigny agree with the E. foliosa of
Audouin and Edwards. In all probability, therefore, the nomenclature might be consider-
ably simplified. The E. Audouinii, Claparede, as given by Carus, 1 is probably referable to
the common species (E. foliosa). This seems to be the dull reddish species figured by
Delle Chiaje (1841). At Naples the common name is " Ti veggo rosso senza spine."

The E. mediterranea of Grube 2 is the same species, though Horst 3 thinks the tips of
the branchige clavate rather than foliate. Baron de Saint- Joseph, who agrees in regard to
the association of E. racemosa, E. Audouinii, and E. mediterranea with the present species,
found that at Dinard specimens of 12 to 15 mm., and having thirty segments, were
distended with ova. The Euphrosyne intermedia, 1888, of this author, rests mainly on
the presence of longer forms of bristles — amongst the dorsal and ventral series — haying
the axial oil-like contents. He thinks they are offensive and contain poison. In all
probability this is only a variety of E. foliosa with longer bristles.

.2. EUPHROSYNE AEMADILLO, Sars, 1851.

Specific Characters. — Caruncle extending to the anterior border of the fifth segment,
with a proportionally long Particulate tentacle between the dorsal eyes. Branchige divided
dichotomously, and terminating in lanceolate processes. The bristles are of considerable
length, the bifid forms having a short spur and quite smooth, the serrated kinds having
the longer arm somewhat flattened, only slightly, though distinctly, curved, the serrations
extending along opposite parts of the fork.

Synonyms.

1850. Euphrosyne armadillo, Sars. Keise i Lofoten og Finmark. Nyt. Mag. f. Naturv., B. vi,

p. 211.
1861. „ „ Forhandl. Vidensk.-Selsk. (Aar, 18.60), vol. viii, p. 55.

1876. „ lanceolata, Mcintosh. Trans. Z. S., vol. ix, p. 395, pi. lxxi, fig. 1.

1886. „ armadillo 3 Langerhans. Zeit. f. w. Zool., vol. xl, p. 253.

1 < Fauna Medit.,' p. 207, 1884.

2 ' Arch. f. Naturges./ 1863, p. 38.

3 <

Notes from the Ley den Museum/ vol. viii, 1886.

EUPHROSYNE ARMADILLO. 239

Habitat. — Dredged in the Porcupine Expedition of 1869 on sandy mud amidst corals
off the west coast of Ireland, in 173 fathoms.

The body appears to be somewhat flattened, and in the injured preparation is about
3 mm. in length, and consists of only about nineteen segments. The dorsum agrees
generally with the typical form, except that the branchise differ considerably in structure,
and thus give a character to the region. Both dorsal and ventral eyes are very distinct,
and the tentacle is comparatively long in the example. On the ventral surface the palpi
form two rounded pads in front of the mouth, and abut on the eyes in front. Pos-
teriorly the vent is indicated by two rounded or globular processes which project on the
ventral surface. The segments in the latter region are much curved, and the tips of
several project beyond the globular anal cirri, the lines of the segment-junctions being in
one or two almost antero-posterior.

Colour is unknown.

The condition of the single specimen is unfortunately indifferent, and it is, moreover,
small (possibly immature), but the following characters were ascertained. The palisade
of bristles dorsally consists of a series of smooth bifid forms with a distinct curve at the
tip, which is slightly hooked, and a short spur at the base (Plate XXXY, fig. 8). The
serrate kind has a proportionally longer spur, the serrations on it corresponding in extent
with those on the longer fork of the bristle, which tapers a little towards the tip (Plate
XXXY, fig. 13). Some present more distinct flattening of the longer limb (Plate XXXV,
fig. 14), and the serrations are less marked, — indeed, only a limited area of similar extent
on each side of the fork shows them clearly, though very minute processes occur on the
longer limb of the fork above the former. The tips of all these bristles are distinctly
curved. On contrasting them with the bristles of Euphrosyne armadillo, Sars, from
Norway (Plate XXXV, figs. 9 — 12), a certain resemblance is apparent in all, but the
tips both of the smooth and serrate kinds are proportionally longer and narrower in the
Norwegian form, and the curvatures differ ; such, however, may be due to age or other
conditions. The longest tips in the case of the smooth bristles occur in the foot.

The branchiae (Plate XXXV, fig. 2) appear to be five or six in number, and when
viewed under a lens have a different character from those of Euphrosyne foliosa, since the
tapering tips are much more slender. They branch from near the base in a similar
manner, the tips being truly lanceolate. The variations, however, seen in the branchiae
of E. foliosa show that no strict reliance on the external appearance of these organs
can be maintained. The only feature of moment is the tufted condition of the tips in
the larger Norwegian examples, which also have proportionally longer terminal pro-
cesses.

Michael Sars describes this species as of a pale yellowish colour, and having nineteen
segments. The narrow caruncle reaches the fifth segment. Cephalic lobe elongate,
narrow; the posterior (dorsal) eyes situated in front of the caruncle, at the base of
the Particulate tentacle, which is conico-acuminate, shorter than the caruncle. Two
short cirri on the dorsum between the pinnae. Branchiae five, rarely six (two or
three of the anterior and posterior segments with fewer), four to ^ive dichotomously
divided, with conico-acuminate tips. Superior edge of pinna with two cirri, ventral with
one. Setae unequally bifid, with serrations in the fork of the dorsal form, while the in-

32

240 EUPHROSYNE ROBERTSOM.

ferior bristles are smooth, with the exception of a single small denticle near the apex.
Length 8 mm., breadth 2\ mm.

It was dredged at Manger, about three miles north of Bergen, on shell-sand.

There is still a degree of uncertainty in regard to the identity of the British and
Norwegian forms, which can only be cleared up by the capture of fresh specimens in this
country.

3. ElTPHROSYNE ROBERTSONI, U. S.

Specific Characters. — The tips of the branchiae are digitate, with only a trace of a
swelling. Dorsal bristles smooth and bifid. The forked and serrate bristles somewhat
resemble those of E.foliosa.

Habitat. — From the Firth of Clyde, whence an example was sent to the British
Museum by the late Dr. David Robertson. It is labelled Euphrosyne foliosa, Aud. and
Ed., No. 1049, 64, 6, 30, 6.

The specimen is upwards of half an inch in length, and shows a similar arrangement
of the caruncle and eyes to E.foliosa, from which, indeed, there is little to distinguish it
in external appearance with the naked eye. The branchiae, however, differ, presenting
stout stems, with dichotomously but sparsely divided tips, which have a nearly uniform
diameter ; for the slight enlargement in some does not alter their character as digitate
processes, slightly lobate and then tapered at the tips (Plate XXXV, fig. 4). They
somewhat resemble the figure of the branchiae of Euphrosyne myrtosa, as given in
Savigny's plate, but little reliance can be placed on it.

The dorsal palisade of bristles x consists of smooth bifid forms, some with longer
(Plate XXXV, fig. 7) and some with shorter tips (Plate XXXV, fig. 6). The bifid
serrate kind (Plate XXXVII, fig. 33) appear to approach those of Euphrosyne foliosa,
though the curves and general character are different.

I have named this species after one of the most persevering and patient of Scotland's
naturalists, who for many years devoted his energies and experience to the fauna of the
Clyde, and to whose influence and example the marine laboratory at C umbrae owes its
existence.

Family II. — Aphroditim.

Annelids of an ovate or oblong form, convex dorsally, with a distinct head
(prostomium), on which are a pair of eyes and a median tentacle, and under which
is a papillose facial tubercle. No lateral tentacles ; two palpi ; tentacular cirri long ;
buccal cirri (ventral cirri of the second foot) moderately long. Proboscis large and
powerful, with four thickened muscular ridges representing teeth, and tough internal

1 I am indebted to Prof. Jeffrey Bell and Mr. Sumner for most courteously making preparations
of the bristles and branchiae and forwarding them for examination.

APHBODITA. 241

lining. Alimentary canal pinnately-branched, the glandular intestinal caeca being long
and complex. Dorsal fimbria small, alternating with the scales, or absent. First foot
bearing three dense tufts of bristles. Elytra fifteen pairs, occurring on the second,
fourth, fifth, thereafter on all alternate segments to the twenty-fifth, and then on every
third segment. Segmental organs (nephridia) opening by a well-marked papilla pointing
upwards between the feet. Nerve-cords median, between the ventral attachments of the
oblique muscles, or in a well-defined epidermal granular layer within the dense cuticle.

Genus V. — Aphrodita, Linnxus, 1735.

Eyes sessile ; dorsum covered with a thick, close felt of matted simple hair ; seta3 of
the ventral division of the foot very numerous, long, silky, and iridescent, and, like all the
other bristles, simple, not barbed or toothed. Intestine with eighteen long and
complex caeca (figs. 16 and 17, p. 250). Nerve-cords in a transversely elongated space
between the ventral attachments of the oblique muscles, and bounded externally by the
basement-tissue and the cuticle.

Swammerdam's account of Physalus, as this form was then called (1758), contains a
notice of the three rows of bristles on the feet, which he considered only papillae. He
thought the scales the branchiae, and was of opinion that the animal was able to
" swell and bloat itself with air." He observed the ramifications of the alimentary canal
and the presence of blood-vessels. He criticised Rondelet for placing it amongst sea-
worms, and was inclined to relegate it to the proximity of the sea-urchins. His figures
are recognisable, though the whole dorsum is rough with tufts of bristles.

Pallas, in his 'Miscellanea Zoologica ' (1766), showed the propriety of removing the
Aphroditae from the Mollusca, with which Linnaeus had grouped them. He indicated
that a more natural classification would be to conjoin the three genera, Aphrodita, Nereis,
and Serpula. He gave a general and fairly accurate description of the Aphroditidse, the
only feature requiring special notice being his observation that branchiae are present on
the dorsum, and that he included in the group Ghloeia and Amphinome (Pleione carun-
culata).

0. F. Miiller, in his ' Zoologia Danica Prodromus ' (1776), included the Aphroditidae
and other Chaetopods under his Helminthica Setosa, the other group being his Helmin-
thica Mutica, in which were Gordius, Ascaris, and Hirudo.

In Gmelin's edition of Linnaeus (1788) l the Aphroditaceans w r ere placed in the
Vermes Mollusca, after Doris, notwithstanding that there was little more than the
general ovoid outline to suggest the relationship.

By Lamarck 3 the Aphroditidae were grouped under his second order of Annelides,
i. e. A. Antennata, along with Nereids, Eunicidae, and Amphinomidae. He simply followed
Savigny.

1 ' Syst. Nat./ p. 3107.

2 ' An. sans. Vert.,' 1818, tome v, p. 304.

242 APHKODITA.

Savigny in his ' Systeme ' l made the Aphroditidas the first family of his Nereidees,
characterised by having the branchia3 in the form of a ridge or papilla situated superiorly
at the base of the dorsal branch of the foot. They are absent from the second, fourth,
fifth, seventh, ninth, and eleventh pairs of feet, and so on until the twenty-third or
twenty-fifth. They determine by their absence that of the superior cirrus, and are
replaced by scales. The scales, when present, number from twelve pairs or fewer to
thirteen pairs or more, and extend to the twenty-third or twenty-fifth segments, and
are followed or not by supernumerary pairs. They are formed of separate membranous
lamellae, the upper thickened, sometimes horny, the inferior delicate, and attached by a
hollow pedicle to the base of the feet without branchias — in a position, however, corre-
sponding to the attachment of the latter. The mouth has a proboscis and four jaws.
The former is cylindrical, massive, striated transversely, and furuished with a fringe of
small tentacles at the orifice. The jaws are horny or cartilaginous, flat, short, more or
less free at the point, and have a vertical motion on each other. The eyes are four in
number, two anterior and two posterior. The antennas are retractile, elongated,
generally complete ; the median of two articulations, the first being short ; the unpaired
the same; the anterior always present and much larger than the others, finely ringed,
conical, and with tapered tips. The feet have either two divisions or these are united —
furnished with aciculi. Cirri prominent, generally composed of two chief divisions ; the
first, short and thick/lies at the base of the other, which is retractile. The superior cirri
are large, extending beyond the bristles, while the latter pass beyond the inferior cirri.
The first pair of feet have the divisions intimately united, without bristles or with
numerous bristles, and the two cirri elongated like tentacles. The second pair of feet
have also a long inferior cirrus, a little larger than the succeeding. The intestine is
provided with numerous ca3ca, which are most distinct in Aphrodita proper. Savigny,
from the foregoing, used the term (Aphroditidge) in its widest sense.

In Cuvier's c Eegne Animal ' 2 (Mem. Edit.) M. Audouin arranged the Aphroditidse
and Polynoidae under the order Dorsibranchiata (corresponding to the Nereidees of
Savigny).

Audouin and Milne Edwards (1834) classed the Aphrodisiens (which included
Aphrodita, Polynoe, Sigalion, and Palmyra) as the first family of their Annelides
Errantes, and they gave a description which defined the somewhat extensive group.
One important feature is the presence of a double row of membranous scales, the elytra
of Savigny, fixed on the dorsum by a pedicle to the superior division of the foot, and ac-
cording to the authors filled with ova at certain periods. They occur in some on
all the feet or on alternate feet, while in others they are absent. They refer to small
processes attached to the under surface of the scales as branchiae. Cirri occur on the
segments devoid of elytra, with the exception of Sigalion, where they are present on all
the segments. The feet are bilobed, each division being armed with a spine, bristles, and
cirri, the last existing on every foot ventrally. In the first segment the dorsal cirri
become tentacular cirri. The antennas are attached to the head, and are three in number

1 ' Systeme des Annelides/ tome i, 3e part., 1820.

2 MM. Audouin, &c, ' Regne An./ 1836-7.

APHEODITA. 243

— a median and two lateral. The eyes are usually four in number and placed in pairs —
one in front of the other.

As the result of their own researches they restored the older name Aphrodisiens for
Savigny's Halithees, and made three principal groups : (1) those in which the elytra
alternate with dorsal cirri and branchiae, (2) those having these organs on the same foot,
and (3) those devoid of elytra. In the first tribe the authors placed Aphrodita, Polynoe,
and Polyodonta. In the second division, which from the elongated body they termed
Aphrodisiens vermiformes, they ranged Acoete and Sigalion ; while in the third group is
placed Palmyre. We shall deal at present only with the first mentioned, viz. Aphrodita,
which was characterised by the authors as furnished with thirty elytra fixed to feet
which bear neither branchiae nor superior cirri, and which alternate regularly (with the
exception of the fourth and fifth segments) to the twenty-fifth segment, with other feet
which have cirri and branchiae. The elytra fixed to the succeeding segments are differ-
ently arranged. Three antennae are present. The jaws are small and cartilaginous or
absent.

A sea-mouse and parts of its digestive system are figured in Tav. iv, fig. 10, of Delle
Chiaje's Memoire (1822), but no description is given.

The same author 1 (1841) speaks of a pair of oval ovaries in Polyodontes maxillosa,
filled with a transparent liquid at the dissepiments and at the bases of the feet, and of an
analogous group of rosy ovaries in Hermione hystrix, and yellowish ovaries in A. aculeata.
These Meckel considered to be small branchiae. In April they had advanced consider-
ably. He did not consider the scales respiratory, as Cuvier, Carus, and Duvernoy had
done, and he supported his opinion by the presence of special branchiae in Sigalion
squamosum. He described the ventral ganglionic chain as a nerve-artery in A. aculeata
and H. hy stria, but had seen vascular trunks in both on the intestine. The blood-vessel
in his figure seems to ensheath the oesophageal trunks and ventral chain. Long before, 3
he had compared the alimentary system of the Aphroditaceans with that in such as
Pleurophyllidia.

Oersted 3 (1843) signalised the Aphroditacea amongst the JSTematode-like Ohaetopods
as having imperfect branchiae (simple). He considered the scales the branchial organs.

Grube's description of the Aphroditea in his ' Familien der Anneliden ' (1851) is
brief but characteristic. He included Palmyra under the same family.

Sir J. Dalyell 4 says that, " though seemingly timid, the Aphrodita is probably fierce
and rapacious, overpowering creatures incapable of resistance ; and there is even reason
to believe that it occasionally devours its own kind." By Aphrodita he means the
Aphroditacea.

Kinberg 5 (1857-8), following on the lines of Audouin and Milne Edwards, grouped
the Aphroditea of Savigny into seven families, the first of which, Aphroditacea, corre-
sponds with our Aphroditidae. These have an oblong wide body, with a rounded head

1 ' Memoire/ vol. i, p. 121.

2 1823.

3 "Ann. Danie. Consp./ 1843, p. 4.

4 ' Powers of the Creator/ vol. ii, p. 163, 1853.

5 ' Eugenies Eesa/ &c, p. 1, 1857-8.

244 APHRODITA.

and a facial tubercle between the palpi and the front of the mouth. Tentacle extending
from the middle of the cephalic lobe. KTo antennas. The sessile or subpedunculated
eyes are situated in front of the middle of the cephalic lobe. The palpi are long, thick,
tapering, and ciliated ; and two tentacular cirri are on each side of the first pair of feet..
The buccal cirri (ventral pair of the second feet) are longer than the succeeding. The
exsertile pharynx has ridge-like transverse processes — ^a^'-cartilaginous, and resembling
jaws. The branchias are in the form of low papillas, situated above and internal to the
bases of the dorsal cirri, and covered by the elytra. They are not always obvious. The
elytra occur on segments 2, 4, 5, 7, 9, &c. The other families were Iphionea, Polynoina,
Acoetea, Sigalionina, Pholoidea, and Palmyracea.

The genus Aphrodita he distinguished as follows : — Eyes sessile (pigment-spots in
pairs). First pair of feet furnished with numerous bristles, and with tentacular cirri.
Dorsal division of the foot distinct from the ventral, low (small) and broad, with strong
sharp spines and capillary bristles, forming a kind of felt on the dorsum ; ventral division
carried outwards, blunt, with numerous bristles, smooth, acute, but neither glochidiate
nor bidentate.

Chenu l (1859) chiefly followed Milne Edwards in placing the Aphroditiens as the
first family of his Annelides Errantes, the Amphinomiens and Euniciens forming the
second and third families.

In the posthumous c British Annelids ' of Dr. G. Johnston, published by the British
Museum in 1865, the first family, Aphroditaceas, included not only the genera pertaining
to the Aphroditidas, but the Polynoidas and Sigalionidas. The author followed in his
description Audouin and Milne Edwards. He gives three species of Aphrodita, viz. the
common form, A. borealis (which is the young of the former) and A. hystrix (Hermione
hystrix). Like Grube, he classified the Annelids under the Eapacia and Limivora.

De Quatrefages 2 included the whole of the group forming the subject of this fasciculus
— with the exception of the Amphinomidas and Euphrosynidas — under the family Aphro-
ditidas, in which the regions of the body are similar while the segments are dissimilar.
They fall under his first order Errantes. The author criticised the classification of
Kinberg, and held that only two families existed in the sub-order, viz. the Aphroditidas
and the Palmyridas, the one characterised by the presence and the other by the absence
of scales. The Aphroditidas form a very natural family of the Errant Annelids. The
head bears two to three antennas and two to four eyes ; w T hile the buccal segment is often
indistinct, and with or without tentacles. The body is more or less covered by the
elytra, and the segments present differences which are repeated with regularity. The
antennas (tentacles) receive their nerves directly from the brain, and their number is at
most three. The external antennas (palpi) are really the tentacles of the buccal ring, and
receive their nerves from a special ganglion. The nerves of the tentacular cirri, again,
come from the first ganglion of the ventral chain, being modified processes of the first
pair of feet. The head bears a kind of caruncle (facial tubercle ?) in front. The eyes in
general are small, resting on the brain, though in some they are pedunculated and sus-
ceptible of movement. The mouth has thick lips.

1 < Encyclop. d'Hist. Nat./ 1859.

2 f Anneles Marins, &c./ 1865.

APHRODITA. 245

The feet are biramous, bearing scales or tentacular cirri, in general only the one or
the other. The ventral cirrus occurs on all the feet. Some have a resplendent covering
of hairs, and a felt-like coating on the dorsum protecting the scales. Those without
such sometimes show a radiate arrangement of the bristles. In the scales he describes
a lacunar system in connection with the general cavity of the body, and therefore he
thinks Savigny was right in associating them with respiration, though he was so far
misled by a balloon-like condition in imperfectly preserved specimens. He does not
regard the elevated and ciliated processes on the dorsum of the feet as branchiae, for
they have no central vessel and no lacunae, and the cutaneous tissues present no special
modification. On the other hand, De Quatrefages saw in the pretended branchial function
of the branching digestive system an analogy with what he had formerly described in the
iEolidae as phlebenterism.

The circulatory apparatus he says agrees with the typical condition, but is difficult
to follow, as the blood is pale. There are dorsal and ventral vessels as described by Trevi-
ranus, and a third considerable trunk accompanying inferiorly the abdominal nerve-chain.

The cephalic ganglia are comparatively large, and the exterior thereof brownish red.
The ventral chain has the ganglia united, though in general the two halves are distinct.
No commissure exists between the lateral ganglia of the first three pairs of feet — a condi-
tion absent in the Polynoidae. The visceral system of nerves consists of a muscular
trunk and a ganglion with a connective joining the brain. It supplies the muscles of the
proboscis.

The only remarks the author makes in regard to the reproduction of the group is
that in a large number of examples of Ajphrodita hystrix he found irregular mounds con-
sisting of eggs enveloped by delicate tissue along the digestive canal and touching the
body-wall. These individuals consequently showed a large number of ova or of sperms
in the perivisceral cavity. Further, in a male he observed sperms escape as a white
thread at the base of the ventral division of the foot about the nineteenth segment.

The classification adopted by De Quatrefages was based for the most part on
external characters, such as the arrangement of the scales, the absence, alternate or con-
tinuous condition of the dorsal cirri, the nature of the antennae (tentacles), and the jaws.

In his general remarks on the Aphroditidae, Claparede 1 corrects the error of
Williams that vibratile cilia are absent from the peritoneal surface of Aphrodita aculeata.
This author was, however, in doubt concerning the vascular system, for though he found
a dorsal and a ventral vessel according to the old observations of Pallas and Treviranus,
yet he could not satisfy himself that they pertained to the vascular system. He makes
a few remarks also about the respiration in the group, stating that in Hermione hystrix,
during the alternation of expansion and contraction, the last pair of scales in the latter
function are raised, and a powerful stream of water sent out. The same is seen, though
to a much less notable extent, in A. aculeata. In the latter species bubbles of air
sometimes accompany the currents, so that Swammerdam had some foundation for the
remark that the Aphroditae swallow (gorgeant) air. He had, however, overlooked the
observation of Sir J. G. Dallyel when he said former authors had not observed these
respiratory movements.

'Ann. Chet, Napol./ 1868.

246 APHBODITA.

In his summary of the Aphroditacea l (1868) Dr. Baird showed that while Audouin
and Milne Edwards included six and Grube seven species, later authors had so increased
the number that Kinberg found it convenient to form most of the older genera into
distinct families. He gives a description of the family Aphroditidas after Kinberg, with
four genera, viz. Aphrodita, Hermione, Aphrogenia, and Lxtmonice. A succinct account of
the species follows under each genus, and he seems to have acquiesced in Dr. Johnston's
view that Aphrodita borealis was a distinct species — a position he subsequently vacated,
this being only the young of A. aculeata. He added one or two new species to the list.
His Lastmonice Kinbergi, however, as he afterwards admitted, is identical with L.filicornis,
Kinberg, a species widely distributed in Northern waters.

Ray Lankester 2 found hemoglobin in the nerve-cords of A. aculeata.

Grube next gave a survey of the family Aphroditidse (which included the whole
series here considered). 3 He divided the group into sub-sections thus : — a. The one
segment with elytra, the other with cirri ; no jointed bristles. These he subdivided as
follows : — a. Between the elytra-bearing segments of the body one segment carrying a
dorsal cirrus ; in the posterior part of the body mostly two bearing cirri, or the elytra
absent. 1. Hermionea (Aphroditacea, Kinberg). 2. Polynoina. The first group, the
Hermionea, which alone concerns us at present, he classified according to the condition
of the ventral bristles and the state of the eyes, (a) All the ventral bristles have simple
tips. In Aphrodita he gives as characters the following: — The ventral bristles in three
rows, short, thick; the dorsal bundle of all the segments bearing the elytra furnished
with longer and stronger bristles and two bundles of fine hair-like bristles, the alternate
segments also with another felted series, under which the elytra lie; two eyes.

Schmarda 4 included the Aphroditidse and Amphinomidse under his Notobranchiate
Chastopods.

In his treatise on the Annelids collected by Semper in the Philippines (1878) Grube
terms the family Aphroditea. Besides the characters previously stated, he mentions
that all the segments bear ventral cirri and two fascicles of bristles, and that the fourth
and fifth segments always carry elytra. The stomach is subcartilaginous, and the
intestine has pinnate ca3ca.

Claus 5 (1880) grouped under the family Aphroditidaa the sub-families Herminioninse,
Polynoinaa, Acoetinaa, Sigalioninao, and Polylepinaa ; while his second family was the
PalmyridaB. Under the Sigalioninas he embraced Psammohjce and Pholoe.

Levinsen 6 (1883) follows Malmgren's classification of the Aphroditida3 in his paper
on the Northern Annelids.

Cams 7 (1884) describes the genus Aphrodita as having fifteen pairs of elytra on
alternate feet, which are destitute of cirri ; the intermediate bearing a cirrus and branchia ;

1 c Journ. Linn. Soc./ vol. viii, 1865.

2 f Ann. Nat. Hist./ 4th series, vol. xi, p. 97, 1873.

3 ' Sitzrmg. d. Schlesischen Gesell./ 1874.

4 'Zoologie/ 2nd edit., 1877, Wien.

5 ' Grundzuge d. Zoolog./ 1880, pp. 490-98.

6 Aftryk, af. ' Yidensk. Meddel. f. d. Nat. Foren. i. Kjobenhavn/ 1882-3.

7 ' Fauna Mediterr./ 1884.

APHRODITA ACULEATA. 247

tentacles three ; long tentacular cirri ; maxillse small or none ; dorsum covered with a
close layer of felt ; ventral spines simple at the apex.

In his ' Annelids of the ' Blake ' ' (1887) Bhlers keeps to the family Aphroditida3, with
sub-families Hermionea, Polynoina, Acoetea, and Sigalionina. The Hermionea he dis-
tinguishes by the facial tubercles and the arrangement of the scales ; and he separates them
from the Acoetea and Sigalionina by the condition of the branchia3 and the structure of
the bristles.

Aphrodita aouleata, Linnmus, 1765. Plate XXIV, figs. 4 and 5.

Body broad, cephalic lobe inflated anteriorly, basal part of the tentacle only half
the length ; facial tubercle with sparsely distributed small globular papillaa ; central
region of the gut differentiated from the complexly lobed glandular csoca ; spines of the
dorsal division of the foot long, piercing the felt; capillary bristles greenish, iridescent
(burnished). Ventral bristles in three rows.

Synonyms.

1554. Physalus, Kondelet. De Piscibus, p. 428.

1602. Scolopendra marina, Aldrovandus. Insect., p. 636, f. 1.

1634. Physalus, Moufet. Tlieatr. Insect., f. 8—15.

1677. Vermis aureus, Oligerus Jacobseus. Acta Haf nise, vol. iii, pp. 87, 88, and 89.

„ „ „ Bartholinus. Act. Haf., vol. iii, p. 88, tab. 88.

1684. Eruca marina Rondeletii pilis in dorso instar colli columbini variegatis, Sibbald. Scot. Illustr.,

vol. ii, p. 32.

1686. Hystrix marina, Eedi. Opusc, vol. iii, tab. 35.

1705. Scolopendra marina, Molyneux. Phil. Trans. Abridg., 1st edit., vol. ii, pp. 833 — 836, pi. xii,

f. 234 and 235.

1714. Eruca echinata marina griseo fusca. Barrelier. Plantse Grail., &c, p. 131, tab. 1284, n. 1.
1734. „ „ „ Seba. Thesaur., vol. i, p. 141, tab. 90, f. 1—3; vol. iii,

p. 9, tab. 4, f. 7, 8 (1758).

1746. Aphrodita nitens, Linn. Faun. Suec, p. 367, No. 1284; Mus. Adolph. Fred., p. 93.
1752. ,, elliptica versicolor, and the Sea Mouse, Hill. Hist. Anim., vol. iii, p. 90.

„ „ subrotundata. Ibid., p. 91.

1756. Mus marinus, Linn. Syst., edit. 1756, p. 79.

1758. Physalus, Swammerdam. Biblia Naturge (transl.), vol. ii, p. 150, tab. 10.

1762. Aphrodita aculeata, Baster. Opusc. Subs., vol. ii, 2, p. 62, tab. 6, f. 1 — 4.

1765. „ „ Gunner. Trondh. Selsk. Skrift, vol. iii, pp. 59—80, tab. 88.

1766. „ „ Pallas. Misc. Zool., p. 77, tab. vii, f. 1—13.

1768. „ ,, Gunner. K. Norske Selsk. Skrift, vol. iv, p. 95, tab. 10.

„ Physalus, Jonston. Hist. Nat., vol. iv, tab. 28.

1776. Aphrodita aculeata, O. F. Miiller. Zool. Dan. Prod., p. 218, No. 2641.

1777. ,, ,, Penn. Brit. Zool., vol. iv, p. 44, pi. xxiii, f. 25, and edit. 1812, vol. iv,

p. 86, tab. 25, f. 1.
1788. „ „ Herbst. Vers., Bd. ix, p. 50, tab. 11.

1790. „ ,, Linn. Syst. Nat., Grmelin, t. i, pars 6, p. 3107.

1791. Aphrodite herissee, Bruguiere. Encyclop. Meth., vol. vi, p. 85.

1806. Aphrodita aculeata, Turton's Gmelin, vol. iv, p. 79.

1807. „ „ Turton. Brit. Fauna, p. 136.

33

248 APHRODITA ACULEATA.

1816. Aphrodita aculeata, Cuvier. Diet, des Sc. nat., vol. ii, p. 282.

1817. „ „ Stewart's Elements, vol. i, p. 387.
1821.
1823.

1825.

Treviranus. Zeit. f. Physiol., vol. iii, 2, p. 157.
Delle Chiaje, Mem. I, 182, Suppl., Tav. iv, f. 12.
Home. Oomp. Anat., vol. iv, pi. xxxix, f. 1, 2*
Blumenbach. Elem. Nat. Hist., p. 245.

1826. Halithea aculeata, Lamarck. Anim. s. Vert., vol. v, p. 307, and 2e edit., vol. v, p. 542.
„ ,, „ Savigny. Syst. des Annel., p. 19.

,, „ sericea. Ibid.

„ Aphrodita sericea. Ibid.

1827. Halithea aculeata, Risso. L'Europ. Merid., vol. iv, p. 412.
„ ,, ,, and aurata. Ibid., vol. iv, p. 413.

1828. „ „ Stark. Elem., vol. ii, p. 140.

,, Aphrodita aculeata, De Blainville. Diet, des Sc. nat., vol. lvii, p. 456, Atlas, pi. ix, f. 1.

1829. „ „ Delle Chiaje, Mem., vol. iv, p. 209, Tav. 68, f. 10—15.

1830. „ „ Regne Anim., 2e edit., vol. iii, p. 206.
„ „ „ Bosc. Vers., vol. i, p. 181.

1831. „ ,, Guerin. Icon, des Reg. Anim., vol. ii, pi. ix, f. 1.

„ Halithea aculeata, Treviranus. Edin. Journ. Nat. and Geogr. Sc, vol. iii, pp. 51, 246.

1832. Aphrodita aculeata, Aud. and Ed. Ann. des Sc. nat., vol. xxvii, p. 402, pi. viii, f. 7.
1834. „ „ And. and Ed. Annel., p. 66, tab. i a, f. 7.

1836. ,, „ Templeton. Loud. Mag. Nat. Hist., vol. ix, p. 234.

1836-7. „ „ Regne Anim., Illust., pi. xvii, f. 2 (Mem. edit.).

1839. „ „ Johnston. Ann. Nat. Hist., vol. ii, p. 429, pi. xxi, and vol. v, p„ 305 (1840).

1840. „ „ Roget. Bridgew. Treat., vol. ii, pp. 102 and 298.
„ „ „ G-rube. Actin. Echin. a. Wiirmer, p. 88.

,, „ horealis, Johnston. Ann. Nat. Hist., vol. iv, p. 370 (young).

1841. „ aculeata. Delle Chiaje, Descrizione, v, 59 — 61, Tav. cix, f. 6— 8 ; Tav. exxxiii, f. 10—

15 (as before).

1842. „ „ Fleming. Ency. Brit., edit. 7, vol. xi, p. 221.

1843. „ „ Oersted. Consp. Annul. Dan., p. 11.

1846. The Sea Scolopendra, Adams. Paul. Mgin., vol. ii, p. 174; vol. iii, p. 344 (1847).

1850. Aphrodita aculeata, De Quatrefages. Ann. d. Sc. nat., 3e ser., vol. xiv, p. 362, pi. viii, f. 3 and 4.

1851. „ „ Williams. Report Brit. Assoc, 1851, pp. 172, 217, 237, and in Ann. Nat.

Hist., ser. 2, vol. xii, p. 405, pi. xiii, f. 5.
,, a ,, G-rube. Earn. Annel., pp. 35 and 119.

Maitland. Fauna Belg., p. 214.

Dalyell. Pow. Creat., vol. ii, p. 170, pi. xxiv, f. 15, 16.

Mettenheimer. Abhandl. Senkenberg. Gesellsch., t. i, 1854, p. 9,
Frankfort (fide St. Joseph).

Thompson. Fauna Ireland, vol. iv, p. 435.

O. G. Costa. Annel. di Napoli, p. 2, Tav. i, f. 1—6.

Kinberg. Freg. Eugen. Resa, vol. ii, p. 3, tab. 1, f. 2.

Danielssen. K. Norske Videns.-Selsk., Bd. iv, Hft. 2, p. 114.

Grube. Die Insel Lussin, &c, p. 77.

Malmgren. Nord. Hafs-Ann. Kgl. Vet. Akad. forh., 1865, p. 52.
j> a » De Quatrefages. Ann., 191, pi. vi, f. 1.

„ Milnesia horealis, De Quatrefages. Annel., vol. i, p. 213.
„ Aphrodita horealis, Johnston. Catalogue Brit. Mus., p. 104, pi. x, f. 1 — 18.
1873. „ aculeata, Sars. Bid. Kundsk. om Christ. Fauna, p. 1.

)i

a

1853.

Ji

1854.

a

1856.

■>i

1857.

a

1858.

it

1859.

a

1864.

a

1865.

?}

APHPODITA ACULEATA. 249

1873. Aphrodita aculeata, Willemoes-Suhm. Zeitsch. f. w. Zool., vol. xxiii, p. 347.
1875. yi „ Mcintosh. Invert., &c, St. Andrews, p. 115.

1879. „ „ Tauber. Ann. Danic, p. 78.

1884. „ „ Y. Carus. Fauna Medit., p. 198.

„ „ „ Webster and Benedict. Ann. Chset. Mass., Kept. Com. Fish and Fisheries,

p. 699.
1886. „ „ Harvey Gribson. Vermes, Liverpool, Proc. Lit. Philos. Soc, vol. xl, p. 148.

1888. „ „ De Saint-Joseph. Ann. d. Sc. nat. (7), vol. v, p. 146 (Polychast. Dinard).

1890. „ ,, Malaquin. Ann. Boulon, 14.

Habitat. — A. aculeata is by no means uncommon in deep water off the eastern
shores, indeed all round the British Islands. It generally frequents a somewhat soft
bottom. A specimen comes from 580 fathoms, on oozy ground in the Faroe Channel,
'Knight Errant,' 1880 ('Challenger Report,' p. 34). After great storms the beach at
St. Andrews is sometimes, e. g. in April, 1857, strewn with multitudes, so that the retiring
tide leaves a line of them for more than a mile. As the species is seldom encountered
within the bay proper, many were probably brought by the currents from the off-shore
grounds, such as near the Bell Rock and south-east of the Island of May. Grube, who
found it in the Adriatic (Lussin?), says that, according to Claparede and Malmgren, A.
aculeata is distributed both in the Atlantic and the North Sea, while Yon Martens found
it at Madeira. It extends to the shores of America (Massachusetts, &c), and a closely
allied species to the north-west coast, though the minute characters of the form from
the latter region have not yet received that attention necessary for certainty.

The head in A. aculeata (Plate XXIV, fig. 6) is smoothly rounded in front, with a
median elevation posteriorly, and thus differs from that of Ldetmatonice filicornis. The
comparatively small, smooth tentacle proceeds from the centre of the anterior curvature,
and has a short basal articulation ; then it gently dilates to the somewhat clavate tip of
the central piece, the terminal portion beyond having a dilated basal region, after which
it narrows and ends in a slightly enlarged tip. The terminal articulation varies, being
longer in some, shorter in others, and often presents a fusiform outline, with the tip
slightly enlarged. A little behind the tentacle is, on each side of the median line, a
blackish or brownish area (in the preparations) — the eye. The palpi are proportionally
shorter than in Ldetmatonice filicornis, and taper gently from their basal articulation to the
tip. One is sometimes longer than the other — from reproduction of a lost organ. They
are papillose, as in L. filicornis, only the papillge are somewhat shorter. The facial tubercle
has sparsely distributed small globular papillse (Plate XXXYI, fig. 23), thus differing
much from L. filicornis. 0. G. Costa's fig. 2, Tav. i, therefore, is not an accurate
representation of the part in this species.

Body. — The body is somewhat ovoid or broadly spindle-shaped, with the broad end
in front. The dorsum is convex, covered with the dense greyish felt in the middle, and
flanked by the gorgeously iridescent green and golden hairs and lustrous brown spines.
The scales are entirely concealed by the felt, while in it many foreign structures are
often fixed. Segments (bristled) forty-three.

The ventral surface is flattened, and the skin is somewhat warty and rough, giving
rise to the somewhat fanciful comparison by Pallas to that of the shark, and often tinged

250

APHRODITA ACULEATA.

of a brownish hue. It is marked by the transverse ridges indicating the segments. The
arrangement of the folds at the mouth is seen in Plate I, fig. 1.

The papilte of the segmental organs commence at the ninth and continue to the
twenty-third foot. There thus appear to be fifteen pairs. The removal of the apertures
of these organs entirely from the ventral surface is a feature of the sub-family, and
distinguishes them at once from the Polynoidse.

Digestive System. — The papilla of the terminal region of the extruded proboscis

Fig. 16.

Fig. 17.

- \ (D

m

/y\

Digestive apparatus of Aphrodita aculeata. From a
well-preserved spirit-preparation. — A. W.

Digestive apparatus of Aphrodita aculeata in a fresh
specimen. The stomach is drawn forward. — A. W.

are somewhat lobate, though usually termed fasciculate (Plate XXXVII, fig. 1). They

differ from those of L. filicornis, yet they sometimes appear as if obscurely dichotomous.

The proboscis itself is a powerful muscular organ, formerly eaten by the natives of

APHRODITA AOULBATA.

251

the Belgian shores, and even in our own country. Only shreds of a fibrillar gelatinous
substance have been found in it. Grube 1 describes six layers in the organ.

About six of the anterior cseca may be considered pregastric (Fig. 16 — in
contraction). The lateral cgeca of the gut are longer than in Lxtrnatonice or Hermione,

Fig. 18.

m.ob„o

Dissection of Aphrodita aculeata to show the segmental organs (nephridia),, s.o.; d.m., dorsal longitudinal muscles;
m.ob.o., outer attachments of the oblique muscles ; m.oh.i., inner attachments of the oblique muscles ; m.b., oblique
muscles ; n.c, nerve-cord.— A. T. M.

and form slender tubes connecting the remarkable caeca near the body-wall and
the long moniliform glandular appendage inferiorly with the central system. These
tubular portions of the casca are less truly dorsal in their origin than those of the genera
mentioned. The woodcut just referred to shows the canal and its appendages in a state
of contraction. In the living and fresh condition the parts often present a different

1 1873.

252 APHRODITA ACULEATA.

aspect (Fig. 17), the caeca forming voluminous organs in the body of the Annelid.
Pallas found nothing in the proboscis (his ventriculus), whereas in the gut he met with
particles of fuci. In many at St. Andrews and elsewhere mud of various degrees of
darkness occurs in the canal and its branches. Swammerdam thought the ramifica-
tions of the intestine anastomosed with each other.

Scales. — The overlapping, irregularly rounded scales x number fifteen pairs, and are of
considerable size. The first two show a narrow external margin beyond the pedicle, but
in the rest the latter occurs at the external border, and it is proportionally large
throughout, the scale being thus firmly adherent over an extensive surface while readily
moved by the muscles of the parts. The surface of the cuticle on the dorsum and the
scale-pedicle is minutely papillose, but the scale proper is smooth, only a few minute
papillae occurring along the edge. They are often coated with patches of a blackish-
brown granular deposit. The size of the scales does not always seem to correspond pro-
portionally with that of the specimen. Many of the older authors, like Swammerdam,
considered these organs the gills.

Dorsal Fimbriated Papillse. — On the dorsum of the sixth foot an elevated ridge at the
posterior border, rather beyond the line of the scale-pedicles, gives origin to a short
process with a thin, flat, fimbriated extremity, generally of three divisions. The terminal
fimbriae of the papillae become more complex and the process longer as we proceed
backwards, the organs appearing, after the sixth, on the eighth, tenth, and every
alternate foot (devoid of scales) to the twenty-sixth, when they occur on the twenty-
seventh and the last, a small one on the twenty-ninth, that is on the foot behind the last
scale. These organs have been interpreted as branchiae by Pallas, Savigny, Kinberg,
and others; while many, such as Cuvier, Carus, Duvernoy, and Oersted, held that the
scales were respiratory organs. The great thickness of the cuticle of these processes, and
the absence of large blood-vessels, as De Quatrefages showed, do not favour the view that
they are special branchial structures, though the lobes of the alimentary caeca come close
to them. The cuticle occasionally may be comparatively thick on the surface of
branchiae, as in certain Buphrosynidae, but it does not attain the great density seen in
these processes of the sea-mouse.

Feet.— The structure of the foot is shown in Plate XXXVI, fig. 10 (representing the
tenth foot), and the organ in the various regions of the body conforms to the same pattern.
The dorsal division bears the beautiful iridescent hairs, which gleam with all the beauty
of a permanent rainbow. A dense and most gorgeous tuft of these, thicker than the
dorsal series, occurs just above the ventral division of the foot, and extends more or less
to the dorsal edge of the spines, where another dense tuft of hairs, less brilliant than the
first series and much finer, occurs, and which form the felt on the dorsum.. This property
of felting does not appear to be due to any roughness of the exterior of these fine hairs,
though fracture may render such rough, but to their flexible and attenuated condition.
The tips of the hairs are often curved (see Plate XXXY, fig. 27). The first-mentioned
series are brittle and gorgeously iridescent, the tips under a lens being wavy, a feature

1 Darwin was of opinion that these were homologous with the wings and elytra; of insects, ff and
it is not improbable that with our existing insects, organs which at an ancient period served for
respiration have actually been converted into organs of flight." Unfortunately, proof is deficient.

APHRODITA ACULEATA. 258

due not to any change in outline, but to the play of light on the organs. The rainbow
lustre is lost on drying these hairs in a spirit-preparation, but is again restored on
immersion in water. De Saint-Joseph (1888) thinks Kriikenberg : was wrong in con-
sidering the felt was like chitin ; it is more like keratin.

The lustrous brown spines spring in a fan-shaped series from the middle of the
foot, and form slightly curved sharp weapons of defence. They end in an acute two-
edged tip, and readily pierce the skin. They have the same brittle chitinous structure
as t^he other spines in this species.

In young specimens the dorsal spines are longer, more tapered and curved, and
meet in the middle of the dorsum so as to guard the back, as in other Aphroditidae.
These and the rest of the spines are also often of a lighter golden hue. The thin and
comparatively even Coating of felt which covers the dorsum of some young specimens
gives a character to such examples on clean ground, as off the west sands at St.
Andrews. A great change ensues in these bristles during growth. Small specimens
from a quarter of an inch upward to an inch appear to be common on muddy ground,
and are densely coated with it, — as, for instance, in St. Magnus Bay, between the
Skerries and Fetlar, and the Fjords of Norway (Canon Norman). On sandy ground, on
the other band, they are beautifully clean, and the long golden spines curve as guards
over the dorsum.

The first foot bears a dense tuft of simple, slightly iridescent, slender, hair-like
bristles, chiefly directed upwards and forwards. In the middle is another fan-shaped
series directed forward and slightly curved (convexity outward). Below is a third
group of similar structure, probably representing the ventral series, directed downwards
and forwards, and with their convexity (for they are stiffish) downwards. On the outer
side of the bases of the first two series are the tentacular cirri. This foot shows no
marked division into dorsal and ventral parts, but has a somewhat clavate outline.

The second foot has the ventral division separated. Dorsally the rounded boss has
externally a dense series of stiff hairs, which stand out in a fan-shaped manner, while
towards the dorsal median line (Plate XXXV, fig. 25) the second or adjoining series are
modified to form the very slender hairs of which the felt is composed. The ventral
division bears smooth spines (Plate XXXVI, fig. 18), and inferiorly the tuft of some-
what stiff, pinnate bristles (Plate XXVI, fig. 17).

The ventral division of each typical foot, as noticed by Baster and Swammerdam,
bears three series of bristles arranged in transverse rows (Plate XXXVI, figs. 10 and 22).
The strongest occur dorsally, and consist of about three powerful dark brown bristles
(Plate XXXVI, fig. 2 c), flattened at the tip, which varies in condition, though funda-
mentally the type is somewhat hastate, as observed in the posterior region of the body
(Plate XXXVI, fig. 20). In front view (a) the tip resembles a broad spear, sometimes
asymmetrical ; in lateral view, again, the tip (b) is more or less tapered, and posteriorly
has a slight dilatation at the commencement of the shaft.

The middle series (Plate XXXV, fig. 25 b) has the same type, but the bristles are
paler and more numerous, about eight occurring in the row. The curvature at the tip is
more distinct. Posteriorly the hastate condition^ evident (Plate XXXVI, fig. 19).

1 ' Vergleichende physiol. Stud./ 2e ser., part 1, p. 54, Heidelberg, 1882, 8vo.

254 APHEODITA ACULBATA.

The lowest bristles are still more slender (Plate XXXV, fig. 24 a), and many present
a remnant of the pilose coat at the tip, as in the figure. Posteriorly the hastate
condition is marked (Plate XXX VI, fig. 19).

The second and third feet thus differ from the succeeding in having, instead of the
most ventral series just described, a tuft of somewhat slender pinnate bristles (Plate
XXXVI, fig. 17), the pinnae in the third foot being stronger. These probably represent
the primitive bristles.

The slender feet at the posterior end of the body have ventrally a series of
elongated bristles, with short spikes somewhat alternately arranged distally (Plate
XXXVI, fig. 3). They end in a tapering tip. These bristles are further modifications of
the pinnate forms.

The warts on the feet and near their bases are often coloured dark brown or
blackish, and in many also the general surface of the feet.

A small specimen (1^- inches) procured in Cromarty Frith on August 31st had the
bases of two anterior feet fixed together by an elongated, hard, brownish mass, which
was sunk in the tissues, and another circular patch was near. Only an indefinite
granular structure was visible with the microscope, and the structure was more or less
calcareous, giving off gas on the addition of hydrochloric acid. Both were firmly fixed
in the skin.

Reproductive Organs. — Pallas represented the ova of Aphrodita as originating in the
perivisceral fluid itself, a supposition in the same category as the notion that the glandular
wall of the alimentary canal in the Oligochgetes gives rise to the perivisceral corpuscles.
At Naples Lo Bianco found the males emitting sperm in March. At St. Andrews
specimens have abundant ova in May. The larvse have not been seen.

The direction of the papilla of the segmental organ would apparently send the
reproductive elements dorsally under the felt, and as streams of water are constantly
pouring through this space, the ova would be duly impregnated and aerated. The
sperm would likewise be rapidly distributed all around.

A specimen off Howth, Ireland, showed Loxosomse, and a delicate creeping Cam-
panularian on the ventral surface and between the feet. In one from St, Andrews
Foraminifera were thickly dotted over the ventral surface along with Balani (small),
great numbers of stalked Infusorians, and an occasional very young mussel.

In the felt of the dorsum many small marine organisms are entangled, from alga3 to
mussels. Annelids, crustaceans, sponges, zoophytes, polyzoa, and spines of Echinoderms.
Small preserved specimens are proportionally broader than the adult, and taper much
more rapidly at either end. Such probably is partly due to rigid contraction. They
occur from an inch downward in the stomach of the haddock, and occasionally in the
stomach of the dab.

Delle Chiaje 1 records a nematode as a parasite in the dorsal felt, and as having a
translucent body and a filiform tail. This is like Phoronis in the test of Cerianthus
from Australia. The nematode, however, may only have lodged temporarily.

The species lives fairly in confinement, but as a rule not for a long time. The
difficulty of supplying it with suitable nourishment — for mud in a confined tank soon

1 c Descrizione/ vol. lii, p. 138.

APHRODITA ACULEATA. 255

becomes odoriferous — is probably the cause of its mortality. In its native sites it seems
to make its way in the mud and sandy mud by aid of its powerful ventral bristles,
whilst its back is laved by currents of sea water under the felt which protects this some-
what delicate surface from direct contact with its surroundings. It appears to be a
limivorous form, pursuing its work in the depths of the sea, where its beautifully irides-
cent hairs can be seen by few admirers. It does not always follow that the reasons for
gorgeous apparel or brilliant phosphorescence lend themselves readily to the inquirer.

The smallest example in my collection was procured on the bottom by Dr. Alford
Anderson on board the ' Garland,' on the 9th August, 1888, on the trawling grounds
near the Bell Rock. It measures 3*5 mm. in spirit. The dorsum is covered with sandy
mud in which are a few fine hairs. The ventral surface forms a proportionally broader
area than in the adult. There are nineteen segments besides the pro- and peristomium,
and there are no signs of the lustrous hairs, yet the larger bristles of the feet are
prominent, though few in number. Each foot has superiorly in the ventral division a
long hastate bristle slightly bent downwards at the extremity, the hastate region having
a coating of fine agglutinated hairs which project beyond the tip. One or two shorter
forms of the same type occur in the next row, the larger having a similar though less
developed terminal coating. The third series of two smaller bristles has smooth hastate
tips. The spine has a long free point. The dorsal felt is already formed as a dense
interlacing series of fibres, which entangle minute particles of mud and sand. The
dorsal spines are still comparatively short and pale, and do not project beyond the felt
and mud of the sides.

The soberly tinted young form is thus a contrast to the adult in the colour of the
bristles, spines, and hairs. Its hues coincide with the surrounding sandy mud.

Baster's account (1765) of this species is, on the whole, careful and characteristic, the
arrangement of the ventral bristles and even their number having received attention.
His structural remarks are also interesting, and he found the male and female elements
in June. His figures are fairly accurate.

Rondelet and Swammerdam called the sea-mouse Physalus, while Bartholinus termed
it the golden worm. Seba, Molyneux, and Barrelier, again, named it Eruca sive Scolo-
pendra marina, Swammerdam was of opinion that it deserved a place near the sea-
urchins, probably from the prominence of its spines.

Pallas' s description of the external and internal structure is excellent. He had not,
however, seen one seven inches long, as Baster reports. He describes the muscular bands,
the perivisceral fluid, the digestive system, and the stomach (ventricidus), which the
Belgian fishermen call mentulam Aphroditse, and eat it boiled — a poor kind of nourish-
ment Ouvier afterwards thought. It somewhat resembles the human uterus, he says,
with its os. He is of opinion that the scales are not branchiae, but that fourteen pairs of
sacculate bilobate organs (the dorsal fimbriated papillae) are. He mentions the pinnate
condition of the intestine, figuring the lateral caeca, and describing their attachment to
the " integument." He thinks that Redi's view of the insertion of the caeca in dorsal
sacs will not bear scrutiny. The caeca communicate with the median gut freely, and
chyme enters and is absorbed, — indeed, he saw particles of algae in them, but nothing
in the ventriculus or oesophagus. He corrects Redi's notion that the nerve-cord and its

34

256 APHRODITA ACULEATA.

ganglia formed a systemic trunk with hearts, the reddish colour being characteristic of
the various parts of the nervous system in this species. The true sanguiferous system is
best seen in moribund fresh examples. In the dorsal region is a membranous space (the
great vein) filled with -turbid lymph, and it gives off twigs between the intestinal pinnae,
and long branches anteriorly. He alludes to the anterior extremity of the great vein
over the ventriculus, and figures it. Under the intestine is a longitudinal vessel with
lateral branches. He noticed the perivisceral fluid and its corpuscles, and found ova in
June in masses therein, and also spermatozoa in males, but did not know how they gained
exit. It is the Soe-Muus and Gold-Muus of 0. F. Miiller in his ' Zoologia Danica
Prodromus ' (1776).

In the first volume of his ' Memorie ' (1823) Delle Ohiaje refers to his figures on
Tav. iv, in which the exterior, the structure of the proboscis, and alimentary canal are
sketched. In his fourth volume (1829) he again recurs to the same form, giving a some-
what indifferent view of the body, several sketches indicating the arrangement of the
nervous and digestive systems, and the three series of bristles on the ventral surface of
the foot.

Treviranus l described the external apertures of the segmental organs in Aphrodita.
Delle Chiaje and he regarded them as openings by which water got into the perivisceral
cavity ; that in reality the ciliary current moves the reverse way.

Audouin and M. Edwards considered this the most beautiful and brilliant of all the
Annelids, stating that it is called the sea-mole and sea-mouse, and that it inhabits the
depths off shore, and also the oyster-banks. Earely is it tossed on shore. The
Aphrodita sericea of Savigny they could not find in the museum, while his A. aurata
is probably a young example of A. aculeata.

Cams and Jourdan 2 figure the body of A. aculeata in vertical section, and also the
alimentary canal, but nothing is added to previous knowledge.

In the Memorial Edition of Ouvier it is said that the flexuous bristles of Aphrodita
shine with all the brilliancy of gold, and change into all the tints of the rainbow. They
do not yield in beauty to the plumage of the humming-bird, nor to the most lively lustre
of precious stones. The gills are concealed by the scales, and are in the form of small
fleshy crests.

Sir J. Daly ell 3 (1852) noticed the habit of elevating, or, as he calls it, recurving
the posterior extremity of the body, and " discharging a stream of water from an orifice
there." None of his specimens fed on any substance offered to them. The account of
the segmental organs of this species by Dr. Thomas Williams 4 seems to rest on a misin-
terpretation of the parts. He also states that the blood-vascular system is absent.

De Quatrefages (1865) gives a somewhat detailed account of its external characters.
He records thirty-nine rings and fifteen pairs of scales, and a length of 16—17 centi-
metres (6 or 7 inches). He mentions an elevated cutaneous fold (facial tubercle ?) in the
middle line running to the mouth. The median antenna is implanted on a caruncle. His

1 Tied, and Trevir., < Zeitsch. f. Physiol./ Bd. hi, 2, p. 157, 1829.

2 c Traite elem. dAnat. comp./ Paris, 1835, pi. v, figs. 24, 25.

3 ' Pow. Creator/ vol. ii, p. 171.

4 < Philos. Trans./ 1858, p. 134, pi. viii, figs. 26, 28.

.APHRODITA ACULEATA. 257

Milnesia borealis (Johnst.) is only the young of A. aculeata in a somewhat imperfect con-
dition.

Grube was of opinion that the A. sericea of Savigny, and A. borealis, Johnst., refer to
this species, and that Risso's form, A. aurata, is the young of the same.

Claparede l describes the peritoneal coat of this species as being the most distinct
example in the group, and he figures the fine stride which characterise it. At
intervals on this surface he found groups (mouchets) of vibratile cilia as in Hermione,
and as Sharpey had long previously observed. He alludes to the ovaries which he found
on the ventral surface at the bases of the feet. They were in bands (boyau) in the
median line attached to a cord. He could not satisfy himself as to the nature of this
cord. He further points out the peculiar structure of the papillae, the bases of which can
be traced to the interior of the palpi. He differs from De Quatrefages, who states that the
first two pairs of ganglia are completely separated from their homologues of the other
side. He considers that these ganglia do not exist ; they are simply the inferior part of
the oesophageal connectives.

A good account of the distribution of the blood-vessels of this species is given by
Prof. E. Selenka. 2 He describes in injected specimens a dorsal and a ventral vessel
running above and below the gut, the former being connected with the trunks of com-
munication — which course in a parallel manner round the gut — by an intermediate
series of inosculating vessels. A fine network is found in the dissepiments and the
membranes in connection with the intestinal caeca, as well as certain vessels which
end blindly over the dorsal muscles, the nephridia, and other parts. He describes also
the growth of the ova in dense masses on the vascular trunks at the bases of the feet.
The larger eggs have a fine membranous capsule with nuclei ; the smaller ova are
brownish red.

Grube 3 (1874) says that in large examples of Aphrodita aculeata forty-three seg-
ments occur, and that Savigny observes that the two dorsal tufts of hairs (felt) come from
the elytra-bearing segments, while Audouin and Milne Edwards held that they come
from the cirrus-bearing segments. Grube's examples were in the former condition.

Cams 4 (1884) mentions that it is called Ti veggo (Claparede) at Naples, a name,
however, given to other forms ; and that an Annelid, Branchiomma vigilans, Claparede, is
parasitic on it.

Dr. Hugo Eisig 5 (1887) makes an interesting comparison, with figures, of the
bristles and hairs of this species with the golden yellow secretion of the spinning glands
of Polyodontes maxillosus.

1 'Aim. Chset./ Nap., 1868.

2 ' Niederlandisches Archiv fur Zool./ Bd. i, Heft 2, p. 33, Taf. iii and iv, 1872.

3 Op. cit., 1874.

4 < Fauna Medit./ vol. i, p. 199.

5 ' Monogr. der Capitelliden/ Naples, 1887, p. 331, et seq., Taf. xxxvi.

258 L^ETMATONICB FILICORNTS.

Genus VI. — L^tmatonice, Kinberg, 1854.

Eyes on short peduncles placed near the anterior border of the head, dorsum
covered with felt. Spines of the elytra-bearing feet glochidiate, other segments with
lateral bundles of stout bristles and a tuft of hair-like bristles. Bristles of the
ventral branch semi-pinnate. Intestinal ca3ca strictly dorsal, arising on each side of the
median dorsal vessel. Segmental organs (nephridia) opening externally by a papilla
directed upwards between the feet. Nerve-cords flattened, less distinctly separated than
in Aphrodita.

1. L^tmatonioe filicornis, Kinberg, 1855. Plate XXIV, fig. 9.

Cephalic lobe rounded, two curved longitudinal lines making it tripartite; facial
tubercle with long mammillate papilla).

Elytra obliquely reniform, with minute cells.
Glochidiate spines with three or four teeth.

Synonyms.

1843. Aphrodita hystrix, Oersted. Ann. Danic. Consp., p. 11.

1850. „ „ Sars. JSTyt. Mag. Naturv., 1850, vol. vi, p. 210, n. 57.

1855. Lxtmatonice filicomis, Kinberg. Ofversigt Kongl. vet. Akad., 1855, p. 382.

1857-8. „ „ Kinberg. Freg. Eugen. Eesa. Zool., vol. ii, p. 7, Tab. iii, f. 7.

1859. „ „ Danielssen. Norske vid. Selsk., 4de Bd., 2 Hft., Throndhjem, p. 114.

1861. Aphrodita hystrix, Danielssen. Nyt. Mag. Nat., vol. xi, p. 49.

1865. Lsetmatonice filicomis, Malmgren. Xord. Hafs-Ann., p. 53.

„ „ Kinbergi, Baird. Proc. Linn. Soc, vol. viii, p. 180, 1865.

„ „ „ De Quatrefages. Anneles, p. 199.

1869. „ filicomis, Mcintosh. Trans. Roy. Soc. Edin., vol. xxv, p. 407.

1873. „ „ Sars. Bid. Christ. Fauna, p. 1.

1875. „ Kinbergi, Ehlers. Ann. 'Porcnp.,' op. cit., p. 31, Taf. ii, f. 1, 2.

1879. „ filicomis, Tanber. Ann. Danic, p. 79.

1885. „ „ Mcintosh. c Challenger J Rept., p. 39.

1887. „ Kinbergi, Ehlers. Annel. U. S. S. ' Blake/ p. 45, pi. vii, f. 6 ; pi. viii, f. 1—5.

1894. „ „ Flor. Buchanan. Proc. Roy. Dubl. Soc, vol. viii (n. s.), ii, p. 170.

1896. „ filicomis, Roule. Camp. d. c Candan ' (Ann. Univ. d. Lyon), p. 442.

Habitat. — This species was first dredged in British seas by Dr. Gwyn Jeffreys off
the Hebrides in 1866, and often subsequently in various grounds off Shetland in depths
ranging from 75 to 100 fathoms, and chiefly on muddy sand. The same persevering
explorer found it off the west coast of Ireland (Yalencia Island). It extends into the

L^ETMATONIOE FILICORNIS. 259

Atlantic on Holtenia-ground or stones and coral, and even to the depth of 1950 fathoms
('Knight Errant'), and is found on the shores of America (Verrill) and Canada
(Whiteaves). Northward it is met with in the Faroe Channel, while Grube gives from
Bohuslan along the Norwegian coast to Finmark. A few examples were dredged along
with Hermione on the oyster-ground off St. Peter Port, Guernsey, but it is absent along
the eastern shores from the Pentland Frith to the south coast.

Head. — The head (Plate XXVII, fig. 2) is somewhat rounded, the median separated
from the lateral region by two curved longitudinal grooves. The tentacle arises between
and rather behind the ocular peduncles above the conical basal segment, and is a
long, slender process gently tapering towards the tip, then slightly enlarging, forming a
constriction, and lastly a bulbous swelling as shown in the figures. In none of the
examples, however, did the tentacle reach the length of the great palpi, as Kinberg says,
nor could Baird's description as "short and conical" apply. Its length, probably from
injury, varies, but the longest are far short of the palpi, so that Kinberg' s were either
recently reproduced or varied in this respect. It agrees in form with a dorsal cirrus.
The somewhat globular ocular peduncles occur on each side of the former, and in the
preparations are devoid of pigment. The long palpi spring from each side of the facial
tubercle in front, and have a smooth basal articulation, from which they gently taper till
near the extremity, when a somewhat elongated dilatation occurs, after which the tip
tapers to a fine point. The whole organ beyond the basal articulation is setose with
long, sharp papilhe, which are finer on the distal region (Plate XXXVII, fig. 2). The
slight enlargement below the tip is not evident in every example, probably from injury,
but it. seems to be present in the most perfect.

The facial tubercle has numerous long, mammillate papilla?, thus differing much
from Aphrodita aculeata (Plate XXXVI, fig. 24).

The body is somewhat flattened, elongate ovoid, about equally tapered at both ends,
though from the contraction of the parts and the condition of the feet the posterior end
often has a slightly broader aspect. The dorsum is covered with a greyish felt, and the
sides are flanked by the proportionally large and iridescent brownish bristles. These do
not overlap the dorsum as in other species, only a few of the shorter bristles spreading
slightly over the outer edge of the felt, 1 which is usually coated with sand, and is so
flexible as to take wrinkles from the edges of the scales. The dorsal felt is formed of a
moderate number of very fine hairs tapering to an attenuate point, and connected
together by a fibrillar gelatinous basis, which has sand-grains, diatoms, and other algae,
mud, and foreign particles of various kinds, e.g. sponge-spicules and anchors of
Synaptge, in it. So far as observed the fine hairs do not terminate in hooked points as in
A. aculeata; they spring from the inner edge of the dorsal division of the foot. The
felt is friable, thicker, however, in the larger and older specimens, and more easily torn
and gelatinous in the smaller. The delicate dorsal cuticle under the felt is smooth, and
the body-wall of this region is much thinner than in Hermione. The long bronzed
spines extend posteriorly somewhat beyond the line of the feet, terminate nearly in a
transverse line with the body, and give a truncated outline to the region in contrast

1 Gruhe thought only mucus was present.

260 L^TMATONICE FILICORJSIIS.

with the anterior end. Kinberg's figure differs, therefore, considerably from British and
other examples— both small and large — that have come under observation; indeed, it is
more or less diagrammatic.

The entire ventral surface, including the feet, is studded with closely set globular
papillas.

The papilla of the segmental organ begins on the seventh foot as a small process,
becomes a free tubular structure on the ninth, and extends to the twenty-ninth foot. The
papilla is comparatively large and long.

Digestive System. — The extruded proboscis has at its extremity a densely villous
border, divided into two by a bare papilla at each side. The long processes (papillae)
are divided dichotomously (Plate XXXVII, fig. 2) in a very regular manner, about five
times from the base. The great proportional size of this organ — for it is more than half
the length of the preserved form — is a noteworthy feature, and indicates its importance
in the economy of the animal.

The pre-gastric cgeca have their terminal appendages even more conspicuous than
in A. aculeata. The intestinal casca arise from the dorsal aspect of the gut close to the
median vessel. They thus slightly differ from those of Aphrodita and Hermione. In
regard to the arrangement of the terminal cseca, they approach those of the former.

In the Zetlandic examples masses of mud and mucus in the stomach were very rich
in Foraminifera, sponge-spicules, radiolarians, diatoms, bristles of Annelids (Spionidas),
and fragments of crustaceans.

Scales. — Fifteen pairs, smooth, somewhat pellucid, and slightly iridescent. They
are more or less rounded or ovoid, and with the exception of the first two are
attached by the centre of the outer border as in Aphrodita and Hermione. While the
first pair are small, they rapidly increase in size, the middle third of the body having
only three or four large scales. They occur on segments 2, 4, 5, 7, and so on to 23, 25,
28, and 31.

Feet. — The long, brownish, iridescent spines which flank the sides are flattened and
flexible, with the tip somewhat abruptly tapered in lateral view (Plate XXXVI, fig. 8),
whereas when seen on the flattened face the sides have four or five recurved teeth (Plate
XXXVI, figs. 4 and 6). The majority have three teeth on each side (Plate XXXVI, fig.
12), but some have four. The barbed bristles have guards or sheaths, or traces of these
in the preparations (as in the figure), but whether they are only perfect in the young con-
dition of the bristle is unknown. They seem to be removed readily. 1 The main part of
the shaft is flat, broad, longitudinally striated, and it somewhat diminishes inferiorly, and
terminates in a flattened, blunt extremity, to which the muscles are attached. For some
distance above the base the shaft is paler, and shows a central band as if from an axial
cavity. The spines present a distinct curvature. About twelve well-marked groups of
the long spines occur in a good example. These spines retain the same essential
structure posteriorly.

The first pair of feet are directed forward, and bear the tentacular cirri (which
are much longer than in Aphrodita aculeata). Each has two tufts of pale, simple bristles,

1 Perfect examples occasionally adhere to the felt.

^

.L2ETMAT0NICE FILICORNIS. 261

which taper to a fine hair-like point. Their outline seems to be smooth, though sandy
debris lodges readily amongst them. The anterior tuft is directed forwards and inwards
as an expanded fan, while the compact posterior one goes inwards and backwards. A
strong spine supports the foot. The bristles are similar but somewhat stronger dorsally
in the second foot. In the third foot a more slender series of bristles occurs to the
exterior of the cirrus, while a stronger series is dorsal. This and the next have
pinnate bristles in the ventral division.

The fourth foot, which is provided with a ventral cirrus, has in the inferior division
a tuft of pale pinnate bristles, like those of Aphrodita. The same pinnate bristles occur
in the fifth foot, adjoining the ventral cirrus, while above them are a few of the ordinary
kind with feathered tip and hook. In the sixth only the latter are present. The typical
forms are shown in Plate XXXVI, figs. 5 and 7.

In the same way the last foot or two have a modification of the ventral bristles, for
the spikes above the hook are short and distinctly separate. Moreover the last foot has
pinnate bristles as in front, along with an ordinary form or two.

The feet appear to be about thirty-four, the dorsal division alternately bearing cirri
and long bristles.

The dorsal cirri spring from the centre of a fringe of the more slender bristles, their
places being taken in the scale-bearing segments by the long hooked spines. In the
case of the cirrigerous feet a group of stout, short, brownish bristles occurs in front of the
cirrus, and a somewhat longer and stronger series springs in the scale-bearing feet on
the dorsal side of the long spines. All are densely covered with a muddy and sandy
investment, but their surface appears to be smooth.

A typical foot, e. g. one bearing the long dorsal spines, has dorsally (from within
outward) the slender hairs which form the felt, then a group of short stiff bristles, fol-
lowed by the papilla carrying the long brown spines, which have a distinct curve. A
tuft of delicate bristles occurs beneath. The scale is fixed posteriorly. After an
interval the ventral branch of the foot bears the semi-pinnate bristles and the ventral
cirrus.

The last pair of feet are rudimentary, and lie in ordinary positions behind the anus
which is dorsal.

The segmental organs (nephridia) point upward between the feet, and are as well
developed as in Aphrodita aculeata.

A Norwegian example, sent by Dr. Merle Norman, 1 shows guards to the front of the
dorsal spines, and a slightly hastate tip beyond the barbs. The ventral bristles seem to
have shorter tips.

A series of small specimens occur from North Unst — from 55 mm. upwards. These
have pale spines and bristles, yet all the bristles and the dorsum are densely coated with
sand-grains. They were captured in July, and probably represent the growth since the
previous spawning season.

Loxosoma3 are common in this species on the feet, ventral surface, and on the
dorsum under the scales.

1 Op. cit., p. 171.

262 L^STMATONIOE PRODUCTA.

Reproduction. — Ova of considerable size are present in examples dredged by Dr.
Gwyn Jeffreys off North Unst in June, 1867.

Kinberg (1857) characterised this species as having a rounded cephalic lobe, tri-
partite from two curved sulci, filiform tentacle longer than the palpi, and the median
elytra reniform. It was procured on the western and northern shores of Scandinavia.

Ehlers (1875) considered the Lsetmatonice Kinbergi of Baird as different from
Kinberg's L. filiformis, but, as previously stated, there is no reason to doubt their
identity.

Ehlers (1887), in his 'Annelids of the United States Expeditions/ considers, as he did
in the former publication (' Annel. of the 6 Porcupine ' ') that he is still dealing with a
new species fide Baird. Only a single form appears to occur in British seas, and it extends
to Norway and the American coast. His figures are the work of an artist, and thus the
scientific accuracy is not beyond doubt, as a glance at the dorsal spine of his pi. viii,
fig. 3, demonstrates. The peculiar flattening of the main part of the shaft, and the
characteristic narrowing of the tip, have been overlooked. This form, however, may be a
variety of L. producta, allied to L. prod., var. benthaliana. His description gives nothing
diagnostic.

2. LiETMATONiCE pboducta,- var. Britannica.

Specific Characters. — Body ovate-oblong, of a pale flesh-tint, and devoid of dorsal
felt. Scales pale, their reticulate cordate structure being better marked than in any
other form. Segments forty-five. Head triangular ; ocular peduncles globular at the
tip, narrowed inferiorly at the peduncle; no eyes. Lateral process at the posterior
part of the head smaller than in the typical forms. Papillse of the angular facial tubercle
more lobate than in L. producta. Grlochidiate bristles with four recurved fangs, besides
the process at the base of the terminal spear- tip. Ventral bristles with stiff er basal and
more slender distal pinnse.

Synonym.

1894. Lsetmatonice producta, Flor. Buchanan. Proc. Eoy. Dubl. Soc, vol. viii (n. s.), ii, p. 169.

Body more or less oblong, of a pale flesh-tint, devoid of dorsal felt, scales pale ; forty-
four bristled segments, which increase to the middle or behind the middle of the body,
and again diminish posteriorly, though the latter region is somewhat blunt. The centre
of the head is more or less triangular, with a prominent swelling, and bearing in front
the two ocular peduncles, which are globular at the tip, but have a distinct peduncular
portion, thus differing from those of var. Willemoesii, which are more or less sessile, and
without a trace of eyes. Those of L. producta are more elongated. The lateral process
towards the posterior part of the head is much smaller than in L. producta from Kerguelen.
It forms a lobate flattened process in the latter, and nearly reaches the base of the
ocular peduncles. On the other hand, in the British specimens it is only a small papilla,

L/ETMATONICE PRODUCTA. 263

touching the base of the lateral cephalic swelling. It appears to be the rudiment of a
scale-bearing process. The ridge from which the tentacle arises extends backwards in
the middle line, and at the posterior border of the head it bends outward on each side
to bear the papilla, viz., the modified scale-bearing process. The papilla? of the facial
tubercle are more lobate than in L. producta from Kerguelen, and proportionally less
elongate. The palpi are, so far as the specimens go, less powerful than in L. producta
from Kerguelen, but the difference in regard to the papilla? or spikes is small. Both
have numerous papillae as in L. filicomis. All that can be said is that perhaps they are a
little more numerous towards the tip of the palpi of L. producta from Kerguelen. The
tentacular cirri are somewhat longer than the tentacle, and their tips are rather more
elongate than in L. filicomis, while they are shorter than in L. producta from Kerguelen.

The segmental papilla? begin on the posterior face of the fifth foot, and extend to the
thirty-eighth. They form flattened processes somewhat more distinct than in L. producta
from Kerguelen. Besides, in the latter form they begin on the tenth foot, and extend to
the thirty-ninth.

The papilla? of the proboscis (Plate XXXVII, fig. 3) have broader tips than those of
L. filicomis, and are less regularly dichotomous. The tips are sometimes divided into
three, and occasionally are ovate, clavate, or irregularly lobate. There is not much
ground for calling the thickened regions of this organ parts representing maxilla?, as
Grube does.

No specimen is in a condition to give a reliable opinion of the arrangement of the
dorsal spines. The papilla? bearing them are a little more prominent than in L. producta
from Kerguelen, but this may be due to their condition, for they have evidently been
subjected to considerable hardships. They probably spread, in the scale-bearing feet,
over the dorsum as in the forms from Kerguelen, the feet carrying cirri having dense
groups of more slender bristles.

The structure of the first two feet agrees with that in the form from Kerguelen.
The other feet are also similar in structure. The tips of such of the long dorsal spines
as were present agreed with the figure (Plate XXXVI, fig. 4), which shows a spine
slightly turned to the left, four recurved fangs occurring on each side, besides the point
at the base of the spear-like tip. The ventral bristles (Plate XXXVI, fig. 5) conform
mostly to the type of L. producta, var. Wyvillei, having somewhat stiffer (stouter) basal
bristles and numerous slender distal ones.

In describing this form in the Annelids of the ' Challenger,' I for some time had the
varieties ranged under separate species, distinguished by certain evident characters.

The reticulate cordate structure of the scales is very well marked (apparently better
than in any other species), forming a series of wavy lines like those on the sand of the
sea-shore.

No parasitic Loxosoma? occur on this form, yet foreign varieties abound with them,
such as L. producta, var. benthaliana, on which the remarkable new gymnoblastic
hydroid, Stylactis vermicola, described by Prof. Allman, occurs on the under surface of
the scales of this inhabitant of the depths (2900 fathoms) of the North Pacific.

The species was first procured by Prof. Haddon during the Royal Dublin Society's
Survey, on a bottom of sand and gravel at a depth of 500 fathoms, fifty-four miles off

35

264 HERMIONE HYSTRIX.

Achill Head, and recognised by Miss Florence Buchanan, 1 who thought it had hitherto
been procured only at Kerguelen ; but it and its varieties have a much more extensive
range, even to the shores of Nova Scotia, near Halifax. The varieties described in the
( Challenger ' were devoid of eyes in the preparations, as this also is.

Genus VII. — Hermione, Blainville, 1828.

Cephalic lobe with a single tentacle and two palpi. Peristomium bears bristles and
two long cirri. Pedunculated eyes fixed to the margin of the cephalic lobe ; maxilke
absent or small (thickened processes). No dorsal felt. Elytra alternating with dorsal
cirri. Dorsal and ventral divisions of the feet distinct. Elytra-bearing feet with
glochidiate dorsal spines, and a tuft of strong bristles over the elytra. Ventral bristles
bidentate.

The pregastric and intestinal casca are shorter than in Aphrodita, and even than in
Ldetmatonice, but they have large terminal dilatations. The latter are shorter than in
A. aculeata. The lateral casca do not approach the median line as in Lseimatonice.
Segmental papillse from the eighth to the twenty- ninth foot. Nerve-cords closely applied
and flattened.

Heemione hystrix, Savigny, 1820. Plate XXIV, fig, 7.

Specific Characters. — Head broad; each ocular peduncle bearing two eyes; facial
tubercle with globular or slightly ovate papillae. Body elongate-ovoid. Segments
thirty-two to thirty-three. Elytra somewhat reniform, furrowed by divergent and
transverse lines, and having large rounded cells. Dorsal spines glochidiate, with three
fangs on each side ; ventral bristles with a spur, and, in the anterior part of the body,
accessory spikes above it. The tip of the bristle is curved.

Synonyms.

1818. Halithea hysbrix, Lamarck. Hist, des An., s. v., vol. v, p. 307.
1820. „ „ Savigny. System, des Ann., Descr. de PEgypte, t. xxi, p. 345.

1823. „ Delle Chiaje. Memorie, vol. i, p. 181, tav. iv, f. 10, 11.

1826. „ hystrix, Eisso. I/Europ. Merid., p. 413.

1828. Hermione hystrix, De Blainville. Diet, des Sc. Nat., ' vers/ p. 457, pi. ix, f. 2.

1834. Aphrodita hystrix, Aud. and Ed. Ann. des Sc. Nat., vol. xxvii, p. 406, tab. vii, f. 1 — 9 ; and

Annelides, p. 70, pi. i, f. 1 — 9.
„ „ „ Cnvier. Eegn. An., vol. iii, p. 207.

1 c Sc. Proceed. Eoy. Dubl. Soc./ vol. viii, pt. 2, No. 15, p. 169, 1893.

HERMIONE HYSTRIX.

265

1836-
1840.

)>
1842.
1843.
1851.

1856.

1857.

))
1858.

}?
1861.
1864.

1865.

868
875
884

888
890

. Eermione hystricella, De Quatref. Reg. an. illust., pi. xix, f. L

„ hystrix, Grube. Actin., Echin., u. Wiirmer, p. 88.

Aphrodita hystrix, Johnston. Ann. Nat. Hist., iv, p. 370, v, f. a — e ; and v, p. 305.
Halithea hystrix, Delle Chiaje. Descrizione, pp. 57 et 105, tav. lviii, f. 10.
Aphrodita hystrix, Oersted. Ann. Danic. Consp., p. 11.

„ ,, Grrube. Fam. der Annel., p. 36.

„ „ Thompson. Fauna of Ireland (partim ?), p. 273.

,, mediterranean, O. Gr. Costa. Fauna d. Reg. d. Napoli, Ann el., p. 8, tav. ii, f . 1 — 1 e.

Hermione hystrix, O. Gr. Costa. Annel. di Napol., p. 5, Tav. i, f. 11 — 14, and Tav. ii, f. 1.

,, „ Kinberg. Fregatt. Eugen. Resa, p. 4, tab. 2, f. 4.

„ hystricella, Kinberg. Ibid., p. 5, tab. 2, f . 1.

Aphrodita hystrix, Danielssen. Nyt. Mag. f. Naturvid, Bd. xi, p. 49.

„ „ Grrube. Die Insel Lussin, &c, p. 77.

Hermione hystrix, De Quatref. Hist. nat. d. Annel., vol. i, p. 206, pi. vi, f . 9 — 14.

„ Kinbergi (?). Ibid., p. 209, pi. vi, fig. 16.

„ hystrix, Baird. Proceed. Linn. Soc, vol. viii, p. 178.

„ „ Johnst. Cat. Brit. Ann., p. 106, pi. ii, f. a — e (a repetition of former).

„ „ Claparede. Annel. Chetop. d. Nap., p. 48, pi. i, f. 2.

„ „ Marion and Bobretzky. Ann. Sc. Nat., 1875, p. 3.

„ „ V. Cams. Faun. Med., p. 199.

,, „ Harvey Gribson. Vermes Liverp., p. 147.

„ „ De Saint- Joseph. Ann. d. Sc. nat. (7), vol. v, p. 146.

,, „ Malaquin. Ann. Boulon., p. 14.

Habitat. — A southern form, 1 abundant at a depth of fifteen to twenty fathoms off
St. Peter Port, Guernsey, amongst debris of shell-gravel, dead oyster- and mussel-shells,
and occasionally at various parts of the southern coast of England and Ireland. It is
common in the Mediterranean. The German exploring ship ' Gazelle ' procured it at
Soleton Bank. It ranges to eighty fathoms.

On the whole it is partial to shell-debris, gravel, and similar regions, and thus fre-
quents rougher ground thanyi. aculeata (Horn ell).

The smooth head (Plate XXIV, fig. 8) is rounded, and the posterior fillet which
bounds it is considerably overlapped in the preparations by the peduncles of the first
scales. This fillet appears to bear no papilla or homologue of the scale-peduncle exter-
nally. The median ridge which runs forward to and ends in the base of the tentacle is
slightly marked, The ovoid lateral swellings are more prominent than in Laetmatonice
filicornis. The ocular peduncles are somewhat clavate, with rounded tips, and each has
dorsally a well-marked black eye, and just in front of it a larger one looking forward and
downward. The facial tubercle has numerous globular or slightly ovate papillae — a few
being longer (elongate-ovate).

All except the basal region of the palpus is " ciliated " with spike-like papillae as in
the foregoing species, this being a character apparently subject to little variation. The
tentacle, the base of which possesses scattered globular papillae, has a peculiarly crenate
outline, slightly enlarged at the tip, and with the clavate terminal process (Plate XXXVII,

1 The notion that it occurs " all over the British area w (Dr. Benham) does not accord with our
experience. It is a southern form.

266 HERMIOXE HYSTKIX.

fig. 5). The tentacular cirri are similar. The dorsal cirri increase in size, but have the
same structure. In all these, minute rounded glands (like papillas) are dotted over the
granular layer of the epiderm, and may be associated with sensation. They are quite
beneath the cuticle, which is somewhat dense. At the tip or bulbous part of the
appendix the cuticle is thin, so that the nervous expansion comes close to the surface.

Body. — About two inches long, large specimens being two and a quarter inches (De
Quatrefages), and is covered dorsally by the fifteen pairs of scales, which largely overlap.
They vary in tint from pale to dark brown, with a slightly iridescent purplish sheen,
and are firmly fixed by their pedicles near the middle of the external border. Under a
moderate power they present a fibrillar or cordate structure, and the exposed surface has
a series of minute papilla, the anterior region alone being devoid of them. The edge is
smooth. The first pair are small, and have the scar for the attachment of the pedicle
near the centre. They again diminish posteriorly, but the structure remains the same.
The Mediterranean forms are generally darker than those from the Channel Islands.

The ventral surface of the body is covered with a firm cuticle studded with minute
papillse, which are slightly brownish, especially on the feet at the commencement of the
posterior third. Anteriorly the buccal fold forms a broad, rounded, finely grooved flap,
diminishing to a nearly cylindrical ridge posteriorly. The minute papillge are numerous
on the edges of the buccal fold. A median and two lateral ridges flanked by the feet
characterise this surface, as in allied members of the group.

The segmental papillae commence on the eighth foot, and continue to the twenty-
ninth, that is, about the fourth foot from the tip.

The papillse of the proboscis are, like the others, somewhat dichotomous in their
division (Plate XXXVII, fig. 4), the tips of many of the processes being broadly and
more or less symmetrically lobate, or having a process at one side so as to be pedate.
They seem to be flattened, and thus in the preparations do not readily separate from each
other. They differ much in outline from the same processes in Lmtmatonice filicornis.

The dorsum is flanked by the lustrous brown spines (Plate XXXVI, fig. 11), which
are directed backwards and slightly outwards. In some views they are brownish, while
in others they have a rich golden sheen. Anteriorly the granular spines stand upwards
with the curve directed towards the middle line, the inner bristles spreading over the
dorsum. The lateral brush, again, on the cirrus-bearing foot is directed outwards and
with a backward curve. As soon as the long spines become prominent the pedicle for
the scales causes a radiate arrangement of the stiff tapering bristles (Plate XXXVI,
fig. 9), which spread over the dorsum and meet those of the opposite side in a symmetrical
and graceful manner. The outer bristles of this series are more erect; the inner are
adpressed, so as to guard the scales. These bristles are of a rich golden colour, with
pale tips.

Three recurved fangs, as a rule, occur on each side of the tip of the spines (Plate
XXXVI, figs. 12, 14, and 15), besides the smaller pair at the angle of the spear-tip.
They are more nearly opposite (though the last two are not so) than in Lxtmatonice
filicornis.

Feet. — The first foot has pale, simple bristles, the tufts being directed forwards and
inwards, and the basal portions of the tentacular cirri are warty like the tentacle.

HEEMIONB HYSTRIX. 267

The second foot has dorsally a tuft of pinnately spinous bristles as in the other
members of the group, for instance, A. aculeata, — little variation occurring in the species.
In the ventral branch are two bristles (Plate XXXVII, figs. 6 and 7), an upper with
a strongly spinose edge above the powerful tooth, and a lower with indications of four
spines, viz. the usual powerful inferior one, the somewhat less spine above it, and
indications of two others beyond, the last being slightly developed. The foot has
numerous globular papillae on its surface.

The third foot, which carries a cirrus, has dorsally and internally a series of
powerful bristles minutely nodular (Plate XXXVI, fig. 9), and with a well-marked curve,
while externally a group of smaller forms of a similar kind occurs. This nodular
development on the surface is peculiar, and forms a swelling below the tip as
if it were a secondary development. The prominent granules or minute nodules
are not affected by potash. The ventral division has often stout bristles with the
spinose tips above the powerful fang, as shown in Plate XXXVII, figs. 6 and 7 before
mentioned; but in some the spines above the fang are only four in number, while
in others they are eleven. At the ventral edge of this division of the foot are
numerous pinnately spinous bristles, as indicated in the dorsal division of the second
foot.

In the fourth (a scale-bearing foot) the dorsal bristles are still curved, and occa-
sionally minutely nodular, but much longer than in the third, and the tips of some are
shaped as in the hooked forms. A few shorter and straighter spines occur externally.
Ventrally the powerful bristles still show spinose tips, but the spines above the inferior
fang are diminishing in number, one having only two.

The fifth (likewise a scale-bearing foot) has dorsally the somewhat broad, granular
bristles, the tips of the amber-coloured ventral still having spines, the number of the
latter above the great fang from above downwards in the three bristles present in the
example being two, three, five.

The sixth foot (cirrus-bearing) has dorsally short, curved, and slightly granular
flattened bristles dorsally and internally ; externally a shorter and more slender form of
similar character. Ventrally three bristles occur, with two, two, and three spines above
the great fang.

The seventh foot has numerous curved, granular bristles dorsally, besides the spines,
some with greatly elongated tips, which, moreover, are somewhat hastate. On the
ventral bristles only two spines in all are present above the fang.

In the eighth foot (cirrus-bearing) the dorsal bristles are curved and granular, with
a tuft of longer and more slender bristles to the exterior (below the cirrus). The three
ventral bristles had each two spines above the fang. The segmental (nephridial) papilla
begins on the posterior border of this foot, completely hidden in the fissure between it
and the next foot.

Dorsally in the ninth foot were numerous curved granular bristles with somewhat
blunt tips. Beyond these stretch the long characteristic brown spines with the glochidi-
ate tips. Ventrally the three bristles have from above downwards one spine above the
fang, two and one. Their shafts are strongly grooved.

The tenth (a cirrus-bearing foot) has dorsally the shorter curved granular spines,

268 HERMIONE HYSTPIX.

and the shorter and more slender bristles externally. Ventrally are three bristles having
above the fang respectively two, one, two spines.

The thirteenth foot has dorsally scales and spines. Ventrally three bristles, the
hooks above the fang being respectively one, two, one.

The fifteenth has scales and spines. Ventrally the two bristles have one spike
each above fang.

The twentieth foot (cirrus-bearing) has longer and more tapered dorsal bristles, the
granules on which are less distinct, especially in the tapered forms. The shorter, broader
forms show the granules distinctly. Ventrally the three powerful bristles have only the
great fang. A typical foot is shown in Plate XXXVI, fig. 13.

The terminal feet have dorsally the granular bristles more pointed and less curved
than in front, moreover the tips are smooth ; while ventrally the tips are more elongated,
and have numerous spines above the great fang. One of the smaller forms is X 90 in Plate
XXXVII, fig. 8. The ventral bristles of this form are much stouter than in Laetmatonice
filicomis.

This species is specially subject to the attacks of Loxosoma, which occurs abundantly
on the ventral surface and cirri, on the feet, mouth, and other parts. Parasitic algas
often grow on the bristles and spines, with mud, sponge-spicules, and other debris
entangled. A remarkable hydroid creeping along the bristles is also present. Hydroids
and Polyzoa, indeed, are not uncommon.

In habit this species appears to correspond with Aphrodita and Lsetmatonice.

Reproduction. — The ova were well advanced in July in those from Herm and other
parts of the Channel Islands. Lo Bianco found the Neapolitan examples with developing
ova in October, and increasing in size in November.

An interesting account of the development is given by Dr. P. von Drasche, 1 of
Vienna, from observations made at the rich Marine Station of Trieste. He found the
ripe forms in October and November, the latter month being probably that in which
deposition of the eggs generally takes place. The egg measures 0*2 mm., and he watched
it through all the stages of segmentation till the embryo moved within the capsule by aid
of cilia. It then issued as a pear-shaped trochosphere, with an equatorial circlet of cilia
and a tuft in front; while a solid mass of deutoplasm occurred in the middle of the body.
The violet-coloured larva soon acquires another ring of shorter cilia behind the first or
pre-oral, an anal ring of cilia appears, and it swims about actively. Two red eyes are
developed in front. The anterior end becomes triangular, and a characteristic broad
cellular collar lies behind the eyes — the mouth, with a tuft of cilia, opening towards (in
front of) its posterior border ventrally. On the fourth day the larva has increased in
size, and the yolk-mass has concentrated into a covering for the alimentary canal. On the
under surface of the snout are five small papillse — organs, however, not homologous with
the cephalic processes of the adult. There are now three segments with bristles. On
the sixth day it is still larger. The alimentary canal is complete. The equatorial belt is
smaller. There are five segments. The feet show a large yellow spine and bristles, the
latter having a structure conformable to the adult type, though the glochidiate forms

1 'Beitr. z. Entwickelung d. Polychaeten/ pp. 7—11, Taf. ii, f. 8—20; and Taf. iii, f. 1—4,

1885.

HERMIONE HYSTRIX. 269

are not yet present. Besides the structures mentioned, the young Hermione has now
three pairs of eyes, and a process (first foot) behind the collar.

Savigny (1820) first distinguished the species, though his description is by no
means diagnostic.

Delle Ohiaje (1823) refers to the condition of the alimentary system in speaking
of the anatomy of Pleurophyllidia, and gives a figure of a moribund example in his
' Memorie.' He thought the papillae of the proboscis were taste-organs. He shows the
globules of the blood. In his ' Descrizione ' he mentions its common occurrence at
Naples, and reproduces the plate from the ' Memorie.'

The Aphrodita hystrix mentioned by Oersted 1 probably refers to Lsetmatonice.

Kinberg (1858) followed De Quatrefages in making two species, viz. H. hystrix and
H. hystricella ; but so far as can be ascertained this distinction rests upon variation, and
Claparede was of the same opinion.

De Quatrefages (Anneles) saw in a male Hermione hystrix the spermatozoa issue in
the form of a white thread at the base of the inferior branch of the foot of the nineteenth
segment. They probably escaped from the segmental papilla between the feet. This
author distinguishes between his H. fallax and this species by the fact that the median
antenna in H. hystrix is least, and that the bristles of the scale-bearing feet have the
points unarmed, while those of the inferior feet are tridentate (instead of the tips
straight and the inferior bristles bidentate as in H. fallax). The bristles of the ventral
division have curved apices, whereas in H. hystricella from the Mediterranean, &c, these
are straight, and Kinberg says so also.

Claparede 2 has already pointed out that De Quatrefages was wrong in thinking that
absolute reliance was to be placed on the number of the denticles at the tip of the spines
or of the ventral bristles, and he also showed the condition generally of the anterior feet
in contrast with the posterior. He enters into the uses of the guard for the tips of the
long dorsal spines, and demonstrates that a process also appears in the developing ven-
tral bristles before they pierce the surface. He alludes to the granular bristles, which
had previously been described by Johnston and Kinberg, and points out the nature of
their tips, with the swollen part beneath the point, as shown in one of the present figures.
He mentions having found in a single example a long, simple bristle in the ventral division
of the foot, probably a pathological phenomenon. He speaks of the warts on the
surface, already mentioned by Pallas, Johnston, and Kolliker. He likewise alludes to
the structure of the peritoneum, and that of the dorsal cirri.

Grube (1874) says De Quatrefages relegated Hermione hystrix to the Mediterranean
and H. fallax to the Atlantic.

Prof. Jourdain, 3 in his account of the histology of the integument and sensitive
appendages of this species, describes the irregular polygons formed by the cells of the
epithelial coat, the nerves which ramify in it, and the thinly distributed nerve-cells. On
the ventral surface, the warts, of which Claparede had formerly given a figure, have a
central pore, through which communication with the granular layer of the epidermis

1 ' Annulat. Danic. Consp./ p. 11, 1843.

2 < Ann. d. Golfe d. Nap./ p. 50, 1868.

3 'Archives Zool. Experiment/ (2), vol. v, p. 91 (1887), pis. hi and iv.

270

POLYNOID^l

occurs, and by which the nerve-supply to the cells and protoplasmic contents of the
sensitive organs takes place. He also notes that the muscular fibres show no strias,
either transverse or longitudinal, and that the elytras, after Haswell's description, have a
double cuticle superiorly and inferiorly, two layers of cells, and an intermediate fibrous
layer. They contain no cavity, but have a nervous plexus. They are evidently organs
of considerable sensibility. Amongst other interesting points he refers to the nerve-
supply of the palpi and dorsal cirri with their ganglia near the base of the terminal
division, the nerve-trunk in each breaking up into a tuft of cells in this region.

Family III. — Polynoid^.

Body more or less elongate; no facial tubercle, convex cephalic lobe; the base of
the tentacle arising from the middle anteriorly ; two lateral tentacles ; four eyes ; palpi
elongate. Peristomium, bearing the first foot, with long dorsal and ventral cirri,
and the ventral cirrus of the next segment long. Pharynx exsertile, muscular,

Fig. 19.

Alimentary apparatus of Harmothoe imbricata. — A. W.

cylindrical, with papillae round the margin ; horny jaws. Intestinal casca shorter than
in the Aphroditidde, slightly branched; first foot bearing only a few minute bristles

POLYNOULE.

271

conforming to the dorsal type. Scales twelve to thirty-five pairs or more ; segments
carrying these devoid of cirri. Segmental organs (nephridia) opening ventrally on a
papilla near the bases of the feet; nerve-cords within the granular layer of the epiderm
and between the oblique muscles. Dorsal bristles with more or less tapered simple tips ;
ventral bristles with simple or bifid hooked tips. Development by trochophores. The
general arrangement of the body-wall in this family may best be understood by con-
sulting the accompanying figure.

Fig. 20.

cli/- ^^ „ dm.

i^mj.so.

Section of body-wall of Polynoe scolopendrina, Sav., in the line of a median dorsal papilla (a), about the
middle of the body, b, bases of dorsal bristles; b' , bases of ventral bristles; c, ova scattered in the
various parts of the perivisceral (ccelomic) space (pv) ; d, intestine; d', portions of intestinal cseca; dm,
dorsal longitudinal muscles; Im, band of longitudinal muscular fibres above the nerve-area; om, oblique
and nearly vertical muscles; vm, ventral longitudinal muscles; nc, nerve-cords; dv, dorsal vessel with
mesenteries at side; v, ventral vessel with mesenteries; vc, ventral cirrus; so, segmental organs
(nephridia). The dorsal cirrus is not shown, nor the perivisceral corpuscles. The drawing was made
by Dr. Masterman from a somewhat contracted example.

Many authors, such as Audouin and Milne Edwards, Johnston, (Erstecl, Grube,
De Quatrefages, and Marenzeller, following Savigny, regarded the Polynoidae as one or
more genera of the family Aphroclitidae.

Kinberg, on the other hand, gave them the position of an independent family, or
indeed two, if we include Iphione, with the characters indicated in the subsequent
summary. In this he was followed by Malmgren and Theel. Olaparede, again, varied the
latter arrangement a little by making them his second Tribe, an equivalent position.

Savigny (1820) placed the Polynoidae as a genus of his family Aphroditae under the
Nereids, the general characters being the oval or oblong body, with its elytra, which
were typically a dozen pairs, branchiae (which he says are easily recognised), head and
its median unpaired, and external antennae, four eyes, and armed papillose proboscis.
With the exception of the remarks on the branchiae his description is fairly good.

By Audouin and Milne Edwards (1834) the Polynoidae were distinguished from the
Aphroditidae by the number of their antennae, by the armature of the proboscis, aud by the
alternation of scales with cirri. Some have large scales completely covering the dorsum,
others have them so minute as to leave the dorsum more or less bare, and in some cases
they are vesiculate. The antennae are five or rarely four; proboscis armed with large
jaws. The simple branchiae occur with the cirri. The first pair of feet terminate in long
tentacular cirri ; the appendages of the last segment form styles. They frequent banks

36

272 POLYNOID^E.

of oysters, and some construct mucous tubes with fragments of shells. Some are phospho-
rescent. They described thirteen species of the genus.

The Polynoidse were included by Kinberg (1857) under his third family, Polynoina.
His diagnosis v^as : — Elongated body ; no facial tubercle ; convex cephalic lobe with the
base of the tentacle from the middle anteriorly; two antennas; four eyes ; palpi thick.
Pharynx exsertile, cylindrical, without long papillae; horny jaws. Scales twelve to
thirty-five pairs. Segments bearing elytra devoid of cirri. He gave six genera, viz.
Lepidonotus, Halosydna, Antinoe, Harmothoe, Hermadion, and Polynoe, the second, third,
fourth, and fifth being new.

Michael Sars (1860) reviewed the condition of the Polynoida3, as shown by GErsted
and the subsequent writers up to date, and gave a list of the northern examples with new
species, viz. P. nodosa, P. asperrina, P. rarispina, and P. scabriuscula.

De Quatrefages (1865) described the Polynoidse as having a very distinct head,
furnished with three antennas and four eyes. The buccal segment is characterised by the
presence of two pairs of tentacles, the superior being bifurcate, the much longer and
larger inferior, simple. The feet are more or less biramous, but the setigeous processes are
united to a common base. Elytra alternating with superior cirri, and covering the back
throughout. Proboscis armed with two pairs of horny jaws.

In his remarks on the tribe of the Polynoidse Claparede (1868) l criticises the great
increase in the genera caused by Kinberg and Malmgren. Thus the former subdivided
the genus Polynoe of Savigny into six, taking as his basis of classification the position of
the lateral antennas ; the number of the scales ; the fact of their covering more or less of
the dorsum ; and the length of the body. Malmgren again augmented the number of
the genera by nearly as many more as Kinberg, so that the original genus of Savigny
was multiplied nearly twenty-fold. Claparede points out that the objections of Sars to
the methods of Kinberg apply equally to those of Malmgren, because he not only
employs the same elements in classification, but adds others of less value, such as the
structure of the terminal parts of the ventral bristles. He shows that, for instance, the
same species of Syllis presents features which would appear to subvert the reliance to
be placed on this method, since the compound bristles are often replaced by simple
bristles. There is, however, more in the systematic study of the bristles than Claparede
imagined. Claparede grouped the Polynoidas as one of the Tribes of his family Aphro-
ditiens, Sav. (sens str.).

In his supplemental volume he correctly indicated, under Hermadion fragile, the
position of the nephridia and their function as channels for the extrusion of the repro-
ductive elements.

Grube, in his ' Annulata Semperiana,' gave the following description of the genus
Polynoe. Body oblong or elongated, flattened and vermiform; segments more or less
numerous, second, fourth, fifth, and every alternate segment to the twenty-third bearing
elytra, the intermediate ones having cirri. The segments after the twenty-third have
scales on every third foot or they are absent, rest bear cirri. Cephalic lobe rounded,
more or less bipartite; two pairs of sessile eyes, three tentacles, two subtentacles (palpi).
Buccal segment with two tentacular cirri, no setse ; two bundles of bristles and two cirri.

1 Ann. Nap.

LEPIDONOTUS. 273

Ventral cirrus twice as long as pinna, seta? simple, two anal cirri. Elytra covering
dorsum or leaving it bare. Pharynx with a crown of simple papilla?. Jaws horny, not
denticulated. He grouped the species chiefly according to the arrangement of the
tentacles, the number of the elytra and their structure, with a few additional particulars.
The account is thus more detailed than that of 1874.

Gr. Darboux 1 fils distinguishes in the dorsal cirrus the cirrophore and the cirrostyle.
The former is an evagination of the entire musculocutaneous envelope. The cirrostyle
is inserted on the cirrophore by a delicate epidermic membrane. A secretion, he says,
fills a glandular pocket at the insertion and causes a strain so that rupture ensues.

Whether as a family or a sub-family the Polynoidas are sufficiently distinguished
from the Aproditidse by the shape of the body, the absence of a facial tubercle, the
diminished size and the chitinous armature of the proboscis, the reduction in size of the
alimentary caeca, and the position of the segmental (nephridial) papilla?.

They are cosmopolitan forms, yet each area has its characteristic species. Thus in
Britain such as Eupolynoe anticostiensis , McL, Melaenis Loveni, Mgrn., and Encranta
villosa, Mgrn., are absent, while they are found in more northern latitudes.

Commensalism is not uncommon in the group ; thus Dr. Baird found Harmothoe
cirrata (?) in the tubes of Ghoetopterus, Gattyana cirrosa is common in the tubes of
Amphitrite, Polynoe scolopendrina in tubes of Terebella nebulosa. Malmgrenia and Acholoe
occur on Echini and starfishes (Astropecten) } and Harmothoe in Euplectellae. Dr. Baird
notes that Mr. Lord found at Vancouver's Island a Lepidonotus coiled under the foot of a
Fisurella, and another on a starfish. Verrill, 2 again, mentions an orange-red Polynoe
which occurs amongst the tentacles of the anemone, Boloeera Tuedide, and another species
with a dark purple proboscis and finely spinulose scales is very abundant among the
branches of Acanella Normani. Dr. H. J. Johnson adds two species to the forms living
as commensals, viz. Polynoe reticulata, in tubes of Amphitrite and Thelepus, and P. gigas
in an Amphitrite, both from the Pacific coast of California. 3

Genus VIII. — Lepidokotus 4 {Leach, 1816), char, emend.
Body short, more or less linear. Anterior part of the cephalic lobe produced into
the bases of the median and lateral tentacles. Palpi smooth or with papilla? in five
longitudinal rows. Three comparatively short alimentary ca?ca directed forwards into the
peri-pharyngeal space. Elytra, twelve pairs, covering the dorsum entirely, and occurring
in segments bearing feet thus : 1, 3, 4, 6, 8, and so on to 20, 22. Bristles of the superior
lobe slender, serrate, shorter than the inferior, which have a smooth portion below the
slightly hooked tip, and then a spinulose region beneath. Nerve-trunks in the granular
layer of the epiderm, between the powerful oblique muscles.

1 C. R., 126 (1878), pp. 257-8.

2 'Albatross' Explorations, 'U.S. Fisheries "Report/ 1885, p. 525.

3 'Proceed. Calif. Acad. Sc./ 1896, 3rd ser., vol. i, No. 5, p. 170, &c.

4 Kinberg (op. cit., 1857) gives as a diagnosis of the genus: — "Anterior part of the cephalic
lobe produced into the bases of the tentacle and antennas ; pharynx with papillee ; jaws ; elytra
covering the dorsum entirely ; body short." Nothing special can be made out of his remarks on the
jaws and tentacular cirri.

274 LEPIDONOTUS SQUAMATUS.

1. Lepidonotus squamatus, Linnaeus, 1788. Plate XXV, fig. 1.

Specific Characters. — Lateral tentacles longer than the cephalic lobe, median tentacle,
tentacular cirri, buccal and dorsal cirri inflated below the apex, smooth. Palpi
elongated and tapering, but in spirit scarcely longer than the median tentacle, the basal
part of which equals the length of the cephalic lobe. Scales ovate and reniform,
studded with chitinous bosses and ciliated on the outer margin. Bristled segments
twenty-six. Dorsal bristles long, tapering, and finely serrated, longer than those of
L. clava. Ventral bristles stout, with a short series of spikes on the distal and slightly
thickened portion of the shaft below the falcate apex. The distal row has two longer
spines. Ventral nerve-cords comparatively small, lying within the dense cuticle and thin
granular layer of the epiderm, and in the area between the attachments of the oblique
muscles ; only connective tissue internally.

Synonyms.

1766. Aphrodita squamata, Linn. Syst. Nat., 12th edit., p. 1084.

1788. „ „ „ „ (G-melin) p. 3107.

1765. „ „ Baster. Opusc, subsec. ii, 2, p. 66, tab. vi, f. 5, A, B, C, D.

1766. „ „ Pallas. Misc. Zool., p. 91, tab. vii, f. 14 a— d,
1768. ,, „ Pennant. Brit. Zool., iv, tab. xxiii, f. 26.
1776. „ punctata, 0. F. Muller. Prod. Zool. Dan., p. 218, n. 2642.

1789. „ „ Abildgaard. Zool. Dan., iii, p. 25, tab. xcvi, f. 1 — 4.

1800. Die gediipfelte Aphrodite, O. F. Muller. Natnrges einiger Wnrm-Arten, p. 170, tab. xiii.

1816 — 1830. Aphrodita squamata, Cnvier. Diet, des Sc. Nat., ii, p. 283.

1820. Polynoe squamata, Savigny. Syst., 22.

1828. Eumolpe squamata, Blainville. Diet, des Sc. Nat., lviii, p. 458, pi. ix, f. 2.

1834. Polynoe squamata, Aud. & Ed. Annel., p. 80, tab. i, f. 10 — 16.

1839. „ „ Johnston. Ann. Nat. Hist., ii, p. 432, tab. xxii, fig. 1.

1840. „ „ Grube. Actin. Echinod. u. Wiir., p. 87.

1843. Lepidonote punctata, CErsted. Annul. Danic. Conspect., p. 12, f. 2, 5, 39, 41, 47, 48.

„ Lepidonotus squamatus, H. Bathke. Beitrage z. Faun. Norweg., 149.
1851. Polynoe squamata, Maitland. Fauna Belg., p. 213.

1855. ? Lepidonote armadillo, Leidy. Mar. Invert. Bh. Is. & N. J., p. 16, pi. ii, f. 54.
1858. Lepidonotus squamatus, Kinberg. Freg. Eugen. Besa, p. 13, tab. iv, f. 15.
1860. Polynoe squamata, Sars. Yid.-Selsk., Forhandl. for 1860, p. 4 (sep. copy), 57 (original copy).
1865. Lepidonotus squamatus, Johnston. Cat. B. M., p. 109, pi. viii, f. 1.
„ ,, Malmgren. Nord. Hafs. Ann., p. 56.

„ Polynoe dasypus, De Quatrefages. Hist. Ann., i, p. 266.

1873. Lepidonotus squamatus, Willemoes-Suhm. Zeitsch. f. w. Zool., xxiii, p. 347.

Yerrill. Invert. An. Yin. S., Bept. IT. S. Fish., i, p. 581, pi. x,f. 40, 41.

1874. „ „ Mobius. Untersuchung. d. Ostsee, p. 112.

Mcintosh. Ann. Nat. Hist., April, 1874, p. 261.

1875. „ „ Mcintosh. Invert, and Fishes St. And., p. 115.
1877. „ ,, Huxley. Man. Invert., p. 227, &c.

1879. „ squamata, Webster. Annel. Cheet. Yirgin., p. 4.

„ squamatus, Tauber. Ann. Danic, p. 79.

LEPIDONOTUS SQUAMATUS. 275

1883. Lepidonotus squamatus, Levinsen. Nord. Annulat., p. 194.

1884. » „ Webster & Benedict. Ann. Mass., p. 699.

„ Polynoe squamata, A. G. Bourne. Trans. Linn. Soc, Zool., ii, p. 349, &c.

1886. Lepidonotus squamatus, Harvey-Gribson. Yerm. Liverp., p. 150.

1888. Polynoe squamata, De Saint-Joseph. Ann. d. Sc. Nat. (7), v, p. 151.

1890. Lepidonotus squamatus, Malaquin. Ann. Boulon., p. 15.

1896. Polynoe squamata, H. F. Johnson. Pacific Annel., p. 166.

Habitat. — Everywhere round British shores, from Shetland to the Channel Islands,
under stones between tide-marks, and stretching to the laminarian and coralline regions
beyond, as well as to comparatively deep water (fifty fathoms), where it is partial to
crevices in old shells, especially univalves covered with coils of Sepulge, and the bottoms
of stones. It is common in the stomachs of the cod and other fishes, and is frequently
tossed on shore after storms along the east coast, as at St. Andrews. It extends across
the Atlantic to the Canadian and American waters — from the St. Lawrence to Cape
Cod, and passes as far south as the Azores, off which (Fayal) it was dredged at a depth
of 450 fathoms in the ' Challenger.' In the north it ranges from Greenland to the
.Norwegian and Western European coasts, and Gould includes it in his list from the
shores of Massachusetts.

Length about 25 mm. ; more rarely 50 mm.

Head (Plate XXV, fig. 2) broadly ovate or rounded, bounded posteriorly by the fold
of the nuchal plate, and anteriorly running into the bases of the median and lateral
tentacles. It is smooth, iridescent, purplish pink, has a longitudinal median furrow,
and bears on its dorso-lateral margins the four black eyes, the first and slightly larger
pair of which are rather in front of the middle line, and the second towards the posterior
border. Anteriorly the base of the median tentacle occupies the centre, and is distin-
guished from the smaller bases of the lateral tentacles, conjoined with it, by the
arrangement of the dark pigment which forms a V with the point posteriorly. The
median tentacle is considerably longer than the lateral, all having the bulbous and more
deeply pigmented region below the pale filiform tip. Beyond the ceratophore the
column is opaque, whitish, toned off gradually to the dark ring at the enlargement. The
tentacular cirri are similar in structure but more slender. All these organs and the
palpi are smooth. In life the latter (palpi) are longer than the median tentacle.
Moreover, the first foot (basal process of the tentacular cirri) bears on its inner edge
towards the front a tuft of six or seven slightly curved and tapered spinous bristles.
These are shorter and thicker than the ordinary dorsal bristles. They are thus
apparently more numerous than in L. claim. The small size of all the appendages of the
head is a distinctive character when contrasted with the latter species.

Body of about twenty-six bristled segments. Dorsum smoothly rounded, with
three or four bars in each segment, at the sides of which are soft elevations upon which
the scales are placed, or which bear cirri at their external borders, the former being more
prominent than the latter. The dorsum is pale, but between the elevations for the last
pair of scales posteriorly a brown central patch occurs, while the segment behind
presents a median, blunt, spear-head of brown, and a lateral ocellate arrangement, the
whole being symmetrical. In the preparations the rigid contraction of the longitudinal

276 LEPIDONOTUS SQUAMATUS.

ventral muscles divides the surface into a median and two lateral regions, the former
having the furrows of the segments from side to side, the latter only at the ed^es, as the
muscular region is smooth. In some the dorsum is rendered dull ochreous by a deposit
of this colour in the grooves, the ridges being pale, and the same ochreous coating is
found on the under surface and inner edge of the scales. A solution of potash makes no
change, but dilute hydrochloric acid slowly removes the deep orange colour and renders
the granules invisible.

Posteriorly the body terminates in two symmetrical basal processes which bear cirri
longer than on other parts of the body, but having the same structure and coloration.
The anus is a small aperture (especially when contrasted with that of L. clava) opposite
the posterior border of the penultimate pair of feet.

Proboscis. — The exserted proboscis shows a series of eighteen 1 papillae around the
margin. These vary somewhat in shape, being conical or hatchet-like in outline, with a
terminal process and a beak, or in shape somewhat like a dactylozoid of Millepora. All
have a trace of dark pigment in the centre. The horny teeth alternate, so that the
upper go slightly to the right of the under pair. The horny ridge on each side of these
organs will also subserve the functions of division of food.

Three caeca from the gut pass obliquely outward to the dorsal wall, the fourth being
nearly transverse. They are larger and less alternate than those of Harmothoe, and of a
darker yellow hue. They are richly glandular, with deep yellow granular cells or masses
here and there, giving a minutely dotted aspect to the surface. Their tips present only
a short lobe in addition to the terminal one. Their arrangement and aspect thus differ
from those of Harmothoe.

The food in the intestine consists of sand-grains, fragments of crustaceans, and
other debris.

Colour. — The dorsum is of an uniform brownish-grey hue in some, or with a lighter
area on each scale, the inner edge of which has, moreover, a dark brownish spot. In
others the central pale area has a nearly complete brownish ring encircling it. In
those from the stomach of the cod the papillae, especially on the anterior scales,
retain their colour, so that they are conspicuous. Again, in examples from Guernsey
and Shetland the larger bosses on the anterior scales were of a reddish amber hue, while
some young specimens were mottled with reddish brown or madder brown. Those from
the Gulf of St. Lawrence had bristles of a dusky golden hue, and the papillae of the
scales were also darker. Under surface iridescent, bluish pink anteriorly, and pinkish
posteriorly. In some from Shetland (Bressay Sound) the bases of the posterior feet
ventrally had a well-marked touch of blackish pigment situated between the nephridial
processes and extending from the tail fully a dozen feet forward, and a series of dark
touches in the median line in each segment, a pale longitudinal streak, however, cutting
each into two.

Scales. —The first pair of scales are rounded, and, like the two or three following,
have large horny tubercles, the darker colour in some specimens making them very
prominent, and each is ciliated for more than half its circumference. The next pair are
reniform, and the succeeding are more or less ovoid. The brownish horny tubercles or

1 De Saint-Joseph says sixteen.

LEPIDONOTUS SQUAMATUS. 277

bosses project as blunt points from the surface in profile. The scales (Plate XXXII,
fig. 1) increase in size posteriorly, the general shape being ovoid, though they are wider
at the ciliated posterior border. The under surface is smooth and iridescent, and shows
the pear-shaped scar for the attachment of the pedicle. In some, however, the surface
presents the ends of fibres torn from the pedicle, and this sufficiently explains why
some are readily removed, and others require separation with a knife. All the scales
considerably overlap each other, and cover the dorsum entirely. The simple filiform
cilia which occur on the edge appear to preserve the same relative length from the first
to the last, and they are often coated with a muddy deposit, and have various micro-
scopic growths. So far as can be observed, these cilia are simple and nearly cylindrical
processes with a smoothly-rounded tip, and a median streak, as if from an axis of
differentiated tissue. Under the action of potash the scale becomes coarsely granular or
areolar, the areas of the bosses or blunt spines being characterised by a more regular
arrangement of granular cells.

In young examples, barely 3 mm. in length, the scales have a few large tubercles,
and the cilia are hirsute, with grains of various kinds, besides being proportionally larger
than in the adult. De Saint- Joseph found that in a small example, 5 mm. long, the
scales were covered with Gramrnatophora marina.

In general structure the scales have externally a more or less chitinised cuticular
layer, with a cellular (columnar) coat beneath, a fibrous stroma passing between the
dorsal and ventral layers, and finely-branched nerves, from the ganglionic mass at the
scale tubercle, terminating in end-organs. The external surface has various chitinous
tubercles or processes, and the edge has cilia.

The chitinous tubercles and spines are placed in the thick cuticular layer, and the
larger are hollow in the centre, and the surface is roughened with small processes so that
they look honeycombed. This is due to the minutely nodular condition on the surface,
and not to scales, as Baron de Saint- Joseph l states. When the dark brownish (Algoid ?)
coating begins to cover their surface, the " bosses " present a reticulated appearance,
since the parasitic growth first invades the hollows between the minute tubercles,
Pigment occurs in the cellular layer beneath. The scar has a complex series of muscles,
some of which pass into the hollow of the organ. The majority of the muscular fibres
are fixed to the scale-tubercle. As Prof. Haswell 3 says, they serve in the Polynoidse for
protection, for the production of phosphorescence, for sensation, and in certain forms
for incubation.

The scales readily separate in ripe examples, and the under surface is as finely
iridescent as in Kaliotis. Captured specimens reproduce their scales. Thus in a month
they reach about a third the size of the original organs, but are pale, and their translucent
condition shows that the tips of the papillas are minutely nodular.

Many parasitic structures occur on the scales, but none more beautiful than
Carchesium, the long tufts of which resemble, under the lens, miniature zoophytes.

Occasionally examples occur with a deep orange coating on the under surface of
the scales and the dorsum of the body. In structure it is minutely granular, as in the
darker (blackish) coating on the bristles and other organs.

1 ' Ann. des Sc. Nat./ 8th ser., v, p. 231. 2 < Ann. Nat. Hist./ 8th ser., x, p. 241, 1882.

278 LEPIDONOTUS SQITAMATITS.

Feet. — The second foot has a small dorsal division, which projects anteriorly, and no
dorsal cirrus. The dorsal bristles are tapering and spinous from the skin outwards, as in
other parts of the body. On the other hand, the ventral division has more slender bristles,
with longer and more tapered spinous tips than elsewhere. When seen antero-posteriorly
they somewhat approach the pinnately spiked forms of the Aphroditidas, though, of
course, differing in character. The long tentacle-like ventral cirrus comes from the
anterior and inner base of the foot, and is directed forwards, thus diverging both in
size and relation to the parts of the foot from the subsequent organs.

The third has become more or less normal in structure, except that the ventral
division has not attained the size of that in the fully-developed foot. The serrated dorsal
bristles spring from an elevation on the upper and anterior border, the only representative
of the dorsal division, if we except the elevated crest bearing the dorsal cirrus. The
ventral bristles are stronger, the serrated tips somewhat shorter, and the points slightly
hooked. The slender inferior series alone still resemble those in the second foot. The
ventral cirrus has now the normal position and form.

The fourth foot, though smaller, is nearly typical, and the short curved dorsal
bristles are also present. The ventral bristles have tips only a little longer than
those of the central feet. The ventral cirrus is in the normal position, but the long
papilla of the segmental organ does not appear till the eighth foot, and it continues to
the penultimate one.

The typical foot (Plate XL1I, fig. 25) is massive, and presents little differentiation
of dorsal and ventral divisions other than the bristles and spines, for the main mass of
the foot is formed by the ventral division. The dorsal bristles rise from a papilla placed
far back on the dorsal edge and in the front of the cirrus, where that is present, and
their tips, as a rule, extend little beyond the bases of the ventral bristles. In structure
they consist of long, tapering, spinous bristles (Plate XXXVII, fig. 13), the spines being
arranged in close rows from base to apex, a few on the dorsal edge being shorter and
stouter, with coarser spikes (Plate XXXVII, fig. 12). The ventral bristles (Plate
XXXVII, figs. 9 and 10) have slightly longer tips than in L. clava, and the two spines at
the tip of the rows are characteristic.

The shape of the foot does not essentially alter posteriorly, and the ventral bristles
of the last foot have tips only a little longer than in the centre of the body, a feature which
almost disappears in that next in front.

The dorsal cirri spring from a dilated base (ceratophore) which has anteriorly and
posteriorly an area in which a whitish granular amorphous substance is deposited.
The deposit, indeed, marks the bulbous region on each side. The column is pale, but at
the dilatation is a blackish band, while the filiform tip is pale.

The dorsal bristles are generally covered with fine mud and parasitic growths. The
general hue of the ventral is golden, but it varies in intensity, some of the northern
forms, e. g. from Norway and the Gulf of St. Lawrence, having darker bristles. In one
large example from St. Abb's Head one side had golden bristles, while in the other all were
of a pale colour. The same form had an enigmatical, hyaline, stalked structure, with
numerous rootlets on a posterior scale. An occasional foraminifer and a small patch of
Lepralia also are met with in large examples.

LEPIDONOTTTS SQUAMATUS. 279

The large deep-water forms (forty fathoms off St. Abb's Head) seem to be softer
than those between tide-marks, and have the scales more or less abraded.

Reproduction. — Large examples from the neighbourhood of St. Abb's Head were
loaded with ova and ripe sperms on August 1st, 1884, and the Irish examples were so in
July. At St. Andrews nearly ripe as well as ripe males and nearly ripe females occur from
the beginning of May to the end of June, and it is probable that the spawning period
is in June and July. The males are distinguished by their pale hue, whereas the ripe
females are of a slate-grey. The spermatozoa have a globular or slightly ovoid head and
a long tail, as in Nereis. No ova could be pressed out of the long segmental (nephridial)
processes with their dilated and truncated ends, but they readily issued from ruptures of
the parietes of the body. The segmental papilla are alike in both sexes, of moderate
length, and with a little dark pigment at the trumpet-like ends.

Young examples of about 3 mm. in length occur in July (' Irish Exped.,' 1886, to
G-weedore). These show proportionally larger cilia, larger and fewer bosses on the scales,
and the cirri have more elongate tips, with scarcely a trace of the enlargement below.
The ventral bristles have proportionally longer tips.

When disturbed in their native haunts, their motions are comparatively slow and
cautious, so different from the restless activity of Harmothoe imbricata or Evarne. They
cling tenaciously, partly by aid of their bristles, to rough surfaces, so that, for instance,
the tubes of Filigrana give way in extracting them, and it is difficult to pull them from
grooves in shells and similar hollows. Fine examples are procured in rock-pools under
large stones that have been little disturbed for many years, where, for instance, Mgirus
jpunctiluceus and patches of Alcyonium occur. They are partial to hollows and crevices ;
thus one thrust itself into the tube of Protula, and being preserved therein has retained
the cylindrical shape. Hitherto, however, they have not been found commensalistic in
the tubes of other annelids, but occupy their sites independently under stones in rivulets
and rock-pools near low-water mark, and for some distance landwards. In confinement,
specimens of Evarne and Harmothoe will occasionally cling to the dorsum, and small
examples may even insinuate themselves along the dorsum under the scales.

The crustacean parasite, Sellius bilobus, Kroyer, 1 is found on this species in northern
waters, while Perigonymus repens* occurred in great beauty on the anterior scales of fine
specimens trawled in thirty-five to forty fathoms off St. Abb's Head.

While they are not infrequently found in the stomach of the cod and other fishes, it
is curious that young green cod in the tanks refused them and other Polynoidse, such as
Harmothoe and Lagisca, while they readily devoured Nereis, Trophonia, and Girratulus ;
the mode in which the Polynoidse curved themselves and kept the bristles prominent,
showed that they were aware of the value of such protective organs. Taken into the
mouth of the green cod they were at once rejected, and fell to the bottom of the tank,
where they were engulfed by Cottus, but with a similar result, viz. immediate rejection.

0. F. Miiller, in his « Wurm-Arten,' calls this species " die gediipfelte Aphrodite
mit rauhen einfarbigen Schuppen."

1 'Naturhist. Tidskrift./ lste E., i, 1837.

2 This zoophyte recalls the loss lately sustained by science in the death of the Rev. Dr. T. Hincks,
whose patient and accurate work amongst the Hydroids and Polyzoa will long he remembered.

37

280 LEPIDONOTUS CLAVA.

Pallas (' Miscellanea Zoologica ') mentions that Linnaeus found it in deep water.
It occurs also with pelagic Fuci, and amongsb the latter and Sertularians at the bottom of
the sea. He gives an account of the external appearance and arrangement of the scales,
and a brief outline of its anatomy.

It is doubtfulif 0. F. Miiller's Aphrodita punctata refers entirely to this form, since
he speaks of sub-crescentic bluish spots.

Audouin and Milne Edwards (1834) describe it as having twenty-seven segments,
and reaching eighteen lines in length. They think that the Polynoe punctata of 0. F.
Miiller comes near P. squamata, and so with the Aphrodita clavigera of M. Freminville ; l
but this cannot be, as M. Freminville found it phosphorescent.

It is difficult to say what H. Rathke's (1837) Polynoe granulosa 2 is. It may be
this species in which a pair of scales has been lost.

Leidy's 3 L, armadillo appears to be allied to this species or to L. clava. His
observation in regard to the unusual number of the tentacles is erroneous.

Kinberg 4 rested jhis specific characters on the comparative length of the tentacles,
and the fact that the inferior bristles were serrated below the apex. He gave a correct
account of the scales. De Quatrefages 5 adds nothing new to the foregoing, and he
includes Lepidonotus clava, Mont., as a synonym, though with doubt. Subsequent French
authors have corrected this error.

2. Lepidonotus olava, Montagu. Plate XXVI, fig. 1.

Specific Characters. — Head similar to that of L. squamatus. The large anterior eyes
are in front of the middle line, and the respective pairs on each side are wider apart.
Anteriorly the broad basal region of the median tentacle is more distinctly separated
from the bases of the lateral tentacles than in L. squamatus, and the transverse diameter
of the three processes is greater. Median and lateral tentacles thicker, and the bulbous
region below the tip marked by a more distinct band of blackish pigment. Palpi with
five rows of minute papillse. Segments twenty-seven, characteristically marked with
pigment on the dorsum ; segmental (nephridial) papillaa large and thick. Scales more or
less circular throughout, and do not quite cover the dorsum, more flexible than in L.
squamatus, and, with the exception of the first four pairs, smooth. The first three have
numerous small tubercles generally distributed, while the fourth pair have a smaller
number. Papilla for the dorsal bristles more prominent than in L. squamatus. The dorsal
bristles are shorter, thicker, less tapered and more curved than in L. squamatus, but
have similar structure. The ventral bristles have shorter curved or falcate tips, with
fewer rows of spikes, and the bare portion is shorter than in L. squamatus.

1 f Nouveau Bulletin de Sc. par la Soc. Philomat./ iii, p. 253.

2 c Beitr. z. Fauna der Krym/ p. 408.

3 ' Americ. Journ. Nat. Sc./ p. 148, pi. xi, f. 54.

4 Op. cit., 1857.

5 Op. ait., 1865.

LEPIDONOTUS CLAYA. 281

Synonyms.

1808. Aphrodita clava, Montagu. Trans. Linn. Soc, ix, p. 108, pi. vii, f. 3.
1824. Lepidonotus clavatus, Leach. Supp. Ency. Brit., i, p. 452.
1826. Polinoe scutellata, Kisso. Hist. nat. Burop. merid., iv, p. 414.

1828. Eumolpe squamata, Blainville. Diet. Sc. Nat., lvii, p. 458, Atlas, f. 2.

1829. Eurnolpe squamata, D. Chiaje. Mem. sulla Storia, iv, p. 155, tab. lvii, f. 8 and 17.
1836. Haliihsea clava, Templeton. Loud. Mag. X. Hist., ix, p. 234.

1838. ? Polynoe squamata, Grube. Anat. u. Physiol. Kiemenw., p. 60, tab. ii, f. 13.
1840. ? „ „ (Sav.), Grube. Act. Echin. u. Wurmer, p. 87.

1860. „ clypeata, Grube. Arch. f. Xaturges., Bd. xxvi, p. 71, taf. iii, f. 1.

1861. „ „ Grube. Ausnug n. Trieste, p. 138, taf. iii, f. 1.

1864. „ n Grube. Insel Lussin, p. 77.

1865. Lepidonotus clava, Johnston. Cat. Brit. Mus., Ill, pi. iv, f. 5 and 6.
„ Polynoe modesta, De Quatrefages. Hist. nat. Annel., i, p. 243.

1867. Lepidonotus clava, Malmgren. Ann. Polych., p. 130.

1870. Polynoe grubiana, Claparede. Suppl. Annel. Chetop., 9 (373), pi. i, f. 2.

1875. Lepidonotus clava, Marenzeller. Sitzb. der k. Akad., 1 Abth., Juli-Heft (sep. Abd.), p. 1.

1876. „ „ Mcintosh. Trans. Z. S., ix, p. 374.

1880. „ „ Langerhans. Zeit. f. w. Zool., xxxiii, p. 273, taf. xiv, f . 2 ?

1881. Polynoe clava, Langerhans. Canar. Annel., Nova Acta, 42, iii, p. 108.

1884. „ „ A. G. Bourne. Trans. Linn. Soc. Zool., ii, pp. 347 — 356, pis. xxiv — xxvi.
„ Lepidonatus clava, Carus. Faun. Medit., p. 202.

1885. Polynoe grubiana, Jourdan. Zool. Anz., viii, p. 128, f. 1, 2.

1887. „ „ , Jourdan. Arch. Zool. Exper., v, pp. 115—120, pi. iv, f. 11, 12, 16, 17.

1898. Lepidonotus clava, De Saint-Joseph. Ann. d. Sc. nat., 8 ser., v, p. 225.

Habitat, — This species is a southern and western form — ranging from Falmouth
and the Devonshire coast to Gairloch, Boss-shire, and extending to Valencia Harbour in
Ireland. None appear on the eastern shores. It is chiefly littoral, occurring under
stones in tide-pools even near high- water mark, as in McNiel's Bay, Lochmaddy, as well
as on oyster-beds under water. It is distributed along the shores of France to the
Mediterranean, Adriatic, and perhaps to the Canaries (Langerhans).

Length about 30 mm. ; breadth 8 mm.

Head (Plate XXXVII, fig. 4) similar in outline to that of L. squarnatus, bounded
posteriorly by the nuchal collar, which has the dark pigment of the dorsum on its edge.
The surface is smooth and iridescent, as in the former species. The eyes are visible from
the dorsum, the larger anterior pair in front of the middle line and wider apart, and two
posteriorly. Anteriorly the broad basal region of the median tentacle is more distinctly
separated from the bases of the lateral tentacles than in L. squamatus, and the transverse
diameter of the three processes is greater than in the latter species, as might be expected
from the larger size of all the appendages. The median and lateral tentacles are
proportionally thicker, and the bulbous region below the tip better marked, especially as
a band of blackish pigment occurs below the enlargement. A little brownish-black
pigment also exists on the columns of these processes and of the tentacular cirri, all these
organs being somewhat stouter and more boldly pigmented than in L. squamatus. The
palpi are somewhat filiform at the tip, and, as shown by Grube, are marked by five rows

282 LEPIDOXOTUS CLAVA.

of elongated papillae, which are conical and taper to a point. They are best developed to-
wards the tip of the organ. Prof. Bourne describes each as furnished with a sensory hair.

Body. — The dorsum is marked in the median line by broad interrupted patches of
blackish-brown pigment, three or four of the most prominent anterior touches having a
pale area in the centre. These touches extend outward to the feet posteriorly ; more-
over, dorsally the bases of the feet have black pigment, which becomes strongly
pronounced after the anterior third, the effect of the mottling of the feet and body
posteriorly being somewhat like that on tortoise-shell.

The ventral surface has a similar arrangement of areas to that in L. squamatus, only
the lateral are marked off from the bases of the feet posteriorly by a slightly elevated and
pigmented border, which terminates at the bases of the caudal styles, and is connected
with touches which extend a short way forward in the median line.

Posteriorly the body terminates in a grooved pedicle for the large symmetrical
caudal styles, all these parts being better marked than in L. squamatus. The anus is
situated dorsally at the base of the pedicle, to which it sends a ridge-like process.

The bristled segments are twenty-six.

Proboscis and Digestive System. — De Saint- Joseph found sixteen papillse along the
margin of the proboscis, but there are eighteen, as in L. squamatus, in all the examples
examined. They are speckled with black pigment. In transverse section the organ is
typical.

Only two cylindrical and rather massive cseca pass forwards into the peripharyngeal
space, and their extremities appear to be devoid of any distinct differentiation. The
stomachal region of the gut anteriorly is remarkably muscular, the fibres at the front end
forming a row of separate dull orange lobes which clasp the proboscis, and which resemble
a series of glands. Their tissue, however, is wholly muscular, the powerful fibres forming
loops in the rounded lobes, which are sometimes made by rupture of the fibres from
the proboscis.

Colour. — The dorsum is of a dull brownish hue speckled with white, and with dark
touches on the scales. The under surface is pale, with some dark touches at the mouth
and the sides of the tail. The segmental papilla has a dark column and a whitish tip in
the posterior half of the body. At the reproductive season the males are pale under
the scales, the females dark grey.

Scales. — The scales (Plate XXXII, fig. 2) of this species are more or less circular
throughout, and do not quite cover the dorsum, or, as Marenzeller says, leave bare
rhomboidal spaces. Some are firmly fixed, others easily separate from the pedicle as in
L. squamatus. They are more flexible than in that species, and, with the exception of the
first four pairs, show only minute tubercles. The first, second, and third have numerous
small tubercles or blunt dumb-bell shaped papillas, distributed generally over the surface,
while the fourth has a smaller number, less distinctly raised above the surface. The
edge of the scale throughout is smooth. They are fixed by the umbilicus, which is
nearer the outer than the inner margin, this area being generally marked dorsally by a
white patch bordered with black pigment. The scales give the dorsum generally a
leaden hue, or in some a dusky brown, speckled with white. Besides the white patch,
which posteriorly is sometimes reniform, at the umbilicus the scales are mottled with a

LEPIDOXOTITS CLAVA. 283

blackish or slightly glistening dull leaden hue, or with more numerous black specks,
which posteriorly do not seem to be connected with papillge. The under surface is
smooth and iridescent, and has the opaque whitish patch of the umbilicus. In spirit
the anterior scales generally show a fold from the umbilicus to the external margin.

The minute structure of the processes of the scales has been specially described and
figured by De Saint-Joseph, 1 who states that the tip is " scaled" and spinous. His
figure, indeed, is regularly and closely diced. So far as can be observed in the British
specimens, it is the tip and neighbouring part of the capstan-like column of the process
of the scale which is thus covered, as recently described and figured. 3

Feet. — On the dorsum of the first foot is a minute tuft of tapered serrate bristles
(Plate XXXVII, fig. 14), and a single large spine which penetrates the skin beneath
the former. Prof. Bourne speaks of four bristles being borne by each division. This
is probably a misprint, as his figure differs.

The second foot diverges from that of L. squamatus in having its comparatively large
ventral cirrus directed more distinctly inwards and forwards, in the smoother foot and
the more translucent bristles. Moreover, while the dorsal series more or less correspond
at first sight, yet the character of the serrations slightly differs from those of L. squa-
matus, being somewhat shorter and finer, and the tips of the bristles are more tapered.
The ventral bristles of L. clava again differ in having their long tips spinous to the apex,
whereas in L. squamatus the spinous tips are shorter, the tip is bare, and in some slightly
hooked.

In the third bristled foot (the second of some authors) the dorsal bristles are less
gradually tapered than in L. squamatus, while the ventral series have longer spinous
regions bare in the upper at the tip— which is more slender than in L. squamatus; indeed,
the whole bristle is more slender than in the latter species, and the rows of spines longer.
Both dorsal and ventral cirri, moreover, are different, being proportionally larger and
more bulbous below the filiform extremity.

In the fully-formed foot (e.g. the tenth, Plate XLII, fig. 26) the papilla for the
dorsal bristles is more prominent than in L. squamatus, and has a distinct black bar in
front, and the bristles are somewhat shorter and more curved (Plate XXXVII, fig. 15).
The tips of the ventral bristles are shorter, the curve more pronounced (Plate XXXVII,
fig. 11), and the bare portion shorter. The stronger and larger hooked tip in L. squa-
matus is a characteristic feature. The segmental papilla is visible on the eighth bristled
foot as in L. squamatus, and is continued to the last foot-
In the terminal foot the dorsal bristles are shorter and more curved than in L. squa-
matus, a feature present throughout the body. The ventral bristles have the curvature of
the shorter tip more pronounced than in L. squamatus. In both species these bristles
preserve great strength.

A marked difference between the species is the diminished size of the body and last
feet in L. squamatus, contrasted with the comparatively large size of all the parts in this
region in L. clava.

1 Op. cit., Aug., 1898, p. 229, f. 4, &o.

2 ' Ann. Nat. Hist./ 1898, p. 108, pi. ii.

284 LEPIDONOTUS CLAVA.

Reproduction. — Fine specimens from Gairloch, Ross-shire, were laden with ripe
ova in February, and had a dull greyish hue from the tint of the eggs.

Habits. — It is a sluggish form.

This is one of the discoveries of the acute and sagacious Col. Montagu on the
southern coast.

Like other Polynoidas it is predaceous, masses of the bristles of Harmothoe imbricata
being found in the intestine, with shreds of skin and other debris.

It is probable that Bisso's Polinoe scutellata 1 is either this or L. squamatus.

Delle Chiaje's figures in the ' Memorie ' 3 would appear to belong to this species ;
the first represents the entire animal, and the second the posterior end enlarged,
though in the text it is termed the anterior end.

The statement by Audouin and Milne Edwards in their ' Anneles ' that perhaps
Montagu's Aphrodita clava approached their Polynoe Imvis, is due to a misap-
prehension. The species much diverge.

Marenzeller, 3 in 1875, reviewed the literature of this species, and showed the
identity of Risso's P. scutellata, Delle Chiaje's and Grube's Eumolpe squamata, and the
P. modesta of De Quatrefages : he also linked on the P. grubiana of Claparede 4 with
Montagu's form ; but the palpi (Unterfuhler) of this species present a different appear-
ance from those examined, since they are studded all over with papilla, whereas those of
L. clava have only rows of pointed papillae, as indicated by Grube, in 1860, in the
Mediterranean examples. Marenzeller procured British specimens from Oumbrae in the
Clyde.

It is doubtful whether the form considered by Langerhans to be a young variety is
this species, since the palpi are smooth and the bristles diverge.

Baron de Saint-Joseph includes the Polynoe dorsalis of De Quatrefages under
this species, but this is doubtful, since P. dorsalis has cilia on the external margin of
the scales. An examination of the specimen, however, may have shown that this is an
error in description. He likewise places Lepidonotus wahlbergi, Kinberg, under the
same head, but the foreign species differs, even under a lens, by the fact that the scales
throughout, that is to the last, have numerous tubercles, those in front forming pro-
minent spines ; and, besides, the scales of L. clava, Montagu, are proportionally larger
and, with the exception of first four pairs, are nearly smooth. Only in the old scales is
there a belt of minute tubercles within the edge, and similar minute processes over the
surface. These are, however, very different from those of L. wahlbergi. This distinction
is clearly shown in the figure 5 in the ' Challenger ' volume. Moreover, the dorsal
bristles are much less tapered and acute than in L. clava, the reverse being the case
with the ventral, which are proportionally more slender and elongate, and with a longer
row of spines than in L. clava. 6

1 ' I/Europ. merid./ p. 414.

2 Op. cit.

3 "Zur Kenntniss der adriatischen Annel./' 'Wiener Akad. Ber./ 1 Abth., Juli-Heft, 1875, p. 1,
sep. Abd.

4 A view still held by Dr. Benham, c Carnb. Nat. Hist./ " Polychset., &c," p. 310, 1896.

5 PI. xi, f. 1.

6 Vide Mcintosh, < Ann. Nat. Hist./ ser. 7, vol. ii, p. 108, pi. ii, f. 12, 1898.

GATTYANA CIEEOSA. 285

Genus IX. — Gattyana (Nychia, 1 Malmgren), Mel.

Lateral tentacles arising below the base of the median, they and the cirri densely
covered with long cilia. Palpi with numerous short clavate papilla? truncate at the tip.
Scales fifteen pairs, minutely spinous, covering all the dorsum, and attached to segments
1, 3, 4, 6, 8, ... . 22, 25, 28, 31. Bristles of the dorsal lobe ranging from stout,
curved, and spinose inner forms to elongated ones with tapering, hair-like tips, finely
spinous. Bristles of the inferior division stouter, with spinous distal regions and simple
hooked tips.

Gattyana oieeosa, Pallas, 1766. Plate XXV, fig. 3.

Specific Characters. — Body of thirty-five to thirty-six bristled segments with a trans-
verse impression in most. Head produced anteriorly into two pointed lobes. Tentacles and
cirri densely covered with cilia. Scales, with the exception of the first pair, somewhat
reniform, with long cilia on the posterior and external margins, smooth to the naked eye,
but with many minute spines on the surface, the larger being external and posterior,
with the tips often bifid, generally pale olive or buff, and frequently with a dark spot at
the point of attachment. Dorsal bristles somewhat slender, with long tapering hair-like
tips and rows of spines. Ventral bristles with simple hooked tips, beneath which is a
spinous region which diminishes in length in the bristles from above downwards.
Ventral cirrus short, with a few short clavate papilla. Segmental process slightly
tapered towards the tip, and of moderate length.

Synonyms.

1766. Aphrodita cirrhosa, Pallas. Miscell. Zool., p. 95, Tab. 8, figs. 3—6.
1780. ,, scabra, Fabricius. Fauna G-roeuL, p. 311, n. 292.

punctata, Fabricius. Ibid., p. 311.

scabra, Bruguiere, Encyc. Method., vers, i, p. 88.

mridis, Montagu. Trans. Linn. Soc, vol. xi, p. 18, Tab. 4, fig. 1.

scabra, Savigny. Syst. des An., 26.

punctata, Savigny. Ibid.
1828. Eumolpe scabra, De Blainville. Diet. d. Sci. Nat., vol. lvii, p. 459.
1834. Polynoe scabra, Aud. and Ed. Ann., 87.
1839. Polynoe mridis, Johnston. Ann. Nat. Hist., vol. ii, p. 437.

1843. Lepidonotus assimilis, CErsted. Annulat. Danic. Consp., p. 13, figs. 3, 6, 14, 32, 33, 37, 38,

45, 46.
„ „ scabra, Ibid. Gronl. Ann. Dorsib., 164, pi. 1, figs. 2, 7, 10, 13, 17 and 18.

1850. Lepidonote scabra, Sars. Reise i Lof. og Finm., p. 209.

1851. Polynoe scabra, Grube. Fam. der Ann., 37 and 120.
1858. Earmothoe scabra, Kinberg. Eug. Kesa, 21.

1792.
1815.

1820.

1 The title Nychia had already been used by Stal for one of the Eemiptera, so that Malmgren's
name (1865) lapses. The term Grattyana may accordingly be appropriately substituted, after the
generous founder of the New Marine Laboratory at St. Andrews.

286 GATTYANA CIRROSA.

1861. Polynoe scabra, Sars. Yid. Selsk. Fork., I860, p. 58.
„ „ scabriuscula, Sars. Ibid., p. 61.

1864. Lepidonotus cirratus, var. parasiticus, Baird. Trans. Linn. Soc, 1864, p. 161.

1865. Polynoe scabra, De Quatrefages. Ann., vol. i, p. 235.

„ Lepidonotus imbricatus, Johnst. Cat. Brit. Annel., p. 118.
1867. Nychia cirrosa, Malmgren. Nord. Hafs-Ann., p. 58, Tab. viii, f. 1.

1871. ,, „ Bhlers. Sitzunsb. phys.-med. Erlangen, 1871, p. 77.

1872. „ „ Sars, Nyt. Mag. f. Natnrv., 19, p. 202.

1873. „ „ Sars. Bid. Christ. Faun., p. 2.

1874. „ „ Mobius. Die Zweite dentsche Nordpolarfahrt, ii, p. 253.

„ „ „ Malm. Groteborgs Kongl. Yet. o. Yitt. Samhalles Handl. ; Ny Tidsfoljd, Haft.

xiv, p. 87.

1875. „ „ Mcintosh. Invert, and Fishes, St. A., p. 115.
„ „ „ Ehlers. ' Porcupine' Annel., 1869, op. cit., 32.

1877. „ „ Marenzeller. Ost.-ung. Nordpol. Exp. (Deutsch. d. k. Akad. Wiss.), 39.

1879. „ „ Theel. Kongl. sv. Yet. Akad. Handl., 16, 3, p. 7.

„ „ „ Tauber. Ann. Danic, 79.

1883. „ „ Wiren. Chastop., 'Vega' Exped., 387.
„ „ „ Levinsen. Nord. Annnlat., 195.

1884. „ ,, Webster and Benedict. Ann. Mass., 700.
1886. „ „ Giard. Bullet. Sc. Nord., Nos. 9, 10, p. 338.

„ Iphione muricata, Harvey Gribson. Yerin. Liverp., 150.

1890. Nychia cirrosa, var. Chsetopteri, Malaquin. Ann. Boulon., 15, pi. 1, f. 7a — Id.

1891. ,, „ Hornell. Trans. Liverpool Biol. Soc, v, p. 230.

„ ,, Trautzsch. Polynoid. Spitzb., Jenaische Zeitsch., 24, 1 Heft, p. 75 ; and Arch.

f. Naturges., 55 Jahrg., 1 Bd., 2 Hft., p. 143.

1896. „ ,, Michaelsen. Polych. Faun, deutsch. Meere (Wiss. Meeresuntersuch., Bd. ii),

p. 7.

1897. Gattyana cirrosa, Mcintosh. Ann. Nat. Hist., ser. 6, vol. xx, p. 167.

1898. Nychia cirrosa, Michaelsen. Gronl. Annel., Bib. Zool., Heft 20, Lfg. 4, p. 120.

Habitat. — This species stretches from Greenland and the American waters to the
shores of Northern Europe, but is numerically less abundant than many of the
Polynoida3. It occurs not infrequently in the deeper water off St. Andrews Bay,
and is tossed on shore after storms. Specimens likewise come from Shetland (J. Gr.
Jeffreys) in sixty fathoms, nine miles off Balta, from Southport Sands (Dr. Carrington),
Hastings, between tide-marks, Channel Islands (Herm), Broadhaven Bay, Ban try Bay,
and Berehaven Bay, in Ireland (A. Gr. More, Professor Had don, and Dr. Scharff), the
specimens found in the first-mentioned region being very large. Fine examples are not
uncommon in the stomachs of cod and haddock at St. Andrews (E. M.). In Shetland
it clings to the branched form of Melobesia in Bressay Sound. Malmgren pointed
out that it was the form called by Baird Lepidonotus parasiticus in the tubes of
Oh&topterus, in the British Museum ; and Hornell has confirmed this in the tubes of the
same form, and in those of Thelepus in the Channel Islands, while at St. Andrews it is
partial to the tubes of Amphitrite figulus, Dalyell.

It comes from the Gulf of St. Lawrence (Whiteaves), the Atlantic in 580 — 630
fathoms (Ehlers, 'Porcupine'), and Canon Norman frequently procured it in Norway
and Finmark. Specimens also occur in Iceland.

GATTYANA OIJRROSA. 287

Length. — A fine example from St. Andrews measured, exclusive of the processes,
47 mm.; and another, 42 mm. in spirit, comes from Broadhaven Bay, W. Ireland
(A. a. More).

The head (Plate XXVII, fig. 5) differs from that in Lepidonotus in the relations of
the median and lateral tentacles, since on each side of the median the head terminates in
a pointed papilla (peak), the lateral tentacles being inferior. The outline of the head is
somewhat ovate, and in life it is rose-red. A conspicuous pair of black eyes occur
posteriorly on the dorsum, while a slightly larger pair are in front at the outer margin,
and rather below the peak on each side of the tentacle. They are not well seen from
the dorsum, and are fitted for anterior vision. The nuchal collar has a prominent
boss or papilla in the centre. The median tentacle arises from a massive base, often of
a slightly brownish tint, as a pale ciliated process about as long as the palpi in the spirit
preparations, though of course the latter are much longer in life, and having a slight
swelling below the filiform tip. The bases of the short lateral tentacles, which are
beneath the massive base of the median, have a brownish bar. They are similarly
ciliated, and have a slight enlargement below the filiform extremity. The cilia
on the tentacles are elongate processes with a slightly bulbous tip, the centre of the
latter and the column being granular.

The pale palpi have numerous small papillae, which are shortly clavate in outline, with
a truncated tip, which has a few minute papilla. These are arranged in six rows —
apparently two dorsal, two ventral, and a lateral on each side, the intervals between the
rows being considerable. Malmgren simply says the surface is densely covered. They
diminish in size towards the tip, and disappear from the filiform termination. Under
a lens in life the organs appear to be smooth, but the minute papillae are visible under
a low power.

Body elongated, somewhat elliptical in transverse section, though more prominent
dorsally than ventrally, gradually diminished in transverse diameter towards the head,
and narrowed somewhat more abruptly towards the posterior end. Bristle-bearing
segments 34 — 36, as Malmgren says for the Arctic forms. The dorsum is marked laterally
by the papillae for the scales, and the corresponding elevations in those feet bearing
cirri. Moreover, each segment has a transverse bar, as it were, imprinted on its
middle, the outer edge of the bar being especially depressed just as the surface of
the segment rises towards the papillae. These are continued to the fourth last segment
posteriorly. On the other hand, the bulging of the body caused by the proboscis
renders them less distinct anteriorly, though traceable in every segment. A tendency
to separation of these bars at the anterior border of the segment is noticeable, while
posteriorly the central half runs to the junction with the next segment. A median
streak passes along the dorsum. Ventrally, the segments show a broader lateral band on
each side, and a narrower median, which is opaque whitish in the centre. A little
behind the line of each segment anteriorly an oblique streak passes to the nephridial
process. The ventral surface of the body is iridescent, and in life it has a purplish
sheen along the median region. In others the pinkish nerve-cords and ganglia
form a well-marked moniliform median band, commencing behind the striated lower lip
as a broad stripe. The feet are dull yellow.

38

288 GATTYANA CIKKOSA.

The segmental papilla is directed between the feet, and is in the form of a slightly
tapered process of some length. The northern forms agree with the British in regard
to this structure.

Posteriorly, the body terminates in two elongated caudal styles, which have a
similar structure to the cirri.

Proboscis. — When extruded, the organ shows on each edge nine terminal papillae
similar in shape to those of Lepidonotus. The horny jaws appear to resemble those of
the genus named, and they are acutely pointed.

Two long, slender caeca pass forward into the peripharyngeal space.

Scales (Plate XXXI, fig. 1) — fifteen pairs. The first pair are somewhat
circular and ciliated all round, the inner and anterior margins having short
and somewhat clavate cilia, the rest being densely covered with similar cilia, which
as usual, are often thickened by parasitic growths. The surface has numerous
small horny papillae scattered over it, so that it is finely spinous. Such a scale
however, is in contrast with the Arctic examples or those from the Gulf of St.
Lawrence, Norway, and Finmark, in which the sharp brown spines form prominent
features in every scale of both large and small examples. Bifid tips also occur on many
of the spines, as mentioned by Malmgren. It is not a question of size in regard to this
difference between the British and the other examples, for in large representatives
from the west of Ireland the same features are present as in the other British specimens.
Moreover, the cilia are more prominent in the northern forms. These features, however,
by no means indicate specific difference, for the general characters remain diagnostic.

The condition of the scales of the British examples is more like that of Gattyana
Amofisdeni, Mrgn., though the spines are not so visible under a lens.

The scales of some have grains of sand and mud attached externally. Old and fine
specimens have a dull olive hue on the dorsum, while the edges are flanked by the paler
bristles and cirri, which are often, however, so coated with parasitic algae and other
structures, and grains of sand, that their normal hne is obscured. The scales of the large
examples from the tubes of Amphitrite have a finely-streaked appearance from the dark
growths on the spines.

Colour. — Dorsum of a pale yellowish-brown or sandy colour, the anterior scales
being somewhat darker. In a considerable number (the majority of those tossed
on shore at St. Andrews) a dark greyish-brown spot occurs at the point of attachment,
and these form a somewhat regular row along each side. The prominent spines of the
Arctic examples are brownish. The same dark spot at the point of attachment is present
in some of these, and a dark touch at the inner border of each scale, so that an
interrupted double band occurs along the middle of the dorsum. Tentacular processes
pale. Head deep pinkish red, or only between the posterior eyes. In some the central
process of the nuchal collar is bordered with dark pigment. Under surface pale,
iridescent. Bristles pale yellow.

None of the British examples appear to show the dark median band of the dorsum
from the nuchal fold backward, with fainter touches of colour in the median depressed
parts of each segment to the tail, where darker pigment again appears, which
characterises some Arctic examples (Godhaven Harbour). Flecks of pigment are also

GATTYAXA CIRROSA. 289

seen along the mid-dorsal line even when no distinct band is present anteriorly and
posteriorly.

Feet. — The tentacular cirri agree with the median tentacle. The basal region has the
usual small tuft of serrated bristles (corresponding to the dorsal) with a spine. They
are somewhat short and rather stout bristles, with rows of short spikes towards the tip.

In the second foot the dorsal bristles are shorter and more roughly spinous than in
the typical foot. The latter is probably partly due to their shortness and greater curvature.
On the other hand, the ventral bristles have much more elongated tapering tips than in
the typical foot. The ventral cirrus has a few short clavate papillae throughout the whole
series.

The third foot presents a considerable change in the dorsal bristles, which now show
the strong series with curved, serrated tips next the body, and the slender series with
tapering tips adjoining the ventral division. The dorsal cirrus is somewhat shorter and
thicker than in the fully developed foot. Both shafts and tips of the ventral bristles are
still much more slender than in the average foot.

In the fourth foot a near approach to the condition in the typical foot occurs,
though both dorsal and ventral bristles are more slender.

In the typical foot the dorsal division is bluntly rounded and turned upwards (Plate
XLII, fig. 27), the spine, which is sheathed in skin, passing out as a slender process
towards the inferior edge of the bristles. The dorsal bristles are rather slender
(Plate XXXVII, fig. 18), have a long tapering tip, with rows of spines, the extremity
being hair-like. The ventral division, on the other hand, has a somewhat triangular edge,
the spine, which is sheathed at the base, and forms an independent process posteriorly,
passing outward at the apex of the triangle. The ventral bristles show tips increasing
in length from below upward, the superior bristles (Plate XXXVII, fig. 16) having
more delicate tapering tips and more numerous rows of spines, while the tips are slightly
hooked, the intermediate series (Plate XXXVII, iig. 17) often having a slight enlargement
below the hooked tip. In large examples from deep water beyond St. Andrews Bay the
smooth tips of the ventral bristles are somewhat shorter and broader. In contrasting
these with the bristles of Arctic examples, the tips (that is, the region beyond the spines)
of the latter are longer, and the bristles, both dorsal and ventral, are in some cases rather
stouter. In the Arctic forms also the cilia on the cirri are somewhat longer.

In the posterior feet the tips of the ventral bristles again become elongated and the
shafts more slender. In the last foot, indeed, the tips are so attenuate that, leaving the
shafts out of sight, the curve of this region is almost the only distinction by which they
can be recognised. The ventral cirri also are more elongate.

A curious ochreous appearance occurs in certain forms from Godhaven, Greenland,
in which the tips of the dorsal bristles are covered with a minutely granular reddish-
brown deposit, probably derived from their surroundings. The same structure coats the
cilia of the cirri and scales.

The figure of the foot in Malmgren's paper does not show the upper spine, and in
the ventral division the conical process for this structure is absent.

Parasites. — Loxosoma is not uncommon on the bristles and feet of the Arctic forms
and those from the Gulf of St. Lawrence.

290 GATTYANA CIRROSA.

In a specimen from Godhaven Harbour (Disco Island) the body was dotted with
rounded or ovoid whitish structures beneath the skin. They consisted of a mass of well-
defined granules, generally of a somewhat ovoid shape and a double outline.

The parasitic crustacean, Sellius bilobus, occurs on the dorsum under the scales in
northern forms, and seems to be a large parasite for so small an annelid.

Fine specimens are procured in the tubes of AmpMtrite figulus (Dalyell) under lar^e
stones near low- water mark of spring tides at St. Andrews. They occupy the anterior end of
the tube. All the examples were large, but the alimentary canal of those examined showed
no recognisable food. The species probably finds in this and other tubes a safe refuge and
sufficient food. It is, moreover, phosphorescent, irritation causing a very pale greenish
or yellowish light to illuminate the scales. The phosphorescence is less vivid than in
Harmothoe imbricata and Polynoe scolopendrina. As a rule, Gattyana occupies a position
close to the mouth of the long tube of the AmpMtrite beneath large stones, so that it is
well protected from marauders, even supposing they were attracted by its light. In the
same way its opportunities for alluring animals are curtailed, so that the remarks formerly
made in this connection still hold. 1

To sum up, the British species differs in the softer and much smoother scales, the
horny papillaB being microscopic, and in the shorter, smooth tips of the ventral bristles,
which tips are likewise more attenuate in the Arctic forms. The innermost bristles of
the dorsal series appear also to be stouter and shorter in the Arctic forms.

Pallas (1776) was somewhat in doubt about this species, which had been sent him in
a rather softened condition by Gronovius from the northern seas, and thought it might be
an older form of his Ajphrodita lepidota, though the structure of the feet differed. Both
description and figures are imperfect, and in fig. 6, Plate VIII, the foot is inverted, but
Malmgren is right in identifying it with this species. The general pallor struck Pallas.
He mentions the " scabrous " condition of the posterior margin of the scales, and the
elongated yellowish dorsal bristles. The description given by Fabricius (1780) is quite
recognisable, and the dull greenish colour dorsally, the pale ventral surface, with the
hispid scales, are characteristic. Savigny (1820) added nothing to the remarks of
Fabricius.

Johnston (1839), both in this paper and in the subsequent remarks in the ' Catalogue,'
gives only Montagu's description. GErsted's figures (1843), rather than his descrip-
tion, show that this was the species to which he referred in both publications. He also
gives its habitat as in the deeps of Greenland. Sars (1861), in his account of the
Norwegian Polynoidx, gave descriptions of both a Polynoe scabra, Fabr., and P. scabriuscula,
n. s. ; but Malmgren has rightly decided that these refer to the same species, considerable
variation occurring between the Arctic and the more southern examples. De Quatrefages
(1865) seems to have had no personal acquaintance with G. cirrosa, but mentions that
Linnaeus gave twenty pairs of scales to his form, and that it requires re-investigation.

This species was procured at a depth of 230 metres on mud during the Austrian

North Polar Expedition. Mobius states that it occurs in Greenland in 4 — 12 fathoms.

In the account of the annelids of Nova Zembla, Theel (1879) says that forms 45 mm. in

length come from the Kara Sea. Yerrill (1879) includes it in his list from Cape Cod

1 ' Ann. Nat. Hist/ 4th ser., ix, p. 1, January, 1872.

EUNOA NODOSA. 291

to the Gulf of St. Lawrence. Finally, Malaquin (1890) describes a variety found in the
tubes of Ghxtopterus at Boulogne.

Genus X. — Eunoa, Malmgren, 1865.

Lateral tentacles arising under the median — below and slightly internal to the peaks
of the head. Palpi with six rows of short cilia ; eyes large, visible from the dorsum. Three
intestinal caeca directed forw r ard into the peripharyngeal space, the ventral long and
narrow, with two or more sacculations at its outer border inferiorly. The other two are
clavate and short. Elytra, fifteen pairs, completely covering the dorsum, and occurring on
the first foot, third, fourth, sixth, eighth, and so on to the twenty-second, twenty-fifth,
twenty-eighth, and thirty-first. Dorsal bristles pointed at the tip (which is bare), then
minutely spinulose in rows ; ventral bristles somewhat longer, and resembling those of
Harmothoe, with a smooth tip, which has a hook and transverse rows of spikes.
External aperture of the segmental organ indicated by blackish pigment ; no papilla.

Note. — This genus approaches Harmothoe, but the peaks of the head in the latter are
close to the median tentacle, whereas an interval exists in Eunoa. The eyes in
Harmothoe are smaller and less visible, the anterior pair being under the point of the
peak, whereas the anterior pair of Eunoa are situated some distance backwards, and are
lateral in position — not ventral. The bristles of the first foot in Eunoa are very distinct.
The remarkable Eunoa hispanica, 1 procured by the c Porcupine ' in 1870 on the Channel
slope, diverges from the other known forms by the great size of the eyes (with their
corneal lens), which exceed by far those of any other example of the family, by the great
length and smoothness of the palpi, and the great length of the ventral cirri and the
foot. The dorsal cirri probably have no enlargement below the tip, if we may judge from
a single lateral tentacle. E. hispanica is scarcely within the British area, but it is
worthy of note in this connection, and may yet be found near the coasts.

1. Eunoa nodosa, M. Sars, 1860.

Specific Characters. — Body broad, flattened, slightly narrowed in front, but much
more gradually and distinctly posteriorly; bristled segments, 36; head about as broad
as long, with a deep notch in front, from which the peaks are clearly marked off on each
side ; eyes large, two anterior to the nuchal fold, and two just in front of the lateral
projection of the head, and thus considerably behind the peaks ; tentacle ciliated, slightly
dilated below the slender tip, longer than the palpi — in preservation ; lateral tentacles
short, of similar shape ; palpi subulate, with six rows of minute papillae ; tentacular
cirri smaller than the tentacle, but of similar structure; scales, 15 pairs, completely
covering the dorsum, with the exception of the first pair, elongate reniform and some-
what thick, external margin densely ciliated, exposed surface rather thickly covered with

1 ' Trans. Zool. Soc./ ix, p. 396.

292 EUNOA NODOSA.

small tubercles and occasional parasitic growths, while a little within the posterior
margin is a row of larger, isolated, rounded tubercles ; dorsal bristles distinctly shorter
than the ventral. The dorsal cirri have the structure of the tentacular cirri shorter
than in E. oerstedi ; ventral cirri smooth. Segmental papilla cylindrical, passing from
the posterior border of the foot, and directed upward between the feet. A dilated
process occurs at the base of the dorsal cirrus.

Synonyms.
1860. Polynoe nodosa, Sars. Vid. selsk. Forhandl., 1860, p. 59.

1865. Lepidonotus pharetratus, Johnst. Cat. B. M., p. 113, pi. 3, f. 17 — 19.

1866. Antinoe zetlandica, Ray Lankester. Trans. Linn. Soc, xxv, p. 377, pi. 51, f . 13, 17, 18, 22, and 23.

1867. Eunoa nodosa, Malmgren. Nord. Hafs-Ann., p. 64, Tab. viii, f. 4, and Ann. Polych., p. 6.
„ Antinoe pharetratus, Parfitt. Cat. Ann., Devon, p. 18.

1872. Eunoe nodosa, Sars. Nyt. Mag. f. Naturvid., 19, p. 202.

1873. „ „ Sars. Bid. Christ. Fauna, p. 2.

1874. „ „ Malm. Goteborgs Kongl. Vet. o. Vitt. Samhalles Handl ; Ny. Tidsfoljd, Haft.

xiv, p. 74.
1876. Eunoa nodosa, Mcintosh. Trans. Z. S., ix, 374, pi. 67, f. 4—8.
1879. Polynoe scabra, Theel. Annel. Nov. Zemb., 7.
„ Eunoa nodosa, Tauber. Ann. Danic, 81.

1883. „ „ Wiren. Chset. Vega-Exped., 387.
„ Harmothoe nodosa, Levinsen. Nord. Annul., 193.

1884. Eunoa nodosa, Webster and Benedict. Ann. Mass., 700.
1886. Eunoe scabra, Marenzeller. Porif., &c, Jan Meyen, p. 11.

Habitat, — North Sea, Lieutenant Thomas ; Zetlandic Sea, Dr. Gwyn Jeffreys ; off
Holy Island, Tynemouth, 25 — 30 fathoms, Professor G. S. Brady ; stomach of the cod,
St. Andrews, E. M.

A fine example, from the collection of the late Dr. D. Robertson, comes from
Cumbrae, on the west coast of Scotland, but none have yet been received from the coast
of Ireland.

It ranges to 690 fathoms on the Channel Slope, ' Porcupine,' and 125 fathoms off
Cape Rosier in the Gulf of St. Lawrence (Whiteaves), and thence to Cape Cod (Verrill).
Malmgren gives the shores of Spitzbergen, Greenland, Finmark, and Scandinavia.

Length 1| in. to 2J in. (Clyde). Some of the foreign examples are about 2^- in.

Head (Plate XXVII, fig. 9). — Nearly as broad as long, with a median sulcus in front,
the sides trending to the peaks, which are free from the base of the median tentacle, the base
of which extends outwards from the sulcus. No specimen had a tentacle, but Malmgren
observes it resembles that of E. oerstedi, which is covered with long cilia with clavate
tips. Moreover, towards the swollen distal region, below the filiform tip, a few large
conical papillae with bifid tips are present — probably as an abnormality. This, however,
may differ considerably from the organ of E. nodosa — which may be less elongate, and
have shorter cilia. The lateral tentacles are short, with small clavate cilia and a filiform
tip. The tentacular cirri are similar to the median tentacle, but shorter. The eyes are
comparatively large, two being situated in front of the posterior border towards the
lateral region, and two just in front of the lateral projection of the head. The palpi

EITNOA NODOSA. 293

are subulate organs marked by six rows of conical papillae, which become larger
towards the tip of the organ.

Body large and broad, somewhat more rounded dorsally than ventrally, slightly
tapered anteriorly, very gradually but much more tapered posteriorly. Dorsally, it is
readily separated from other allied genera by the presence of the peculiar expansion
at the base of the dorsal cirrus. The bristled segments are thirty-six in number. The
shorter dorsal bristles at once distinguish the species when compared with E. oerstedi.
Dorsally, the segments are generally marked by a transverse bar often dimpled at the
outer edges, and by irregular papillae internal to the scale-bearing process. Ventrally, the
usual median groove marks the centre.

Posteriorly the body terminates in two caudal styles, which have a similar structure
to the cirri.

None of the spirit preparations show colour along either dorsal or ventral surface,
except a few touches on the area below the median and lateral tentacles; thus the species
is in contrast with E. oerstedi. The only pigment visible consisted of a brownish
hue on the lateral tentacles, and a brownish ring below the dilated region of all the
cirri.

Proboscis. — No example of E. nodosa has an extruded proboscis. E. oerstedi shows
nine dorsal papillae and nine ventral, and the teeth are powerful. In E. oerstedi three
gastric caeca pass forward to the dorso-lateral wall of the peripharyngeal space, the
fourth being nearly transverse. The longest is the ventral, which forms a narrow tube
anteriorly and extends to the fourth segment in front of the stomach. The posterior
half, however, is widened by two or three sacculations at its outer edge. The second
and third are shorter and wider clavate caeca, terminating in the corresponding spaces
behind the first. The stomach, if we may so term the anterior dilated part of the
intestine, has thick muscular walls, and its inner surface has a closely arranged transverse
series of firm glandular lamellae, interrupted at short intervals, so as to be crenate or
papillose. These ridges become less and less distinct as we proceed backwards, until
towards the vent the thin wall shows only the distinctly isolated papillae as in Halosydna
gelatinosa. It might be considered that the foregoing appearances of the stomach were
due to the condition of the preparation, but such did not seem to be the case.

The external opening of the segmental organ has no papilla, but is marked in
E. oerstedi by a blackish pigment-speck. In one example two specks were present on a
posterior foot. The genus thus diverges from HarwMhoe.

Scales. — The scales (Plate XXXII, fig. 3) of this form are distinguished from
E. oerstedi by their more leathery consistence, by their characteristically reniform
outline, by their longer and more abundant cilia on the outer edge, and by the
divergent character of the tubercles, which are much more developed in E. oerstedi. The
anterior scales, as usual, are rounded, but the typical scales are reniform with the free
parts studded with small horny tubercles and a row of from six to nine much larger
blunt horny tubercles, or short cones, some of which, a little within the posterior border,
present a bluntly spinose condition at the tip. The anterior and inner margins are smooth,
while along the posterior border a few isolated short cilia occur, gradually increasing in
length till they terminate at the outer edge in the long cilia with the clavate tips. In

294 EITXOA NODOSA.

E. oerstedi, again, the anterior edge of the typical scale (anterior third) is smooth, but
otherwise the entire margin has short clavate cilia scattered at intervals amongst the
tubercles and spines, the outer edge having these no more prominent than the inner.

Considerable variation exists in regard to the size and distribution of the smaller
spines over the surface. Thus in the figure from the specimen in the British Museum these
are comparatively small both towards the inner border and along the anterior edge, but,
on the general surface, the size increases while the number diminishes. The contrast
between the outer and the inner borders is marked, the former having fewer and larger
spines. The number of the large papillae along the posterior edge is nine to eleven,
which is more than usual, and some of them show spinous tips. The cilia are
confined to the outer edge. 1 An increase in the size of all the spines takes place in
an example from Spitzbergen, the larger forms on the general surface being proportionally
few. Only five large tubercles are present, with traces of spines at the tip of one or two ;
some sparsely distributed cilia occur along the posterior border as well as the denser
series of the outer edge. In fine specimens from the Gulf of St. Lawrence the propor-
tions of the various spines agree rather with the British forms, but a few short cilia are
found along the posterior border in addition to the longer outer series.

Colour. — Wo fresh example has been seen, so that all that can be said is that the
scales are marked with reddish-brown or madder-brown touches, darkest in the anterior
scales. Malmgren observes that the scales have a violet-brown hue, generally with a
white spot in the middle.

Feet. — The first foot — bearing the tentacular cirri — has a few (about three) bristles
which conform to the type of the dorsal, being stout, slightly curved, and spinous on the
convex margin, with a short, smooth tip, which affords a contrast to the more elongate
tip in E. oerstedi.

The second foot is distinguished, as usual, by its long ventral cirrus, and by
the diminution and modification of its parts. The dorsal division has bristles of
the ordinary type. The tip of these is short and smooth, and the curvature of the inner
bristles is marked. The same foot in E. oerstedi has much longer and less curved bristles,
and the smooth region at the tip is longer and has a different outline.

Ventrally is a group of much more slender bristles, with long spines on the distal
region and a needle-like, smooth extremity (Plate XXXVII, fig. 20). The tip of the
corresponding bristle in E. oerstedi is less elongate, and, instead of the finely-pointed tip,
it has the outline of a narrow knife-blade (Plate XXXVII, fig. 21).

In the third foot the dorsal bristles are smaller than in the typical foot, but have the
same structure. The ventral bristles, again, though slender, already show shorter tips,
which, further, are less acute than in the first foot.

The changes in the bristles have considerably advanced in the fourth and fifth feet,
so that the typical arrangement (Plate XLII, fig. 28), in which the longest dorsal
bristles reach a little beyond the spine of the ventral, is soon reached. These bristles
(Plate XXXVII, fig. 26) are shorter and more distinctly curved internally, the longer and

1 A large Zetlandic example, the Antinoe zetlandica of Prof. Eay Lankester, corresponds in the
main with the foregoing, though the large spines posteriorly extend along the outer border, and,
like the former, the cilia are confined to the outer edge.

EUNOA NODOSA. 295

straighter forms being external, that is, next the ventral. The tips have the character
shown in fig. 24, Plate XXXVII, viz. less tapered than in E. wrstedi (Plate XXXVII,
fig. 25) — with a shorter smooth portion and more boldly marked spinous rows.

The ventral bristles (Plate XXXVII, figs. 22 and 27), on the other hand, have pro-
portionately longer spinous regions superiorly than in E. wrstedi ; the tips are more
distinctly hooked, and the bare portion is broader on an average than in the species just
mentioned (Plate XXXVII, fig. 23). The latter is not what might have been expected
from the condition anteriorly. The ventral cirrus extends a little beyond the inferior
border of the foot.

Posteriorly both dorsal and ventral bristles become more attenuate, as well as smaller,
in conformity with the diminished feet, but they do not assume the elongate and slender
condition observed anteriorly.

Two very large examples from the stomach of a cod agree with the fine Zetlandic
specimen in having the smooth tips of the dorsal bristles somewhat shorter. On the other
hand, young specimens (about J of an inch) have longer tips to the dorsal and ventral
bristles, and the scales are more rounded and have proportionally thicker cilia.

While the scales in the examples from Spitsbergen agree in shape and structure
with those from the Gulf of St. Lawrence and Britain, the dorsal bristles have
assumed a more elongated condition, so as to resemble those of E. aerstedi, being
more tapered, and with a longer bare portion at the tip ; yet the general curve of these
bristles is more pronounced than in E. wrstedi. The tips of the ventral bristles are also
proportionally longer than in the ordinary type. The typical characters of E. nodosa
are seen in examples from the Arctic seas (Greenland), where the species attains a large
size. The distinctions between the two forms are therefore evident, but whether two
species should be formed may be to some an open question.

A parasitic sponge, with long processes, occurs on a scale of the example from
Cumbrae.

Habits. — So far as can be ascertained, the present form is only found in deep
water off the British shores, and that but rarely. It is more abundant in the northern
seas of both Europe and America. Loxosomse abound on the feet and bristles of those
from the Gulf of St. Lawrence.

Reproduction. — The only example of the genus observed with fully developed
reproductive elements is a large female of E. oerstedi, from Greenland, distended with
ova, and probably procured in July.

This is one of the many forms that marine zoology owes to the industry and keen
observation of the elder Sars, who patiently explored the shores of his native country
for so many years and with such remarkable success.

It is difficult to know to what Sir J. Dalyell (1853) 1 refers under the name of
Aphrodita squamata, from Shetland, unless it be this species. Theel (1879) noted that
Eunoa nodosa and E. aerstedi, Malmgren, were the same species, viz. the Lepidonotus scabra
of (Ersted, an opinion coinciding with the remarks made by myself many years previously.
He procured large examples (70 mm.) at a depth of 90 — 200 metres in the Kara Sea (Nova

1 ' Pow. Creat./ ii, 166, pi. 24, f. 3, 4.

39

296 EITNOA TRITONI.

Zembla). The processes on the scales vary considerably in size, but he did not
sufficiently discriminate between the forms.

The Polynoe islandica of Dr. Hansen (1882) 1 is probably either a variety of this
species or E. oerstedi, in which the tentacles are smooth. His P. arctica* I would also
be inclined to unite with the same form. Nor does any other conclusion seem to present
itself with regard to his P. assimilis* his P. spinulosa* or his P. foraminifera. 5 From the
inexperience of the artist, Dr. Hansen's figures are not reliable, the tentacular cirri, for
instance, springing as a single trunk which becomes bifid, a condition which can only
occur as an abnormality.

Wiren, in his account of the annelids of the 'Vega' Expedition, expresses the same
view as Theel and the writer, and he gives notes on five varieties, with figures of
three examples of scales, which differ considerably in regard to the papillge.

2. EUNOA TRITONI, Mcintosh.
1898. Eunoa Tritoni, Mcintosh. Ann. Nat. Hist., Aug., 1898.

Specific Characters. — Head somewhat pyriform, with the broad end posteriorly,
a median furrow and two minute peaks close to the base of the median tentacle.
Eyes equal, of moderate size, placed on each side of the lateral eminence. Lateral
tentacles short and densely ciliated. Palpi with rows of minute clavate cilia. Body
normal ; segmental papillae commence on the sixth foot and continue almost to the
posterior end. Scales reniform in outline, densely fringed with long cilia on the outer
edge. The surface of the scale has a series of capstan-like tubercles with a minutely
nodular surface towards the posterior border, the general surface being studded with
minute spines. Dorsal and ventral divisions of the foot have each a long process for the
spine. The dorsal bristles are long, little tapered at the tip, which forms a short blunt
cone invaded by the spinous rows so closely that the tip of the bristle is almost reached.
Ventral bristles with slender shafts, spinous tips of the average length, and a long and
rather broad terminal region, with a well-marked hook.

Trawled on board H.M.S. 'Triton,' at Station 8 (Faroe Channel?), 22nd August,
1882, in 640 fathoms.

A comparatively large species, about 34 mm. long.

Head. — Somewhat pyriform, with the broad end posteriorly, a median furrow in
front, and two minute peaks close to the base of the median tentacle. Eyes of moderate
size, equal, visible from the dorsum, placed on each side of the lateral eminence, and thus
towards the middle of the region. The anterior pair look slightly forward and outward.
Median tentacle absent. The lateral are comparatively short and densely ciliated. The
palpi have rows of minute clavate cilia. Tentacular cirri absent.

1 c Norweg. N. Atlantic Exped./ p. 24, pi. i, figs. 15—21.

2 Ibid., p. 27, pi. iii, f. 1—5.

3 Ibid., p. 27, pi. i, f. 22—26.

4 Ibid., p. 28, pi. i, f. 6—10.

5 Ibid., p. 29, pi. i, £. 11—14.

EUNOA TKITONI.

297

Body. — Convex dorsally, flattened ventrally, slightly tapered anteriorly, and more
gradually posteriorly. Bristled segments about 40. The bases of the dorsal cirri show
an expansion. The markings on the dorsum correspond with those of allied species.
The segmental (nephridial) papillae are visible on the sixth foot, and are directed
upwards between the feet. They continue almost to the posterior end.

The scales are more nearly allied to those of E. nodosa than to those of E. cerstedi, being
somewhat reniform in outline, densely fringed on the outer or narrower edge with loner
cilia ending in a probe-point (Fig. 21). Posteriorly these gradually diminish and run into

Fig. 22.

Fig. 21.

Q •

Outer edge of scale of JEunoa tritoni.

JEunoa tritoni.— Dorsal bristle to right and ventral to left.

the short cilia which occur at intervals along this border to the inner edge. Besides the
cilia a series of eight or nine capstan-shaped tubercles, with a minutely nodular
surface, project from the posterior border, besides others within it, and, in addition,
two or sometimes three much larger tubercles of a similar shape are ranged a little
within the posterior border. These larger tubercles are thus fewer in number
and proportionally larger than in E. nodosa. Just within the densely ciliated outer
border are a series of elongated papillae, with trifid or multifid tips (Fig. 21). They

298 LAGISCA FLOCCOSA.

extend along the outer border, and also form a group in the neighbourhood — within the
anterior edge. The anterior border of the scale is smooth, with the exception of an
isolated group of one to five or more long cilia some distance from the outer edge. The
entire surface of the scale is densely studded with minute spines, smallest in front and
increasing in size posteriorly. The developing scales are of various degrees of smooth-
ness, the youngest being quite smooth.

Both dorsal and ventral divisions of the foot bear a long pointed process for the
spine. The dorsal division has characteristic bristles (right of Fig. 22), which somewhat
approach those of Harmothoe haliaeti, though quite distinct. They are of considerable
length, slightly curved, and not much tapered at the tip, which, indeed, ends in a short
blunt cone, and in some shows a slight fold or differentiation on the spinous side. The
short bare cone at the tip is much broader than that of H. haliaeti, and has a different
character. The spinous rows so nearly approach the tips that they in some almost
reach as far as the point of the bristle, and are as close as in the species just
mentioned. The ventral bristles (left of Fig. 22) have somewhat slender shafts, spinous
tips of the average length, and a long and rather broad terminal region with a well-
marked hook. The ventral line of the smooth tip is slightly convex.

The dorsal cirri are of considerable length — with a filiform tip. They are densely
covered with long cilia having probe-pointed tips. The ventral cirri have only a few
scattered and very short clavate cilia.

This fine species appears to be confined to deep water.

Genus XL — Lagisca, Malmgren, 1865.

Body rather narrow, attenuate posteriorly. Cephalic lobe produced into lateral
peaks. Lateral tentacles attached below the median. Eyes four; two lateral Justin
front of the middle line, two in front of the nuchal border. Scales fifteen, covering the
dorsum— except the last few. Bristles of the dorsal lobe with acute tips and finely
serrated. Bristles of the ventral lobe simple superiorly and inferiority, rest bidentate,.
and all with long spinous rows. Papilke of proboscis f. Two slender cseca go forward
into the peripharyngeal space. Segmental (nephridial) papilla commences as a minute
process on the fifth foot, and extends nearly to the last foot. It is much smaller than in
Harmothoe.

1. Lagisoa floccosa, Savigny, 1820. Plate XXVI, fig. 2.

Specific Characters. — Body rather narrow throughout, slightly diminished anteriorly,,
and much more so posteriorly. Median and lateral tentacles, and the tentacular cirri
rather short, densely ciliated and with filiform tips ; little or no enlargement below the
latter. Byes large, posterior in front of the nuchal border ; anterior lateral in position,
and just in front of the middle line. First pair of scales rounded, the others reniform or
somewhat ovate, mottled greyish brown, often with a white spot in the centre, densely

LAGISCA FLOCCOSA. 299

covered with minute spines (visible under the microscope), and having posteriorly a series
of soft globular papillse arranged at intervals just within the margin. Cilia on the outer
border — very few and short. Dorsal bristles with acute tips and finely serrated ; ventral
bristles with the tips simple superiorly and inferiorly, rest bidentate. Alternate spinous
rows long, though the bristle-tip is proportionally short. Dorsal cirri with numerous
and rather short cilia. Ventral cirrus smooth.

Synonyms.

1820. Polynoe floccosa, Savigny. Syst. Annel., pp. 22, 23, and 27.

1828. Eumolpe floccosa, Blainville. Diet, de Sc. Nat., vers., 459.

1834. Polynoe floccosa, Aud. and M. Edw. Ann., 88.
1851. „ „ Grube. Fam. d. Ann., 37 and 120.

1865. ,, „ De Quatrefages. Ann., 236.

„ „ semisculptus, Johnst. Cat. Brit. Mus., p. 116, pi. v, f. 1 — 11, and pi. vi, f. 4 — 6.

1867. Lagisca propinqua, Malmgren. Ann. Polych., p. 9, Tab. i, fig. 3.

1870. Polynoe floccosa, Grube. Archivf. Naturges., 287.

1873. Lagisca propinqua, Sars. Bid. Christ. Fauna, p. 3.

1875. Polynoe floccosa, Mcintosh. Invert, and Fish., St. A., p. 116.

„ Mcintosh. Trans. Z. S., ix, 380, pi. 68, f. 6, 8, 10.

1876. Lagisca propinqua, Mcintosh. Trans. Z. S., 375, pi. 67, f. 12 — 14.

1879. „ „ Tauber. Ann. Danic, 81.

1880. „ „ Langerhans. Die Wurmfauna Madeiras, Zeit. f. w. Zool., xxxiii, p. 274,

Taf. xiv, fig. 3c.

1882. Polynoe semdsculpta, Hansen. Norweg. N. Atl. Exped., 26, pi. iii, f. 16 — 20.

1883. Harmothoe propinqua, Levinsen. Nord. Annul., 192.
„ „ semisculptus, Ibid.

1886. Lagisca propinqua, Harvey-Gibson. Verm. Liverpool, 148.

„ „ „ Langerhans. Zeit. f. w. Zool., xl, 251, Taf. 15, f. 6.

„ Polynoe floccosa, Harvey-Gibson. Yerm. Liverp., 150.
1891. ,, (Lagisca) propinqua, Hornell. Op. cit., p. 234, pi. 13, f. 1, 7, 9.
1896. Lagisca propinqua, var. abyssorum, Roule. Camp. d. ' Caudan/ 444.
1898. Harmothoe semisculptus, Michaelsen. Gronland. Annel., Bib. Zool., Heft 20, Lfg. 4, p. 121.

Habitat. — Everywhere on the British shores from Shetland to the Channel Islands,
where it is the chief form between tide-marks. It also ranges from 600 fathoms to low-
water mark. It extends to the Gulf of St. Lawrence in Canada, as well as to Norway
and Sweden. It was procured likewise during the ' Vega ' Expedition, and apparently
also is found in Greenland.

Length 1-J inches, breadth about \ inch.

Head (Plate XXVII, fig. 14) with a median furrow anteriorly, which passes on
each side of the base of the tentacle to the peaks. The black eyes are comparatively
large, the posterior pair being dorsal in position and somewhat nearer each other than the
anterior pair. The latter are more or less lateral in position just in front of the prominent
median region. They are slightly larger than the posterior pair. Both pairs show in
some specimens a cuticular lens, like a central speck. The median tentacle is rather short,
with a long filiform tip and a slight swelling below the latter. It has short clavate cilia.
The lateral tentacles are also short with similar cilia, and a long filiform tip. They have

300 LAGISCA FLOCCOSA.

a dark bar at the base, then a pale region with another bar below the filiform tip. If
colour is to be relied on, the pale part would be homologous with the slightly swollen
region in other forms. The tentacular cirri are similar to the median tentacle. All these
cirri have the slightly enlarged part pale, with a brown band on each side, and the
column below is brownish. The palpi show minute papillse-like serrations along the
sides under a high power, probably partly due to the rigid contraction. They thus
differ materially from those of Harmothoe imbricata with the minute clavate cilia.

Th e Body is narrower than in Harmothoe irnbricata, and the line of bristles on each
side is straight and trim ; moreover, it has a general firmness which is characteristic.
The anterior end diminishes from the eighth or ninth bristled segment towards the head,
while posteriorly the diminution, which is gradual, begins shortly behind the middle, and

Fig. 23.

Papillae and teeth of the proboscis of Lagisca floccosa.

the body terminates in a process bearing the two caudal cirri beneath the vent. The
number of bristled segments is about forty-two. Dorsally the latter are marked from
the head backward by a median series of brownish touches, which posteriorly show a
somewhat symmetrical arrangement in the centre of each segment, consisting of an
anterior, narrow, curved bar, a fan-shaped region divided by a median streak and a short
transverse bar posteriorly. A prominent dark patch also occurs on the papilla (homo-
logue of scale-papilla) and a little pigment at the base of the dorsal bristles. The ventral
surface is pale and iridescent ; the nephridial papilla is smaller than in Harmothoe irnbricata,
and passes into the fissure between the feet. A considerable portion of the tail is devoid
of scales, and it is this region which has the pigment on the dorsum best developed.
The segmental (nephridial) papilla commences on the sixth foot, and extends to the
posterior feet as a very minute process.

Broboscis. — The extruded proboscis (Fig. 23) has the usual teeth, the lower pair biting
to the right of the upper, nine flattened conical papillas dorsally, and nine ventrally at the
margin. The skin-folds at the mouth are marked by brownish pigment. The pre-gastric
casca are short and small. The ventral alone is conspicuous in the preparations, for the
second proceeds little beyond the stomach, and the third is nearly transverse.

Scales (Plate XXXII, fig. 5) — fifteen pairs. First pair rounded, the others reni-
form or ovate-reniform, the last pair being ovoid, of a brownish-red colour, or sometimes
slightly purplish marbled with grey. They generally show a pale area over the scar for
the pedicle, and from the depth of the colour this is best marked anteriorly, the pigment
in the posterior scales being broken up into a series of touches and granules. The

LAGISCA FLOCCOSA. 301

entire surface, with the exception of a small area at the anterior and inner border, is
densely covered with minute spines, which are longest towards the outer border. The
posterior edge, again, has just within the margin a series of soft globular papillae
arranged at intervals and visible under a lens. Some are slightly clavate in outline,
while the tip in others is truncate or slightly nodular. They are few in number on the
last scales. A few short slender cilia occur at the outer or posterior margin. The
under surface is iridescent, and the scar for the pedicle is near the hilus or indentation,
where it is present, on the outer border. The first pair of scales have a few short cilia
on the inner border, and a well-marked series of clavate cilia on the outer and anterior
margin. The large globular papillae on these are from four to six in number. A variety
procured by the ' Knight Errant ' at Station 2 (1832) is pale, with few and small tubercles
on the posterior border of the scales.

Feet. — The first foot has two dorsal bristles, one curved and less pointed, the other
straighter and more acute. The spinous rows are more distinct than in the typical foot.
The tip in both is mioute.

The second foot has a dense tuft of dorsal bristles, generally more curved and with
more distinct spinous rows than in the typical foot, and the bare portion at the tip is
proportionally broader and better marked. The slender ventral bristles have rows of
long spines and simple tips, those at the ventral edge of the series approaching the
pinnate type seen in other forms. They do not project beyond the dorsal bristles.

The third foot has the dorsal bristles less curved and the tips more acute. In the
ventral division the bristles have increased in strength, and, while the upper and lower
series have simple tips, the rest have the secondary spur. The tip of the smooth ventral
cirrus is long and filiform.

In the fourth foot the ventral bristles have become more prominent by the increase
of the fleshy part of the foot, and some simple tips still occur superiorly, and a more
numerous series inferiorly. The spinous rows are long. The dorsal bristles are
elongating, the long outer ones having acute tips, the inner broader smooth tips.

In the typical foot the dorsal division has long and very slightly curved bristles with
gently tapered acute tips throughout, the sharpest tips as usual being external (on a slide
next the ventral). They are thus easily distinguished from those of Harmothoe imbricata,
and also by the much closer spinous rows. Bristles of the same length are decidedly
more slender than in H. imbricata. One of the longer forms is represented in Plate
XXXVIII, fig. 2. The tip tapers to a smooth, blunt point, and immediately below the
latter very fine and close spinous rows occur. A glance at the latter in rapidly examining
specimens is one of the most satisfactory points in discrimination. The ventral division
bears superiorly a series (Plate XXXVIII, fig. 1) with long spinous extremities, more
slender and with longer spinous rows than in H. imbricata. In a few the smooth tips
have no secondary process. The next series has a small secondary process, and the
spinous border gradually diminishes in length. The bifid tip differs from that of
H. imbricata, especially in regard to the minute size of the secondary process. Inferiorly
the tips are again simple. Posteriorly the structure of the dorsal bristles remains
nearly the same, but the ventral become attenuate, the bifid tip being visible, but the
secondary process is minute. The dorsal cirri have a dark bar above and below the

302 LAGISCA FLOCCOSA.

slight enlargement towards the tip, and the latter is attenuate. The cilia are numerous
rather short and clavate, and extend beyond the distal pigment-bar. They become more
slender posteriorly, but have the same structure. In these translucent organs the areoke
of the hypoderm (epidermic granular layer) are very visible. The long first ventral
cirrus is ciliated like one of the dorsal, and some of the succeeding ordinary forms
show a few cilia. These gradually disappear and leave the cirrus smooth throughout
the greater part of the body.

Some from Guernsey have the dorsal bristles covered with a reddish-browm or dull
orange granular coating, from the surface of which minute algoid filaments project.
Such would seem to be parasitic. A similar growth occurs on the dorsal bristles at
St. Andrews.

In my notes of November, 1873, at St. Andrews, this species (and the preparation
accompanies) is described as phosphorescent, but recent examination, both when laden
with ova in January and in the warmer months, such as July, has not borne out this
description. In the preparation the bristles, especially the dorsal, are covered with a
parasitic granular growth.

Though this form is more plentiful in the south, the size of some dredged off Balta
is quite as large as elsewhere.

In a specimen from Lochmaddy in which five of the posterior feet of the right side
had been removed, long papillse from the dorsal border (cirri?) and shorter papillae
from the ventral indicated regeneration.

Reproduction. — A specimen from St. Andrews in November was laden with ripe ova,
so that the breeding season would seem to be in winter, as in the case of Harmothoe.

Variety. — A very distinct variety comes from the ' Porcupine/ 1869, 173 fathoms,
on muddy sand, bottom temperature 49*6° ; from 45 fathoms, eight miles N.W. of Cape
Sagres, ' Porcupine,' 1870 ; off the Hebrides, 1866, Dr. Gwyn Jeffreys ; West Sands, St.
Andrews, 1867 (R. M.) ; and 90 — 25 fathoms, 30 — 50 miles west of Valencia, Ireland
(Prof. Haddon). Abundant in various parts off Shetland, Dr. Gwyn Jeffreys, 1867-8.

The pigment on the scales is similar ; that on the naked segments posteriorly, how-
ever, diverges, for it forms a single fan with a posterior bar only. Moreover, the
papilla for the dorsal bristles has black pigment. No appreciable change is observable
in the structure of the scales. The distinguishing feature is the condition of the dorsal
bristles, which, instead of forming the elongated tip so characteristic of the typical
form, end bluntly, so that the whole tip is broad, with a very short, bare portion at the
tip. The spinous rows are as closely arranged as in the normal form. The ventral
bristles and cirri do not differ from the typical example.

A form dredged by the 'Porcupine' in 173 fathoms, 1869, and in 160 fathoms, 17th
August, 1870 (No. 42), differs from Lagisca floccosa and its variety in having no distinct
bosses or globular papillse on the scales posteriorly, while considerable papilla3 occur
anteriorly ; the scales, moreover, having much larger spines, which show a tendency to
be arranged in rows in the reniform scales, the largest being near the posterior border.
In the first pair of scales the spines are larger, and the short cilia extend at intervals all
round, while a few short cilia occur along the outer and posterior edge. The ventral cirri
have a few clavate cilia. The bristles of the dorsal branch of the foot (Plate XXXVIII,

LAGISCA ELISABETHS. 303

fig. 3, an average form) have the spinous rows much more distinct than in L. floccosa or
its variety, and the smooth portion at the tip is small and acutely pointed. The ventral
bristles do not much differ. The anterior pair of eyes are proportionately much larger
than in L. floccosa.

The Polynoe foliosa of Savigny (1820) seems to come near this species.

De Quatrefages (1865) describes it as having a small head, almost quadrate, a long
thick median antenna, the lateral small and slender. Segments 40 — 42. Scales large,
much imbricated and decussate, rounded, smooth, not ciliated, caducous. He procured
it, 42 mm. long, on the oyster-banks at St. Yaast. When living it is of a brownish-white
colour (reddish brown ?), and thus is readily distinguished from its congeners.

Grube found a species at St. Vaast, where De Quatrefages had met with his, with
only fifteen pairs of elytra, which, however, had cilia on their border ( u Am Aussenrande
gefranzte Elytren besitzt "). If such be so, then the species differs from Lagisca floccosa,
in which no cilia are present on the scales.

Hornell says his specimens possess fifteen pairs of elytra, with only subglobular
processes on the margin (whereas P. floccosa has clavate). He thinks Malmgren's artist
exaggerated the spikes on the dorsal bristles, making them too coarse. His specimens
agreed in colour with Malmgren's L. propinqua, but differed from mine. He figures one
of the globular papillee near the margin of the scale.

L. Poule found what he thinks a variety at depths ranging from 650 metres to
1700 metres in the Atlantic, with small eyes and pale scales. Its relationship to other
closely allied forms has yet to be determined.

2. Lagisca Elisabeth^, 1 Mcintosh, n. s.

Specific Characters. — This species has 30—35 segments, but a considerable portion
of the tail is absent. Head curiously mottled with black. Eyes black, nearly equal; a
pair at the anterior border, and the other on the lateral prominence of the head.
Tentacles and tentacular cirri ciliated, and the ventral cirri have also short clavate cilia.
Palpi with a dense series of minute papillaB with enlarged tips. Scales, probably fifteen
pairs, scabrous, greyish mottled with black, densely spinous, with one or two large conical
processes posteriorly, and the outer and posterior edge fringed with club-shaped cilia.
Dorsal bristles with well-marked spinous rows, and a minute bare portion at the tip.
Ventral bristles mostly bifid. Akin to L. floccosa.

Synonyms.

1875. Lagisca propinqua, Mcintosh. Invert, and Fish., St. A., p. 115.

1876. „ „ Ibid. Trans. Zool. Soc, ix, p. 375, pi. lxvii, figs. 12—14.

Habitat. — Procured from the debris brought by fishing-boats from the off-shore
waters, St. Andrews, 1870 (E. M.).

Head (Plate XXVII, fig. 3). — The head is curiously marked, for a pale band of
considerable breadth occurs posteriorly — boldly defined by the blackish collar. A pale

1 Named after the best benefactress in Marine Zoology my museum ever had.

40

304 LAGISCA ELISABETHS.

belt runs from this forward in the median line to the base of the tentacle, which is
blackish ; and the anterior areas of the head thus mapped off are brownish red, with dark
grains along the edges. The anterior pair of eyes are lateral, lying in front of the
brownish-red border of the region. The posterior pair are large, black, widely separated,
and situated on the pale band of the region. The median tentacle is absent. The
lateral tentacles are short, brownish at the base, with a filiform tip, and furnished with
long clavate cilia. The palpi have a dense series of minute papilla^ with enlarged tips.
The tentacular cirri have a blackish patch at the base, a light brownish one in the
slightly dilated portion near the tip, then a whitish ring, and lastly, a dark brownish
one at the base of the filiform termination. They possess long cilia with globular ends.

Body. — The body is typical, that is, slightly narrowed in front, more so posteriorly,
about 13 mm. in length, and mottled with blackish pigment on the dorsum in a
remarkable way. The dorsum of the first four segments is mainly blackish, then each
segment presents near the posterior border a median bar, which by-and-by becomes a
speck. Symmetrical touches of pigment mark the lateral regions of each segment, and
two blackish specks occur on the base of the foot. A well-marked median band of
black characterises the dorsal lip; the ventral surface is otherwise pale. The pale scale-
pedicles are very distinct, and proportionally large.

The dorsal cirri agree in colour with the tentacular cirri, and besides the longer cilia
with globular ends on the column of the organ, shorter cilia proceed upwards rather
beyond the lower third of the extremity. The ventral cirrus is subulate, with sparsely
distributed and short clavate papillae. The first has a dense coating of cilia, with large
globular tips.

Scales (Plate XXXII, fig. 6) probably fifteen in number, though the last pair
had disappeared in the specimen. The first pair are rounded, densely spinous, and
ciliated almost entirely round ; the cilia being longest externally, and with characteristic
globular ends. The rest of the scales are more or less reniform, becoming ovate pos-
teriorly. The general colour of the scales is dark greyish with a blackish patch in the
centre, the pigment being broken into fragmentary portions. The outer and posterior
edge is ciliated, as indeed is the greater part of the circumference. They commence as
short, almost baccate processes, and towards the outer edge are more elongate, the
extremities being nearly globular, the series again diminishing to terminate in short
papillae. With the exception of the covered portion of the scale the surface is densely
covered with minute and rather blunt spines, a few of which towards the outer and
posterior edge become larger acute processes, or bluntly conical papillae. A considerable
portion of the inner region of the scale is curiously reticulated, so that the spines are
grouped in areas — a condition visible under the lens as well as the microscope over the
greater part of the scale. The anterior and outer border has larger spines than those
on the general surface, as shown in the figure. From the extreme roughness of the scale,
mud and debris lodge in the crevices.

Feet. — The first foot has two curved bristles which correspond in structure to the
dorsal type, though the smooth tips are somewhat broader.

The dorsal bristles of the second foot are nearly typical, except that the smooth tips
are larger. The ventral are more slender than the subsequent forms, but show a bifid tip.

LAGISCA JEFFREYS!!. 305

The dorsal division in the typical foot bears a somewhat dense mass of rather short
pale bristles with a slight curvature. The tips are short and by no means acute
(Plate XXXYIII, fig. 4, representing one of the longer forms). The spinous rows are
much more distinct and longer than in Lagisca floccosa, and the shape of the bristle
differs. The ventral division has translucent bristles with moderately long shafts. The
tips of the superior series (Plate XXXVIII, fig. 5) are long and somewhat tapered, with
rather distant rows of long spines, the smooth terminal region being minutely bifid.
The tips gradually become shorter and stouter inferiorly (Plate XXXYIII, fig. 6), the
strongly curved terminal hook and the secondary process with its characteristic angle
of incidence being noteworthy. Some of the latter bristles show traces of an outward
curve between the secondary process and the first row of spines. Towards the ventral
border the secondary process diminishes with the general size of the bristle, but a
minute trace occurs in almost all.

The dorsal bristles in most feet are densely coated with debris and minute
filamentous algoid growths, and in some cases the tips of the ventral are likewise
encrusted with a parasitic structure showing minute rods.

This species approaches the Polynoe aspera of Hansen l from the Norwegian North
Atlantic Expedition, and of Theel, 2 from Nova Zembla, but differs in regard to the palpi,
which are smooth in the northern form, and also in the shorter tips (bare) to the dorsal
bristles. It may be that further examination will show they are identical.

3. Lagisca Jefpreysii, 3 n. s.

Specific Characters. —Length 16 mm. or more. Head more elongate than in Lagisca
floccosa, with blunt anterior peaks. Posterior pair of eyes just in front of the collar,
anterior pair somewhat further forward than in L. floccosa, and lateral in position.
Median tentacle absent ; lateral subulate, and slightly beneath the base of the former.
Tentacular cirri slender, and with a series of clavate cilia which commence only when
the basal third of the process is reached. Palpi have rows of minute papilla with
nodular tips. First scales rounded and minutely spinous, rest ovate-reniform, with a
distinct fold from the scar of attachment to the hilus at the anterior border, speckled
throughout the posterior half with pale specks as if variolated. The outer border has
long cilia. Dorsal bristles of moderate length, with well-marked spinous rows, and a
very short smooth tip. They are nearly straight, Yentral bristles with rather short
spinous rows, and a short bare tip with a strongly curved hook at the end, and a
secondary process— directed nearly straight— distally. Yentral cirrus with clavate cilia.

Habitat— Dredged in sixty fathoms, nine miles off Balta, in 1868, by Dr. Gwyn
Jeffreys.

1 'Nat. Mag. f. Naturvid./ 24, p. 4, 1877, and < Norsk. Nordh. Exped.,' vii, p. 5, pi. ii f 11—
15, 1882.

2 < Annel. N. Zemb./ p. 10, pi. i, f. 1—4, 1879.

3 After the late Dr. Gwyn Jeffreys, a veteran explorer of the Zetlandic seas. The name was
formerly given to the succeeding species— now associated with P. extenuata, Grube.

306 LAGISCA JEFFKEYSII.

Length 16 mm., but the specimen had the posterior extremity regenerated, and
therefore was probably considerably longer.

The head (Plate XXVII, fig. 10) is more elongate than in L. floccosa, and the
peaks in front terminate bluntly. The posterior pair of eyes are of moderate size, and
lie just in front of the collar; the anterior pair are somewhat further forward than in
the species mentioned, are larger than the posterior pair, have a lens-like centre, and
are more or less lateral in position. The median tentacle is absent. The lateral tentacles
are short and subulate, being lateral in position, and only slightly beneath the bases of
the former. They have a series of cilia with clavate tips. The tentacular, like the dorsal
cirri, are rather slender, with a filiform tip and a series of clavate papillae, which do not
commence till the basal third of the process is reached, and extend within a short
distance of the filiform tip. They a,re in moderate number, and diminish at each end of
the series. The palpi are pale brown, and have rows of minute papillae with expanded
nodular tips. Two of these rows are dorsal.

The body is somewhat narrowed anteriorly, and posteriorly appears to have been
recently regenerated after the twenty-fourth foot. The only colour exists in the dorsal
fold of the mouth, which is brownish.

Scales (Plate XXXII, fig. 7). — The first pair of scales are rounded, and under a lens
are minutely speckled as if variolated. This condition, however, is due to hypodermic
structure, and does not affect the surface. The latter has a series of minute spines,
which densely cover the outer and posterior part of the scale. Even in these scales a
distinct fold occurs anteriorly, but no cilia on the edge. The other scales present
are rather large, ovate-reniform in outline, and have a distinct fold from the scar of
attachment to the hilus at the anterior border. They are faintly brownish in hue (in
spirit), and on a dark surface — under a lens — are speckled throughout the posterior half
with pale specks which simulate pustules. The outer border has a well-marked series of
long cilia, which are not dilated at the tip. The smoothness of the edge of the scale,
with this exception, is noteworthy.

Feet. — The second foot shows rather short and straight dorsal bristles, with distinct
spinous rows as in Lagisca floccosa, var., from the 'Porcupine,' 1 with a very short
smooth portion of a bluntly conical shape at the tip. The ventral bristles are slender,
with elongate spinous regions and attenuate tips.

In the third foot the ventral bristles are stronger, and the bifid condition distinct in
many — the secondary process passing nearly straight towards the tip.

In the typical foot the dorsal bristles (Plate XXXVIII, fig. 7) are of moderate
length, with well-marked spinous rows and a very short smooth tip. They are nearly
straight, with the exception of a few at the inner border of the tuft. Loxosomse and algoid
growths are common on these bristles. The ventral division has a dense group of pale
bristles with elongated shafts, and, though the upper forms (Plate XXXVIII, fig. 8)
have long tips, on the whole with rather short spinous regions, as shown in one from
the middle of the foot (Plate XXXVIII, fig. 9). These bristles have very distinct spinous
rows, a short bare tip with a strongly curved hook at the end, and in the majority a
secondary process which is directed nearly straight distally, so that they differ quite from

1 Vide p. 302.

LAGISCA EXTENUATA. 307

those of Lagisca floccosa, which have longer spinous regions and a different curve at the

tip. The ventral cirrus has well-marked clavate cilia. The segmental papilla is minute.

In the presence of the pale specks on the scales it resembles the rare Polynoe nivea

of Sars, 1 but the absence of cilia in the latter species and other characters distinguish it.

4. Lagisca extenuata, Grube, 1840 (?).

Specific Characters. — Length about one and a quarter inches in spirit. Head with a
deep median groove and two prominent peaks ; eyes comparatively large, two in front
of the nuchal collar, and two larger in front of the middle line, and lateral in position.
Median tentacle somewhat long, scarcely dilated below the filiform tip, and with
moderately long cilia. Lateral tentacles short, with attenuate tips. Tentacular cirri
similar to the median tentacle. Palpi have rows of minute conical papilla?. Scales fifteen
pairs, rounded in front, reniform posteriorly, densely covered with minute spines, and the
free edge is profusely ciliated. Dorsal bristles strong, slightly curved, and closely
spinous, with a smooth spear-shaped tip ; ventral with a bifid tip and close rows of
spines, the tips of some at the ventral edge being simple. Dorsal cirri somewhat slender,
with numerous and slightly tapered cilia with bulbous tips. Ventral cirri stout, with
short cilia. Segmental (nephridial) papilla scarcely distinct.

Synonyms.

1840. Polynoe extenuata, Grube. Actio. , Echin., u. Wurmen p. 86 (?).
1861. „ cirrata (0. F. M.), Grube. Ausflug nach Triest., p. 81.
1865. Lepidonotus Leachii, De Quatrefages. Ann., p. 258 ?
„ „ dumetosus, Ibid. Annel., t. i, p. 259 (?).

1867. Lagisca Ehlersii, Malmgren. Ann. Polych., 9.

1868. Polynoe extenuata, Claparede. Ann. Nap., 70, pi. ii, fig. 2.
1870. „ „ Supp., p. 372.

1875. „ „ Marion and Bobretzky. Ann. Sc. Nat., 1875, p. 6.

„ Lagisca extenuata, Marenzeller. Zur Kennt. Adriat. Annel., p. 5, Taf. i, f. 1.

1876. „ Jeffreysii, Mcintosh. Trans. Z. S., ix, p. 397, pi. lxxi, f. 8, 9, 11, and 12; pi. lxxiii,

f. 17 and 18.
1884. ,, extenuata, V. Cams. Fauna Medit., 202.
1888. „ „ De St. Joseph. Ann. d. Sc. Nat. (7), v, p. 180, pi. viii, f. 52—54.

1890. „ „ Malaquin. Ann. Boulon., 22.

1891. Polynoe (Lagisca) extenuata, Hornell. Op. cit., 12, pi. xiii, f. 4 and 8.
1898. Lagisca extenuata, De St. Joseph. Ann. d. Sc. Nat. (8), v, p. 237.

Habitat.— A common Mediterranean form procured in the ' Porcupine ' Expedition
of 1869, in the tube of an Eunice, in 173 fathoms off the west coast of Ireland, and also
in a free condition on the same ground — muddy sand. Next year (1870) it was dredged
at the depth of 690 fathoms in the Atlantic (Channel slope).

It is also a tidal form on the eastern border of the Irish Sea.

1 ' Geol. og Zool. Eeise/ 1862, p. 39.

308

LAGISCA EXTENUATA.

Head somewhat hexagonal (Plate XXVIII, fig. 2), with a deep median groove at
the base of the tentacle, and two prominent peaks, each of which is at a distance from
the tentacles. The eyes are comparatively large, and, judging from the conditions of the
largest example, increase with age and depth of water. Two slightly smaller are situated
in front of the nuchal collar, and look dorsally, and two somewhat larger rather in
front of the middle of the head, and lateral in position. In the example with the large
eyes (from 690 fathoms) the anterior pair show traces of a corneal lens. The median
tentacle is somewhat long, scarcely dilated below the filiform tip in the preparations, and
covered with moderately long cilia having a slightly bulbous tip. The lateral tentacles are
short, but also have attenuate tips. The tentacular cirri are similar to the median
tentacle. None of these organs show any signs of pigment in the preparations. The
palpi are of moderate length, and have rows of minute conical papillae, as De St. Joseph
also found, whereas Marenzeller observes that their upper surface is smooth. In the
British forms one prominent papillose ridge occurs dorsally.

Body. — Somewhat narrow and elongated, slightly tapered towards the head, and
very gradually towards the tail. Forty-five segments bear bristled feet, and the tail is
not quite complete. Behind the distension caused by the included proboscis, the
segments are boldly marked by the lozenge-shaped transverse bars which stretch
between the bases of the scale-pedicles, or their homologues in the other feet. Each
segment thus shows an anterior and posterior paler belt, and a median darker area.
Traces of brownish pigment occur posteriorly at the bases of the feet and on the
lozenge-shaped transverse bars. In some the brown pigment is broken into a number
of symmetrically placed touches, two of which are on the bases of the feet, and
the rest in interrupted bands across the segments. Ventrally traces of pigment occur
on the lips/ and the median iridescent depressed band is well marked. The conical
segmental (nephridial) papillse at the posterior border is directed upward between the
feet. A considerable portion of the body posteriorly is uncovered by scales.

De St. Joseph states that there are only seventeen papillas at the margin of the
proboscis, but so far as can be seen there are nine in each half.

Fig. 24.

Edge of scale of Lagisca extewuata, Grube.

LAGISCA EXTENUATA. 309

Scales (Plate XXXII, fig. 8) somewhat thin, in number fifteen pairs. They are
rounded in front, reniform or irregularly rounded posteriorly, and of a uniform greyish
hue in the preparations, though some are quite pale. Their surface is nearly smooth to the
naked eye, but under a lens the whole is densely covered by a series of minute, pointed,
slightly brownish spines, and the free portion of the edge is profusely ciliated (Eig. 24),
the cilia being pellucid, tapering structures terminating in a bulbous tip. Marenzeller
speaks of the network formed on the surface by the pigment; and Hornell mentions that
the surface is mapped out into separate areas, each containing several spines. The
under surface is iridescent, and the scar for attachment is situated near the outer and
anterior border. Marenzeller shows, besides the spines on the edge of the scale,
cylindrical papillas, but these have not been observed in our specimens. The first scale
is ciliated all round with the exception of the short covered portion.

De St. Joseph often found a white touch in the centre of the scales, which were
marked with brown, grey, and whitish. Hornell's figure of the scale differs from the
specimens referred to here.

Feet. — The base of first foot shows about two bristles, which conform to the type of
dorsal tuft, though they are considerably smaller. In the next foot the dorsal bristles
have attained great strength, though they are shorter than the typical forms, and much
broader towards the tip, which is little tapered. The bristle is therefore proportionally
more powerful than the typical form. They have a close series of spinous rows. The
contrast between the massive dorsal and the slender ventral bristles is marked, and the
tips of the longest of the latter extend only a very little beyond the extremities of the
dorsal. Their tips seem to be brittle, and while the inferior are more elongated and
slender, the upper forms present the characteristic short broad tips of the typical bristles.

In the third foot the size of the ventral bristles has largely increased, and they
stand out nearly half their length beyond the dorsal. The tips now approach the
normal, except that those of the inferior series are longer, and the upper slightly shorter.

In the typical foot (Plate XLII, fig. 30) the dorsal bristles form a powerful fan, the
inner and outer borders of which have more slender forms, while the greater number
consist of strong, slightly curved and tapered bristles, with somewhat closely arranged
spinous rows, and a smooth spear-shaped tip (Plate XXXVIII, fig. 10), the latter having
a tendency in some to follow the shape of the same region in the ventral bristles
(Plate XXXVIII, fig. 11). Others, again, show a sharper tip, and at the inner border
of the tuft are one or two with a slender tip. The ventral division commences with
a series having very long spinous regions, and a short and characteristically shaped bare
tip (Plate XXXVIII, fig. 12). The rows of spines are much finer and more dense than
in Harmothoe imbri cat a— indeed, in this respect they approach Acanthicolepis asperrima.
Those following the upper series show a distinct secondary process beneath the tips
(Plate XXXVIII, fig. 13). The length of the spinous tips diminishes in the usual manner
towards the ventral edge of the foot, the bare portion at the tip in several of the lowest
showing no secondary process. The papilla above the spine is long and filiform, and the
ventral cirrus extends beyond the bases of the adjoining bristles, and has sparsely
distributed and short clavate cilia. All the bristles are of a pale yellowish hue.

Posteriorly the dorsal division diminishes in length, and disappears in some of the

310 LAGISCA EXTENUATA.

terminal feet. At the tail the tips of the attenuate ventral bristles become much
elongated, some, however, maintaining the typical outline.

The dorsal cirri spring from the posterior border of the foot, and in those feet
without scales a well-marked papilla, the homologue of the scale-peduncle, occurs at the
inner border of the basal process of the cirri. They are somewhat slender, and have
numerous and slightly tapered cilia with a bulbous tip. The cilia are shorter interiorly
above the base, and again dis tally, and they cease about the middle of the filiform tip.
Posteriorly the cirri become longer and more slender, and the cilia more attenuate.
The caudal cirri are also slender. The long first ventral cirrus is stout, with a distinct
enlargement below the tip, which is much less filiform than the dorsal. The cilia, which
are considerably shorter and more uniform in length than those on the dorsal cirri,
extend from the base beyond the middle of the terminal slender region. They are
slender processes with a bulbous tip.

Loxosomm occur at the bases of the bristles and algee with other forms on them.
In Grube's original description of Polynoe extenuata 1 the scales are entered as
smooth at the edge, and with minute warts. The inferior bristles are a fourth longer
than the upper. Colour brownish grey above, bluish iridescent beneath. Elytra
caducous. There is nothing to give certainty.

Baron de St. Joseph (1888) found one in the tube of Serpula vermicularis, but it
may have simply crept there for temporary shelter. He is of opinion it is closely allied
to Lagisca floccosa.

Hornell (1891) describes apparently the same form from the Liverpool district,
though his figure of the dorsal bristles is rather indefinite. Its relation to Evarne impar,
to which he refers, is less marked than to Lagisca floccosa. The figure of the scale given
by this author differs from those procured by the ' Porcupine. 5

This form would seem to be nearly allied to Grube's P. extenuata as described
by Marenzeller in his ' Adriatic Annelida.' 2 He does not give the minute details nor
describe the special condition in the tips of the upper bristles, which are longer in his
figure and more distinctly bifid than in the British examples. On the whole the latter
would seem to be a well-marked variety, in which the secondary process of the ventral
bristles was less developed.

De St. Joseph (1878) , 3 after Langerhans, would make Lagisca rarispina and L.
propinqua, Malmgren, varieties of this species — a view which the preceding descriptions
and figures will sufficiently criticise. He had the advantage of the specimens in the
Parisian Museum in identifying Lepidonotus Leachii and L. dumetosus, both of De
Quatrefages, with Lagisca extenuata.

i ' Actin., Echin., u. Wiir./ p. 86, 1840.
2 Op. oit., 1898 ; p. 238.

AOANTHICOLEPIS ASPERRIMA. 311

Genus XII. — Aoanthicolepis {Norman MS.); 1 Dasylepis, Malmgren, 1867.

Body elongate-oblong or sublinear. Head produced anteriorly into two pointed lobes
on each side of the median tentacle, below the base of which the lateral tentacles spring.
Byes as in Eunoa. Scales eighteen pairs, coarsely spinous, overlapping each other
and covering the entire dorsum. Dorsal bristles stronger than the ventral, tapering
from the middle to the tip. Yentral bristles with short, bare, hooked tips, above the
spinous rows, with a minute secondary process in some towards the upper third of the
series. Segmental eminence distinct, papilla somewhat long and slender anteriorly, short
posteriorly.

Aoanthicolepis asperrima, Sars, 1860.

Specific Characters. — Body rather broad and thick, tapered slightly anteriorly, and
still more posteriorly. Segments about fort}^-one. Byes large, visible from the dorsum,
two in front of the nuchal fold, and two somewhat lateral in position about the middle
of the head, or perhaps a little anterior to it. Palpi of moderate length (in spirit), with
minute cylindrical papillae. The other appendages of the head have a slight enlargement
below the long tapering tip, the region above and below the enlargement having long
cilia. Tentacle and dorsal cirri similar. The ventral cirri are of moderate length, and
have short cilia. Scales eighteen pairs ; the first pair rounded, the rest more or less
reniform, roughened, with long and strong horny spines, especially towards the free
border. The tips of the spines are often bifid.

Habitat. — The only British locality seems to be the Frith of Clyde, where it was
obtained by the veteran naturalist of that region, Dr. D. Robertson. It ranges to
Norway, where it was procured by Sars, and lately by Canon Norman in his productive
dr edgings.

Synonyms.

1860. Polynoe asperrima, Sars. Forh. Vid. Selsk. Christ., 1860, p. 59.
1865. Lepidonotus pharetratus ? Baird. Johnst. Cat., B. M. Suppl., p. 340.
1867. Dasylepis asperrima, Malmgren. Ann. Polych.,, p. 7.
1873. „ „ Sars. Bid. Christ. Faun., p. 2.

1876. „ „ Mcintosh. Trans. Z. S., ix, p. 374, pi. lxvii, f. 9— 11.

1879. „ „ Tauber. Ann. Danic, 82.

1883. „ „ Levinsen. Nord. Annul. Vidensk. Meddel. f. d. Nat. For. i Kiobenhavn,

188 and 195.

The length of the example in the British Museum is about an inch.

The head (Plate XXVII, fig. 6) is somewhat longer than broad, with a well-marked

1 As the title Dasylepis has been pre-occupied by Pander for a Silurian Ganoid (1856), the
name suggested by Canon Norman, viz. Aoanthicolepis, may be substituted.

41

312 ACAXTHICOLEPIS ASPERRIMA.

median groove, which trends outward in front to terminate in the peaks on each side of
the median tentacle. The eyes are large and black, two being situated a little in front
of the nuchal fold, and slightly lateral in position, and two in front of the lateral
projection of the head, and also more or less lateral in position. Both pairs are thus
only fully seen in side view. JMo example has a median tentacle. The lateral tentacles
arise under its base, and are short organs with a trace of an enlargement below the
slender tip, and somewhat closely ciliated. The palpi, which are of moderate length,
show minute cylindrical papilla in certain parts, but seem to be mostly smooth!
Segmental papilla slender and somewhat long anteriorly, short posteriorly; directed
upwards between the feet.

Body flattened and proportionally broad, slightly narrowed towards the head, and
more distinctly posteriorly, where it is terminated by the two subanal cirri. The dorsum
presents no feature of note. Ventrally the segmental papilla is slender and small,
arising from an elevation close to the posterior border of each foot. These rounded
elevations form a moniliform series along each side of the body. The cirri are
comparatively slender and short.

The colour in the spirit-preparations has mostly disappeared, only one showing a
few brownish transverse bars anteriorly on the dorsum, and a slightly brownish hue on
the anterior folds of the mouth. The palpi have a curious greenish-grey colour, while a
trace of brown remains on the cephalic appendages, the dorsal cirri, and the first (long)
ventral cirrus. The anterior scales are brownish grey, with the brown spines standing
prominently outward.

Scales (Plate XXXII, fig. 4). — The first pair are more or less rounded, and studded
chiefly at the margin with the horny spines, the intervening spaces being often coated
with mud and parasitic growths. The scales generally are by no means thick, but are
covered with the chitinous papilla or bosses, small towards the anterior edge, but rising
into the long spinous processes posteriorly (Plate XXXIII, fig. 3). These processes
terminate in a bifid, trifid, quadrifid, or quinquefid tip. , The central axis of each is
cellulo-granular. From the inner to the outer margin also a few slender cilia with
clavate tips are present. The majority of the scales are more or less reniform, but
posteriorly the last pair become ovoid, with the same structure of spines and cilia.
The under surface is smooth and iridescent, with the scar for attachment somewhat
nearer the anterior than the posterior border, and more distant from the inner than
the outer margin. The scales are easily removed.

Feet. — The first (with the tentacular cirri) has three bristles, which conform to the
type of the dorsal bristles, only they are proportionally shorter and more curved.

In the second foot the dorsal bristles are stout and have boldly marked spinous rows
which in lateral view follow slightly oblique lines across the shaft. They are formed
by a thin chitinous plate split into the spines or teeth, somewhat after the fashion of the
ctenidial rows in Pleurobrachia, and increase in size from the dorsal to the ventral
surface of the bristle. The terminal region of the bristle is smooth. The ventral
bristles are very slender, with long tips having alternate spinous rows, the extremities
being slender and slightly hooked. The long ventral cirrus has moderately long clavate
cilia.

The usual gradations occur till the typical foot is reached (Plate XLII, fig. 29).

HARMOTHOE. 313

It is characterised by the long and strong dorsal bristles (Plate XXXVII, fig. 28),
which almost reach as far as the tips of the ventral. They are for the most part stiff,
straight, and gently tapered bristles, those at the inner edge of the tuft being shorter
and slightly curved. The spinous rows are well marked from a short distance above
the free edge to the extremity, which is pointed and smooth. The ventral division of
the foot is less acute than in front, the tip being almost fan-shaped, with the spine at the
upper border. The bristles have long straight shafts, while the tips are slightly dilated
at the commencement of the spinous region, have somewhat short bare tips of moderate
breadth, a well-marked hook at the extremity, and in some, especially at the upper
third of the series, a minute spur beneath (Plate XXXVII, figs. 29 and 30). The
great density of the dorsal bristle-tufts gives the animal a woolly appearance, and in
some they are tinted of a ferruginous hue from adherent growths. The ventral cirrus
does not reach the tip of the fleshy part of the foot, and has short clavate cilia.

In the terminal feet a similar condition to that in front exists, though the dorsal
bristles as a whole are more tapered. Their spinous rows are very distinct. The
ventral bristles in the last foot are attenuate. Parasitic algse are frequent on the
bristles.

The segmental (nephridial) papilla becomes distinct about the sixth foot.

Reproduction. — One example (Canon Norman's) from Norway, in July, 1878, shows
numerous ova in the perivisceral space.

The presence of a minute tooth below the tip of the ventral bristles shows the
proximity of the present species to the genus Harmothoe and its allies. This spur had
escaped the notice of Malmgren.

It is allied to Harmothoe areolata, Grube, but the latter differs in the more regular
arrangement of the surface of the scale, the fewer though larger and less acute spines,
the exquisite reticulation around them, and the much more numerous cilia of the outer
and inner borders. It seems to be rare in Britain, though the example from the Clyde
is of comparatively large size.

Acanthicolepis asperrima was first described by the elder Sars from specimens pro-
cured a few miles north of Bergen, a region which has been rendered classic to zoologists
by his labours and by those of his successors. Recently no one has done more in
collecting the annelids of the same region than Canon Norman.

Genus XIII. — Harmothoe (Kinberg, 1857), 1 char. em.

Body not much elongated. Lateral tentacles fixed below the median. Palpi with
rows of minute truncate papilla3. Eyes four ; two at the peaks in front, two posterior
on the dorsum in front of the collar. Scales fifteen pairs, covering the whole of the

1 Kinberg described the genus thus : — Cephalic lobe narrow anteriorly. Base of the tentacle
occupying the fissure of the cephalic lobe. Bases of the antennae fixed under the base of the tentacle,
Pharynx with f papillae and jaws. Elytra fifteen, covering the dorsum to the end of the body.
Body not long.

314 HARMOTHOE IMBRICATA.

dorsum. Dorsal bristles strong, with well-marked spinous rows and a smooth portion at
the tip. Ventral bristles with simple hooked tips, superiorly and inferiority; rest with a
well-marked secondary process. Segmental papilla long. Papillse of proboscis f.
Pre-eastric casca long- and slender.

1. Harmothoe imbrioata, i., 1767. Plate XXVI, fig. 3.

Specific Characters.— Body elongate-ovate, narrowing more distinctly posteriorly
than anteriorly. Head somewhat ovate, with the median furrow in front, and terminating
on each side in a blunt peak. The posterior pair of eyes are of moderate size, dorsal in
position, and are alone visible from above. The anterior pair lie under the peaks in
front, are somewhat larger, and look outward and forward. A trace of a cuticular lens
is occasionally seen in these. Tentacle of moderate length, arising from an enlarged dark-
coloured base (ceratophore) ; the proximal part of the column (ceratostyle) brownish with
a dark belt below the pale enlarged region, to which the filiform tip is attached. A few
rather short clavate cilia occur on its surface, the filiform tip being smooth. The lateral
tentacles are beneath the former, and are about half their length, with a very slight
swelling below the filiform tip. They have proportionally more numerous clavate cilia
than the median tentacle. The tentacular and dorsal cirri agree with the latter in
outline, and have a considerable number of clavate cilia. Palpi elongated and tapering,
with rows of somewhat truncate clavate papillse. Ventral cirrus of moderate length,
with a few short clavate cilia. Segmental (nephridial) papilla comparatively long.
Scales fifteen pairs, and, with the exception of the rounded first pair, ovate-reniform
or obliquely ovate. Though smooth to the naked eye, they are minutely spinous
under the microscope, and the outer margin has somewhat short cilia. Only in large
specimens is there a row of brownish subglobular papilla within the posterior margin.
Dorsal bristles strong, with rather distinct spinous rows, and a well-marked smooth
region at the tip. Ventral bristles with an elongated spinous region superiorly, a short
spinous region inferiorly, and a simple smooth tip with a hook. All the rest have a well-
marked secondary process beneath the hook.

Habitat. — Distributed between tide-marks and the adjacent region everywhere round
the shores of Britain, from Shetland to the southern coast of England, but it is com-
paratively rare in the Channel Islands (Guernsey), where its place is occupied by
Lagisca floccosa. It extends also to the depth of 75 to 96 fathoms (' Porcupine'
Expedition, 1869) and to 125 fathoms off the west coast of Ireland. It ranges to
Spitzbergen, Greenland, Iceland and Scandinavia, to the Adriatic and Mediterranean,
as well as to other European shores, and is also found in America (Verrill) from
Cape Cod to the St. Lawrence. According to Marenzeller it extends to Japan, and
Grube records it from northern and eastern Siberia, and from Sitcha and the Sea of
Okhotsk, and various other parts of the Arctic Sea.

It is common in the stomach of cod and haddock (E. M.). It has also been found
in the tube of Terebella nebulosa, Bressay Sound, and beside Poly cirrus in an old shell
in the same region.

Length. — On each side of 1J inches, and reaching 2.

HARMOTHOE IMBRICATA. 315

Synonyms.

1766. Aphrodita lepidota, Pallas. Miscell. Zool., p. 94, Tab. 7, fig. 15 ; Tab. 8, LI, 2.

„ ,, imbricata, L. Syst. Nat., twelfth edit., vol. i, p. 1084.

1768. „ violacea, Strom. Kongel. Norsk. Yidensk. Selskabs. Skrifter, Deel iv, p. 366.

1776. „ cirrata, 0. F. Miiller. Prod. Zool. Dan., p. 218, n. 2644.

„ „ violacea, idem. Ibid., p. 218, n. 2645.

„ „ lepidota, idem. Ibid., p. 218, n. 2643.

„ Die flache Aphrodite, Martini. Allgem. G-eschichte der Natur, iii, p. 132.

„ Aphrodita imbricata (Ziegelrticken Aphrodite), Martini. Ibid., iii, p. 151.
1780. „ violacea, Fabricius, 0. Fanna Grronl., p. 308, n. 290, Tab. 1, fig. 7.

1792, „ „ Brugniere. Encycl. Method., vers, i, p. 89.

1800. Die flache Aphrodite, 0. F. Miiller. Naturges. einiger Wnrm-Arten, p. 180, Tab. 14, figs. 1 — 5.
1820. Polynoe cirrata, Savigny. Syst. des Ann., 26.

1827. „ violacea, Bory de St. Yincent. Beproduced in Tableau Ency. Method., p. 135, pi.

Ixi, f. 30—33.

1828. Eumolpe cirrata, De Blainville. Diet. Sc. Nat., vol. lvii, p. 459.
1830. Aphrodita cirrata, Bosc. Hist, des Vers., 169, 2nd edit., 183.
1834. Polynoe cirrata, Audouin and Edwards. Annel., p. 86.

1840. „ „ Johnston. Ann. Nat. Hist., ii, p. 434, Tab. 22, f. 2.

1843. Lepidonotus cirratus, CErsted. Grronl., Ann. Dors., p. 14, f. 1, 5, 6, 11, 14, 15.

„ „ „ idem. Annel. Dan. Consp., p. 13, fig. 43.

„ Polynoe cirrata, Bathke. Fanna Norweg., 150.
1851. „ „ Maitland. Fauna Belg., 214.

1853. Aphrodita varians, Dalyell. Pow. Creat., ii, 168, pi. xxiv, f. 11, 12.

1854. Polynoe cirrata, Thompson. Fauna, Ireland, 173.

1860. „ „ Sars. Yid. Selsk. ForhandL, 1860 (sep. copy, 5).

1865. „ ,, De Quatrefages. Ann., i, p. 232.

„ Lepidonotus cirrosus, idem. Ibid., i, p. 261. (?)

„ Polynoe cirratus, Johnston. Cat. Brit. Mus., 114, pi. viii, fig. 2.

„ Harmothoe imbricata, Malmgren. Nord. Hafs. -Annul., p. 66, Tab. ix, fig. 8.
1867. „ „ idem. Ann. Polychast. Spets., p. 154 (sep. copy, p. 9).

1871. „ „ Ehlers. Sitzb. phys.-med. Soc. Erlangen, Heft 3, p. 77.

1873. „ „ idem. Op. cit., Heft 5, p. 7.

„ „ „ Sars. Nyt. Mag. f. Naturvid., 19, p. 203.

„ „ „ idem. Bid. Christ. Fauna, iii, p. 3.

„ Polynoe cirrata, Mobius. Jahresb. Com., 1871, p. 111.
„ „ „ Kupffer. Ibid., p. 150.

1874. „ „ Mobius. Die Zweite deutsche Nordpolarfahrt, 1869, p. 253.

„ Harmothoe imbricata, Malm. Goteborgs Kongl. Yet. och Yitt. Samhalles Handl., Haftet

14, p. 74.

1875. „ „ Ehlers. Annel. f Porcupine/ 1869, op. cit., p. 32.
„ „ „ Mcintosh. Invert, and Fishes, St. A., p. 116.

„ Polynoe cirrata, Mobius. Jahresb. Com., 1872, p. 166.

1876. Harmothoe imbricata, Mcintosh. Trans. Z. S., ix, p. 398.

1877. „ „ Hansen. Nyt. Mag. f. Naturvid., 24.

1878. „ „ Lenz. Jahresb. Com., 1874-6, Anhang., p. 12.

1879. „ „ Marenzeller. Siidjapan. Annel. Denkschr. d. Kaiserl. Akad. Wiss. Wiexi,

xli, p. 17, Taf. ii, f. 1.
Tauber. Ann. Danic, 80.

316 HARMOTHOE IMBRICATA.

879. Polynoe imbricata, Theel. Kongl. sv. Yet. Akad. Handl.,Bd. xvi, 3, p. 9.
1881. „ cirrata, Horst. Mederland. Archiv Zool., 1881, Suppl. Bd. i, p. 5.

» « « Pelsener. Bull. Soc, Boy. Malacol. Belg., xiv, p. lxxxix.

1883. « « Kallenbach. Inaugur. Dissert. Eisenach, 1883.

„ Harmothoe imbricata, Levinsen. Nord. Annulat., 194.

„ Polynoe imbricata, L. Wiren, Chsetop. ' Yega ' Exped., &c, p. 389.
1884. „ cirrata, Cams. Fauna Medit., i, 201.

„ Harmothoe imbricata, Webster and Benedict. Ann. Mass., 701.
1886. » „ Harvey Gribson. Yerm. Liverp., 149.

1888 - « » De St. Joseph. Ann. Sc. Nat., 1888, p. 161, pi. vii, f. 21.

1889 - » , » Trautzsch. Jenaiscbe Zeitscb. f . Nat., xxiv, p. 66, and Arch, f . Naturges.,

55 Jahr, Bd. i, Hft. 2, p. 136, pi. vii, f. 1.

1890. „ „ Malaquin. Ann. Boulon., 21.

1891. Polynoe (Harmothoe) imbricata, Hornell. Polycha3ta, Liverpool Dist., p. 231, pi. xiii, f. 2.

1896. Harmothoe imbricata, Michaelsen. Polych. Fauna, p. 11.

„ Polynoe (Harmothoe) imbricata, Roule. Camp. d. c Caud./ 443.

1897. Harmothoe imbricata, H. P. Johnston. Pacific Annel., Calif or. Acad. Sc, 181, pi. vii, f. 37.
I 898 - „ „ Michaelsen. Gronl., Ann., p. 121.

The head (Plate XXV, fig. 6) is broader posteriorly than anteriorly, and shows only
a single pair of eyes, which are of moderate size and just in front of the nuchal collar.
The other and somewhat larger pair are placed under the anterior peaks, and present
a trace of a cuticular lens. They look forward, outward, and slightly downward.
Some specimens have the anterior eyes a little more lateral in position, so that they are
partially seen from the dorsum, and occasionally an additional eye lies midw^ay on one
side, as in an example from the stomach of a haddock at St. Andrews and in another
from Shetland (Fig. 25) ; while in a third from the latter region (Bressay Sound) the

Fig. 25. Fig. 26.

/ v \

Fig. 25. — Head of Harmothoe imbricata with an additional eye on the right, and the anterior pair more than
usually evident.

Fig-. 26. — Head of Harmothoe imbricata with three posterior eyes, while only one anterior is visible from the
dorsum.

anterior eye on the left has moved backward, so as to resemble the position in Lagisca
floccosa (Fig. 26), and in another from Lochmaddy the right anterior eye approaches the
posterior even more closely. A still more remarkable variation exists in a young
example from the same region in which both anterior eyes have moved backwards, so
that they occupy the position of those in Evarne. The median tentacle is of moderate
length, with a brownish column, a dark belt below and often encroaching on the whitish
enlargement, and another dark belt beyond it at the base of the filiform process. The
lateral tentacles are only slightly enlarged below the filiform tip, and have a few short
clavate papillae on the surface. The tentacular cirri are similar in shape and colour to the
median tentacle. The palpi have rows of short and somewhat truncate clavate papillae, the
terminal surface showing in many minute processes.

HAEMOTHOE- IMBRICATA. 317

The body, which consists of thirty-seven bristled segments, presents the normal out-
line, viz. narrowed somewhat abruptly anteriorly, but very gently posteriorly. 'The
dorsum is often boldly pigmented from the first to the last segment with dark touches,
which stretch across each segment between the pedicles of the scales or the corresponding
papillaa in the other feet. In the anterior segments two of these touches occur, viz. a
large one in the transverse "mark" of the segment, the pigment even invading the
scale-pedicle, and a narrow one behind the former. In some a narrow belt in front of
the larger bar is also present. A pale longitudinal line cuts the large median bar just
mentioned into halves, but the smaller anterior and posterior pigment-belts remain
entire except in a few posteriorly. The dorsum of the foot has also a sprinkling of dark
pigment extending to the bases of the bristles. In some the pigment of the dorsum is
much increased, so that the entire surface between the peduncles for the scales is
blackish, enlivened only by the lines of the segment-junctions, and the pale lines
around the transverse central pigment-bar, for no median longitudinal stripe occurs:
The ventral surface is pale, with the exception of the anterior folds of the mouth, which
have bands of dark pigment. The large terminal anal cirri are similar in structure to
the dorsal, which, again, agree with the tentacle.

In the British specimens of Harmothoe imbricata the segmental papilla is so long
as to merit the name of a cirrus throughout the greater part of the body, the process
diminishing anteriorly towards the sixth foot, where it commences, and posteriorly in
the terminal feet. So far as could be observed no sexual differentiation of the organ
occurs either at the breeding season or subsequently. It is similar in both sexes. The
segmental organs (nephridia) were clearly described in the Polynoidse of Prof. Haswell,
and shortly afterwards by Prof. A. G. Bourne.

Digestive System, — Proboscis. — Nine papillse occur dorsally and ventrally in the
extruded proboscis, the lower teeth of which bite to the right of the upper. The dorsal
papillas are slightly tinted with dark pigment, and the dorsal wall joining the organ to
the mouth is similarly coloured.

The first (dorsal) gastric cascum is a long slender tube which passes forward to the
body- wall, and terminates in a bulbous region composed of two or three lobes. The
narrow part of the canal is whitish like the wall of the gut, while the bulbous
terminal portion is slightly yellowish or buff, like the glandular cseca of the gut, with
which it agrees in structure. The second is nearly as slender. The two succeeding
increase gradually in size. The next passes nearly transversely out, and begins the
series of deep yellowish lateral glandular casca.

The contents of the gut show sponge-spicules, foraminifera, star-fish plates, bristles
of annelids, sand, and mud. Occasionally, however, portions of other annelids, such as
Nereis pelagica, occupy the intestine. It is a voracious form.

Scales (Plate XXXII, fig. 10) fifteen pairs. The first pair are rounded, minutely
dotted all over with short blunt spines, and have at the posterior border a series of
somewhat clavate papillas of variable number (six to twelve), which are visible under
a lens. The outer and posterior margin bears short clavate cilia. The minute blunt
spines increase in size at the outer and posterior borders. The rest of the scales are
reniform or obliquely ovate, minutely spinous, the outer margin being furnished
with short clavate cilia, and in certain specimens with large and small subglobose or

318 HARMOTHOE IMBRIOATA.

clavate papilla. These papillae were well marked in a series of comparatively small
pale examples brought by Canon Norman from Finmark. The larger coloured speci-
mens showed them less distinctly. The spines are larger in certain arctic examples,
e. g. from Davis Strait. The cilia differ much in length even in large specimens. Some
of the scales in spirit-preparations adhere with great tenacity ; others separate readily.
Their colour varies much, being, to take the order of Malmgren, bluish grey,
greyish, brownish, or dark brown with a purplish or violet lustre, reddish brown
(Vandyke brown), almost black, or greatly variegated. A striking variety has pale buff
scales bordered with a narrow belt of dark brown. Others agree with the foregoing
except that the scale is darker brown (Bismarck), with a small white patch in the centre
and a few darker grains in front, the whole forming a beautiful series. Some have the
outer half of the scales whitish, the inner blackish grey or brown, so that the animal
has a broad blackish or brownish band down the centre. Occasionally the pale fawn-
coloured scales are minutely and uniformly speckled with small brown touches. A few
again are almost white. Some have a broad brownish-red belt, almost a third the
breadth of the scale, curving within the posterior border, and as these are arranged
symmetrically the effect is agreeable. A developing scale shows minute clavate cilia
before the spines appear.

Development of the Scales. — These seem to be reproduced with considerable rapidity,
forming miniature scales on the end of the peduncles, with a trace of brownish pigment
over the scar. They consist of a superficial cuticle, which is quite smooth, and of a
subjacent cellular layer within which the connective tissue rapidly proliferates.

Feet. — The first foot has often only a single bristle, conforming to the dorsal
type.

The second foot has a short ventral lobe and a long ventral cirrus with clavate
papilla?. The dorsal bristles are smaller, shorter, and more curved than usual, but
are otherwise of the normal structure. The ventral are much more slender than the
normal, and the spinous region is proportionally long and hispid, while the smooth tip
is slender and simple.

The third foot has much stronger dorsal bristles than the second, and the ventral
bristles have also increased in size. In these the upper and lower groups still have
simple tips, while the median series have bifid tips, but the region is more slender than
in the typical foot, and the secondary process small. The spinous rows are also more
hispid (i. e. have longer spines) than in the typical foot.

As we proceed backward the characters of the typical foot are acquired (Plate XXX,
fig. 1). It shows dorsally a strong series of somewhat straight or only slightly curved
bristles with well-marked spinous rows, and a smooth spear-shaped tip of some length
(Plate XXXVIII, fig. 14). In examples from Greenland the tips of these (dorsal)
bristles are so elongated as to be almost sabre-shaped. In specimens from Cornwall,
again, the spinous rows are finer, and the bare portion at the tip somewhat differs. In
young examples the front edge is slightly bevelled. The ventral division bears a series
of strong bristles, every one of which has a bifid tip. In the upper series the smooth
portion at the tip is slightly curved outward, and the secondary process is short and
sharp (Plate XXXVIII, fig. 15). The terminal hook is well marked. In those with

HARMOTHOE IMBRICATA. 319

short tips from the middle of the foot (Plate XXXVIII, fig. 16) the same relations of the
secondary process occur, and thus they are easily discriminated from the bristles of
Lagisca floccosa. The ventral cirrus has a series of short clavate cilia.

In the terminal feet the dorsal bristles remain stout, but their tips are more
tapered. The slender ventral bristles, on the other hand, have the spinous region
greatly elongated, but the delicate tips in most cases retain the bifid condition, only one
or two at the ventral border being simple.

Varieties. — A variety procured by Canon Norman in Norway has shorter bristles,
but though the rows of spikes on the dorsal bristles are somewhat finer, in all other
respects it corresponds. In another from Loch Portan, Lochmaddy, the dorsal bristles
immediately above the ventral are slender, and in this form also the scales have more
numerous and longer cilia. The colours are brighter, and the under surface and sides
pinkish.

A pale variety, having somewhat rougher scales with longer cilia, occurred between
tide-marks at Lochmaddy.

The large arctic examples, such as those of Dr. Walker, have a distinct tendency
to elongation of the tips of the bristles, both dorsally and ventrally ; while a variety
from St. Andrews presents more elongated dorsal bristles than in the typical example.

Few annelids are more abundantly distributed, yet the structural variation of the
individuals is not great.

Habits. — H. iwibricata lurks under stones in pools and moist places between tide-
marks, among sponges and in their crevices, in tubes of Terebella and Chseiopterus,
in empty acorn-shells, and in almost any convenient crevice. It is one of the most
plentiful forms between tide-marks, and is also dredged abundantly in the laminarian
region and beyond it, in the crevices of old shells, tangle-roots, and stones. It clings
closely to such surfaces, and apparently tries to escape observation. Young examples
have been found in old shells with Polycirrus. It is most active and restless when
disturbed, and wriggles violently, leaving fragments of the body or separated scales in
the hands of the captor.

It is a somewhat delicate animal in confinement. Thus it suffers rapidly in a
bottle with other marine animals on the collecting ground ; indeed, if the search be
prolonged few are alive, while other marine forms survive. Sir J. Dalyell, however,
retained them until the discarded scales were reproduced, being scarcely distinguishable
from the original scales in about six weeks or two months. It swims with an undu-
latory motion in the water, but, like the loach, soon sinks to the bottom. As in allied
species, it often strikes the glass vessels with its jaws, making sounds heard at a
considerable distance.

It is brilliantly phosphorescent, discharging bright bluish-green or greenish scintil-
lations from the point of attachment of each scale, and thus under irritation the flashes
are arranged in pairs along the body or in a double moniliform line. The separated
scales also continue to gleam for some time, chiefly at the scar for attachment, that is
near the great ganglion of the region. If severely pinched it wriggles through the water,
emitting sparks of greenish or bluish-green light from the foregoing points. The phos-

42

320 HARMOTHOE IMBRIOATA.

phorescence appears to be less vivid during severe weather and when confined for a night
or two in shallow vessels — a feature probably due to nervous prostration, or it may be
associated with approaching maturity. Specimens placed in a weak solution of picric
acid in sea water are not luminous, and the scales are not at first thrown off.

Parasites. — The crustacean parasite Herpyllobius arcticus* Stp. Ltk. (Silenium
Polynoes, Kr.), occasionally occurs in arctic examples attached to the dorsum. Levinsen
also found another crustacean parasite, viz, Selio'ides Bolbroei, Lev., on one from
Greenland. 2

On the dorsum, under the scales of a specimen from the tidal region at Balta
Sound, Shetland, numerous examples of a fine Loxosoma in various stages of growth are
found.

A peculiar warty growth appears on the tentacular cirrus of an example from the
Gulf of St. Lawrence, and a series of minute whitish tubercles in an arctic example from
Bessels Bay.

The bristles harbour many parasitic algae, besides mnd, foraminifera, and sponge-
spicules. At St. Andrews a small Sabella, Syllis, and Pholoe have been found amongst
the bristles, and young mussels occasionally fix themselves to the dorsal bristles.

A translucent Ascaris, fully half an inch in length, occurred in the peri-pharyngeal
space of a female. The slightly truncated snout showed one or two blunt papillae, while
the pointed posterior end had a few acute papilla. Though outside the gut it had free
access to the perivisceral fluid, and, moreover, the long anterior casca passed forward on
each side of the space.

Development. — The broad outlines of the development of this species have been
known for a considerable time, and it may therefore form the type for the group.
Comparatively small species are occasionally found mature, or carrying ova under the
scales.

Michael Sars, 3 as early as 1845, described the occurrence of ripe examples of
Harmothoe in February and March. He noticed the mature females exhibited a change
of colour, becoming pale rose behind the anterior fourth. This was due to the eggs
which covered the dorsum. He thought that the ova passed out by a small aperture
above the feet. He watched the development of the egg from early segmentation to the
movement of the embryo by cilia, and its slightly greenish coloration. The larvas
(monotrochous trochophores) escape after two weeks, swim freely, and bear two eyes,
with a pre-oral band of cilia. Max Miiller, Desor, and others have also described the
development. Desor, however, fell into the error of supposing that the larva escaped
from a similar ciliated investment to that of Linens obscurus.

Max Miiller, 4 in the young of either this or an allied form, pointed out that new
segments' were interpolated posteriorly. He described the dimorphism of the bristles,

1 'Kgi. Danske vidensk. Selsk. Skrifter naturv.-mathem. afd./ Bd. v, 1861; and Kroyer,
< Naturh. Zidskr./ 3 die E., Bd. ii, 1863, &c.

2 < Yidenskah. Med. fra den nat., &c./ i Kiobenh., 1887.

3 " Zur Entwicklung der Anneliden," ' Arch. f. Naturges./ 11 Jahrg., v. 1, 1845 ; and ' Ann. Nat.
Hist./ 1845, p. 188 (vol. xvi).

4 "Ueb. d. Entwicklung u. Metamorph. d. Polynoen," ' Mull. Archiv/ Jahrg. 1851.

HARMOTHOE IMBEIOATA. 321

the condition of the alimentary canal, and the development of the feet. His species
had two pairs of eyes.

Claparede 1 gives the early stages of a Polynoe up to the formation of feet and
bristles. His example had eleven pairs of feet.

Yiguier 3 refers to a pelagic Polynoe of sixteen segments, but which, he says, showed
no larval appearance. The feet and the ventral bristles are comparatively long. It is
in all probability a post-larval Polynoe from the Bay of Algiers, as Marenzeller also
thinks.

Dr. W. Michaelsen 3 describes a pelagic polynoid from Ceylon (Drieschia pelagica),
in which the foot is simple, with very long hair-like bristles, and only a few shorter
thicker forms. These apparently represent the ventral division of the foot, and only a
single spine is present. The tentacular cirri and dorsal cirri are very long.

Marenzeller 4 gives an account of a pelagic form of twenty-four segments procured
by the Prince of Monaco, in 48° 50' lat. N., and 21° long. E. of Greenwich, under the
name of Nectoclideta Grimaldii. It is characterised by the great elongation of the inferior
bristles. Scales absent, palpi and cirri smooth.

Fewkes 5 described a young Polynoe with only three pairs of feet.

Too little is yet known of the pelagic forms described by Michaelsen and Marenzeller
to speak with certainty of their precise relationships. The presence of only a single
spine and the simple nature of the foot in Drieschia are features which diverge from the
ordinary types.

Prof. Y. Hacker of Freiburg gave in 1896 6 the results of his studies of the larvae of
Polychseta at Naples. In the Aphroditidse he describes three stages of a Polynoe, viz.
the trochophore, metatrochophore, and nectochgete 7 stages. In the Aphroditaceans
the trochophore moves at first by aid of its cilia, and then the bristles develop secondarily
and enable the post-larval stage to assume great activity during its pelagic life. His
species was probably Polynoe reticulata.

The ovaries of Harmothoe imoricata form a series of lobulated organs stretching
from the seventh foot (which has a segmental [nephridial] papilla, as has also the sixth)
to the posterior end. They are small anteriorly in the region of the proboscis, though
as a rule they do not reach this part, but attain in January a considerable bulk through-
out the rest of the body, again diminishing posteriorly.

The eggs become prominent in November, being coarsely granular, with a distinct
nucleus and nucleolus (Fig. 27). They seem to have a hyaline connecting substance,
to which perivisceral corpuscles attach themselves. They vary in size, the smaller being

1 'Beobach./ 1863, p.80, Taf. viii, f. 1—11.

2 < Archiv. de Zool. exper./ vol. iv, 1886, p. 416.

» ' Jahrb. d. Hamburg. Wiss. Anstalt/ ix, 2, 1892, p. 6, figs. 15—18.

4 c Bullet, de la Soc. Zoolog. de France/ 1892, p. 173.

5 'Bull. Mus. Comp.Zool. Harv. Coll. Carnb./ v, 11, 1883-5,

6 < Zeitsch. f. w. Zool./ Bd. lxii, pp. 74—168, Taf. iii— v.

7 Nrixeiv, swimming or pelagic ; and x aiTr l> bristle.

322 HABMOTHOE IMBRICATA.

about an eighth the size of the larger, — the latter, however, being nearly equal, and their
nuclei contain several nucleoli.

Some females retain a considerable number of ova till the middle of May, though
the body is by no means distended. Many have been discharged.

Fig, 27.

Developing ova of Harmothoe imhricata. x 280. November.

Besides the larger ova are many minute forms attached to the germinal tissue, so
that the spawning period is prolonged, or the minute ova retained or absorbed.

About the middle of February specimens with masses of ova under the scales are
common. The slightly pinkish eggs are attached to each other and the surface by a
transparent mucous secretion, so that they do not readily fall off. They form a dense
layer under the scales, and in some are almost invisible. The process would appear to
be protective, giving them the shelter of the adult, and enabling them to escape the
attacks of predatory crustaceans or other forms. They have a diameter of about *56
to '78 mm. ; the zona is delicate and translucent, yet resists some pressure. In the
perivitelline space are a few granular cells, such as those found in various eggs of fishes
(e.g. the gurnard). In structure the yolk is minutely granular, the figure showing the

Fig-. 28.

Segmenting ovum of Harmothoe imhricata.

egg cleft into two spheres (Fig. 28). The eggs are so opaque that section alone reveals
the structure. In confinement the females carrying ova readily throw off their scales a
feature probably due to the absence of nourishment and the condition of the water.

The males have their spermatozoa fully developed at the end of January and
beginning of February. These consist of simple tapering rods with a very attenuate
filament from the broader end (Plate XXVI a, fig. 1). They thus differ in shape from

HARMOTHOE IMBRICATA. 323

those of Lepidonotus squamatus (Plate XXVI a, fig. 2). When punctured the tissues of
a ripe male heal in a day or two, and the animal regains activity.

In the first week of March the trochophores swarmed in the vessels containing the
adults, congregating like the copepods and Nauplii at the margin next the light. The
early trochophores (Fig. 29) present a less developed ring of cilia, but soon (Fig. 30)

Fig. 29. Fig. 30.

FlG. 29. — Early trocliopliore of Harmothoe imbricata viewed from the anterior end.

Fig. 30. — Trocliopliore of Harmothoe imbricata with fully developed pre-oral ring of longer cilia in rapid movement,

viewed as in Fig. 29, from above.

these organs assume a characteristic appearance, and the larvse are very active. They
are bluntly conical at both ends, and devoid of eyes. When compressed under a cover
the ring of cilia (pre-oral) (Plate XXVI A, fig. 3) has a very regular rotular aspect.
Those captured by the bottom tow-net in the Bay at this time were somewhat larger.

On the 28th March the larvae in confinement had for the most part gone to the
bottom, and, moreover, had assumed a greenish appearance, as indicated in the figure.
In the same plate (XXVI a) the older trochophores with a pair of eyes are shown in
various positions in figs. 4, 5, and 6. In the first-mentioned figure the mouth forms a
projection to the left.

On the 25th April the intermediate stages occur in the bottom tow-nets, viz. larger
and more advanced forms having a bluntly rounded anterior end, a prominent ventral
surface, and minute feet, with or without traces of bristles, the feet being marked only by
crenations. The head is bluntly rounded, with two or three pairs of eyes, and the body
is somewhat pear-shaped (Plate XXVI a, fig. 7). The digestive apparatus is indicated by
the opaque granules in front and the paler posterior region.

Very soon (indeed, in the nets of the same date) the bristles make their appearance in
the feet (Plate XXVI a, figs. 8 and 9), though the head still remains blunt. In the first-
mentioned figure the foot-papillaa are minute, and project very little in lateral view, but
as transparent objects they show traces of the bristles. The latter project by degrees
from the ventral division of the foot, and present a somewhat broad spinous tip with
distinct spinous rows and a well-marked terminal hook, but no secondary process.
When they become more elongated, so that the shaft is clear for a considerable distance,
a trace of a secondary process occurs below the hooked tip. Minute scales have also
made their appearance. No anal papilla (cirri) are yet visible, though in front the
tentacular cirri are represented by papillae. A line of pigment occurs at the ciliated
ring, and the eyes present a rudimentary lens.

324 HARMOTHOE IMBRICATA.

As shown in Plate XXVI a 5 fig. 9, the feet by-and-by assume a lobate form, and
the bristles lengthen, while two rounded papilla? mark the anal cirri. The body is com-
paratively broad and short, and the anterior end truncated.

At a stage a little subsequent to the foregoing the head is better differentiated, and
the lateral tentacles appear as two rounded papilla? on each side of the median dimple.
There were ten feet in the example, and the scales were well formed. The tentacular
cirri had a single large (the largest in the animal) curved dorsal bristle projecting at the
base, with prominent rows of spines, and one or two smaller bristles just appearing
beyond the surface. The organs themselves showed indications of developing cilia, as
also did the dorsal cirri. Inferiorly the palpi formed two large rounded discs on each
side of the head, projecting laterally, but not yet reaching the front. No trace of a
median tentacle was observed in this specimen. The dorsal bristles were all curved,
with prominent rows of spines, and the tips of the ventral had also very distinct
spinous rows. The bristles project prominently (nectocha?te stage). No caudal styles
were present.

In Plate XXVI a, fig. 10, the next stage is represented. The head is less bluntly
conical, and the two tentacular cirri are leaf-shaped, and soon the tip of the powerful
dorsal bristle at their base projects nearly straight outward. The body is more elon-
gated, the feet more prominent, and the bristles project more than half the diameter of
the body on each side. The caudal cirri are now broadly lanceolate. Minute scales are
present.

The dorsal bristles resemble the ventral more closely than in the adult. Moreover
they approach those towards the posterior end of the adult rather than the median, a
fact which may be connected with the growth of new segments between those adjoining
the head and the tail. Thus it is that the anterior and posterior segments of the adult
show the primitive condition of the bristles, the most modified and the most typical
being those in the region between them.

In the stage shown in Plate XXVI a, fig. 11, the median tentacle (which may have
been lost in the previous preparation) projects conspicuously in front, and the caudal
cirri with tapered extremities are distinct.

In early stages, in which eight feet and four pairs of scales occur, the eyes on each
side are nearer each other than subsequently, and a smaller third black speck is observed
on each side in front of the rest on the under surface near the site of the adult eye,
though the peaks are not yet formed.

In the stage figured in Plate XXVI a, fig. 12, the head is still further defined, the
eyes being arranged obliquely on each side (the posterior pair nearer each other than the
anterior). The median and lateral tentacles are distinct, and the palpi project promi-
nently forward, with tapering extremities. Their margins are crenatecl, but distinct
papilla? were not observed. The tentacular cirri show short clavate cilia. The scales
are larger, and have short clavate papilla? projecting from the outer border, the last
pair of scales having the largest cilia. The feet are deeply cleft and prominent, and the
slender shafts of the bristles have become greatly elongated, so that they project far
outwards on each side. There were nine bristled feet in the example. In this instance

TBOCHOSPHEEE. 325

the caudal cirri were absent (had fallen off), and a distinct papilla occupied the centre
of the caudal region.

When fourteen feet are present a great advance has been made in regard to the
head, cirri, and feet. The head shows a blunt peak on each side, and the median
tentacle is proportionally large and long, while the palpi have not yet attained full
development, though they show the rows of minute papillae. Two large black eyes are
situated posteriorly, and two occur on the mid-lateral region, and are w 7 ell seen from the
dorsum. A considerable interval occurs betw T een the eyes of each side. All the cirri
have large clavate cilia. The posterior scales show large clavate papillae with palpocils
at the tip. The bristles at this stage are long, but they do not show distinctive
characters, though the attenuate condition of some of the dorsal suggests an approach to
Gattyana (Nychia). The specific identity of those captured in the tow-nets could not,
of course, be absolutely determined.

A different trochophore appears in the nets in June, characterised amongst other
things by its black and brown pigment-belts at the ciliated ring, the former tint
being in front of the latter (Plate XXVI a, fig. 15). This form closely approaches
Dr. Hacker's species (Taf. iii, fig. 2) from Naples, though certain points of difference
are present.

In this species the tips of the bristles seem to be less elongated, and the pre-oral
lobe is curiously marbled with black pigment. When scales form, the body is marked
in a somewhat tessellated manner, with transverse bars of reddish-brown pigment. At
this stage also the anterior end (ventrally) has two rounded, pigmented, lateral regions,
with a patch of black in the centre. A little later the black pigment bounds a median
central region in front, and there are six eye-spots, two large rounded eyes on each side,
and a black crescentic eye to the outer side of the anterior.

The body is shorter, the feet more closely arranged, and the scales more persistent.
The latter have traces of low papillae on their outer edge, and the surface is roughly
areolated, probably from papillae, as might occur in the young of L. squamatus. When
the bristles are viewed from behind the spikes are alternate. The tips are much shorter
than in the previous form, and the shafts stouter. Large diatoms often occur in the
intestine.

In the stage (Plate XXVI A, fig. 18) with the developing palpi the head is rounded
in front, and there are often three or four additional eye-spots besides the four normal.
The single spine of the first foot is prominent, and also two curved bristles. A band of
black pigment bounds the anterior border of the snout.

There are eight feet, the dorsal bristles of which present very fine serrations, and
thus differ wholly from the previous forms. The ventral bristles have short tips with
rather closely arranged spinous rows. The caudal end terminates in a pointed pygidium
without cirri. In the early stages the bristles of the first foot are largely developed in
proportion, and must be especially useful in protecting the head. The cirri in this form
present a somewhat ovoid outline.

As soon as the palpi have developed (Plate XXVI A, fig. 19) the head shows a deep
notch in the centre, the two large black eyes on each side being situated far back, the

326 TBOCHOSPHEEE.

anterior eye being behind the middle line. From the first foot three strong curved
bristles project forwards at the side of the palpi. These bristles have well-marked
spinous rows. The palpi have tapering extremities. The dorsal bristles are slender
and tapering, with distinct rows of spines, which from the dorsum appear as opposite.
The tips of the ventral bristles are simple, and the spinous region shorter than in the
first species, and from the dorsum the spinous rows seem to show an alternate arrange-
ment.

When eight pairs of feet are present, and the tentacles, palpi, and the anal cirri
fairly formed, the dorsal bristles show a tendency to finer rows of spines, so that they
approach the condition in such as Lepidonotus and Gattyana.

At the stage in which the two caudal cirri appear as two broad short processes there
are at least four scales, and the tentacular cirri are present.

The head is truncated in front, and has a curious spectacle-mark ventrally (Plate
XXVI a, fig. 16), a black spot being situated in each lateral dilatation. The spectacle-
mark is outlined with black and touched with reddish brown, the rest of the head (in
spirit) being pale. It is bounded by the black belt at the ciliated ring, while behind
is the reddish-brown pigment, followed by streaks of black, which also occur at the
segment-junctions. The caudal cirri are yellowish. On the dorsal surface (fig. 15),
again, are two black eyes, obliquely situated on each side anteriorly, a long interval
occurring between them and the black belt of the ciliated ring. Behind the black is
a definite reddish line. The bristles project considerably on each side, so that the stage
is a nectochgete one.

At a slightly later stage (Plate XXVI a, fig. 17) the palpus arises on each side
at a black spot, which at first marks the budding organ. The growth of the various
appendages, however, is irregular, since in this instance the caudal cirri were less
advanced than in those without elevation of the area of the palpus. Coincident with
the projection of the palpus is that of the median tentacle, which forms a rounded boss.

Pallas (1766) made his description of Harrnothoe imbrieata (his Aphrodita lepidota)
from an incomplete and probably young specimen (as Malmgren says) with fourteen pairs
of scales, the latter being partly ciliated, though in the description this is not alluded
to, and the ventral view mistaken for the dorsal (pi. v, fig. 2). He thought it might
readily be distinguished by its longitudinal black band on the dorsum. It occurs, he
says, frequently in the North Sea, between Britain and Belgium.

The Polynoe fulgurans of Ehrenberg, 1 one third of a line long, may be the pelagic
young of this or an allied form.

De Quatrefages considered the Polynoe fas ciculos a and the P. maculata of Grube 2
as pertaining to this species, but the descriptions are so vague that doubt exists.

This was the only member of the group procured in the Siberian Expedition of
Gerstfeldt, as described by Grube. De Quatrefages, in his work on the c Anneles '

1 c Das Leuchten des Meeres/ 1835.

2 < Acfcin., Echin., u. Wurm./ p. 87, 1840.

HARMOTHOE SPINIFERA. 327

(1865) says it has forty-two to forty-four segments, but adds nothing of interest. H.
Lenz (1878) agrees with Mobius in considering that this form included Lxnilla glabra,
Antinoe Sarsii, and Evarne impar ; but, as already mentioned, this was due to insufficient
acquaintance with the group.

Theel (1879) observes that while it is common on the west of Nova Zembla, he
found none in the Kara Sea. He also records his dissent from the view of Mobius and
Lenz, who included under this species Evarne impar, Antinoe Sarsii, and Lsevilla glabra,
the second being supposed to be the young stage of II. imbricata. I have already alluded
to this in the ' Challenger Report/ 1 The habits, for instance, of Evarne impar quite
differ from those of H. imbricata, the former being much more irritable and active.

Kallenbach 2 gives an account of the minute structure of the scales of this species.

Hornell (1891) alludes to the varied coloration of Harmothoe imbricata, e. g. as
striped with a median black band edged with pale border. He considers Theel is correct
in stating that only the largest examples possess any notable rounded processes on the
scales.

2. Haemothoe spinifeea, Ehlers, 1864.

Specific Characters. — Body narrow and elongate. Head elongated from before
backward, and the anterior peaks produced at the sides of the broad base of the median
tentacle. The anterior pair of eyes lie under these peaks, but are also seen from the
dorsum. Posterior eyes at the nuchal collar, dorsal, and nearer each other than the
anterior pair, from which they are separated by almost the entire length of the head.
Palpi of average length, brownish, with rows of small truncate papillae, which are some-
times bifid at the tip. Median tentacle with a broad base (ceratophore), a brownish
column (cerato style), a pale tip with little or no enlargement beneath, and numerous
clavate papillao. The lateral tentacles are inferior, subulate, and brownish. Tentacular
and dorsal cirri (which are short) brownish, furnished with a few clavate papillae, and
slightly enlarged below the pale tip. Ventral cirrus with a few clavate papillae. Scales
fifteen pairs, almost smooth at the margin, only a few small clavate cilia occurring in
some. A limited area towards the inner edge shows a series of spikes. First pair nearly
circular, brownish ; succeeding anterior scales reniform, blackish, with a metallic lustre ;
posterior scales mottled like granite. Bristles comparatively short, the dorsal slightly
curved, finely serrated, and with a short broad tip like a paper-scraper ; ventral bristles
with short spinous regions, and the smooth terminal part boldly bifid.

Synonyms.

1864. Polynoe spinifera, Ehlers. Die Borstenwiirmer, p. 96, Taf. iii, figs. 1 — 4 and 6.

1875. „ crassipalpa, Marenzeller (?). Sitzb. d. k. Akad., 1875, p. 6 (sep. Abdr.).

1876. Harmothoe Sibbaldii, Mcintosh. Trans. Zool. Soc, ix, p. 378, pi. lxviii, f. 1 — 3.
1880. Polynoe spinifera, Langerhans. Zeit. f. w. Zool., xxxiii, p. 275.

1888. Harmothoe spinifera, De St. Joseph. Ann. des Sc. nat. (7), v, p. 171, Taf. xiv, f. 4.

1 P. 58.

2 ' Inaug. Dissert./ Jena, 1883, p. 23.

43

328 HARMOTHOE SPINIFERA.

Habitat. — Several examples were dredged in 6 to 10 fathoms, amongst tangle-roots,
in Bressay Sound, Shetland, classic ground to marine zoologists since the days of Edward
Forbes. A form apparently identical occurs in the British Museum, from chinks in the
rocks, Polperro, Cornwall. The same form is found in the Adriatic.

Length about seven-tenths of an inch.

The head (Plate XXVIII, fig. 3) is elongated from before backward, and since the
large anterior pair of eyes are carried outward, almost at the tips of the peaks, beneath
which they are placed, and are visible from the dorsum, the condition is diagnostic.
They are, moreover, wider apart than the posterior pair, and look forward and outward.
The smaller posterior pair are dorsal, situated near each other, and almost touch the
nuchal collar (in spirit). The two pairs of eyes are thus separated by a long antero-
posterior interval. The median tentacle has a broad base, a brownish column, a pale tip
with little or no enlargement beneath, and well-marked clavate cilia. The lateral ten-
tacles are inferior in position, small, subulate, and with similar clavate cilia. The
tentacular cirri are also brownish, furnished with numerous clavate cilia, and slightly
enlarged below the filiform tip. The palpi are brownish, with rows of small blunt papillae,
which are sometimes bifid at the tip.

The body, which consists of thirty-seven segments, is comparatively short and of
more uniform diameter than usual, being only a little narrowed anteriorly, and somewhat
more posteriorly. Transverse markings occur on the dorsum between the scale-pedicles
and their homologues. No pigment remained on the dorsum in the preparations, though
it is probable traces of such were present during life. On the ventral surface traces of
brownish pigment occur at the sides of the prominent fold in front of the mouth. The
lateral prominences at the bases of the feet are well marked, but the segmental processes
are minute.

Scales (Plate XXXIII, fig. 4), fifteen pairs. The first are small, nearly circular, and
their light brownish colour contrasts strongly with the succeeding. The surface is
studded with short blunt spines, which are especially distinct at the outer border, where
there are also a few short clavate cilia. The rest of the margin is smooth. The second
pair are reniform, the exposed parts being almost uniformly black, with a silky sheen and
a smooth margin, except at the outer border, where five or six very short clavate cilia
occur at intervals. The short blunt spines are distributed over the posterior two thirds
of the surface. The third scales are also blackish with the same metallic sheen, but
have a few minute pale points besides the microscopic blunt spines. The pale specks
increase in size in the succeeding scales, and the pigment becomes paler, the posterior
pair, indeed, being mottled like granite. The general shape of the posterior scales is
irregularly rounded or ovoid, with the scar for attachment towards the anterior and
outer border. The penultimate and last' pairs are much elongated from before back-
wards. The number of the minute blunt spines diminishes posteriorly, so that they are
chiefly confined to the outer border, where an occasional short clavate cilium is observed.
The definite pale areas are due to the absence of pigment in the areola of the epiderm
(hypoderm), while in the dark parts each areola is deeply pigmented. If the P. spinifera
of Ehlers is the same form the scales had a greyish- violet sheen.

HAPMOTHOE SPIXIFEEA. 329

Feet. — The first foot shows dorsal bristles, which are little altered from the typical
form.

The second foot has dorsally a series of short broad bristles, the tips especially
being characteristic. Yentrally the shafts of the bristles of the region are stout, and
the spinous tips well marked. The superior bristles have very short smooth tips, which
are bifid. There is thus less change than usual in the foot.

In the typical foot the superior branch bears a short cirrus, the tip of which (in
spirit) reaches the extremity of the bristles. It is almost cylindrical, except near the tip,
where a gradual diminution occurs. The surface has rather numerous stout clavate cilia,
which are best developed on and near the slight enlargement below the filiform tip. The
ventral cirrus has an enlarged base, reaches a little further than the insertion of the
inferior ventral bristles, and has a few stout clavate cilia.

The dorsal branch of the foot has comparatively short and not very stout bristles,
slightly curved, and finely serrated. The smooth tips are peculiar, being fashioned like
a blunt Esquimaux harpoon or paper-scraper, as represented in one of the larger
examples (Plate XXXVIII, fig. 17). The spinous rows at the distal part project
characteristically at a greater angle than usual, but are generally obscured by dense
granular parasitic growths and much

The ventral bristles have comparatively stout shafts and short spinous regions.
The smooth tip is also short. Every bristle is boldly bifid. The superior series (with
longer spinous rows) have a very short smooth region (Plate XXXVIII, fig. 18), the
longer terminal part being only diminished on its dorsal edge — not hooked, while
the secondary process is stout and goes straight to a point, thus differing from the
attenuated divisions of Parmenis ljungmani. The smooth tips of the middle and inferior
bristles (Plate XXXVIII, fig. 19) are somewhat longer and have a slight hook, but the
secondary process is straight. The spinous regions in these are proportionally
short.

Reproduction. — The specimens procured in July had the ova well advanced, so that
their spawning period would seem to be in autumn at latest. It is interesting that
the examples of Ehlers and De Saint-Joseph also carried eggs. The latter found a
specimen of 7 mm. in the same condition.

This species presents considerable differences from Parmenis ljungmani, Malmgren.
The body is larger and broader, the segments thirty-seven instead of thirty-five or
thirty-six; the head is more elongated, and the arrangement of the eyes different. Thus,
for instance, the smaller anterior eyes in the northern form are situated at an incurved
region of the head just in front of the middle. The dorsal bristles are thicker than the
ventral, and have the peculiar tips indicated, whereas in P. ljungmani the tips are simply
tapered. The ventral bristles have short spinous regions in both, but in the Zetlandic
form the tip is less hooked, and the secondary process is short, stout, and straight.
There are points of similarity between the forms, but there is no warrant for uniting
them.

The Polynoe spinifera of Ehlers approaches this species very nearly, and he probably
overlooked the papillas on the palpi. It is possible the small size of his example (7 mm.)
may have been the cause of certain variations. The figures and remarks of Langer-

330 HARMOTHOE ZETLANDICA.

hans 1 point to the same conclusion. In the southern forms the pallor of the first pair of
scales is characteristic. The Polynoe crassipalpa of Marenzeller from the Adriatic is
a very closely allied form, but, if the author's figures and descriptions are to be relied on,
the British form is distinct, the tip of the dorsal bristles alone being characteristic. In
Marenzeller's specimen also the large anterior eyes are not so near the tip of the peak as
in the British form. Levinsen (1883) seems to think this closely approaches Harmothoe
ljungmani, but the foregoing remarks demonstrate the differences.

3. Harmothoe zetlandioa, Mcintosh, 1876.

Specific Characters. — Body sublinear, narrowed anteriorly and posteriorly ; bristle-
bearing segments thirty-five to thirty-six. Head elongated from before backwards, with
two acute peaks in front. Tentacles and cirri without apparent enlargement below the
tip, and with sparsely distributed short clavate cilia ; lateral tentacles inferior. Palpi
minutely papillose. Eyes all visible from the dorsum, the larger anterior pair at the
base of the peaks. Scales fifteen pairs, covering the dorsum, ovate or oval with the
exception of the first pair, which are rounded, with few and indistinct papillae. Dorsal
bristles strongly curved, sharp-pointed, and with distinct rows of spines, the sharp tip
being bare ; ventral bristles with short spinous regions and smooth tips, the latter being
hooked and having a long straight secondary process, which is parallel and closely applied
to the other division, — that is, no gap is present.

Synonym.

1876. Harmothoe zetlandica, Mcintosh. Trans. Zool. Soc, ix, p. 379, pi. lxviii, f. 4 and 5; pi. lxix,

f. 1.

Habitat. — Dredged in 5 fathoms amongst the tangle-roots in Bressay Sound,
Shetland, July, 1871.

The specimens are about half an inch in length.

Head (XXVIII, fig. 1) somewhat elongated from before backwards, terminating
anteriorly in two pointed lobes on each side of the median tentacle. The eyes are all
visible from the dorsum ; the smaller posterior pair lie in front of the nuchal collar, while
the larger anterior pair are situated at the base of the peaks and look laterally and
f/V v dorsally. They are not so far forward as the anterior eyes of Harmothoe spinifera.

The cephalic appendages are somewhat short. The median tentacle is not much, if at
<\P> all, enlarged below the filiform tip, and is furnished with a few clavate cilia. In some it

is deeply pigmented at the base. The lateral tentacles are short, enlarged at the base,
but tapering at the tips after the manner of the ventral cirrus, and with sparse but
distinct clavate cilia. The palpi are gently tapered from base to apex, and, while smooth
or only wrinkled under a low power, show minute papillae under a high power. The
tentacular cirri taper from base to apex, have no enlargement below the latter, and have
clavate cilia like those of the median tentacle.

1 c

Zeit. f. w. Zool./ xxxiii, p. 275.

HAKMOTHOE ZBTLANDIOA. 331

Body elongate and somewhat narrow, consisting of about thirty-five bristle-bearing
segments. The segments dorsally present no peculiarity, while inferiorly the depressed
median region proceeds from the buccal fold to the tail. The segmental papillae seem
hardly to project beyond the elevation. The body is of a pale or dull straw-colour, the
translucent scales showing only a few pale touches. Posteriorly the anus projects on a
median process.

Proboscis. — The extruded organ presents nine papillee along each edge, and the teeth
are pale brown.

Scales (Plate XXXII, fig. 15). — The species was formerly stated to have fourteen pairs
of scales, but a re-examination of the broken specimens points to the probability that fifteen
pairs are present. They cover the dorsum, are rather thin, translucent, and soft, and
seem to be smooth under a lens ; but under a high power short clavate papilla occur
sparsely along the outer and posterior border, and over the usual area externally. The
papillae along the outer and posterior border are not to be confounded with the
appearances found in the granular area of the epiderm. They are much more minute
than those of allied species. Only a microscopic papilla here and there on the
same border projects beyond the smooth outline. The first pair are small and round,
the size increasing posteriorly to the twelfth, when a diminution again occurs in those
behind. Only a few show a slightly reniform outline, the majority being more or less
ovoid.

Feet. — The first foot shows bristles of the dorsal type.

In the second foot the dorsal bristles are curved, and have well-marked spinous
rows. The ventral bristles, again, differ from the typical forms in the proportionally
longer and more finely spinous regions, and in the simple tip, which is but little hooked.

The third foot has more or less become typical, except that the ventral bristles are
more slender.

The typical foot (Plate XXX, fig. 2) presents dorsally a series of divergent, stout,
sharp-pointed, and brittle bristles, with well-marked transverse spinous rows. The
spinous region has a distinct curve. The smooth portion at the tip is of considerable
length, has a slight bend, and tapers gently to the point (Plate XXXVIII, fig. 20, and
front view, Plate XXXVIII, fig. 20 a). The ventral bristles (Plate XXXVIII, fig. 21)
have superiorly a short spinous part of five or six rows, and a smooth terminal region
which forms a well-marked hook. The secondary process is remarkably long, and passes
straight outward parallel with the former, and in the upper bristles reaches as far as the
terminal hook.

In the terminal feet the dorsal bristles alter little, except that they become smaller
and the spinous rows more prominent. The ventral bristles, again, have shorter shafts
and much more slender and elongate spinous regions and simple tips.

The bristles throughout are pale yellowish, and the dorsal are often loaded with
algoid and other parasitic growths.

The dorsal cirri, like the tentacular cirri, taper from base to apex, which is filiform,
and reaches the extremity of the ventral bristles (in spirit). Their surface has sparsely
distributed short clavate cilia. The ventral cirri are tumid above the basal region, then
taper to the extremity, and have a few clavate cilia.

332 HARMOTHOE LJUNGMANI.

When contrasted with the young of Harmothoe imbricata the head of this form is
much more elongated antero-posteriorly, and the four eyes are visible from the dorsum,
whereas in H. imbricata the posterior pair only are generally seen. The tentacles and
other cephalic processes are different. The body is narrower and longer, and the bristles
are much smaller, so as to alter the outline ; and their structure, as well as that of the
scales, is essentially different. From the Parmenis ljungmani of Malmgren, 1 to which it
is allied, it differs in the structure of both dorsal and ventral bristles as well as in that
of the scales.

Allied forms to both this and the succeeding species (Parmenis ljungmani) are the
Polynoe vasculosa of Olaparede, 2 and the Polynoe crassipalpa of Marenzeller. 3 They,
however, differ in regard to the shape of the head and the structure of the bristles.

This species takes the place of the closely related Parmenis ljungmani of the Outer
Hebrides, and the representatives of both appear to be few. It is probable that, like
allied forms, it may be commensalistic.

4. Haemothoe ljungmani, Malmgren, 1867.

Specific Characters. — Body small, sublinear, bristled segments thirty-five to thirty-
six. Head elongated antero-posteriorly, somewhat narrower in front, and with two short
peaks. Eyes visible from the dorsum ; two at the posterior border, and two larger at the
side in front of the middle line. Tentacles and tentacular cirri short and tapering, with
sparsely distributed clavate cilia. Palpi elongate, subulate, with minute papillae. Ten-
tacular and dorsal cirri alike. Ventral cirri enlarged at the base, similarly ciliated.
Scales fifteen pairs ; first pair suborbicular, the others obliquely ovate or ovoid, increas-
ing in size till the two last pairs, with very few minute papillae along the outer and
posterior border, while the tubercles on the surface are larger and more numerous than
in H. zetlandica. Dorsal bristles stouter than the ventral, rather short, curved and
little tapered, with close rows of spines which extend to the tip. The ventral bristles
have short spinous tips ending in a smooth and characteristically curved claw, while the
secondary process leaves a gap between it and the base of the hook.

Synonyms.
1867. Parmenis ljungmani, Malmgren. Ann. Polycheeta, p. 11, pi. i, fig. 2.

1875. Harmothoe Macleodi, Mcintosh. Invert, and Fish., St. A., p. 116.

1876. „ „ Ibid. Trans. Zool. Soc., ix, p. 882, pi. lxix, f. 2, 3.

1888. „ „ De Saint Joseph. Ann. d. sc. nat. (7), v, p. 168, pi. vii, f. 37 — 40.

Habitat. — Between tide-marks under stones at Lochmaddy, North Uist, and in the
stomach of the cod, St. Andrews (E. M.). Shores of France — Dinard.
Length about 14 mm. ; breadth across bristles 3 mm.

i ' Annulat. Polychast., &c.,' p. 11, pi. i, fig. 2, 1867.

2 ' Annel. Chetop. Nap., Suppl./ p. 12, pi. i, f. 4.

3 " Zur Kenntniss d. Adriat. Annel./' < Sitz. der k. Akad. Wissensch./ April 1, 1874, p. 6,Taf. ii.

HARMOTHOE LJUNGMANI. 333

Read (Plate XXXVIII, fig. 4) elongated from before backward, terminating in two
peaks anteriorly, and with eyes similarly placed to those in R. zetlandica, viz. a pair of
smaller eyes on the dorsum in front of the nuchal collar, and a larger pair on the sides
anteriorly in front of the middle line of the head, and at the base of the peaks. All are
visible from the dorsum, though the main vision of the anterior eyes is lateral. The palpi
have rows of minute papillae, and are similar to those of R. zetlandica, as also are the
tentacles and tentacular cirri — which have clavate cilia. All are comparatively short, and
taper gently to the extremity without an evident enlargement below the tip (in spirit).

Body elongated, rather narrow, slightly tapered in front, more so posteriorly, and
with comparatively short bristles, the segments bearing these being from thirty-five to
thirty-six. In regard to external structure and coloration it agrees with the foregoing
form.

Scales (Plate XXXII, fig. 14) amount to fifteen pairs, are thin and semi- translucent,
have a very few minute cilia along the outer and posterior border, while the papillae,
which are generally scattered over the surface, are larger and more numerous than in R.
zetlandica. Along the anterior edge somewhat smaller papillae occur in rows. In shape
they correspond with those of the latter, the first pair being circular, the rest more or
less ovoid and increasing in size to the twelfth or thirteenth.

Feet. — In the first foot the upper division bears a few short dorsal bristles similar
in type, though more curved than the succeeding.

In the second foot the dorsal bristles are already numerous and of considerable
length. The curvature is well marked, especially at the tip, which shows no smooth
portion, the spinous rows passing quite to the extremity. The bristles of the inferior
division present superiorly almost the typical structure with the somewhat broad short
tip, and the long secondary process. The bristles of the middle division have longer tips,
very slightly bifid, while in the inferior series the tips are elongated, simple, and with
long spinous rows, which in some views are more or less pinnate.

As we proceed backwards the bristles gradually assume the typical form, the dorsal
becoming longer and stronger (Plate XXXVIII, fig. 22) as well as less curved. Moreover
the tip is but slightly tapered, and the spinous rows pass to the extremity and are closely
arranged, except in a few of the short forms next the body. The ventral division has
boldly bifid bristles (Plate XXXVIII, fig. 23) throughout, the entire tip being rather
broad and the spinous region short. The inner line of the hook forms a different angle
from that in R. spinifera, and leaves a wider gap distally. The secondary process
diminishes in length from the superior to the inferior series of bristles, so that in the
latter it scarcely reaches to the middle of the peculiarly curved terminal hook. The
spinous region of the tips of the inferior bristles is very short. The bristles generally
are faintly yellowish, and the dorsal form a favourable site for the development of
parasitic growths.

The dorsal cirri are comparatively short, and taper almost from the base to the
slender tip. Sparsely distributed clavate cilia occur on the surface below the latter.
The ventral cirri are enlarged at the base, and have also a few clavate cilia.

When the species was described in 1876 it was thought to differ from the Parmenis
ljungmani of Malmgren, on the grounds of the proportional strength and structure of the

334 HARMOTHOE ANTILOPIS.

dorsal bristles, and in the pale semi-translucent scales, but the ventral bristles approached
each other closely. A re -examination of all the characters and of Malmgren's description
now inclines me to unite them, especially as in the generic characters he states that the
dorsal bristles are shorter and thicker than the ventral, while in the specific he says they
are much shorter and a little more slender than the inferior bristles. Moreover, if his
figure of the dorsal bristle be supposed to be that next the ventral series (always the
most slender and elongate), the distinctions more or less disappear. The only doubtful
point is the possible existence of a species agreeing in all respects with that from
Lochmaddy, but having more elongated dorsal bristles. Levinsen seems to be of the
same opinion.

Few individuals of this form would seem to exist. Only a single example was
obtained at Lochmaddy, and one in the stomach of a cod at St, Andrews. Baron de St.
Joseph found one in the tube of Lanice conchilega at Dinard. With the two foregoing it
forms a special series — closely approaching each other and such forms as the Polynoe
crassijpalpa of Marenzeller.

5. Harmothoe antilopis, Mcintosh, 1876.

Specific Characters. — Length about three-quarters of an inch or a little more. Head
broad behind, peaks truncated in front, two widely separated eyes posteriorly, and a
larger pair laterally and below the short peaks, not visible from the dorsum. Median
tentacle and tentacular and dorsal cirri moderately long, slightly enlarged below the
filiform tip, and covered with numerous cilia with slightly bulbous tips. Lateral
tentacles inferior. Palpi of average length — with clavate papillas. Body, moderately
elongated, of considerable breadth, bristled segments 33 — 35. Scales fifteen pairs,
mostly reniform, entirely covering the dorsum, with long cilia externally and
posteriorly and shorter on surface, which also has large horny papillae ending in
spines with hispid tips. Dorsal bristles elongated, with smooth tips and very
distinct spinous rows (resembling the horns of an antelope), ventral with bifid tips,
scarcely hooked, and in some the distal part of the spinous region is narrower than the
smooth tip. Ventral cirri rather slender and tapering, with short clavate cilia. The
segmental papillae are but slightly marked.

Synonyms.

1876. Earmotlwe antilopis, Mcintosh. Trans. Zool. Soc, ix, p. 383, pi. lxix, f. 4 — 6.

1877. Evarne mazeli, Marion. Eev. des Sc. nat., April 15th, 1877.

1879. „ antilopes, Marion. Ann. des Sc. nat., 1879, p. 13, pi. xv, f. 1 to If.
1884. Polynoe antilopes, Cams. Faun. Medit., p. 200.

Habitat. — First procured at Lochmaddy, North Uist, in 1865; off the Hebrides
(J. G. J.); on muddy sand at 173 fathoms, 'Porcupine,' 1869; < Porcupine,' 1870, 576
fathoms, on the Channel slope ; 227 fathoms outside Gibraltar; and 358 fathoms in the
Atlantic. It is included in the Mediterranean fauna also by Professors Marion and Cams

HARMOTHOE ANTILOPIS. 335

Its range is thus extensive, but the numbers hitherto have been few, generally a single
example at each station.

Length from three quarters to nearly an inch.

Head (Plate XXVIII, fig. 13) somewhat broad behind, with the anterior angles
ending in very short peaks. Two comparatively large and somewhat widely separated
eyes occur near the posterior border. The larger anterior pair are not visible from the
dorsum, being situated laterally and below the truncated angles of the snout. In most
specimens in spirit the pigment of the eyes has become very faint. The median tentacle
is moderately long, slightly enlarged below the filiform extremity, and covered with
numerous cilia with very slightly enlarged tips. The lateral tentacles are inferior, have a
slight enlargement below the tip, and are also supplied with clavate cilia. The tentacular
cirri are similar to the median tentacle, the cilia being continued about a third up the
filiform termination. The palpi are of moderate length, with rows of minute clavate
papillae, which toward the termination are dilated and then constricted below the slightly
granular tip.

The body is moderately elongated, of considerable breadth, not much tapered
anteriorly and only a little more so posteriorly, and with the usual markings dorsally
and ventrally. Bristled segments from thirty-three to thirty-five. The segmental
papillae do not appear to be produced as tubular processes, only the projecting " boss "
or elevation indicating this structure.

The scales (Plate XXXII, fig. 16) amount to fifteen pairs, and entirely cover the
dorsum. The first pair are more or less rounded, the rest reniform, the outer lobe being
larger than the inner. They are fringed on the outer and posterior borders as well as
studded over the surface with cilia, which become very long on the former border, and
the tips are slightly bulbous. They are often coated with a granular parasitic growth.
The horny papillae or tubercles on the surface are large and boldly marked, the dilated
tips forming blunt processes or spines truncated at the tip, which is hispid with sharp
processes, and in favourable specimens they resemble a rosette of spikes. The appear-
ance is thus characteristic. A pale brownish coloration occurs on the dorsal surface
where the scales overlap, but otherwise they are pale with the exception of the minute
brownish spines. The under surface is smooth and iridescent.

Feet. — The first foot has a few short bristles of the normal dorsal type, and very
slightly curved. The second foot has a considerable tuft of short dorsal bristles, only
the inner being curved, those next the spine being more or less straight, with the well-
marked spinous rows from which the name of the species is derived. The ventral series
of slender bristles has long spinous regions superiorly and shorter inferiorly. All have
simple tapering tips. These bristles are proportionally shorter and stouter than the
typical series. The bristles gradually elongate and the ventral series become bifid, so
that in the typical foot (Plate XXIX, fig. 14) the dorsal division carries a conspicuously
long and strong series, the inner bristles being shorter and more curved as well as more
abruptly pointed at the tip, which has a short smooth portion; while the outer and
inferior series are long, very gently tapered, and slightly curved, with a short, conical,
bare portion terminally. All are distinctly marked by transverse spinous rows at rather
wide intervals, whence the name of the species, from the resemblance of these organs to

44

336 HAKMOTHOE HALIAETI.

the horns of certain antelopes, such as Hippotragus oryx. One of the larger bristles is
represented in Plate XXXVIII, fig. 24. The ventral series again forms a dense tuft of
fine and by no means long bristles, commencing superiorly with a group of about three,
with elongate spinous regions, tapering to an acutely pointed simple tip. The adjacent
series has also long, slender, spinous regions, but the tips are bifid, the part which is
usually hooked having scarcely a trace of a curve, and the secondary process being
closely applied (Plate XXXVIII, fig. 25). The tips become shorter and stouter
inferiorly, and the secondary process more apparent. Though a slight curve is present
the tip is scarcely hooked (Plate XXXVIII, fig. 26, representing one of the stoutest
forms). The upper third of the spinous region is even narrower than the bifid part
behind the hook at the tip, a peculiarity seldom seen. Inferiorly the simple tips as well
as the bristles themselves become shorter.

Posteriorly (in the last feet) both dorsal and ventral bristles become much more
slender as well as smaller. The dorsal, however, show even in the least developed the
well-marked spinous rows. The ventral bristles have long and finely tapered spinous
regions and simple tips.

The dorsal bristles are often loaded with parasitic growths, one of the most
beautiful being a handsome stalked infusorian.

The dorsal cirri stretch beyond the bristles, and have a similar structure to the
median tentacle. The ventral cirri are rather slender and elongate, with a filiform tip.

Reproduction. — All that can be said about the reproduction of the species is that in
an example procured in early autumn (August) numerous ripe eggs occurred between
the feet and on the dorsum under the scales. Though the body was ruptured it is
probable these escaped, at least in some cases, naturally. A ripe form also was dredged
in the Minch in July.

This species approaches Evarne, but a glance at the head and the arrangement of
the eyes at once differentiates it, while the structure of the scales and the characters of
the dorsal and ventral bristles still further emphasise the distinctions. The Polynoe
spinifera of Ehlers is a closely allied form, but the anterior pair of eyes, if the figure is to
be trusted, quite differ, since they are visible from the dorsum. The details of the bristles
are not given. In the same way, while admitting the relationship between the species
and Evarne as above mentioned, I do not think with my esteemed colleague, Prof.
Marion, for the reasons indicated, that it should be placed in that genus. It is true that
in the French naturalist's figure the anterior pair of eyes are dorsal, but such was not
the case in the examples observed here. The character of the spinous rows of the
dorsal bristles differs from that of either Evarne impar or E. Johnstoni.

6. Harmothoe Haliaeti, Mcintosh, 1876.

Specific Characters. — No complete example has yet been found. The dorsal bristles
are rather long, slender, and slightly curved, with conspicuous rows of spines, and a very
short, bare portion at the tip. The ventral bristles have slightly hooked tips, and the
secondary process is either straight or bends outward a little at the tip. The ventral

HAEMOTHOE FRASER-THOMSONL 337

cirrus is somewhat enlarged at the base, filiform at the tip, and with rather long papillae
sparsely distributed.

Synonyms.

1876. Harmothoe Haliaeti, Mcintosh. Trans. Zool. Soc, ix, p. 384, pi. lxix, f. 7 — 10.

1885. „ ,, idem (partim). ' Challenger 3 Annel., p. 96.

1886. ,, „ Harvey-Gibson. Yerm. Liverp., 149.

1891. Polynoe (Harmothoe) Haliaeti, Hornell. Op. cit., 232, pi. xiii, f. 5.

Habitat. — The first specimen was procured in the Minch in July by Dr. G-wyn
Jeffreys, who did so much in dredging examples of the British Annelids.

In a posterior foot the dorsal branch bears a series of rather long, slender, slightly
curved, bristles with conspicuous rows of spines (Plate XXXVIII, fig. 27). Such y /
bristles, when viewed antero-posteriorly, present a much narrower aspect than when /> /C <■

seen in profile. The arrangement of the spinous rows is alternate, as in the ventral
bristles. Only a very small portion of the tip is smooth — a feature of moment when it
is contrasted with the larger and longer tip of H. Fraser-Thomsoni. The dorsal bristles
are often coated with a brownish granular substance and algoid growths. The superior
ventral bristles have elongate spinous portions and slender tips (Plate XXXIX, fig. 1).
At first the bifid tips are almost straight or very slightly curved, but they soon become
more characteristic (Plate XXXIX, fig. 2). The fades of the tip is even more
diagnostic in the inferior series (Plate XXXIX, fig. 3), where the spinous region is
distinctly curved. In the superior group the secondary process is nearly straight, but
in the others it bends outwards at the tip, which has a distinct hook.

The inferior cirrus is enlarged at the base, slender and filiform distally, and
furnished with rather long papillae sparsely distributed.

Hornell found an allied form at Port Erin, Isle of Man, in fifteen fathoms. He
describes the scales as densely fimbriated round the border after the manner of
Lepidonotus squamatus, so that there is need for further investigation as to the condition
in this respect. Unfortunately this specimen could not be found in the collection at
Liverpool, and the others are incomplete.

7. Haemothoe Feasee-Thomsoni, 1 Mcintosh, 1396.

Specific Characters. — Head somewhat resembles that of Lagisca, having a pair of
widely separated eyes posteriorly, and a larger pair on the lateral eminence. Median
tentacle absent. Palpi of moderate length, with rows of minute papillae. Body of
considerable length and breadth ; bristled segments thirty-nine to forty. Dorsum has
touches of brown pigment posteriorly as in Lagisca. The lateral segmental eminences
are prominent, but there is no process. Scales mottled brown, fifteen pairs, covering
the dorsum; first small and rounded, rest more or less ovoid; border smooth, anterior

1 Named after my early and valued friend, the late Dr. Fraser Thomson,, of Perth.

338 HABMOTHOE FRASER-THOMSONI.

and inner half studded with small horny papilla, outer and posterior areas have
sparsely distributed, large tubercles, with an interrupted row along the posterior border.
Dorsal bristles stout, moderately long, and slightly curved, with closely arranged spinous
rows and a short, smooth tip. Ventral bristles bifid, the secondary process coming off
at an angle. Dorsal cirri appear to be fusiform — from the gradual nature of the
dilatation and the long filiform tip, and have clavate cilia. Ventral cirri slender, with a
few clavate cilia.

Synonyms.
1885. Harmothoe Haliaeti (partim), Mcintosh. l Challenger ; AnneL, p. 96.
1896. „ Fraser-Thomsoni, Mcintosh. Sc. Proceed. R, Dub. Soc, vol. viii, n. s., p. 401.

. { [ f\ Habitat.— Dredged in the 'Knight Errant' in 1880, in the Atlantic, at a depth of

fifty-three fathoms, and procured by Prof. Haddon in the Royal Irish Academy's

Expedition, July 15th, 1886, in ninety-three fathoms off the south-west coast of Ireland
— along with Malmgrenia castanea, which occurred on Spatangus raschii.
^ ^ The head (Plate XXVIII, fig. 7) somewhat resembles that of Lagisca, having a pair

of well-marked and widely separated eyes posteriorly, and a larger pair still more widely
separated on the anterior lateral prominence, looking laterally and dorsally. Both pairs
are conspicuous from the dorsum. The head terminates anteriorly in sharp peaks on
each side of the median tentacle. The lateral are brownish, subulate, and with a long
filamentous tip. The palpi appear to be of moderate length, and to have rows of minute
papilla3, but they had been dried.

The body is of moderate breadth and of considerable length, narrowed abruptly in
front, but — from the middle — gradually diminishing posteriorly. The bristled segments
seem to amount to thirty-nine or forty. The brownish pigment of the dorsum posteriorly
is very prettily arranged in lozenges and touches, as often seen in species of Lagisca.
The lateral eminences for the segmental apertures are well marked, and a small papilla
projects between the feet.

The scales (Plate XXXII, fig. 11) number fifteen pairs, and are thin, translucent, and
cover the dorsum. The first pair are small, rounded, and studded with minute
spines and tubercles. Few seem to be reniform, the majority being ovoid. The border
is smooth throughout. The anterior and inner half is densely covered with small horny
papillas or tubercles, while the outer area and that behind the scar for the pedicle have
large tubercles sparsely distributed. The small tubercles are grouped with a few
larger spines along the outer edge, while the posterior border is marked by about eight
large blunt spines or tubercles. Some of the latter under pressure project a little
beyond the posterior border, but no trace of cilia exists. Such scales thus differ quite
from those mentioned by Mr. Hornell, so that two species are involved. The scales are
mottled with brown, best marked round the scar for attachment.

Feet. — The first foot is not in a condition for description.

The second foot presents a dense tuft of short, strong, and very slightly curved
dorsal bristles with smooth tips, and distinct though closely arranged spinous rows.
Ventral cirri somewhat tumid at the base, and with a few comparatively long cilia.

HAKMOTHOE MARPHYS.E. 339

The ventral bristles form two groups, a stronger upper series and a more slender inferior
group, both with longer and more tapered tips than the typical. The tips of most of the
upper series are spear-shaped, though in one or two a double contour is seen. The rows
of spines are boldly marked. The inferior group has very long tapering tips, which are
hair-like in their attenuation, and the spinous rows are finer than in the superior
bristles.

In the third foot the dorsal bristles have increased in size and strength, and the
terminal bare region is more distinct. While the upper and lower ventral bristles have
simple tips, the median show bifid tips. The feet gradually assume the typical condition
as we proceed backwards.

In the fully formed foot (Plate XXIX, fig. 15) the dorsal branch bears a series of
moderately elongate and stout dorsal bristles with closely arranged spinous rows, only a
short portion at the tip being smooth. The curvature is slight (Plate XXXIX, fig. 4).
The ventral bristles are of average length, the upper series with long spinous regions
and simple tips, the next with shorter spinous rows and bifid tips (Plate XXXIX, fig. 5).
The tips in the stouter forms are slightly hooked, and the secondary process comes off at
an angle. Only in the upper forms with the longer distal regions does the secondary
process form a small angle. At the ventral edge of the series the spinous region
becomes much shorter, but almost all are bifid.

These bristles differ from those of HarmotJioe Haliaeti.

The dorsal cirri are somewhat slender and elongate, with a slight swelling — marked
on each side by a brown bar below the long filamentous tip. They are sparsely covered
with clavate cilia, the longest of which scarcely attain half the diameter of the thickest
part of the cirrus. The posterior cirri are very slender throughout. The ventral cirri
are rather small and slender, and have a few short clavate cilia.

Loxosomae occur on the skin of the dorsal division of the feet, and also on some of
the dorsal bristles (Plate XXXIX, fig. 6).

8. Harmothoe marphysj:, Mcintosh, 1876.

Specific Characters. — Head elongated, rounded in front, widest posteriorly. Eyes
small, the larger anterior pair further apart and situated laterally in front of the middle
line. Posterior pair in front of the nuchal collar. Median tentacle short, with clavate
cilia; lateral short, with filiform tips and a few clavate cilia. Palpi (in spirit) short and
stout, with delicately tapered tips. The tentacular and dorsal cirri are comparatively
short and slender, and the tips finely pointed. A few clavate cilia occur on the surface.
Body somewhat elongate, about three quarters of an inch, and with thirty-two to thirty-
three bristle-bearing segments. The feet increase in bulk after the twelfth (sexual).
Scales thirteen or fourteen pairs (it may be fifteen), smooth under a lens-, but under a high
power showing widely separated horny papillas. In shape they are rounded in front,
reniform or ovoid posteriorly. Dorsal division of the foot little developed, and the tuft
of bristles minute. Ventral bristles with short spinous regions and mostly with bifid

340 HARMOTHOE MARPHTSiE.

clavate cilia. Colour pale brownish inclining to buff, with a red patch on the head
and a purplish hue over the proboscis. Under surface pinkish with a broad streak of
carmine.

Synonyms.

1876. Earmothoe marjphysse, Mcintosh. Trans. Zool. Soc, ix, p. 384, pi. lxix, f. 11—14, and

pi. lxx, f. 7.
1890. „ „ Malaquin. Ann el. Boulon., 21.

Habitat. — In the galleries of Marphysa sanguinea in Guernsey, and from chinks in
the rocks, Polperro (British Museum).

Head (Plate XXVII, fig. 11) rather elongated from before backward, rounded in
front instead of having the usual peaks, and with the widest part behind the middle. Eyes
small ; the larger anterior pair wider apart, and situated laterally in front of the middle
line at the edge of the red patch on the head. The small posterior pair lie in front of
the nuchal collar and behind the red patch. A slight median groove runs forward to
the base of the median tentacle, the column of which is absent. The lateral tentacles are
inferior (i. e> below the rounded anterior border of the head), short, with filiform tips, and
have a few short clavate papillae. The palpi are short and stout, with delicately tapered
extremities. No papilla) were visible, but they may be present in section. The tentacular
cirri are comparatively short and slender, with finely tapered points. On the surface
are a few clavate cilia.

Body somewhat elongate, with the feet greatly developed posteriorly. Bristle-bearing
segments thirty-two to thirty -three. The segments after the twenty- third (bristled),
however, are in process of renewal. The markings on the dorsum are less distinct than
usual, as the feet are separated by deep clefts. The feet increase in bulk after the twelfth
bristled pair, becoming larger and longer, a feature which may be connected with repro-
duction. An elevated line occupies the middle of the dorsum in the preparation, while
ventrally the median region has a depressed line at each side with a cushion-like ridge
between. The segmental eminence is well marked, and a distinct conical process
extends between the feet. The process is evident about the seventh foot, and continues
almost to the posterior end.

The colour is pale brownish inclining to buff, with a red patch (from the ganglia)
on the head, and a purplish hue just behind (clue to the proboscis), while a faint median
streak occurs on the dorsum. The cirri are pale brownish, pellucid, the two caudal
styles being darkest. The under surface is pinkish, with a broad streak of carmine in
the centre.

The scales (Plate XXXII, fig. 13) appear to be thirteen or fourteen pairs (possibly
fifteen), covering the dorsum and easily separated, but the specimen is not in a condition
to give accuracy in this respect. In front they are rounded, then reniform, and even
somewhat quadrate posteriorly. They are rather thin, apparently smooth and pellucid,
the anterior only having a pale brownish patch on a whitish portion. Under the
microscope, however, a belt of small papillae (spines) occurs within the anterior bay of
the scale, in front of the scar. Other pale points which resemble these (by transmitted

HARMOTHOE MARPHYSJB. 341

Light) are dotted over the lateral area, but they seem to be the end-organs in connection
with the numerous nerves.

Feet, — The first foot has a strong spine and a bristle or two like the dorsal.
The second foot has a considerable number of short dorsal bristles, little tapered.
In the ventral division is a group of comparatively short, strong bristles, with short
spinous regions, and a somewhat elongated smooth portion at the tip which ends in a
probe-point. The length of the spinous region diminishes in the usual manner from
above downwards.

Gradually the feet assume the complete form, many of the ventral bristles, indeed,
in the third foot being bifid, but the dorsal division, instead of becoming more prominent
as in the majority of the Polynoidee, increases very little.

In the fully formed foot (Plate XXIX, fig. 16) the dorsal division is marked by a
long bluntly conical process bearing the spine, above which a short tuft of comparatively
few bristles projects from a small eminence. These bristles (Plate XXXIX, fig. 7, one
of the longer) are slightly curved, delicate, and translucent, with minute rows of spines.
The ventral division consists of a somewhat long and obliquely truncated foot, the spine
occurring at the upper angle in a pit between two fleshy lobes — a larger upper and a
smaller inferior. The bristles have moderately long shafts and short spinous regions.
The upper examples have more elongated spinous regions and simple tips (Plate
XXXIX, fig. 8), while a distinct secondary process is observed in the succeeding forms
(Plate XXXIX, figs. 9 and. 10, the latter being seen from the front). Toward the
ventral edge of the group the spinous regions become very short and the tip simple
(Plate XXXIX, fig. 11). In large examples from Polperro the dorsal edge of this division
of the foot is curiously wrinkled in the preparations. The spinous region of the ventral
bristles is often coated with parasitic growths.

Posteriorly both dorsal and ventral bristles become more slender and elongate, the
tips of the ventral forming long hair-like processes.

The dorsal cirri are pale brownish and pellucid, the caudal styles being darker.
The cirri are simple tapering processes, with sparsely distributed clavate cilia, which are
longest towards the base of the filiform tip. The ventral cirrus is somewhat tumid at the
base, and has a very few comparatively long clavate cilia.

Reproduction. — The specimen procured in July at Guernsey carried nearly ripe ova.
Habits. — A single specimen occurred in each gallery of Marphysa sanguined, and the
examples from Polperro, " from the chinks of rocks," may have had a similar relationship.
When placed in an open vessel beside Marphysa it clung to the body of the latter near
the head.

Baron de Saint-Joseph procured his Harmothoe picta, 1 an allied species, in the tube
of Lanice conchilega at Dinard, on the French coast. Another species (E. arenicolse) he
found clinging to a lobworm. It is curious that this also was a ripe female. His
figures of the bristles are not so strictly drawn as is necessary for accurate diagnosis, but
the species closely approaches H. spinifera and H. ljungmani. It is, however, larger,
viz. 25 mm. long.

1 ' Ann. d. Sc. nat.' (7), v, p. 172, 1888.

342 HARMOTHOE LUNULATA.

9. Harmothoe lunulata, Belle Chiaje, 1841.

Specific Characters. — The head approaches that of Harmothoe marphysse, only it is
less elongated and the eyes are larger. The tentacles and tentacular cirri are brownish ;
palpi smooth. Body about three quarters of an inch long ; ventrally, with a series of
brown spots, which, as a rule, commence as four rows somewhat behind the middle.
Occasionally by union they form bars at the junction of each segment. A prominent
segmental eminence and papilla. Scales fifteen pairs, smooth round the margin, but
with a dense cluster of minute horny papillae on an area in front of the scar. Brown
pigment variously arranged— some having a ring of brown or a V-shaped pattern on the
scale, while the outline of the pigment in others has the shape of the shell of Pandora.
Dorsal bristles are better developed than in H. marphysse., being long, tapering, slightly
curved, and finely spinous. The ventral bristles, again, form a fan, and the tips of
almost all have a secondary process. The dorsal cirri are comparatively short tapering
organs, with sparsely distributed short clavate cilia. The ventral cirri have similar cilia,
and are slender, their tips reaching beyond the base of the nearest bristles.

Synonyms.
1841. Polynoe lunulata, Delle Chiaje. Descriz. e not., vol. v, p. 57, pi. cxliv, f. 5, 6.
1865. „ maculosa, Carrington. Proceed. Lit. and Phil. Soc, Manchest., iv, p. 177.

1867. Monocolea tessellata, Costa. Ann. d. Mus. Zool. d. v. Univ. d. Xapoli, i, p. 82.

1868. Polynoe lunulata, Claparede. Annel. Chet., Naples, p. 63, pi. ii, fig. 1.

1875 - „ „ Panceri. Atti E. Accad. Napoli, vol. vii, p. 13, Tav., f. 1—3.

„ Harmothoe lunulata, Mcintosh. Invert, and Fish., St. A., p. 116.
1876. „ „ idem. Tr. Z. S., vol. ix, p. 385, pi. lxix, f. 16—20.

1884. Polynoe lunulata, V. Cams. Faun. Medit., i, 200.
1886. Harmothoe lunulata, Harvey Gribson. Verm. Liverp., 148.
1891. Polynoe {Harmothoe) lunulata, Hornell. Op. cit., p. 236,

Habitat. — Yery generally distributed throughout British waters— from Shetland to
the Channel Islands, and from the west coast of Ireland to the east coast of Scotland.
It ranges from the tidal rocks in the Channel Islands to 120 fathoms off the south-west
coast of Ireland. It extends to the Mediterranean.

Head (Plate XXVII, fig. 8) resembles that of H. marphysse, only it is less elongated
and the eyes larger. The median tentacle is brownish, has a filiform tip and sparsely
distributed clavate cilia. The lateral tentacles are subulate, with a filiform tip and
clavate cilia. The tentacular cirri have the same form as the median tentacle, and also
the same brownish colour. The palpi are smooth.

Body about three quarters of an inch in length, slightly tapered anteriorly, and
much more so posteriorly. The chief feature of moment is the presence, in the majority,
of a series of brown spots, which commence as four rows somewhat behind the middle.
In some the sets are united so as to form two rows of bars at the junction of each
segment ; and this confluence sometimes occurs posteriorly even when the four rows are
distinct in front. The segmental eminence is well marked, and a small cylindrical
papilla projects between the feet.

HAPMOTHOE LITXITLATA. 343

Scales (Plate XXXII, fig. 12). — Fifteen pairs, somewhat thin, and entirely covering
the dorsum. The first pair are rounded with a broad transverse bar of brown pigment
in front. The succeeding are reniform, and the posterior more or less ovoid. They are
finely veined, and as smooth under a lens as in H. marphysse, but show a dense group of
horny papillse in front of the scar, that is, near the hollow of the reniform kinds. The
coloration of these scales is varied. Thus some of the Zetlandic examples are faintly
tinged with brown toward the posterior border ; in others each scale has a brownish
ring, or the brown pigment forms a bold border on the inner third, and sends a process,
in some cases enlarged near the termination, toward the centre of the scale, so as to
simulate a V. The spot or enlargement is at the scar. In the forms from St. Andrews
the pigment assumes the shape of the shell of Pandora, with a spot (over the peduncle)
corresponding to the hinge anteriorly. The colours are for the most part retained after
immersion in spirit. The persistent attachment of the scales is not a feature character-
istic of the Zetlandic forms, for they readily fall off.

Feet. — The first foot has the usual spine and a few bristles resembling the dorsal.

The dorsal division of the second foot bears a larger group of bristles than in
H t marphysse, with similar fine rows of spines, and the bristles are generally larger and
stouter. The ventral series are fairly developed, the upper and lower with simple, the
median with distinctly bifid tips, though the secondaiy process is slender.

In the third foot both dorsal and ventral bristles are longer and approach more
nearly to the typical forms. They are also more numerous.

In the typical foot (Plate XXX, fig. 4) the dorsal division is more developed than in
H. marphysse, and the bristles attain a size never seen in that species, though it has to
be borne in mind that the environment may have considerable influence in modifying
these and other parts. They are long, tapering, slightly curved, and finely spinous
bristles (Plate XXXIX, fig. 12, in profile, and Plate XXXIX, fig. 13, antero-posteriorly,
so as to show the alternate disposition of the spines, both representing the longer and
more tapering forms next the ventral). Some, however, are even more finely tapered
than the forms shown. The spines proceed almost to the tip, a mere trace of a terminal
smooth portion being present. Those next the body are much shorter and less tapered
(Plate XXXIX, fig. 14). Parasitic infusoria and slender algaa are common on these
bristles. The ventral division of the foot is somewhat less elongated than in
H. marphysx, and the bristles form a more regular fan. Superiorly the spinous region is
longer, and the tips more finely tapered (Plate XXXIX, fig. 15) ; the secondary process,
which appears to be present in all at this edge, being very closely applied to the larger
division, which is scarcely hooked. In a variety from Guernsey some of these bristles
have simple tips. The fissure between the divisions becomes more evident as the
spinous region diminishes in length, but it again is less distinct ventrally, some at the
extreme verge having a mere trace of this process. A bristle from the middle of the
ventral group is drawn in Plate XXXIX, fig. 16. Their colour throughout is very pale
yellow. Microscopic filamentous algas occur on them.

The dorsal cirri are rather short, tapering organs, with scarcely a trace of an
enlargement below the filiform tip, and the short clavate papillae are sparsely distri-
buted. The ventral cirri are comparatively slender, and the tips reach considerably

45

344 HARMOTHOE LUNULATA.

beyond the bases of the nearest bristles. They have the same sparsely distributed short
cilia.

Habits. — Earmothoe lunulata is a very active species amongst the laminarian roots,
and displays as much irritability as Evarne impar. It is also sometimes found as a
commensalistic form in the tube of Poly cirrus. It is brightly phosphorescent, glowing
when irritated at the bases of the feet for a considerable time, and giving off flashes
when immersed in spirit, as well as generally breaking in pieces.

The Zetlandic examples of the species are somewhat elongated, have longer feet, and
longer and more delicate pale bristles ; moreover the dorsal and ventral cirri are longer
than in the southern forms.

It is evident that the species approaches E. marphysx very closely, though the cirri
of the latter are shorter and smoother, the bristles of the dorsal branch much shorter
and less conspicuous, and those of the ventral division shorter and more slender. More-
over those of the superior ventral series have no bifurcation at the tip, the closest
approach to the latter condition being in a variety of E. lunulata from St. Peter Port,
Guernsey, which had an indistinctly bifid tip in one or two of its superior bristles.

I have united it, after Claparede, with Delle Chiaje's form, though in his original
description of the species he gave it a single caudal style and fourteen pairs of scales-
characters also illustrated in his somewhat stiff figure, which shows the middle line of the
dorsum quite bare from end to end.

Claparede, who studied the species at Naples, correctly described it as having fifteen
pairs of scales, and pointed out the pinkish hue of the ventral median line anteriorly—
from the nerve-cord — on which he found no ganglia. He gives the total number of
segments as thirty-seven, and shows that the last bears two large terminal cirri. At the
bases of the feet are vibratile rosettes as described by Bhlers, and they are about four in
number. In the preserved examples the two whitish masses he mentions on the feet
are not visible. He also notes the apparent absence of blood-vessels, and describes the
nerve-cord as having a median and two lateral bands corresponding to the nerve-cells,
of which he gives some further details. He is of opinion that Costa's P. tessellata is the
same species. He describes the scales as finely granular and without horny papillae, but
in his figure the greater part of the surface (all but the anterior third) shows such
papillas. This, therefore, is a point on which further evidence is required, especially as
the distribution of these horny papilla in the British examples is so well defined.
Should the evidence prove that the Mediterranean form is different, then Dr. Carrington's
name, E. maculosa, stands. Ife first found the species in this country stranded on
Southport sands.

Prof. Panceri l experimented with this species and others in regard to luminosity, and
came to the conclusion that it emanates entirely from the scales, and that it is connected
with the remarkable terminations of the nerves in the subcuticular granular layer of the
epiderm.

Grube's P. maculata 2 seems to be an allied form, but the description of the cirri
differs.

1 ' Atti B, Accad. di Napoli,' 1875, p. 13, Tav. 304.
Actin., Echin., u. Wiirm./ p. 87, 1840.

2 f

HARMOTHOE SETOSISSIMA. 345

10. Haemothoe setosissima, Savigny, 1820. Plate XXV, fig. 4.

Specific Characters. — Head somewhat short and broad. The posterior eyes are
nearer each other than in H. imbricata, while the anterior are larger and more visible
from the dorsum. Median tentacle rather long, with just a trace of a dilatation below
the filiform tip, and its surface has sparsely distributed short clavate cilia. Palpi appa-
rently smooth, but show minute papillas under a high power. Body somewhat elongated ;
segments thirty-eight, either pale or variegated rather prettily with brown. Segmental
eminences and papillge distinct. Scales somewhat adherent, thin, but fairly tough ; first
pair rounded, rest reniform-ovate, their surface densely covered with minute papillae;
colour pale or brownish, sometimes with a broad ring. Feet with a long acute process
above the spine. Dorsal bristles long, gently tapered, with very close rows of short
spines, and the point rather blunt except in the external forms. Ventral slender, with a
long spinous region and a very short, smooth, bifid tip. Dorsal cirri elongated, tapering,
and with sparsely distributed short clavate cilia ; ventral slender and rather elongate,
also with a few short clavate cilia.

Synonyms.
1820. Polynoe setosissima, Savigny. Syst. Annel., p. 25.
1828. Eumoljpe setosissima, Blainville. Diet, de Sc. nat., lvii, p. 459.
1834. Polynoe setosissima, Audouin and Edwards. Annel., 90, pi. i, f. 18.
1836. „ „ De Quatrefages. Beg. an. illust., pi. xix, f. 2.

1863. „ longisetis, Grube. Archiv f. Naturges., xxix, p. 37, taf. iv, f. 1.

1864. „ „ idem. Die Insel Lussin, &c., p. 78.

1865. Lsenilla glabra, Malmgren. Nord. Hafs-Ann., p. 73, Tab. ix, f. 5 ; and Ann. Polychast.,

p. 136.
„ Polynoe setosissima, De Quatrefages. Ann. I, 229, pi. vi, f. 17.
„ Antinoe semisculptus, Baird (partim). Journ. Linn. Soc, viii, p. 192 (?).

1869. Harmothoe Malmgreni, Ray Lankester. Tr. Linn. Soc., xxv, p. 375, pi. li, f. 11, 25, 28.
„ Polynoe longisetis, Mcintosh. Trans. R. S. E., xxv, p. 406, pi. xv, f. 3.

1870. „ laevigata, Claparede. Ann. Nap., Sup., p. 14, pi. i, f. 3 (?).

1875. Lsenilla glabra, Ehlers. 'Porcupine, 5 1869, op. cit., p. 32.

,, „ setosissima, Mcintosh. Invert, and Fishes, St. A., p. 116.

1876. „ „ idem. Trans. Z. S., ix, p. 387.

1882. „ glaberrima, Hansen. Norweg. N. Atlantic Exp., 29, Tab. iii, f. 6 — 11.
1886. „ setosissima, Griard. Bullet. Sc. Nord, 339.

1890. „ „ Malaquin. Ann. Boulon., 23.

1891. Polynoe {Lsenilla) setosissima, Hornell. Op. cit., 235.

1898. Harmothoe longisetis, De St. -Joseph. Ann. d. Sc. nat., viii, ser. v, 234, pi. xiii, f. 21.

Habitat. — Not uncommon between tide-marks, under stones at Herm, and also in
the tubes of Ghdetopterus ; tossed on shore amidst sponges, sea weeds, and corallines at
the west sands, St. Andrews (E. and R. M.) 5 in the stomach of the cod and haddock at the
same place (E. M.), at Plymouth, Moray Frith, and various parts of the British coast.
Ehlers found it ranging in the ' Porcupine ' to 767 fathoms on Holtenia- ground with mud
and globigerina-ooze. Extends to the Mediterranean.

346 HARMOTHOE SETOSISSIMA.

Length one and a half to one and three quarters inches.

Head (Plate XXVIII, fig. 5) somewhat like that of Harmothoe imbricata in outline,
only the anterior peaks are less acute and the widest region is further forward, so as to
give the head a more distinctly ovate form. The posterior pair of eyes are nearer each
other, and the larger anterior pair are more visible from the dorsum. The median tentacle
is of moderate length, with just a trace of a dilatation below the filiform tip, and furnished
with sparsely distributed short clavate cilia. The lateral tentacles are short and tapering,
and have a very few of the same small clavate cilia. The palpi are smooth under a lens, but
show very minute papilla? under a high power. The toughness and extensibility of these
organs are well seen in this species. The tentacular cirri also show a trace of an enlarge-
ment below the filiform tip, and have sparsely distributed short clavate cilia. These and
the tentacles are sometimes tinted brownish.

Body rather elongate, slightly tapered in front and more distinctly from the anterior
third backward, and having thirty-eight bristled segments. In some the dorsum is
pale, but in others, as in the drawing, it is barred with brown throughout, the pattern in
each segment posteriorly being beautifully defined. The under surface is pale. The
segmental prominence is distinct, and a considerable papilla projects between the feet.
The body terminates in two long, tapering, caudal styles.

The pre-gastric cseca agree with those in Harmothoe imbricata.

Scales (Plate XXXIII, fig. 5) fifteen pairs, entirely covering the dorsum; the first
pair rounded, the succeeding reniform-ovate. The anterior cling somewhat firmly to the
pedicles. The surface is densely covered with minute chitinous papilla? (really minute
spines), so that it appears like shagreen under the microscope. Towards the outer
border a few short clavate cilia project from the edge and also appear on the neigh-
bouring surface. The scales are comparatively thin, but of considerable toughness.
They increase in size posteriorly. Their colour is often pale, but in some it is brownish,
and in the coloured figure the papilla? of the scales and other organs are infested by a
blackish parasitic growth which greatly alters the aspect.

In two fine examples procured after a storm on the west sands at St. Andrews
(E. M.), the scales had a broad marginal belt in the exposed portions, leaving a large
ovoid pale region in the middle. The dorsum besides was richly tinted with brown.
Coloured examples, indeed, are more common in the north than in the south.

Many Loxosomse occur on the scales of a specimen from the Moray Frith.

Feet. — The first foot has a very strong spine and a group of about four short
bristles of the dorsal type.

In the second foot the dorsal bristles form a dense tuft, the closely arranged spinous
rows and the rudimentary bare portion at the tip being diagnostic, though the bristles
have not attained the length of the typical forms. The ventral division consists of a
series of slender bristles with long, 'tapering, spinous regions, and simple slender tips.
Moreover the shape of the foot is peculiar, the dorsal division in lateral view forming a
high crest, while the ventral is distinguished by its long, acute cone projecting
horizontally above the spine,

In the third foot the bifid condition of the ventral bristles is more evident, and the
dorsal are longer. The conical process of the ventral division is larger. By easy stages

HABMOTHOE SETOSISSIMA. 347

the typical foot (Plate XXX, fig. 12) is soon reached, showing a ventral division
terminating superiorly in a long conical process above the spine. The dorsal bristles
(Plate XL, fig. 5) spring from an oblique eminence, and are characterised by their great
proportional length, slight curvature, and gently tapered extremities, those next the
body having somewhat blunt tips, those next the ventral being more acute. The tips of
the blunt forms scarcely show a bare portion, the rows of spines being continued to the
tip, but the acute bristles next the ventral have a minute bare portion which sometimes
presents a slight keel. The spines are short and the rows very close, so that the bristle
is at once distinguished from that of Harmothoe imbricata, irrespective of the length of
the tip. The ventral bristles are numerous, long, and slender, the spinous region of the
upper forms being of great length, the rows closely arranged, and the spines rather
short. The tips are bifid, the terminal hook is small, and the secondary process makes
a very slight angle. Moreover the smooth region is remarkably short, the spines
passing up to the fork (Plate XL, fig. 6). Both sets of bristles are of a lustrous pale
yellow, almost like those of Ghloeia, and thus afford a contrast with the dull straw-
coloured organs of Harmothoe imbricata. In shape the ventral lobe of the foot is
peculiarly pointed, the base being bevelled superiorly, and with a projecting fold
inferiorly.

In the terminal feet the dorsal bristles are few in number, more slender and
elongate, the tip pointed, and the rows of spines distinctly wider. The ventral are also
few in number, slender, with long, tapering, spinous regions, and attenuate, simple tips.

The dorsal cirri present scarcely any enlargement below the filiform tip, are pale
throughout, have short clavate cilia sparsely scattered over the surface, and they are
longer distally than proximally. The ventral cirri are long subulate organs, the tip
extending considerably beyond the bases of the bristles.

The dorsal bristles are the seat in some of a peculiar blackish fungoid growth, as
indicated in the coloured figure. This minutely granular structure finds a suitable site on
the spiny ridges of the dorsal bristles, and thus both shaft and tip are barred with
black in a characteristic manner. This coating can readily be removed by pressure, and
no evident change is apparent in the bristle. It is rendered brown by hydrochloric acid,
while caustic potash does not seem to alter it much. This growth also affects the
spinous region of a few of the ventral bristles.

Habits. — This is another example of the fact that species which frequent off-shore
waters in the north are found between tide-marks in the south, as in the Channel
Islands. It is a comparatively hardy form, and the example from which the coloured
drawing was made lived for weeks in the centre of Perthshire, though towards the
termination of the period many of the ventral bristles were shed.

After careful consideration this form has been associated with Savigny's Pohjnoe
setosissima.

Audouin and Milne Edwards (1843) assigned forty segments to this species, and
they and Savigny found their examples at Havre.

De Quatrefages (1865) included it under his fourth group, in which the body is more
or less elongated, and covered by fifteen pairs of scales. He describes the species as
provided with a simple triangular head; the median antenna large, about as long as the

348 HABMOTHOE SETOSISSIMA.

tentacle. The lateral antennas small. Forty segments. Scales whitish, obliquely ovoid;
sometimes spotted, with the entire margin ciliated. He points out that Savigny and M.
Edwards described individuals devoid of scales. Moreover he readily distinguished it
by its pale colour ; he thinks that Cuvier was wrong in saying Savigny found it at
Havre.

Baird found it in tube of Ghaetopterus insignis from Beaumaris.

This species was carefully described by Grube 1 from specimens procured at the
larger and smaller Lussins and at Crivizza. He noticed its relationship to P. setosissima
of Savigny, but the latter had shorter palpi and two additional papillae, viz. twenty, to
the border of the proboscis. He 2 says the specimens so named in the Parisian Museum
were Laenilla glabra, Mgrn., and Eva/rue impar.

Malmgren's artist represents the anterior pair of eyes on the dorsum, and of the
same size as the posterior pair.

The Polynoe laevigata of Claparede, from Naples, approaches this form ; indeed, there
is little to differentiate it. Claparede considered the P. tentaculata of De Quatrefages as
closely allied.

In Prof. Pay Lankester's early remarks on this species the essential points were
omitted.

Ehlers alludes to the variations of the scales exhibited by the specimens from
different depths in the ' Porcupine ' Expedition of 1869.

Dr. Hansen's (1882) P. glaberrima appears to be this species. Unfortunately, while
placing comparatively little weight on the minute structure of the bristles, the
diagnostic characters relied on by this author would not appear to have been of much
avail.

It is difficult to make out to what species Audouin and Milne Edwards' Polynoe
Ixvis is to be referred. Malmgren includes it under his Laenilla alba. Prof. Giard
(1886) considers that Polynoe laevis is not a variety of Polynoe setosissima, since the
scales in the former are smooth, whereas in the latter De Quatrefages distinctly says
they have a margin fimbriated all round, but he corrects himself in a subsequent
communication. 3 He does not think the P. laevigata of Claparede is this species.

Hornell (1891) considers that it is the Laenilla glabra, Mgrn., and found it
invariably present in the tube of Chaetopterus in Herm.

Baron de Saint- Joseph 4 (1898) in a recent paper doubts if Prof. Giard and I are
warranted in connecting this species with Savigny's form, which had lost its scales.
The description given by MM. Audouin and Edwards, however, though imperfect, comes
nearest this species. He also is doubtful about the inclusion of Claparede's P. laevigata
as a synonym, but there is no valid reason for altering the view already expressed.

1 " Beschreibung neuer oder wenig bekannter Anneliden/' ' Arch. f. Naturges./ 1863, p. 37.
3 < Arch. f. Naturges./ 1870, p. 287.

3 < Bullet. Sc. Nord/ pp. 340, 341.

4 Op. cit., 1898, p. 236.

HARMOTHOE AREOLATA. 349

11. Harmothoe areolata, Grube, 1860.

Sjoecific Characters.— Body elongate, slightly tapered anteriorly, more so posteriorly,
and terminating in two caudal cirri. Bristled segments 37 to 39. Head somewhat
ovoid, with two elongated peaks in front, and clearly separated by an interval from the
base of the median tentacle. Posterior pair of eyes on the dorsum and in front of the
nuchal fold, while the anterior pair under the peaks are almost invisible from the
dorsum. The median tentacle is of moderate length, gently tapered, little if at all
dilated below the filiform tip, and densely ciliated. The lateral tentacles are short, and
similar in structure. The tentacular cirri agree with the tentacle. Palpi of moderate
length, gently tapered from base to near the apex, which is short and filiform. They
have rows of short clavate papilla?, which diminish in size from base to tip, which is
bare. Scales fifteen pairs, imbricate, covering the dorsum and extending over the
bristles. They vary in shape from the rounded first pair to the reniform kinds, and the
D-shaped posterior pair are studded with horny papilla? of various sizes, from the small
rounded processes to the elongate forms which rise into bluntly conical spines pos-
teriorly. Margin ciliated posteriorly from the inner to the outer border. Those on
the outer border are densely grouped and long. Dorsal cirri of moderate length,
tapered and densely ciliated almost to the tip. In many the distal half of the process is
dilated, so that the organ is broadly spathulate, with the filiform tip projecting at the
distal region. Feet have a dorsal tuft of curved bristles which are densely spinous,
with a smooth tip, which has a streak as if a rudimentary tooth were present. The
ventral bristles are rather slender, with tapering bifid tips and well-marked spinous
rows. The ventral cirrus reaches beyond the base of the bristles, and has somewhat
short clavate papillae.

Synonyms.

1857. Polynoe areolata, 0. Gr. Costa. Annel. Nap., Tav. if, figs. 2 — 2j.

1860. „ „ Grube. Arch. f. Naturges., 1860, 72, pi. iii, f. 2.

1861. „ „ idem. Ein Ausflug n. Triest u. Quarnero., p. 139, Taf. iii, fig. 2.

1864. „ „ idem. Die Insel Lussin, p. 78.

1865. „ „ De Quatrefages. Ann., i, 232.

1866. Antinoe nobilis, Eay Lankester. Trans. Linn. Soc., xxv, p. 375, Tab. li, f. 1 — 9.
1868. Polynoe areolata, Claparede. Ann. Chet. Nap., 71, pi. ii, f. 5.

1876. Earmotlioe areolata, Mcintosh. Trans. Z. S., ix, 381, pi. lxviii, f. 3, 12, 14.

1884. Polynoe areolata, V. Carus. Fauna Medit., i, 200.

1886. Evarne areolata, Griard. Bullet. Sc. Nord, 341.

1888. Harmothoe areolata, De St.-Joseph. Ann. d. Sc. nat. (7), v, p. 169, pi. vii, f. 41 — 43.

1890. „ „ Malaquin. Ann. Bonlon., 18, pi. i, f. 1, 2.

Habitat. — First found by O. G. Costa in the Mediterranean, afterwards by Prof.
E. Grube in the same sea and the Adriatic, and in Britain by Prof. Ray Lankester close
to and in the tubes of Terebella nebidosa in Herm, by Mr. Cooper in the tubes of
Chsetopterus, and by the author frequently under stones at Herm.

350 HARMOTHOE AREOLATA.

The head (Plate XXVII, fig. 15) of this species is distinguished by its somewhat
ovoid outline and the prominent peaks in front. The eyes are of moderate size — two on
the dorsum posteriorly in front of the nuchal fold and quite within the margin of the
head, and two at the outer and under surface of the peaks, so that they look forward
and slightly outward. The median tentacle is of moderate length, and tapers from the
base towards the tip, which is filiform and densely covered with rather long cilia having
slightly bulbous tips. Scarcely a perceptible swelling in the preparations occurs below
the tip. The cilia diminish in length distally, but extend along part of the filiform tip.
The lateral tentacles arise below and somewhat external to the median, are short, and
taper from the base to the filiform extremity. Short cilia occur on the surface and on
the filiform extremity as small papillae.

The tentacular cirri are similar to the median tentacle. The palpi are of moderate
length, gently tapered from the base to near the apex, which is filiform, though short.
They have densely crowded rows of clavate papillae — more slender than in Harmothoe
imbricata, and which gradually diminish from below upwards.

The body presents the usual slight convexity dorsally with the transverse depression
in each segment — especially marked behind the proboscidian region. Moreover much
of the body is of uniform breadth. Ventrally a central depressed area occurs in spirit,
with lateral elevations, from which the rather small segmental processes project between
the feet. Posteriorly it terminates in the caudal cirri.

Proboscis. — The proboscis is less massively muscular than in Lagisca or Harmothoe
imbricata, and the horny teeth are smaller; indeed, they may be called minute. The
horny ridges are also less prominent, The terminal papillae of the organ agree with
those in Harmothoe.

In the peripharyngeal space only one short pregastric caecum goes forward to the
second segment in front of the stomach. The second is nearly transverse. The
reduction of the caeca is thus a marked feature.

The scales (Plate XXXII, fig. 17) are somewhat stiff and leathery, especially in
front. The first pair are more or less rounded, and thickened by the numerous large
chitinous areas terminating in the blunt spines, which are minutely nodular with short
chitinous spikes. The surface of the scale is further cut into definite areas by a series of
chitinous reticulations which enclose the space occupied by each bluntly conical spine,
and consequently in this (the first scale) irregular hexagons, pentagons, or similar
figures occur in the central region round the large bosses, while the margin is cut into
longer areas with intermediate small triangles — for spines — at the edge. Moreover the
right or upper differs from the left or under scale, the former corresponding more
particularly with the description, while the latter in the region overlapped by the former
is thinner, and has numerous smaller reticulations enclosing a diminishing series of horny
papillae from the central region of larger spines to the edge. The latter is also smooth,
but, with this exception, the edge has cilia all round, and of considerable length as well
as more numerous at the outer margin. The right or upper first scale shows a
continuous series of marginal cilia. The succeeding scales are more or less reniform or
somewhat ovate, according to their position, while the last are almost D-shaped. Their
general structure is similar, viz. the anterior and inner edge, that overlapped, being more

IIAPMOTHOE AREOLATA. 351

or less thin and smooth ; then the small papillae stud the surface, and gradually increase
in size towards the posterior and outer edges. The border is formed of the long ridges
terminating in the marginal, blunt, spinous papilla. The smaller rounded bases of the
chitinous papillae internally are so closely arranged that they resemble the shields in
certain regions of an armadillo. Moreover, whereas in the first scales the chitinous lines
which bifurcate to form the triangular areas of the margin are simple, in the rest they
form a complex and continuous series of reticulations somewhat resembling those in the
skins of holothurians (Plate XXXIII, fig. 3). The minutely nodular or caespitose
condition of the chitinous spines is a marked feature throughout. The posterior
border has cilia of considerable length internally, then they diminish along the
posterior edge, again elongating on approaching the outer border, where they form a
long, dense fringe. The bulbous tip is minutely granular in its centre, and is continuous
with the central axis of the cilium. The under surface of the scale is iridescent, with a
well-marked scar for attachment somewhat nearer the outer than the inner border. The
scales separate readily. Occasionally a small patch of Gellejpora is found on a scale.

Colour. — The dorsum has a light brownish or amber hue from the scales, the
posterior part of each of the latter being somewhat darker. The pinkish hue of the
body occasionally shines through these. The cirri are pale. The under surface is
iridescent bluish-pink, as in Polynoe scolopendrina. Ray Lankester speaks of the head as
being violet, and the palps deep madder-brown. In spirit they are dark greyish, streaked
with the lines of cilia. Towards the posterior end the dorsum has various transverse
streaks of dark greyish (in spirit). In rather more than the posterior third also a dark
touch occurs on each side of the base of the dorsal cirri, and there is a tendency for the
elevations in a line with the tubercles for the scales to have a touch of the same colour.
The pigment thus seems to be connected with the cirrigerous feet, and consequently is
best developed behind the last scale-tubercle. On the ventral surface the posterior
third presents dark pigment touches on the segmental eminences, and the tint increases
in depth posteriorly, spreading outward on the bases of the feet. A touch or two of
pigment also occur on the anterior folds of the mouth.

Feet. — The modified dorsal bristles of the first foot have smooth tips. The second
foot is small, with a flipper- or paddle-shaped fleshy region dorsally, the inferior margin
terminating in the long papilla for the spine. The bristles, though short, are typical in
structure. The ventral division is somewhat conical, terminating in the pointed region
for the spine, and from the tip of this process a long cutaneous papilla projects. The
bristles are slender with elongated curved spinous tips.

The third foot shows a considerable increase in the size of both dorsal and ventral
bristles. The ventral are still proportionally slender, but the majority have bifid tips,
only the lower short forms having simple tips. The ventral cirrus has large clavate cilia.
The papilla beyond the spine is present.

The fourth foot approaches the typical condition except that the number of simple
bristles inferiorly is greater.

In the typical foot (Plate XXX, fig. 3) the superior division has a somewhat con-
spicuous tuft of elongate, slightly curved bristles externally, and shorter and more curved
forms internally (Plate XXXIX, fig. 17). The spinous rows are dense. The tip has a

46

352 HARMOTHOE AKEOLATA.

short smooth portion with, in many, a slight though distinct streak, best marked in the
shorter bristles, in which it forms a groove at the extremity. The external bristles (those
next the ventral division) form a contrast to the inner from their sharp tapering tips and
slight curvature. In the older specimens these bristles are often densely coated with
parasitic growths, such as algaB and infusorians, besides mud. The ventral branch has a
series of rather elongate slender bristles with tapering tips, which, as usual, diminish in
length from above downward. The tips superiorly are attenuate, and one or two show
no secondary process, but this soon appears, again to disappear, in the shorter inferior
forms. One of the attenuate superior bristles with a bifid tip is shown in Plate XXXIX,
fig. 18. The spinous rows are rather prominent. A shorter bristle again, with a slightly
abraded tip, is given in Plate XXXIX, fig. 19. Inferiorly the bristles have slender,
short, spinous extremities with a minute secondary process, and, at the ventral edge of
the series, the tips are simple. The nearly cylindrical papilla above the ventral spine is
well marked; and the ventral cirrus reaches beyond the base of the bristles, and has
somewhat slender clavate papillse.

Comparatively little change ensues in the bristles of the terminal feet. The dorsal
are more slender and proportionally longer, so that they extend almost to the tip of the
longest of the ventral division. The ventral division has the same type of bristles as
in front, except that they are more slender. The elongated forms usually found in this
region were not observed, but the specimens may have been recently injured. The
papilla above the ventral spine continues to the last foot.

The dorsal cirri are of two types, viz. those of the usual kind, with just a trace of a
dilatation below the elongated filiform tip, and densely coated with cilia, which are long
except at either end of the series ; and secondly, those with a greatly enlarged distal half,
so that the organ resembles a tennis-racket with the filiform tip appended to it, and
coated with cilia as in the first form. This condition of the cirri was first noticed by
Ray Lankester in specimens from Herm, for Grube's original ones had only the slender
cirri. No connection between the sexual or other condition of the specimens and this
state of the cirri has yet been observed. In one instance these enlarged cirri were found
in the tubes of Chastopterus, and were forwarded as parasites.

Reproduction.— A specimen of good size from Herm carried ova at the end of July
and beginning of August.

Habits. — Ray Lankester, who placed it as a new species under the genus Antinoe,
thought it fed on Terebella nebulosa, and I found some procured in Herm had fed on
Eunice and other forms. It would not always seem to be an inhabitant of tubes of other
annelids, but occurs in a free condition under tidal stones. De Quatrefages gave this
form a position near Harmothoe imbricata, but added nothing to Grube's remarks.

G-iard (1886) includes it under the genus Evarne on account of the form of the head,
and the presence of chitinous protuberances on, as well as on account of the structure of,
the elytra. Yet the proboscis, the margin of the elytra, and the cirri present certain
characters which are almost sufficient to give generic distinction.

De Saint Joseph (1888) seems to doubt my diagnosis in regard to Ray Lankester's
Antinoe nobilis, but it rests on a careful examination of specimens kindly sent me by
Lankester himself, and a survey of the same collecting grounds on the Channel Islands.
There is no doubt on the subject.

EVAPNE IMPAP. 353

Genus XIV. — Evarne, Malmgren, 1865.

Body rather small, ovate-oblong, attenuate posteriorly. Lateral tentacles fixed
under the median, head deeply incised in front, and with prominent lateral peaks.
Eyes four, generally large, the anterior pair on the projecting lateral region;
the posterior in front of the nuchal collar and widely separated. Palpi densely
covered with minute papillae. Scales, fifteen pairs, spinulose, ciliated or smooth
at the edge. Dorsal bristles shorter than the ventral, and with wide rows of spines ;
ventral bristles very long, with minutely bifid tips. Segmental papillae well marked,
but not long.

1. Evarne impar, Johnston, 1839. Plate XXYI, fig. 5.

Specific Characters. — Head wider than long, with prominent peaks anteriorly. Eyes
large ; posterior pair in front of nuchal collar, widely apart ; anterior pair on projecting
lateral region. Median tentacle strong and moderately long, densely ciliated, madder-
brown, with a slightly dilated whitish region, marked off by bars, below the filiform tip.
Lateral tentacles inferior, subulate and tapering, with short clavate cilia. Tentacular
cirri similar to, but more slender than, the median tentacle. Palpi of average length,
with tapered extremities, and densely covered with minute clavate papillae. Body some-
what thin and flattened, broadest at the anterior third, thence tapering to the tail.
Segments thirty-eight to forty, greenish-brown on the dorsum, darkest in front, the pigment
forming somewhat regular bars and touches. Arch of pigment in the cirrigerous seg-
ments terminates laterally in two small touches of dark olive, besides other touches at
base of cirri. Ventral surface pale or brownish. Segmental eminence and papilla well
developed, commencing on the sixth bristled segment. Two long caudal styles. Scales,
fifteen pairs, mottled with brownish pigment, often with a yellowish speck in the centre.
First pair subcircular, rest renif orm and then ovoid ; external margin densely ciliated ;
surface mostly covered with small horny papillae, with larger ones towards the outer
and posterior borders, while along the latter are a few large pyriform or globular papillas.
Dorsal bristles translucent, rather acutely pointed, with wide rows of spines which cross
the shaft at right angles; ventral bristles with elongate tapering spinous regions and
minutely bifid tips ; the latter diminish in length inferiorly, and some at the ventral edge
have simple tips. Dorsal cirri like tentacular cirri; ventral cirri subulate, the slender
tips reaching the ventral bristles, and with numerous short clavate cilia.

Synonyms.

1839. Polynoe impar, Johnston. Ann. Nat. Hist., ii, 436, pi. xxii, f. 3 — 9.

1840. Lepidonote impar, Oersted. Annul. Dan. Consp., 13.
„ „ „ Grube. Fam. Annel., 36.

1873.

1875.
1876.
1879.
1886.

354 EVARNE IMPAR.

1863. Polynoe impar, Claparede. Anat. u. Entwickel., 60.

1864. „ „ Kolliker. Kurz. Bericht., p. 15, pi. vi, f. 4—6.

1865. Lepidonotus impar, Johnst. Cat. B. M., 112, pi. viii, f. 3 — 9.
„ Evame impar, Malmgren. Nord. Hafs-Ann., 71, Tab. 9, f. 7.

„ De Quatrefages. Anneles, 226.
„ Sars. Bid. Christ. Fauna, p. 4.
„ Mcintosh. Invert, and Fishes, St. A., p. 116.

idem. Trans. Zool. Soc., ix, 386, pi. lxx, f . 1—3 ; ibid., p. 398,
„ Tauber. Ann. Danic., 81.-
„ Marenzeller. Porif., &c, Jan Meyen, p. 11.
„ G-iard. Bullet. Sc. Nord, i, 16, with figs.
1888. Harmothoe impar, De St. -Joseph. Ann. d. sc. nat. (7), v, p. 162.

1890. „ „ Malaquin. Ann. Boulon., 18.

1891. Polynoe (Evame) impar, Hornell. Op. cit., 232, pi. xiii, f. 3 and 6.

1896. Harmothoe impar, var. Pagenstecheri, Michaelsen. Polych. Fauna, p. 7, pi. i, f . 1.

1898. „ „ „ „ De St.-Joseph. Op. cit., 1898, 231, pi. xiii, f. 14—20.

Habitat. — Everywhere distributed round the British shores — from Shetland to the
Channel Islands, and ranging into the Atlantic to the depth of 690 fathoms, as well as
extending (fide Verrill) to the shores of America, from Cape Cod to the St. Lawrence.
It occurs not only in purely salt water, but in such lochs as Loch Portan, Lochmaddy,
which receive a stream of fresh water, small examples are likewise found.

Length about an inch. The finest examples in my collection are from the west
coast of Ireland and the Irish Channel.

Head (Plate XXVII, fig. 13) somewhat wider than long, with the usual median
groove, which widens out anteriorly to join the prominent lateral peaks. The eyes are
large and visible from the dorsum, the posterior pair widely apart, and situated imme-
diately in front of the nuchal collar; the anterior pair, which are scarcely larger than the
posterior, being on the projecting lateral region, and thus little removed from the former,
while they are separated by a considerable interval from the peaks. In an example from
Whalsay, Shetland, the anterior pair of eyes were nearer the posterior than usual. In
those from deep water the eyes are somewhat larger, but forms between tide-marks
show considerable differences in this respect, some having large, others small eyes.
Again, a young specimen 6 or 7 mm. long, procured on a thick mass of Flustra off Fermain
Bay, Guernsey, had no eyes. The median tentacle is of considerable length and strength,
generally of a madder-brown colour, and with a pale dilatation marked off by bars below
the filiform tip. It is rather densely covered with clavate cilia, some of which equal in
length the diameter of the process. The lateral tentacles are inferior, subulate and
tapering, and have numerous short clavate cilia. The tentacular cirri are similar to,
though smaller than, the median tentacle. Palpi of average length — with a tapered
extremity, and densely covered with minute clavate papillse, which occur from the base to
the commencement of the filiform tip. These papillaa are proportionally large.

Body moderately elongate, somewhat thin and flattened, the broadest part being
about the anterior third, and thence tapering to the tail. The hue of the dorsum is
greenish-brown, darkest in front, and forming somewhat regular bars and touches along
the back. Some, again, have the dorsum very prettily and symmetrically barred with

EVAHNE IMPAR. 355

brown after the condition in Lagisca. In the cirrigerous segments the arch or band
of pigment in the middle of the body terminates in two small touches of dark olive-
green, and other touches occur at the base of the cirri. Nine or ten of the terminal seg-
ments show these specks on every foot. The proboscidian region is often deeply
pigmented. The ventral surface is pale and iridescent, only a tinge of brown occurring
on the ridge in front of the mouth. A variety from Bressay Sound, Shetland, has the
ventral surface dull olive throughout, with lighter bars, while the dorsum is much
variegated with brownish-olive and dark touches. Rarely the anterior half of the mouth is
dark olive, and a double row of dark touches occurs at each side of the median depres-
sion, whi]e the dorsum is pale madder-brown. In some large examples the elevated ridges
posteriorly show pale olive pigment, which in each segment presents two transverse bars and
a pale centre. The median longitudinal region is pale. In others similar pigment extends
forward to the proboscidian region. The segmental papilla is evident on the sixth foot,
and continues nearly to the posterior end. In size it is comparatively large, though it is
not long, and points between the feet. The body terminates posteriorly in two long
caudal styles, which are much larger than the adjoining dorsal cirri.

Proboscis. — The extruded proboscis shows a range of nine papillaB of the usual
dactylozoid shape dorsally and ventrally. No pigment occurs in the centre of these
organs. The pregastric and other cseca follow the arrangement in HarmotJioe.

Scales (Plate XXXII, fig. 18) fifteen pairs, in the smaller thirteen to fourteen pairs 5
somewhat rough in aspect, mottled with brownish pigment and often with a yellowish
speck in the centre, best marked posteriorly. The first pair are suborbicular, the
succeeding reniform, and the posterior more or less ovoid, though angles are occasionally
formed. The external margin is densely ciliated, the cilia having bulbous tips and
varying in length. Moreover many occur on the neighbouring surface of the scale, and a
few are scattered within the posterior margin. With the exception of the inner and
anterior area the entire surface is covered with small horny papilke, which often increase
into small spines widely scattered towards the outer and posterior borders, while along
the latter are the large pyriform or globular papillas so characteristic of this species.
Some of these occur even on the first scales. The summit of each is occasionally roughly
papillose, or with a series of large conical spines. These large papilla are sometimes
absent, — as, for instance, in those from the tidal region at Lochmaddy, in an example
from 690 fathoms (Station 1, ' Porcupine, 5 1870), and in others from Station 6 and outside
Gibraltar, while the cilia along the border are longer. Young specimens, as a rule,
have smaller processes along the posterior border. Malmgren's figure shows the low
rounded bosses very well. The scales are somewhat thin, and the under surface is
smooth and glistening. The cilia on the scales are frequently overgrown with a granular
parasitic structure.

In a large specimen dredged by Dr. Gwyn Jeffreys off Valencia the scales were
devoid of tubercles. Considerable variation, indeed, exists, for some of those from Herm
have shorter cilia with more distinctly globular heads, a feature accompanied by smaller
dorsal bristles.

Feet. — The first has a few comparatively short dorsal bristles, slightly curved, and
with closer and finer serrations than the typical form. A very short conical portion at
the tip is smooth.

356 EVARNE IMPAR.

The second foot has a dense tuft of dorsal bristles which, though shorter than the
typical forms, already show the chief characters in regard to the rows of spines. They
have a more distinct curvature and less tapered tips than the succeeding forms. One or
two at the inner side (next the body) have the finer rows and aspect of those at the base
of the tentacular cirri. The ventral bristles have elongated spinous regions, the upper
and middle with slender bifid tips, the lower with simple tips.

The foregoing, therefore, requires little change to assume the typical form (Plate
XXX, fig. 7), which has dorsally a fan-like group of translucent bristles, springing from
a prominent process, with rather acutely pointed tips, the bare portion being short. The
smooth tip is formed like a knife, the dorsal edge being bevelled to the ventral (next
the spines), and this shows in some a differentiation as in the figure. In lateral
views the somewhat wide spinous rows pass nearly at right angles across the
long axis and stand out prominently at the edge (Plate XXXIX, fig. 20), a condition
probably due to the nearly opposite condition of the rows. The aspect is thus different
from the more powerful bristles of Rarmothoe antilopis with the curved and oblique
spinous rows, which project very little at the edge. Such groups of bristles (dorsal) form
a gradational series under pressure, those next the body being proportionally thicker
and shorter — with more closely arranged spinous rows, the series rising to the much
longer and more tapered central, and then declining to the more slender forms adjoining
the ventral. Some of the latter are very slender — with finely tapered tips. Occasionally
these bristles are more curved and less tapered than usual.

The ventral bristles are moderately elongate, and superiorly have a long, slender,
tapering spinous region — the rows appearing as oblique bars — and a minutely bifid tip
(Plate XXXIX, fig. 21). The spinous region becomes shorter and stouter, and the rows
of spines more delicate, while the tip is more distinctly hooked, and the secondary
process, which passes nearly straight out or with a very slight angle, is more evident
(Plate XXXIX, fig. 22, representing one of the lower median). Inferiorly the bristles
diminish in length, have a well-marked hook at the tip, and delicate oblique lines from
the fine spinous rows. The bristles throughout are pale, with a faint straw-colour. The
shape of the fleshy part of the foot is characterised by the length of both dorsal and
ventral processes for the spines.

Posteriorly the bristles become shorter and more slender, the dorsal being propor-
tionally long and nearly straight, with well-marked rows of spines which are nearly
opposite, so as to differentiate these bristles from others. The ventral are also very slender,
and the tips in the terminal feet are so attenuate that it is difficult to make out the bifid
condition. Thecate infusoria are common on the dorsal bristles.

The dorsal cirri are finely tapered, and present a slight enlargement towards the
filiform tip in life, whilst in spirit-preparations this is very evident, and more opaque than
the rest; and they are densely covered with long cilia with bulbous extremities, visible
under a lens. The ventral cirri are subulate, and also have a filiform tip which reaches
the bases of the ventral bristles. They have numerous short clavate cilia on the
surface. •

A curious appearance occurs in both dorsal and ventral bristles of a specimen pro-
cured by Canon Norman in 1879 in Norway (Lervig Bay, three to twenty-five fathoms),

EVARNE IMPAR. 357

and is probably due to imperfect or peculiar preservation. Crystalline masses are present
in the centre of the bristles, and in the case of the ventral distend in some the bases of
the spinous region.

Reproduction. — Specimens procured in Norway in summer (Canon Norman states)
bear large ova. Small examples (about 6 mm.) from Loch Portan, Lochmaddy, had
advanced ova in August. A little larger example had advanced ova in Whalsay on
July 17th. A ripe male again occurred at ninety fathoms off North Uist in July.

No ripe forms have been seen at St. Andrews, though numerous small ova are present
in the middle of May.

Habits. — The haunts of this species are similar to those of other Polynoidae. It
occurs under stones between tide- marks, in the crevices of tangle-roots, and in the
cavities of shells — both bivalve and univalve. It was also included in the annelids pro-
cured from holes in the telegraph cables in 1876, and which were thought by Dr.
Carpenter to be borers in the gutta-percha, It is of course free from suspicion on this
head.

It presents a rougher aspect than Harmothoe imbricata, from which it likewise
differs in colour, and it is much more tapered posteriorly. It is also more lively, being
active and irritable, as well as frequently breaking in pieces if molested. The movements
are more graceful than usual in the group, and the long caudal styles are generally
carried nearly in contact. It is likewise brilliantly phosphorescent. Like others of the
family, it bears confinement well, and can be transmitted inland and kept for months in a
small vessel of sea water.

In this, as in other forms, palpi and other appendages as well as the posterior part
of the body are readily reproduced.

Parasites. — Levinsen mentions Herphyllobius crassirostris, Sars, as a Crustacean
parasite fixed to the body of Evame impar in Norway.

Dr. Johnston's Lepidonotus pellucidus may be a young example of Evame.

Claparede (1863) drew special attention to the palpocils on the cilia of the cirri, and
showed how general these organs were in the annelids.

De Quatrefages (1865) followed Johnston in assigning only thirteen pairs of scales
to this species. Prof. Giard (1886) does not think that the P. articulata of Claparede
is identical with this form, as I did in the ' Zoological Transactions.' He also says
P. spinifera, Ehlers, is an allied but not identical form, since the position of the eyes is
different, and the elytra are not fringed as in E. impar. Remarks on these forms are
made elsewhere.

Hornell (1891) records examples from Southport of 33 mm. in length, and thirty-
seven segments, whereas Malmgren gives only thirty- five and De St. -Joseph thirty-
eight.

Varieties. — A variety from 358 fathoms in the 'Porcupine' of 1870 shows some
shorter ventral bristles, the dorsal being pointed. It is a female.

In young specimens procured in the ' Triton ' in 1883 by Dr. Gwyn Jeffreys only
the intermediate horny papillae occurred on the scales along with the cilia. The eyes
were invisible in the preparations.

Dr. Michaelsen describes a variety from Heligoland, Harmothoe impar, var.
Pagenstecheri, of which he gives a special description. By his courtesy a careful

358 EVARNE JOHNSTONI.

examination of the microscopic preparations has been made. The form agrees with the
typical British examples. Malmgren's artist had omitted the cilia on the ventral cirrus,
while in the description they are not mentioned. Baron de St. -Joseph, in a recent
publication, 1 alludes to the same form, and appears to agree with the original author in
thinking it necessary to make a variety.

Another well-marked variety was procured between tide-marks, Herm, under a stone
in August, 1868, and also at Lochmaddy, North TJist. While in regard to the general
outline, the shape of the head, the position of the eyes, and the arrangement of the
pigment on the dorsum, it agrees with the ordinary form, the dorsal bristles are shorter,
though they do not deviate from the typical structure ; and the same may be said of the
ventral. .The scales, however, vary, since the cilia on the outer border are shorter than
usual, and have large ovate heads containing granular epidermic elements ; some of these
also occur within the posterior border, while along the whole of the latter is placed a
short series of cilia with large globular heads and short stalks, the terminal ones only
towards the inner border becoming ovoid instead of globular. The general surface of
the scale is studded with larger tubercles ; and few of the others, except towards the inner
region, and one or two of those near the posterior border, assumed considerable pro-
portions, while none were so large as usual. A similar variety in regard to scales was
procured by Dr. Grwyn Jeffreys in 1868 in ninety fathoms ofE North Unst, Shetland, but
the ventral bristles were more slender, both somewhat approaching those of E. Johnstoni,
though the dorsal were more tapered towards the tip. The specimen was a male.

2. Evarne Johnstoni, Mcintosh, 1876.

Specific Characters. — Distinguished from E. im/par by the deep brownish hue of the
dorsum, and the brownish-purple proboscis ; eyes more minute, the anterior pair not
visible from the dorsum. Scales with fewer and larger horny papillae, longer, fewer,
and more delicate cilia along the posterior and outer borders. Dorsal bristles more
slender and less tapered distally ; ventral more slender, and with longer bifid tips.

Synonym.
1876. Evarne Johnstoni, Mcintosh. Trans. Z. S., ix, p. 398, pi. vii, f. 13—18.

Habitat, — Dredged at Station 3, ' Porcupine, 5 1870, in 690 fathoms in the Atlantic ;
Station 115, West Ireland, August 20th, 1890 (Mus. E. Coll. Science, Dub.). Ranges to
Norway. 2

Length about 9 mm.

1 ' Ann. d. sc. nat./ viii ser., v, p. 231, 1898.

2 Canon Norman, 1879, Stat. 30—34, 41 and 44.

EYAENE JOHNSTONI. 359

The head (Plate XXVII, fig. 7, from a large example, and fig. 12, small example),
has the same form as in E. impar, and is pale throughout in the preparations, the absence
of pigment at the base of the tentacle being noteworthy. The eyes appear only as
minute black points ; two lie at the posterior border of the head, almost hidden by the
collar; two laterally in front of these, as in the ordinary species. A variety dredged
by the 4 Porcupine ' in 1870 at 690 fathoms has very large eyes (Plate XXVII, fig. 7),
the anterior pair having a lens-like corneal thickening. Adult Norwegian examples
show still further increase in these organs.

Body apparently similar in shape to U. impar, viz. abruptly diminished in front and
gently narrowed from behind the anterior third. The dorsum has a deep brownish hue,
with a tinge of purple in front — from the proboscis. Behind the latter the pigment is cut
into bars by the pale belt at the junction of each segment. The ventral surface is pale,
the margins of the oral aperture alone being deeply tinged with brown. The segmental
eminence is distinct, but the papilla is minute.

In the examples the proboscis is more or less extruded, and the usual number of
conical papilla (nine) fringe each lip.

Scales. — A single reniform scale occurred in the vessel, and from what is observed
in the examples from Norway it would seem to belong to the specimen. The surface,
with the exception of the inner fifth, has rather large conical horny papillae, often with
blunt spinous tips, while the adjoining external and part of the posterior border have a
few long and very slender cilia, with a somewhat fusiform tip. The contrast, therefore,
with the more densely and minutely spinous scale of U. impar is marked ; the cilia,
moreover, on the outer edge of the scale of the latter are more numerous, larger and
longer, and globose at the tip.

Feet. — As in E. impar, the second foot presents shorter dorsal bristles with less
tapered tips, but otherwise they are of similar structure to the succeeding. The ventral
bristles of this foot do not project more than the dorsal, and hence are short as well as
slender, the long attenuated spinous region ending in a hair-like tip.

In the typical foot (Plate XXX, fig. 6) the dorsal division bears longer and more
slender bristles than in JE. impar, the slight tapering towards the tip being a noteworthy
feature, and the rows of spines are even more distinctly marked. The smooth terminal
portion is decidedly shorter than in the latter, and in some cases it presents a slight mucro
at the extremity, then a shallow notch, and another elevation a little above the first row
of spines. A long clear shaft projects beyond the foot before the rows of spikes appear,
so that the bristles are comparatively long. One of the stronger bristles is represented
in Plate XXXIX, fig. 23, while the tendency to differentiation of the tip is observed in
fig. 24, Plate XXXIX. The superior ventral bristles have tips so attenuate that it is
difficult to make out their structure; but the bifid condition is present, with the
exception perhaps of the first. The next series have much longer and stronger distal
regions, with extremely delicate and translucent bifid tips ; the terminal hook is short
and very slightly curved, and the secondary process is rather short and broad, and passes
far up, while the rows of spines are distant and well marked. The tips of the succeed-
ing (lower) bristles become broader and shorter, but the character of the termination
remains the same. Toward the inferior edge the tip is simple, only a faintly developed

47

360 EVARNE HUBREOHTI.

hook being present. All are very translucent and delicate. One of the elongated forms
near the dorsal edge of the fascicle is shown in Plate XXXIX, fig. 25, and a more highly
magnified tip in Fig. 27, the arrangement of the spines and the short bifid tip being
characteristic. The bristles from the middle of the foot are exceedingly translucent and
very faintly serrated (Plate XXXIX, fig. 26, and a tip more highly magnified in Plate
XXXIX, fig. 28). In large examples from Norway the dorsal bristles show more
acutely-tapered tips, but otherwise the characters closely approximate.

The dorsal cirri have a filiform tip without enlargement, and rather long clavate
papilla3 sparsely distributed. The ventral cirrus is slender and elongate with a finely
tapered tip, which in the preparations ends in a slight enlargement. Short clavate
papillas — sparsely distributed — occur on the surface. The tip reaches considerably
beyond the base of the lowest bristles.

The species is broadly distinguished from E. impar by the deep brownish hue of the
dorsum and the brownish-purple proboscis, by the structure of the scales, which have
only large horny papillse and few and slender cilia, by the longer, more delicate, and less
tapered dorsal bristles with shorter tips, and by the more slender ventral bristles with
much more finely tapered tips.

The capture of a variety during the ' Porcupine' Expedition of 1870 at 690 fathoms,
in which the eyes are large — especially the anterior pair, which have a lens-like corneal
thickening — and still more the presence in Norwegian waters of a form closely resembling
Evarne Johnstoni in colour, but differing in having very large eyes and in other minute
particulars, raise the question as to how far such species vary with age, sexual con-
ditions, and environment. The younger Norwegian forms agree with the description of
E. Johnstoni, except that some have larger eyes (without a lens-like thickening). In the
larger and older forms, however, the eyes still further increase in size, the lens-like corneal
thickening of the anterior pair leaving only a rim of black pigment round the edge. The
posterior pair are also provided with the central lens-like thickening. The dorsal bristles
are broader, more acutely pointed, as well as somewhat more closely spinous ; and the
smooth portion at the tip is longer, so that it is dagger-shaped. In the ventral bristles
the tips are bifid even to the ventral edge of the foot. Age, therefore, considerably
alters the foregoing organs. The scales differ from those of E. impar, and agree with
those of E. Johnstoni.

The prominent question therefore is, can Evarne impar pass insensibly — by epigamy,
for instance — into E. Johnstoni, as in many other groups of annelids ? So far as present
observations go, a negative reply would seem to be most in harmony with the structure
of the scales, the eyes, bristles, and the geographical range. Further knowledge may
show the propriety of union, but at present it is better to leave the question as it is.

Prof. Giard (1886) finds a third and closely allied species of Evarne commensalistic
on Cucumaria pentactes off Brittany and the neighbouring shores of France.

3. Evarne Hubrechti, 1 Mcintosh, n. s.
Specific Characters. — Head comparatively small, and in the adult the eyes are very
large; both pairs more or less lateral in position. Median tentacle long, tapering,
1 Named after the distinguished Professor of Zoology at Utrecht.

EVARXE HITBRECHTI. 361

smooth, and the tentacular and cirriform appendages are also smooth. The palpi are
large, slightly tapered, and smooth. Body rather short and broad, about thirty-one
bristled segments, much tapered posteriorly, and furnished with a caudal rudder in the
shape of a thick style with a membranous flap, tapered from base to apex. In structure
it is delicate and brittle, and is tinted of a deep madder-brown both dorsally arid ventrally.
Segmental eminence marked, but no distinct papilla is visible. Scales, fifteen pairs, thin
but tough, rounded in front, rest large and ovate, completely covered with minute spines,
but with a smooth edge ; some cling firmly to the peduncles. Feet of considerable
length, dorsal division rudimentary ; bristles translucent, large, long, and acutely pointed,
with wide rows of spines like an JEquisetum ; ventral division large, ovate at the tip, and
having a broad fan of very long slender bristles with long spinous regions and acute tips,
mostly bifid. Yentral cirrus comparatively long and slender.

Habitat. — Procured by the ' Triton ' in August, 1882, at 600 fathoms ; at Station 2,
500 fathoms ; in the tow-net at 300 fathoms, and again at the surface.

Length about 14 mm.

Head (Plate XXVIII, fig. 6) comparatively small, broadly ovoid, with a median
groove and two prominent anterior peaks. Both pairs of eyes are dorso-lateral in
position, their largest surface in the preparation being lateral. Their large size and
the limited area of the head leave little of the lateral region free, viz. a small portion at
the peaks, the narrow line of separation between the pairs, and the brief space between
the last and the nuchal collar. No distinct trace of a corneal opacity is present. From
the contour of the head the anterior pair look forward and outward, the posterior out-
ward and upward. Both pairs are partially seen from the dorsum. In young examples
the eyes are considerably smaller. The long median tentacle is pale, slightly enlarged
immediately above the basal region (in spirit), then it tapers to the filiform tip. Its
surface is quite smooth, the median nerve-trunk being readily seen through its translucent
granular layer. The lateral tentacles are pale, subulate, and small, with a filiform tip
and a madder-brown basal region. They likewise are quite smooth. The palpi are
smooth, of moderate length, and somewhat thick, with comparatively short tapering tips.
They have a faint tinge of madder-brown at the base. The tentacular cirri, the upper of
which is, as usual, larger than the lower, have a similar form to the median tentacle.

Body covered by the large scales ; of about thirty-one bristled segments, and com-
paratively short and broad. It is slightly narrowed in front, and tapers somewhat
rapidly posteriorly. The dorsum has a madder-brown hue, with transverse elliptical
markings in the middle, paler on the feet. Posteriorly the segments have very beautiful
patterns, the madder-brown ellipse being surrounded by a pale and somewhat crenate
line. The entire under surface is madder-brown, with a pale median band, and iridescent,
the darker region in front showing fine metallic lustre. The feet are also slightly tinted
of the same brownish hue, the ventral cirrus being thus rendered conspicuous, and
between its base and the body a pale transverse line occurs. The segmental eminence is
marked, but no distinct papilla is visible— a feature perhaps associated with the pelagic
habits of the species.

Posteriorly the body terminates in a single comparatively thick style, which arises
beneath the anus and to the left. It remains nearly cylindrical from base to tip— which

362 EVARNE HUBRECHTI.

is somewhat bluntly rounded. The remarkable feature about it. however, is the presence
of a broad web or lamina attached to its lower surface, and which tapers from base to
tip. The organ, therefore, forms an efficient rudder. The body is delicate and brittle,
and the length of the bristles still further adds to the characteristic appearance.

Scales (Plate XXXIII, fig. 1), fifteen pairs, thin but tough, entirely covering the
back, comparatively large and apparently smooth to the naked eye, but minutely granular
under a lens. The first pair are rounded, the succeeding more or less ovate, and all with
a smooth border. The entire surface is studded with minute horny spines, which increase
in size from the inner to the outer border, and especially posteriorly. No part of the
scale is free from them. It sometimes happens that a few of the larger conical spines
project beyond the posterior border under examination, but otherwise the edge is smooth.
In shape almost all the spines are acutely conical. The scales are pale, and the scar for
the pedicle has an unusually large area behind it, from the great development of the
posterior region. The scales, as a rule, adhere firmly. In a mounted scale a number of
rounded bodies of various sizes, with a distinct capsule and granular contents, occurred,
along with certain rectangular bodies composed of two halves, like Desmids. Their
nature is enigmatical.

Feet. — The first bears a short, slightly curved bristle or two of the dorsal pattern,
and resembling the shorter forms next the body in the typical foot.

The second foot presents a double ellipse — one for each division — with the project-
ing papilla for the spine, the inferior having in addition a fleshy digit-like process
above the spine. A few of the inner bristles of the dorsal tuft are short and curved like
those of the tentacular cirrus, each, however, having a smoothly pointed tip. The chief
bristles are long, translucent, and tapering, with a long bare tip marked by wide and
slightly oblique spinous rows, so that in some views it resembles an Equisetum of glass.
The ventral bristles are slender and elongate, with long spinous regions tapering to a
delicate tip, which is bare, and provided with a terminal hook. The filiform tip of the
long ventral cirrus extends beyond the bristles.

The next foot leads to the typical shape, which is peculiar, since in a ventral view
the tip is rounded. In profile, again (Plate XXX, fig. 10), the outline of the ventral
division is comparatively deep and rounded, with the spine near the middle, and the bristles
project as a broad fan. Dorsally the bristles (Plate XL, fig. 1) are remarkable for their great
length, diaphanous nature, and brittleness — conditions pointing to a pelagic habit. The
dorsal lobe itself forms only a small boss or eminence, and is thus in marked contrast
with the same region in Uvarne impar. From the inner side of this eminence spring
a few short, slightly curved tapering bristles, with closer rows of spikes, next which
are some shorter straight ones (Plate XL, fig. 3). The rest are large straight bristles
(Plate XL, fig. 2), tapering almost from the base to the smooth and sharp tip. The
spinous rows are short, slightly oblique, and very wide, as in an Bquisetum, the appearance
of the whole being diagnostic. The ventral series consist of elongated and slender
bristles (Plate XL, fig. 4), with long and finely tapered spinous regions superiorly. They
terminate in a long smooth tip, which is bifid and slightly hooked. The bifid condition,
however, is difficult to detect in some, and interiorly the tips are simple. The bristles
can be followed in the diaphanous foot inwards to the muscular boss at the spine.

EVARNE ATLANTICA. 363

Posteriorly the dorsal eminence disappears, and the bristles become more slender,
but still retain their characteristic structure. The tips of the slender ventral series are
extremely elongate, but traces of the bifid condition are still to be found in many. The
foot is thus practically formed of the ventral lobe.

In young examples the bristles, especially the ventral, are proportionally longer,
and at this stage — 5 to 6 mm. — two were pelagic at the surface.

In the development of these bristles the tip is practically complete on issuing from
the foot, and additions are made posteriorly as it pushes out.

The dorsal cirri have the form of the tentacular cirri, and their filiform tips extend
beyond the bristles. The ventral cirri are slightly brownish (madder), elongate,
subulate organs, the slender tips of which extend beyond the fleshy part of the foot.
They are perfectly smooth.

The delicacy of the body, the structure of the feet and bristles, as well as the
rudder-like caudal style, show that this species has peculiar habits, probably more or
less pelagic. What relation it may hold to other forms is unknown, and epigamy, or the
epitocous condition, may yet be clearly demonstrated in the family.

4. EVARNE ATLANTICA, Mcintosh, 1897.

Specific Characters. — Head like that of JEJvarne impar, but the cilia on the tentacles
and the papillge on the palpi seem to be smaller. The cirri generally are more slender.
Dorsal bristles longer and less curved than in E. impar, and their rows of spines closer.
Ventral bristles have longer tips than in E. impar.

Synonym.
1897. Evame atlantica, Mcintosh. Ann. Nat. Hist., ser. 6, xx ; p. 168.

Habitat. — Dredged at Rockall (Station 3a) by the Royal Irish Academy's Expe-
dition, June 15th, 1896.

A fragment of about fifteen segments of the anterior end.

The head resembles that of U. impar, Johnst., in general outline, but differs in
having somewhat smaller eyes. The tentacles and palpi also are similar, though the
cilia on the former and the minute papilla3 on the latter are less bold. The cirri
generally are a little more slender.

The body is thicker and more massive than in U. impar of the same size, and both
dorsally and ventrally in the preparation has a pinkish skin-colour. The arrangement of
the bristles at the side of the body is more trim. No segmental papilla is observable,
though the eminence is distinct. In this respect it agrees with & impar of the same
size, in which the papilla only becomes noticeable about the twelfth bristled foot. In
large examples it is evident on the seventh bristled foot.

Feet. — The first foot (bearing the tentacular cirri) has a few short bristles conforming
to the dorsal type, though with somewhat closer rows of spines. In the second foot the
dorsal bristles are longer and less curved than in R impar, and while there may be room
for doubt concerning the proportionate distances of the spinous rows, there can be none

364

EVARNE ATLANTICA.

about the length of the smooth portion at the tip, which is diagnostic of this form and
also of Evarne Normani} The present species differs from the latter, again, in the more
tapering extremities of these bristles, and in the closer rows of spines. The spinous tips
of the ventral series are somewhat longer than in E. impar, and in this respect approach
E. Normani.

In the typical foot the lower lobe is more pointed and the ventral cirrus longer than
in E. impar, so that it projects as far as the tip of the foot. The translucent dorsal
bristles (Fig. 31) are considerably longer and less curved than in the typical species, and
their rows of spines much closer, the latter character also distinguishing them from
E. Normani. The ventral bristles (Fig. 32) approach those of the latter species more
closely than those of Evarne Johnstoni, being somewhat shorter than those of E. Normani,
which, again, have stronger tips than those of E. Johnstoni.

Fig. 31. Fig. 32.

IS!

Fig. 31. — Tip of a dorsal bristle of Evarne atlantica. x 350.
Fig. 32. — Tip of a ventral bristle of Evarne atlantica. x 350.

No scales are present. In all probability they approach those of E. Normani,
By the lengthening of the tips of the ventral bristles and their general slenderness
this species and E. Johnstoni come near Antinoe and allied forms.

1 Op. cit., 173, pi. iii, fig. 13.

ANTINOE SARSI. 365

Genus XV. — Antinoe, Kinberg, 1857.

Lateral tentacles arising under the base of the median tentacle ; cephalic lobe with
lateral peaks anteriorly. Palpi subulate, densely covered with minute papillse. Dorsal
bristles long and tapering, with well-marked spinous rows. Ventral bristles long and
slender, with elongated spinous regions and hair-like tips. Segmental papillae distinct
though not large.

This genus approaches Harrnothoe.

1. Antinoe Saksi (Kinberg), Malmgren, 1865.

Specific Characters. — Head about as long as broad, with two acute anterior peaks.
Two posterior eyes, nearer each other and smaller than the anterior, which are situated
within the dorsal margin of the widest part of the head, and considerably removed from
the anterior peaks. Median tentacle brownish, tapering, with a long filiform tip and
numerous long clavate cilia. Lateral tentacles with short clavate cilia. Palpi long and
tapering, with minute clavate papillae, often with processes on the blunt tips. Tentacular
cirri with very long slender tips and numerous clavate cilia of moderate length. Body
somewhat broad, most distinctly tapered posteriorly, segments thirty-eight ; dorsally of a
brownish hue — in various transverse belts posteriorly. Scales, fifteen pairs, ovoid or
rounded, soft, rather opaque, and with cilia of moderate length on the outer and
posterior borders, while some show minute conical spines on the surface. Dorsal
division of the foot with a prominent spine, and rather long, slightly curved tapering
bristles with well-marked spinous rows, which in profile occupy half the bristle, only a
minute portion of the tip being bare. The ventral division has a very prominent
spinigerous lobe, and a series of slender bristles — with long spinous regions and hair-like
tips. Dorsal cirri long, tapering, with a filiform tip, and the surface has numerous short
clavate cilia. The ventral cirrus is subulate, rather long, and with short clavate cilia.
Segmental papilla begins on the fifth foot.

Synonyms.
1865. Antinoe Sarsi (Kbg.), Malmgren. JNTord. Hafs-Ann., p. 75, tab. 9, fig. 6.

1871. „ „ Elilers. Spitzb. Annel., Ann. Nat. Hist., 4th ser., viii, p. 53 (transl. from Sitzb.

d. Phys.-med. Soc, Erlangen).

1873. „ „ Sars. Bid. Christ. Fauna, p. 5.

1874. „ „ Mcintosh. Ann. Nat. Hist., April, 1874, p. 263.

1875. „ „ Ehlers. Annel. < Porcup./ 1869, p. 33, Taf. ii, f. 3 and 4 (Zeitsch. f. w. Zool.,

xxv, p. 53).
1879. Polynoe Sarsi, Theel. Annel. Nova Zemb., p. 16.

„ Antinoe Sarsi, Tauber. Ann. Danic, 80.
1886. „ „ Marenzeller. Porif., &c, Jan Meyen, p. 12.

Length of largest about If inches.

Habitat. — In the 'Porcupine' Expedition of 1869 it occurred at No. 28 (July
5th, 1869), in 1215 fathoms on mud.

366 ANTINOE SAKSL

It extends to various parts of the Arctic Sea, to the Gulf of St. Lawrence (where it
is common), and to the shores of America (Yerrill).

Head (Plate XXVIII, fig. 10) about as long as broad, with two acute peaks in front.
The two posterior eyes are much nearer each other and smaller than the anterior pair —
which are situated within the dorsal margin of the widest part of the head, and therefore
considerably removed from the anterior peaks. These eyes in the Canadian forms are
further forward, more lateral in position, and present in some a lens-like central thickening,
and both they and the posterior pair are larger than in the European forms, which have
eyes of moderate size. The median tentacle is brownish, with a slight swelling below
the very long filiform tip. It has numerous long clavate cilia. The lateral tentacles are
small and subulate with a tapering tip, and have short clavate cilia, The palpi are long
and tapering, furnished with minute and somewhat clavate papilla, which often have
processes at the tip. A few of the papillae occur on the tapering extremity. The
tentacular cirri have very long slender tips and numerous clavate cilia of moderate length,
a few shorter forms occurring on the filiform region distally. These organs are boldly
areolar, and, like the tentacle, are tinted brownish.

Body. — Somewhat broad, tapered a little in front, and more distinctly so posteriorly;
bristled segments thirty-eight. The colour is brownish dorsally, generally arranged in
belts across the segments, which posteriorly show a broader belt in the middle and a
narrow one in front and behind. The Canadian preparations are darker, the tubercle on
the cirrigerous segments being dark olive, so that the tints are striking. As usual, the
pigment is broken up in the terminal segments, and a pale band occupies the median
line. The under surface is pale. The segmental elevations are well marked and project
posteriorly as distinct processes, while the papilla are directed nearly straight backward
instead of between the feet. A marked feature ventrally is the prominence of the
spinigerous lobe of the ventral division of the foot.

Proboscis. — The extended proboscis has nine conical papillae dorsally and ventrally.

Scales (Plate XXXIII, fig. 17). — All the Norwegian examples show that the ovoid
or rounded scales have their surface covered with small conical spines ; indeed, in the
Arctic examples these are visible under a lens. The spines in the posterior scales (those
alone present) attain larger dimensions externally and anteriorly, and three or four larger
conical processes are found on the posterior border. Cilia, of considerable length in the
latter case, occur on the outer and posterior borders and invade the neighbouring surface.
They are slender and translucent, with clavate tips.

On the other hand, the scales of the Canadian examples were more or less smooth,
soft, somewhat reniform, and furnished with cilia of moderate length on the outer and
posterior borders and on the neighbouring surface. The opacity of the scale, however,
may hide the spines if they are few. At any rate, in the specimen from Barent's Sea
numerous conical papillae were present. Unfortunately many have no scales, and at best
they are few in the preparations.

Feet. — The first foot bears a strong spine and about three bristles of the dorsal type,
very slightly curved, and with boldly marked spinous rows. The spine forms a powerful
projecting process in large specimens.

The second foot is also marked by the great prominence of the dorsal spine, and has
a series of somewhat strong slightly curved bristles with well-marked spinous rows, which

ANTINOE SARSL 367

in profile occupy only about half the bristle, the inferior half being thus smooth and
translucent. Only a minute portion of the tip is bare. Yentrally are a series of slender
bristles, with long spinous regions and capillary extremities.

In the typical foot (Plate XXXI, fig. 2) the dorsal spine is still very prominent, and
the bristles are long, slightly curved, and with a moderately acute tip. The spinous
rows are distinct, and leave only a very short portion of the tip bare (Plate XL, fig. 7).
The ventral are slender, elongate, with long spinous regions and remarkably attenuate
tips, which form long hair-like processes (Plate XL, fig. 8). The foot has a long process
for the spine superiorly. In the terminal segments the character of the bristles alters
little, except as regards size.

In the Canadian examples thecate Infusoria are common on the dorsal bristles,
and, in some, Loxosomx. Infusoria also frequent the British form, and a cluster occurs
in one on the projecting spine of the ventral division of the foot.

The dorsal cirri are long and tapering, with a filiform tip, while the surface has
numerous short clavate cilia. The ventral cirrus is subulate and rather Ions' and
tapering, with numerous short clavate cilia.

In certain respects, e. g. the slenderness of the ventral bristles in comparison with
the dorsal, and in the clearly separated rows of spines, Antinoe approaches Evame.

Reproduction. — None of the specimens give any light on this process.

Habits. — So far as present knowledge goes, this species would seem to frequent
deep water only.

Ehlers (1871), in his account of the Spitzbergen annelids, gives a very interesting
description of two well-marked colour-varieties of this species. The more abundant
form presents the brown coloration which Malmgren states distinguishes the Spitz-
bergen from the Baltic form. In the other the scales are bordered with greenish-grey
on the inner and posterior margins, with a darker spot at their junction, the ventral
surface and feet being colourless. The dorsal surface of each segment is banded with
greyish-green, the pigment being variously developed in different examples. This
variety attains even larger dimensions than the first, viz. 46 mm. in length and
24 mm. in breadth, as against 35 mm. and 17 mm. He contrasts these two varieties
with that now living in the Baltic, which is smaller, possesses a greenish dorsal
coloration, and scales with brownish margins, — that is, it has an intermediate position.
He is of opinion that it has retained, since the glacial period (when the Baltic was
united with the Arctic Sea), the original coloration, from which the two races in the
Spitzbergen Sea have been developed by differentiation.

Ehlers found a Nucula 8 mm. by 6 mm., and 4 mm. in thickness, in the intestine of
one of the largest greyish-green varieties. The same author (1875) describes it from
the ' Porcupine ' Expedition of 1869 at 1215 fathoms, and gives good figures of the
bristles. His specimens had respectively twenty-nine and thirty-five segments.

Hansen found it in the collection made by the Swedish Expedition of 1876.

Marenzeller (1877) includes this form also in the annelids of the Austrian North
Polar Expedition from a depth of 240 metres on mud.

Verrill (1879) enters it in his list from Cape Cod to the St. Lawrence, where
Mr. Whiteaves dredged it in 1873.

48

368 ANTINOE EINMARCHICA.

Several specimens occur along with Uvarne Johnstoni in Norwegian collections
(1870) by Dr. Merle Norman. Whether any relationship of an epitocous kind exists
with this or other forms is unknown.

Mobius records it under the name of Polynoe cirrata on the authority of Theel, who
gives P. cirrata as a synonym of the species, but the latter has since seen that there
are no grounds for this arrangement.

2. Antinoe finmarohica, Malmgren, 1867; an. var. A. Sarsi.

Specific Characters. — Head as in Antinoe Sarsi, and the body has a similar outline,
with thirty segments in the largest example (less than -| inch in spirit). Scales with
short and sparse cilia on the outer a,nd posterior borders, and the minute spinous
papillae are chiefly distributed on the anterior half of the scale as well as externally and
internally. Feet similar to the foregoing in general structure, but the dorsal bristles
have slightly narrower intervals between the rows than in Antinoe Sarsi, and they are
longer, less curved, and more acutely tapered at the tip. The ventral bristles are
decidedly stouter; the tips, instead of being capillary, have a long delicate portion of
appreciable breadth, with a slightly curved point.

Synonyms.

1865. Antinoe finmarchica, Malmgren. Ann. Poly., p. 13.

1876. „ „ Mcintosh. Trans. Z. S., ix, pp. 387 and 400, pi. lxxii, f. 1, 2.

Habitat. — Dredged off the west coast of Ireland (Donegal) during the ' Porcupine '
Expedition of 1869 in 20 to 420 fathoms ; and also by the Expedition of the Royal
Irish Academy in 1886 at 93 fathoms.

Head differs from that of A. Sarsi in that the eyes are proportionally larger, and
the anterior pair are more lateral in position. The other organs present no feature of
moment in the preparations,

Body of similar outline to the foregoing species, and of thirty segments, but, as the
largest specimen is under \ inch in spirit, it may be immature.

Scales (Plate XXXIII, fig. 8) show a definite structure in contrast with A. Sarsi,
but changes may occur in course of growth. They have a similar outline, but the cilia
on the outer and posterior borders are short and sparse, and the minute spinous papillge
are chiefly distributed in the anterior region in front of and on each side of the scar for
the pedicle. The rest of the area posteriorly seems to have less distinct papilla, but
towards the posterior margin the cilia appear on the surface. Some of the papillae have
the summits curiously truncated. Small Norwegian examples of A. Sarsi show the
same arrangement of the horny papillae, so that the question is still open.

Feet. — The general structure of the fleshy part of the foot (Plate XXX, fig. 13) is
the same as in the previous species. The dorsal bristles are long, translucent, sharply
tapered from the commencement of the spinous portion to the tip, and with slightly
narrower intervals between the spinous rows than in the former (Plate XL, fig. 9), the

AXTINOE MOLLIS. 369

latter figure representing one of the straighter forms next the ventral, while an outer
curved one is given in Plate XL, fig. 10. In contrast with those of A. Sarsi they are
longer, less curved, more acutely tapered at the tip, and with more closely arranged
spinous rows, and this is the case whether we compare them with adult or with young of
the same size. The ventral bristles, again, are decidedly stouter than in A. Sarsi, and
superiorly, instead of those with capillary tips, are bristles with a very long and delicate
portion of appreciable breadth, distinctly spinous, and ending in a slightly curved point.
The spinous rows continue nearly to the latter. The tips gradually (Plate XL, fig. 11)
diminish in length toward the inferior edge, and the spinous rows are closer. Some,
e. g. those from the south-west coast of Ireland, show a tendency to have long spines at
the tip as in Antinoe mollis (Plate XL, fig. 13), but the feature is indistinct. Young
specimens of A. Sarsi of the same size show the fine hair-like tips of the adult.

The dorsal cirri have somewhat fewer cilia than in Antinoe Sarsi.

This form seems to frequent deep water. It is probable that further observations
may show that it is only a variety of the foregoing, the points of difference between
them, and it may be other forms, being due either to epigamy or other sexual variation.
In the meantime the distinctive characters will ensure its ready identification.

3. Antinoe mollis, G. 0. Sars, 1871.

Specific Characters. — Head somewhat broader than long (in spirit), with blunt
anterior peaks. Byes larger than in A. finmarchica, the anterior pair being situated
laterally at the base of the peaks, and thus are not visible dorsally. The posterior pair
are separated by a wide interval from the anterior pair. The palpi show minute papillse
under a high power. Lateral tentacles with short clavate cilia. First pair of scales
rounded, rest ovoid ; outer and posterior borders with slender clavate cilia, which are
longest externally; entire surface closely covered with short, blunt, and rather soft
spines. Dorsal bristles rather large, long, slightly curved, less acutely tapered than in
Antinoe finmarchica, and with close rows of spines. Ventral cirri with short clavate cilia.

Synonyms.
1871. Lsenilla mollis, Gr. 0. Sars. Viclensk.-selsk. Forhandl., 1871, p. 406.
1873. „ „ idem. Bidrag Christ. Fauna, p. 7, pi. xiv.

1876. Antinoe mollis, Mcintosh. Trans. Zool. Soc., ix, p. 400, pi. lxxii, f. 3, 4.
1879. Lsenilla mollis, Tauber. Ann. Danic., 81.
1883. Harmothoe mollis, Levinsen. Xord. Annul., 191.

Habitat.— Dredged on a bottom of muddy sand at a depth of 20 fathoms off Donegal,
and 370 and 420 fathoms at various points of the Irish coast in the ' Porcupine ' Expe-
dition of 1869. Dredged at 257 fathoms in the Expedition of 1870, in the Atlantic. A
similar form occurs in Norwegian waters, and Yerrill includes it in his American list.

Head (Plate XXVIII, fig. 14), in spirit, somewhat broader than long, with blunt
anterior peaks. The eyes are larger than in the previous form, the anterior pair being
situated at the base of the peaks laterally, and thus are not visible from the dorsum,
as well as separated by a wide interval from the posterior pair, which are smaller,

370 PHYLLAXTINOE.

and placed in front of the nuchal collar. The palpi appear to be smooth under a
low power, but when magnified 350 diameters are minutely papillose. The median
tentacle is absent. The lateral are small, somewhat enlarged at the base, and with a
filiform tip. They have a few short clavate cilia. According to Sars the tentacular
cirri also have a few clavate cilia.

Body has forty-three bristled segments (Sars). The only available British example
is fragmentary. The segmental eminence is distinct, but the indifferent preparation
showed no differentiated papillae.

Scales have, so far as present, a somewhat ovoid outline (Plate XXXIII, fig. 9),
with the exception of the first pair, which are rounded. The outer and posterior borders
have slender clavate cilia, which are longest externally, becoming short and sparse as
we proceed inward. Their entire surface is closely covered with short, blunt, and rather
soft spines, and thus they differ in certain respects from the figure and description
of Sars.

Feet. — The first foot has a sharp spine and a group of four small bristles conforming
to the dorsal series.

The typical foot presents dorsally a series of rather large, long, yellowish bristles,
which, though as conspicuous in size as those of A. finmarchica, are much less acutely
tapered, and have closer rows of spines. They are slightly curved (Plate XL, fig. 12).
The superior ventral bristles have somewhat shorter tips than in A. finmarchica, and
the rows of spines are not so distinctly separated. The tip is similar, viz. slightly
hooked ; but the spines become so elongated towards the tip that they project on each
side like a series of filaments (Plate XL, figs. 13 and 14).

Xo dorsal cirri are present, but the ventral cirri are long and filiform with short
clavate cilia.

This form appears to present the following differences from the species described by
Sars :— The anterior pair of eyes lie in front of the middle of the head, and are not visible
from the dorsum, whereas in the Norwegian form they are situated on the median lateral
eminence, and therefore considerably removed from the front. They are also seen from
the dorsum. The posterior pair of eyes are less widely separated in the British form.
It is difficult to compare the bristles, since some of the finer characters are lost in the
kind of plate adopted by Professor Sars, but they closely approach each other. The
scales, on the other hand, present certain divergences, e. g. in the presence of only
minute cilia in the northern form.

Genus XVI. — Phyllantinoe, Mcintosh, 1876.

Head short and broad; base (ceratophore) of the median tentacle passing far
backward between the prominent anterior peaks. Anterior pair of eyes much larger
than the posterior; pairs separated by a brief interval.

Dorsal bristles short, stout, translucent, with prominent and close rows of spines,
and a short, acute, smooth tip. "Ventral bristles extremely slender ; spinous region long
and tapering, tips simple.

In the absence of accurate information concerning the relations of such rare forms

PHYLLANTINOE MOLLIS. 371

to the epitocous condition of species perhaps elsewhere described, it has been deemed
prudent to continue the generic distinctions. The large size of the eyes and the
slenderness of the ventral bristles may be associated with sexual changes.

Phyllantinoe mollis, Mcintosh, 1876.

Specific Characters. — Head short and broad, the base of the median tentacle passing
far backward between the prominent and acute anterior peaks. Eyes large, especially
the anterior pair, the pigment in which is crescentic in the preparation, and placed on
the median lateral prominence. The posterior pair are smaller, and lie in front of the
nuchal collar. The pairs are separated by a brief interval. The fragmentary example
has about forty bristled segments, the body being elongate, and tapering much pos-
teriorly. The dorsum has brownish pigment, which posteriorly is regularly arranged in
the segments. Segmental eminence prominent, but papilla minute. Dorsal bristles
short, stout, translucent, considerably carved, with prominent and somewhat close rows
of spines, the tip having a short, acute, smooth region. Ventral bristles translucent,
extremely slender ; the spinous region rather long and tapering, with a short smooth
portion at the tip, which is simple and slightly bent. The spinous rows are distinct but
somewhat close.

Synonym.
1876. Phyllantinoe mollis, Mcintosh. Trans. Z. S., ix, vii, p. 401, pi. lxxii, f. 5 and 6.

Habitat. — Dredged in 539 fathoms in the Atlantic during the 'Porcupine 5 Expe-
dition of 1870.

Length. — The fragmentary specimen measured about | inch.

Head (Plate XXVII, fig. 18) in the preparation is shorter than broad, the base of
the median tentacle passing a considerable distance backward between the prominent
and acute anterior peaks. The four eyes are visible from the dorsum. The anterior
pair are unusually large, crescentic in form, and situated on the median lateral promi-
nence of the head. Their form would suggest the occurrence of a lens-like corneal
thickening in the concavity in front, but the preparation is uncertain on this point.
These eyes look forward, outward, and slightly upward. The posterior pair are smaller,
but still of comparatively large size, and lie in front of the nuchal collar. They are
more or less rounded, with traces of lenses. The pairs of eyes are separated by a brief
interval, as in Evarne. All the cephalic appendages are absent.

Body. — The fragmentary example showed about forty segments, besides head and
tail. It is characteristically elongated, and tapers much posteriorly. The whole body
is soft and delicate, and the feet comparatively short. The dorsum has a brownish
colour throughout, and the pigment posteriorly is somewhat regularly disposed in the
segments. The segmental eminence is prominent, but so far as could be made out the
papilla is short and small. The ventral surface has the usual pale iridescence.

No scales are present.

Feet.— The structure of the feet could not fully be made out, as the condition of the
specimen was unfavourable, but they seem to have proportionally short fleshy lobes.

372 SCALISETOSTTS COMMUNIS.

Anteriorly the pedicle for the scale is large, the ventral branch of the foot projecting
further than the dorsal as a soft process. Few bristles occurred in any of the feet.
The dorsal bristles (Plate XL, fig. 15) are short and stout, translucent, considerably
curved, with prominent yet somewhat close rows of spines, and an acute, smooth portion
at the tip. The ventral bristles (Plate XL, fig. 16) are translucent and extremely
slender both as regards shaft and tip, the latter being long and tapering to the bare
region at the point, which is simple, and in some is slightly bent. The spinous rows are
distinct, but somewhat close.

So far as ascertained, no form approaches this species.

Genus XVII. — Soalisetosus, Mcintosh, 1885.

Anterior border of the head smoothly rounded. Eyes placed close together on each
side, while the right and left pairs are widely separated and placed far back. Palpi
smooth. Proboscis somewhat thin. Body of moderate length. Scales cover the dorsum
in front, but leave the centre bare posteriorly. Segmental eminence distinct, but the
papilla is small. Bristles transparent as crystal ; dorsal slightly curved, and with 5 — 8
very distinct rows of spines. Ventral bristles slender, elongate, with close spinous rows,
and hooked and bifid tip. Longitudinal ventral muscles present a well-marked convexity
in section. The ventral area is of considerable width, nerve-cords flattened and indefi-
nite. Cuticle and subjacent granular layer attenuate. A strong band of fibres connects
the insertions of the powerful oblique muscles, and thus bounds the cords internally.

Professor Griard 1 gave an amended description of Kinberg's genus Hermadion to suit
the present forms, but the name Scalisetosus had already been published. It is perhaps
well to remove them altogether from Kinberg's genus.

1. Soalisetosus communis, Belle Chiaje, 1841. Plate XXVI, fig. 6.

Specific Characters.— Head touched with yellowish or whitish pigment, anterior lobes
smoothly rounded ; eyes of each side placed close together, while the right and left pairs
are widely separated and placed far back ; anterior pair larger, and wider apart. Median
tentacle long, slightly dilated below the filiform extremity, and with rather numerous
short clavate cilia. Lateral tentacles long, with filamentous tips, and similarly coated with
cilia. Palpi smooth. Tentacular cirri resemble the tentacles. Body \ to | inch long ; seg-
ments twenty-two to forty-two. Colour pale yellowish, faintly marked with brown
towards the posterior end— especially between the feet ; or speckled with brownish
spots arranged transversely, and numerous yellowish or whitish specks. Occasionally the
dorsum is blackish, interrupted by numerous transverse lines. The ventral surface is
pale. Segmental eminence prominent, but papilla minute. Scales fifteen pairs,
rounded or ovate, minutely dotted on the ventral surface, outer and posterior portions

i (

Bullet, scient./ No. 1, 1886, p. 7.

SOALISETOSUS COMMUNIS. 373

dorsally studded with a series of small papillse or short cilia, and, besides, with a
smaller number of much larger clavate processes. The outer and posterior borders
of the scale have a close series of short cilia with globular extremities. The
transparent scales show the nerve-distribution minutely. They have numerous yellow
patches, or are dotted with white specks, or are of a purplish-brown hue, especially
at the inner border. The bristles are crystalline in their transparency. The dorsal
bristles are slightly curved and not much tapered, have about eight rows of spines,
and the tip is notched. The ventral bristles are slender and elongate, the shaft
dilated superiorly, and with a spinous frill. From this the spinous region passes
with a slight bend to the dorsum, and ends in a well-marked hook with a secondary
process apart from its base. The dorsal cirri are rather long, with a slight dilatation
beneath the long filiform tip. They have similar cilia to the median tentacle. The
ventral cirrus is also rather long, but smooth.

Synonyms.

1841. Lysidice communis, Delle Chiaje. Discriz. e not., 'hi, p. 104, tab. 103, f. 1.

1853. Aphrodita velox, Dalyell. Pow. Creat., h, 169, pi. xxiv, f. 13.

1864. Polynoe pellucida, Ehlers. Die Borst., p. 105, taf. in, f. 5, 7 — 13 j taf. iv, f. 1 — 3.

1868. Her madion fragile, Claparede. Ann. Chet. Nap., 73, pi. v, f. 2.

1869. Lepidonotus pellucidus, Mcintosh. Trans. R. S. E., xxv, p. 408, pi. xv, f. 2.

1870. Hermadion fragile, Claparede. Supplem., Ann. Chet. Nap., p. 16, pi. ii, fig. 2.
1875. „ pellucidum, Mcintosh. Invert, and Fish., St. A., p. 117.

„ ,, fragile, Marion and Brohretzky. Ann. sc. nat., 1875, p. 6.

„ „ pellucidum, Marenzeller. Sitz. der k. Akad. (sep. Abd.), 13.

1880. „ „ Langerhans. Z. f. w. Zool., xxxiii, p. 271. * * %

1884. „ „ V. Cams. Fauna Medit., 203.

1886. „ „ G-iard. Bullet, sc. Xord, i, p. 12.

1888. „ „ De St.-Joseph. Ann. d. sc. nat. (7), v, p. 177, pi. viii, f. 51.

1890. „ „ Malaquin. Ann. Boulon., 22.

1891. „ „ Hornell. Op. cit., p. 237.

Habitat. — Though not abundant it extends from Shetland to the Channel Islands
in deep water, in the latter region between tide-marks, and on both the eastern and
the western coasts. It ranges to the Mediterranean and Madeira. It is commensalistic
on star-fishes (Hornell).

Head (Plate XXVII, fig. 12). — ■ Yellowish in some, or with white pigment-specks
posteriorly and at the eyes ; somewhat trapezoidal, the anterior lobes being smoothly
rounded instead of forming the usual peaks. The eyes of each side are placed close
together, while the right and left pairs are widely separated and placed far back. The
anterior pair are considerably larger than the posterior, and are wider apart. In the
British examples no lenses are present in them, but Claparede shows such in the
Neapolitan. The median tentacle is somewhat long, slightly dilated below the filiform
tip, and with rather numerous short clavate cilia. The lateral tentacles are long with
slender filamentous tips, and the column has the same short cilia. The palpi are smooth.
The tentacular cirri resemble the median tentacle. The ganglia form two distinct ovoid
masses in the head.

374 SCALISETOSITS COMMUNIS.

Body somewhat elongated, narrowed posteriorly, from ^ to f inch in length, though
sometimes longer. Segments from twenty-two to forty-two. Colour pale yellowish
throughout, in some faintly marked with brown towards the posterior end, especially
between the feet. In others the dorsum is speckled with brownish spots, arranged trans-
versely, and numerous yellowish specks. The naked portion behind the scales (including
the bases of the feet) is speckled with yellowish and whitish. In some Scotch examples the
dorsum is of a blackish hue, interrupted by numerous fine transverse lines, most marked
at the segment-junctions. In others a distinct band of dark pigment, interrupted by
numerous pale striae, most marked at the segment-junctions, passes along the dorsum.
The ventral surface is pale. The segmental eminence is prominent, but the papilla is
very small. Posteriorly the body terminates in two caudal styles.

Proboscis. — In extension this organ in the preparation shows apparently thinner
walls than in the ordinary forms. A wide canal thus exists in the centre, and the teeth
of the dorsal and ventral arches are widely separated. The usual nine papillae occur on
each arch. The teeth are pale brown (amber-coloured).

Scales (Plate XXXIII, fig. 7) cover the dorsum in front, but leave the centre bare
posteriorly. The normal number seems to be eighteen pairs, but few examples are
complete in this respect. They are more or less rounded or ovate translucent organs,
so that their minute structure is readily shown. The entire scale is minutely dotted, and
its areolae so arranged that the whole has the aspect of a transparent plate. The dots,
as Ehlers stated, appear to be in the cuticular layer of the under surface of the scale.
The outer and 'posterior portions of the scale, moreover, are studded dorsally with a
series of small papillae (modifications of cilia) of a cylindrical or clavate outline, many of
which run iii the lines of the finely branched nerves. In life these structures have
minute papillae or palpocils at the top, so that they probably perform the function of
special tactile organs, as indeed the able description of Ehlers would lead us to expect.
Only a small portion at the anterior and inner area of the scale is devoid of them, and
the nerves seem to terminate there in special end-organs. In addition, a series of much
larger clavate processes occurs thinly scattered over the same region (outer and
posterior), the largest sometimes projecting at the posterior border. They are readily
seen under a lens, standing out like minute pillars from the surface of the scale. These
receive nerve-twigs from the main branches, as shown by De St. -Joseph. The outer
and posterior borders, further, are beset with a closely arranged series of short globular
cilia, which also occur on the surface adjoining the edge. Many of these have micro-
scopic papillae (modifications of palpocils) at the tip, and .all show a minutely granular
central area — connected by the pedicle with the central connective tissue.

The nerve- supply of these organs is clearly outlined, as originally described by
Ehlers, and subsequently by Langerhans. At the posterior border of the scar for the
pedicle an enlargement (ganglionic) occurs, from which a main stem, rapidly breaking
up into branches, proceeds posteriorly, giving off twigs in a somewhat dichotomous
manner to the greater part of the scale, while a considerable branch goes forward, or
obliquely forward and outward, to complete the distribution. The smooth pale area at
the anterior and inner border has the smallest branch, probably because its functions
are less prominent. The nerve-twigs terminate in peculiar granular swellings — men-

SCALISETOSUS COMMUNIS. 375

tioned by Ehlers. Every .little papilla has its nerve-twig, and the granular enlargements
are connected with the system like fruit on the tips of a much ramified wall-tree.

In life the scales are quite translucent, sometimes presenting towards their anterior
and posterior borders numerous yellow patches at the papillae, or, as in the figure,
simply dotted with white specks. In others they have a faint purplish-brown tinge, 1
enlivened by orange and yellowish specks. The brownish tinge in some is towards the
inner border.

This active and irritable annelid frequently throws off its scales, but, as in other
cases, they are readily reproduced, an interesting account of their development having
been given by Ehlers.

De St. -Joseph found a number of specimens of Pedicellina belgica, van Beneden,
on the under surface of the scales.

Feet. — The first foot has a spine, but no bristles in the available specimens. De
St. -Joseph thinks this is the normal condition, but it is uncertain.

The dorsal division of the foot (Plate XXX, fig. 9) forms a short and broad process
bearing a series of translucent slightly curved bristles (Plate XL, fig. 17), which are not
much tapered distally. The rows of spines (or frills), about eight in number, are wide
apart inferiorly, while the tip of the bristle is notched as if from a minute claw or
secondary process. So homogeneous are these bristles that the usual longitudinal striae
have disappeared, and they resemble crystalline structures. In the ventral division
the spine forms the apex of a somewhat conical lobe, the upper slope, however, being
the shorter. The bristles (Plate XL, fig. 18) are equally transparent, but are slender
and elongate, the shaft terminating superiorly in a dilatation with a spinous frill, which,
like those of the dorsal, seems to be continuous. Erom this part the spinous region
passes with a slight bend towards the dorsum, and ends in a w r ell-marked hook with a
small secondary process — apart from its base beneath. The tip is flattened and not
much tapered, while the oblique rows of spines are somewhat adpressed. Little
change takes place in their structure in the posterior feet. The bristles seem to have
few parasites.

The dorsal cirri (Plate XL, fig. 19) are comparatively long, with a slightly dilated
portion beneath the long filiform tip. A yellowish belt, or in some a white ring,
marks the dilated portion. A few minute clavate cilia occur on their surface, and to
these nerve-twigs proceed. The nerve in the centre of the cilium appears to form an
enlargement in the opaque region at the base of the filamentous tip, and then continues
onward to the extremity. The posterior cirri are longer and more slender, and the
dilatation below the filiform tip is distinct.

The ventral cirri are rather long, smooth and tapering, their tips extending beyond
the bases of the bristles.

Reproduction. — Specimens procured in Bressay Sound in July showed ripe eggs,
and they were extruded in masses on rupture of the body. A male captured off Peel,
in the Irish Sea, bore ripe spermatozoa the same month.

Development. — A young form measuring a little over 1 mm. occurred amongst the

1 De St. -Joseph calls it roseate.

49

376 SCALISETOSUS COMMUNIS.

debris from the haddock-hooks on the 16th September, 1889. The injured specimen had
nine or ten bristled feet. The head showed two broad lobes in front, with a deep median
notch. Only the ceratophore of the median tentacle remained. The lateral tentacles
formed two short subulate organs. The palpi were fairly developed, with tapering tips.
The tentacular cirri had long slender tips, readily distinguished from those of allied
forms, and all these organs were smooth. A pair of black eyes— very widely separated—
were situated at the posterior part of the head. The teeth of the proboscis were clearly
visible as four hollow, pointed chitinous processes, with a small spur near the base, and
with long homy limbs for the attachment of muscles. The dorsal bristles were recognised
by their characteristic scalariform structure, from three to six or seven rows of spines being
present. They were more slender than in the adult, and tapered to a delicate tip. The
ventral bristles had the prominent) basal spur, and the long slightly curved and more
finely spinous tip. They projected considerably on each side, as became the nectochaete
stage.

The Aphrodita velox of Dalyell (1853) is in all probability this form, its translucency,
irritability, and general aspect being characteristic. His example was scarcely half an
inch long.

Kinberg's Hermadion (185 7) had the following characters :— Head broader posteriorly.
Posterior eyes distant from the anterior. Elytra fifteen, not covering the posterior
part of the body. Inferior bristles serrate below the apex. Foot elongate. There is
little that is diagnostic in this, and certainly the present genus requires a more precise
definition. His species were quite different from the British, and came from the Straits
of Magellan.

Ehlers (1864) gave a long and careful description of the species from Quarneo in the
Adriatic, detailing the essential characters, though his figures of the bristles and scales
needed improvement. His specimens were small, only from 7 to 9 mm., and of twenty-
two segments. The ciliated processes on the dorsum of the foot he considered the
external part of the segmental organs ; and as his examples bore reproductive elements
(summer), he was the more certain of the function of these organs. Haswell and Bourne
have both pointed out, in other species, the true segmental organs, which are on the
ventral surface. Ehlers thought the species had relationship with Kinberg's genus
Hermadion.

Olaparede (1868) published a fairly accurate account of this species from Naples,
pointing out the palpocils on the cilia of the cirri, and the ciliated cushions on the dorsum
of the feet. He also describes the remarkable facility with which the distribution of the
nerves can be followed. Its irritability struck him, and it cast off both scales and cirri.
He, like Ehlers, placed it under Kinberg's genus Hermadion mainly because the scales
only overlapped in the anterior region of the body.

In his supplement to the 'Annelids of Naples ' (1870), the same author recognised
that his Hermadion fragile was Delle Chiaje's Lysidice communis, which that author
had represented with only the anterior pair of scales, and to which his artist had
added a pair of cirri on every segment — of which the figure shows no less than
about sixty. In this communication Olaparede describes the segmental papilla
at the inner border of the foot ventrally, and shows a membrane investing the

SCALISETOSUS ASSIMILIS. 377

developing ova. He could not say that this membrane represented the wall of the
segmental organ ; indeed the absence of cilia inclined him to think it did not. He also
drew attention to certain cell-masses, with yellowish concretions in the centre, in the
intestinal diverticula — which he thought excrementitious. Prof. Giard agrees with me in
considering the distinctions of this form and 8. communis insignificant.

Marenzeller (1876) found 8. communis — in the Bay of Muggia at Trieste at a depth
of 18 metres, on Ophiothrix alopecurus — with thirty-three segments, and measuring 10 mm.
His example had fourteen pairs of elytra. He also mentions the advantages of the species
for the study of the nerves of the elytra, and alludes to the characteristic structure of the
bristles.

Langerhans (1879) records large examples (2 cm.) from Madeira. In connection
with the scales he discourses on tactile hairs and rudimentary organs of various animals.

Hornell (1891) found all his specimens as commensals on Echinoderms — one in the
ambulacral groove of Astropecten irregularis, another on Orossaster papposa, and the third
on Ophiothria? rosula. In one the pellucid scales had posteriorly a crescent of orange,
while the first pair had an orange belt all round.

2. Scalisetosus assimilis, Mcintosh, 1875.

Specific Characters. — Head similar to that of 8. communis, the eyes occupying the
same position, and the larger anterior pair having lenses. Median tentacle long, smooth,
slightly dilated below the filiform tip ; lateral tentacles short, also slightly dilated below
the tip. Palpi smooth. Tentacular cirri similar to the median tentacle. Body narrow
and elongated, with a brownish-black median band from the nuchal collar to the posterior
end. It is widest at the posterior part of the proboscidian region, though even there
less than a third of the breadth of the dorsum. Segmental eminence prominent, but a
special papilla could not be made out. Scales even more delicate and transparent than
in S. communis, minutely punctate on the under surface. The outer and posterior borders
and neighbouring surface have short clavate cilia — less numerous than in the former
species. The finely branched nerves and nerve-endings are similar. From the dorsum
the tips of the feet are blunt, and viewed laterally the dorsal eminence for the bristles
is large, while the spine has a broader border of granular epiderm. The dorsal bristles
are smaller than in 8. communis, slightly curved, the spinous rows being less prominent
and covering a shorter region of the bristle. The tip is bluntly rounded, bearing a minute
terminal claw and a small secondary process with a notch between. The ventral division
shows a broad terminal lobe with a bluntly rounded margin in front of the tuft of
bristles. The latter, as in S. communis, has the distal end of the shaft expanded, with a
small spinous collar, and from this the spinous region, with a slight bend to the dorsum,
tapers to the tip, which turns bluntly round to the spiked side and ends in a small hook ;
then, after an edge directed obliquely backward, a secondary process, lateral in position,
occurs. The rows of spines are extremely fine. The dorsal cirri are apparently shorter
than in the previous species, but of a similar shape— smooth, with a long filiform process
at the tip. Only one was observed.

378 SCALISETOSUS ASSIMrLIS.

Synonyms.

1875. Hermadion assimile, Mcintosh. Invert, and Fish., St. And., p. 117.

1876. „ „ idem. Trans. Zool. Soc, ix, 387 and 400, pi. lxx, f. 4—6.
1886. „ „ Harvey-Gibson. Yerm. Liverp., 151 and 348.

1891. „ „ Hornell. Op. pit., 237.

Habitat. — First found at St. Andrews, by my sister; afterwards on the west coast of
Ireland, in 80 fathoms, eighteen miles west of Skellig, by Dr. Gwyn Jeffreys ; south
of England and off the Spanish coast in the ' Porcupine ' Expedition. Oommensalistic
on Echinus esculentus (Harvey-Gibson).

Head has a similar outline to that in the former species, and in the preparations is
about as long as broad. The anterior lobes are smoothly rounded, and the eyes have the
same position and proportions, the larger anterior pair showing lenses. Both are
visible from the dorsum, and have a long portion of the head in front. The median
tentacle arises between the rounded anterior lobes, is long, smooth, slightly dilated
below the filiform tip, and minutely dotted under a low power, though this seems to
disappear in the mounted preparations. The lateral tentacles are short (in spirit) and
slightly enlarged below the filiform tip. The palpi are quite smooth. The tentacular
cirri are also smooth, slightly enlarged below the extremity, and are shorter than those
of 8. communis.

Body somewhat narrow, about f inch or upwards in length, very slightly tapered in
front, and very gradually diminished posteriorly. It is distinguished by a brownish-
black median band, which commences behind the head and continues to the tail. It is
widest towards the posterior part of the proboscidian region, though even there occu-
pying less than a third of the arch of the dorsum. In the preparations a darker band
occurs at the segment- junction, and a pale belt just in front. The ventral surface is
pale. The segmental eminences are prominent, but special papilla3 could not be
made out.

Attached to the ventral surface were several long Pedicellarise, probably from an
Echinus, and some were also fixed to the feet.

Proboscis. — The proboscis seemed to agree with the preceding in structure, but in
the preparation the papillse were somewhat clavate instead of acute.

Scales (Plate XXXIII, fig. 6), fifteen pairs, even more delicate and transparent
than in the previous form, and similar in shape. The cuticle of the inferior surface is
minutely dotted throughout as in S. communis. The outer and posterior borders have
short clavate cilia, less numerous than in the latter, and they also occur on the neigh-
bouring surface. The finely branched nerves arise from a similar ganglion at the
posterior border of the scar. 1

Feet. — In looking at the feet of the two species from the dorsum a decided difference
is observed in the terminal region of the ventral division, which in 8. assimilis (Plate
XXX, fig. 15) is somewhat blunt and rounded, whereas it is acute in 8. communis. In
profile this divergence is more pronounced, since the fleshy part of the dorsal lobe is
larger in 8. assimilis, and the spine has a broader covering, and instead of the acute cone

1 The thickenings observed in these scales do not appear to indicate papillas in all cases.

MALMGBEMA CASTANEA. 379

in the ventral division of the other species, it is here a broad terminal flap or lobe with a
bluntly rounded margin in front of the tuft of bristles.

The bristles are as translucent as in the former species, but are scarcely so long in
proportion. The dorsal are smaller than in 8, communis, and slightly curved, the
spinous rows being less prominent, and covering a much shorter region of the bristle
(Plate XL, figs. 20 and 21). About seven or eight are visible. The tip is bluntly
rounded, with a minute terminal claw and a small secondary process — with a notch
between.

The ventral division has a fan-shaped series of slender bristles, which, as in the
former case, expand at the end of the shaft, where a small collar of spines occurs, the
finely spinous region, with a slight bend to the dorsal edge, tapering to the tip, which
turns bluntly round to the spiked side and ends in a small hook ; then follows an edge
directed obliquely backward between this and the secondary process, which is lateral
(Plate XL, fig. 22). So fine are the spines on the terminal region that they are scarcely
visible, but the oblique lines in lateral view are distinct. The whole bristle thus charac-
teristically differs from that of 8. communis.

Habits. — This is evidently a commensalistic form on an Echinus, for Pedicellarige
frequently adhere to the skin and processes.

"What relationship the Bermadion echini of Professor Giard l has to this species
remains to be seen. It is evidently a closely allied form, but the minute characters of the
bristles are not given with that distinctness which is necessary for critical diagnosis.
It is interesting in this respect, that 8. assimilis was found by Harvey-Gibson near Port
Erin, Isle of Man, coiled round the peristome of Echinus esculentus, protected by the
peristomial spines. This author gives some interesting structural details. Hornell also
(1891) found one on the spines of Echinus esculentus near Liverpool Bar.

Genus XVIII. — Malmgeenia, Mcintosh, 1876.

Head somewhat pyrif orm, with the narrow end in front, devoid of peaks, the median
and lateral tentacles springing from the front as in Lepidonotus. Eyes large, nearly
forming a square. Palpi, tentacles, and cirri smooth. Body of moderate length and
breadth. Segmental eminences fairly developed, but without papillae. Scales, fifteen
pairs, smooth with the exception of a small group of papillas at the anterior curve.
Dorsal bristles translucent, short, and with faint spinous rows. Ventral bristles trans-
lucent, with rather short distal regions and five rows of spines ; the tip hooked, and a
secondary process beneath.

1. Malmgeenia castanea, Mcintosh, 1876.

Specific Characters. — Head somewhat pyriform with the harrow end in front,
without peaks, the lateral and the median tentacles springing from the front as in
Lepidonotus. Eyes large, nearly forming a square ; anterior pair in front of lateral

1 ' Bullet, so. Nord/ i, 1886, p. 8.

380 MALMGRENIA CASTANEA.

prominence, posterior in front of collar. Median tentacle of moderate length, smooth,
as are also the lateral tentacles, tentacular cirri and palpi, which are of moderate length.
Body about § inch long, of thirty-six to forty-one segments, with madder-brown pigment
posteriorly on the dorsum, and more sparingly on the ventral surface. Segmental
eminences fairly developed, but without evident papillse. Scales, fifteen pairs, adherent,
smooth, with the exception of a small and somewhat triangular group of papillse at the
anterior curve in those of a reniform shape ; variously bordered with madder-brown.
Dorsal bristles translucent, somewhat short, slightly curved, little tapered, and with a
rather abrupt point; spinous rows faint. Ventral bristles translucent, with short spinous
regions ; fine rows of spines, a well-marked hook at the tip and a secondary process after
an interval.

Synonyms.

1868. Eunoa. Report Brit. Assoc, 1868, p. 337.

1876. Malmgrenia castanea, Mcintosh. Trans. Zool. Soc, ix, 376, pi. lxvii, f. 15 — 19.

1886. Malmgrenia castanea, Harvey-Gibson. Verm. Liverp., 149 and 345.

„ Lsenilla castanea, Giard. Bull. Sc. Nord, i, 3.
1891. Polynoe (Malmgrenia) castanea, Hornell. Op. cit v p. 235.

Habitat. — Dredged by Dr. Gwyn Jeffreys off North Unst, Shetland, in 1867, in
90 and 96 fathoms, on Spatangus purpureas, and again in 1868, attached near the
mouth of the same Echinoderm, living on a bottom of shell-sand in 85 fathoms, twenty-
five miles north-north-east of Unst. The same veteran explorer of our seas found it in
80 — 125 fathoms, fifty miles west of Valencia, and in 110 fathoms, thirty miles west of
the Blasquet, south-west Ireland, in 1870. It was also dredged off St. Peter Port,
Guernsey, in 5—7 fathoms, in 1868. Mr. Hornell procured a few on the same host
in the Liverpool district in 20 — 22 fathoms, and Professor Herdman at the Isle of
Man. The Royal Irish Academy's Expedition of 1886 also procured a small example
in 480 fathoms. Professor Giard found it on the shores of France on the same
Echinoderm.

Head (Plate XXVIII, fig. 15) somewhat like that of Lepidonotus and Halosydna in
so far as the anterior border of the head runs into the base of the lateral tentacles. The
head is pyriform, broad and rounded behind, and narrowed in front. A pair of large
eyes lie in front of the nuchal collar, and a still larger anterior pair in front of the
lateral prominence, looking forward and outward. They are thus separated by a
considerable interval, and in the preparations nearly occupy the corners of a square. A
trace of a lens appears in the centre of the anterior pair, which are only partially visible
from the dorsum. The smooth median tentacle is moderately developed, and has a slight
swelling below the tapering tip (in spirit). The lateral tentacles are short and subulate,
with brownish pigment above the base. The palpi are smooth and of moderate length.
The tentacular cirri have a similar structure to the tentacle, most being somewhat
fusiform in outline.

Body about f inch in length, and having from thirty-six to forty-one bristled
segments. In most of the preparations it is pale anteriorly, but marked with madder-
brown pigment on the posterior segments. The colour varies considerably. The

MALMGBEXIA OASTANBA. 381

under surface of the body is iridescent pinkisb. In some a considerable amount of
brown pigment occurs posteriorly on each side of the median groove, and on the
segmental eminences, the site of the papilla being indicated by a darker speck. On the
whole the body is rather firm and broad, tapering a little anteriorly and more so
posteriorly. It terminates in two tapering caudal cirri, often of a deep brownish hue.
The segmental eminences are fairly developed, but no distinct elongation of the papillae
occurs.

Scales (Plate XXXIII, fig. 10), fifteen pairs, covering the dorsum, only a brief
portion of the tail being uncovered. The first pair are rounded, the succeeding reniform,
then ovoid, while the large posterior scales are irregularly quadrate. They are smooth
glistening organs under a lens. Under the microscope they present a somewhat trian-
gular group of papillae on the anterior concavity of the reniform scales, but in the
posterior scales these almost disappear. They are surrounded by a madder-brown belt,
with a tendency to the development of a denser portion in the anterior band. In
addition to broader marginal belts, the posterior scales have a general sprinkling of the
brown pigment. In some, especially those near the mouth of Spatangus purpureas,
all the exposed portion of the scale is tinted of a deep madder-brown. The scales
seem to adhere with considerable tenacity. The nerve-supply is similar to that in
Scalisetosus communis, but it is considerably obscured by pigment.

Feet. — In those examined only a spine occurred in the first foot.

In the typical foot (Plate XXX, fig. 5) the dorsal lobe is not much developed, form-
ing a small process from which the somewhat short bristles project. They are very
slightly curved, and taper a little towards the tip, which ends in a short point (Plate
XL, fig. 23). They are translucent, finely striated longitudinally, and with faint spinous
rows which go almost to the point. The dorsal spine projects in its sheath only a short
distance towards the lower edge of the bristle-bundle. The ventral bristles have slender
translucent shafts with a median axis, and striae distally as well as in the spinous region.
The ordinary appearance of one of the superior ventral bristles is shown in Plate XL,
fig. 24, scarcely a trace of the secondary process being visible below the well-marked
hook at the tip, and it altogether disappears ventrally (Plate XL, fig. 25). In the
developing form the secondary process is clearly indicated. The spinous rows are
close, and leave only the short terminal region bare. In specimens from Yalencia (south-
west of Ireland) this process is very distinctly seen, for instance, when the bristle is
slightly turned round (Plate XL, fig. 26). The secondary process is less marked in
specimens from the Channel Islands and Shetland, though indications are present in all.
In viewing the body from the ventral aspect the feet after the thirteenth become con-
siderably larger, and continue so till the thirtieth. This, as in Harmothoe marphysde,
may be associated with reproduction.

The dorsal cirri are of moderate length, smooth, slightly enlarged towards the end
of the column (in spirit), and with a filiform tip. They increase in length (as usual)
posteriorly. The ventral cirri extend to the bases of the bristles, and are subulate and
smooth.

In a small example dredged at 480 fathoms off the south-west coast of Ireland
both dorsal and ventral bristles are shorter and proportionally thicker, the former being

382 MALMGKENIA ANDREAPOLIS.

little tapered, and the latter with shorter spinous regions. The structure of the scales
corresponds with the typical form. The body is purplish throughout.

Habits. — They cling to the test near the mouth of Spatangns purpureas, and are thus
commensalistic forms.

Prof. Harvey-Gibson and Mr. Hornell found them on Astropecten irregularis —
between the rows of pedicels — at a depth of 20 fathoms in the Liverpool district.

The former gives an account of various structural features of this species.

2. Malmgkenia andreapolis, Mcintosh, 1875.

Specific Characters. — Head less pyriform than in M. castanea, with anterior peaks
more or less adnate. Eyes smaller than in M. castanea ; anterior pair wider apart than
the posterior. Median tentacle incomplete in all ; lateral tentacles small and subulate,
with two brown rings at the base. Palpi smooth. Tentacular cirri brownish, with a
few clavate cilia. Body elongate. Scales, fifteen pairs (?); first pair rounded, rest reni-
form or irregularly rounded. Those after the second pair with a brown ring more or
less complete, and at the sixth or seventh pair a V-shaped mark and a spot become
distinct. A belt of small papilla? (microscopic) occurs along the central region of the
anterior border and extends to the inner corner in the reniform scales. Tips of feet are
blunt and bifid ; dorsal division less developed than in M. castanea ; bristles slender,
slightly tapered, with a probe-like tip, and minute serrations on the edge. Ventral
bristles long, translucent, with a tapering spinous region which is simple superiorly —
ending in a distinct knob, the next series with a secondary process beneath the claw-
like tip, and inferiorly a single knob. A few clavate cilia occur on the stout, brownish,
and tapered dorsal cirri. The ventral cirri are slender and tapering, reaching only a
little beyond the bases of the bristles, and have a few short clavate cilia.

Synonyms.

1875. Malmgrenia andreapolis, Mcintosh. Invert, and Fishes, St. A., p. 117.

1876. „ „ idem. Trans. Zool. Soc, ix, p. 377, pi. lxvii, f. 20—23.

Habitat. — Not uncommon in the debris of the fishing-boats from the off-shore
(E. and R.), on the west sands after storms (E. M.), and in the stomachs of cod and
haddock at St. Andrews (E.).

Head (Plate XXVIII, fig. 8) tinted with brown on each side in the preparations,
the pale median groove marking off the symmetrical coloured areas on each side. The
anterior peaks are more or less adnate, but still visible at the origins of the lateral
tentacles. The head is less pyriform than in M. castanea, the eyes are smaller, and the
anterior pair are wider apart than the posterior pair. The anterior eyes look forward and
outward. The median tentacle is incomplete in all. The lateral are small and subulate,
with two brown rings at the base ; they are not in a condition to show cilia if they are
present. The palpi are smooth. The tentacular cirri are brownish, and have a few
clavate papillse. The pigment under the median tentacle is better marked than in the
former species.

MALMGRENIA AXDREAPOLIS. 383

Body rather elongate, more than an inch in length, and having about 36 — 37
bristled segments. In some the posterior region (about a dozen segments) is prettily
mottled in the preparations with dark brown pigment both dorsally and ventrally.
Occasionally the brown bars, sometimes with a pale centre, are best marked on the
ventral surface. Very little tapering of the body occurs anteriorly. The ventral surface
is for the most part pale and finely iridescent. The segmental eminence is prominent,
but a special papilla cannot be made out.

Scales (Plate XXXIII, fig. 11) probably fifteen pairs, but no specimen is complete.
The first pair are rounded, with a broad belt of madder-brown round the edge and a
spot in the centre, though in some the latter joins the outer portion of the ring. The
rest are reniform or irregularly rounded. The second scale in some has a brown ring
round the exposed part, and a patch near the outer border anteriorly, representing the
spot in the centre of the first pair and that of the scales behind. Those after the second
pair have a brown ring more or less complete, the broadest part beiug toward the inner
margin, and the spot at the anterior leg of the V-shaped mark gradually becoming more
evidently separated. About the sixth or seventh pair the V-shaped mark and the spot
become distinct. Posteriorly a tendency to the obliteration of the ring is observed, and
the spot becomes connected with the remnant of it at the inner border. In a few
the pigment in the posterior scales occurs in detached specks.

The scales appear to be smooth under a lens, but under the microscope a belt of
small papilla3 occurs along the greater part of the anterior and outer borders (where
the curve is). This belt is continued in the reniform scales round the anterior and inner
corner.

Feet. — As in the former species, the specimens seem to have lost the bristles in the
first foot.

In looking at the feet from the dorsum it is observed, in contrast with if. castanea,
that the tips are blunt and somewhat bifid, though the posterior process is less prominent
than the anterior flap. In profile, again (Plate XXXI, fig. 3), the foot has a greater
depth from above downward in proportion to its length, and thus the terminal cone is
shorter. The dorsal division is less developed than in the former species, and bears a
series of slender, slightly tapered, inconspicuous, translucent bristles, with a peculiar tip,
which forms a kind of rounded knob (Plate XL, fig. 27, representing one of the larger
bristles), of much interest when contrasted with the ventral forms, since it demonstrates
how closely the same type holds in both divisions. The serrations are minute, and leave
only a short portion of the tip bare.

The superior bristles in the ventral branch are long and translucent, have a long,
tapering, spinous region, with a distinct knob, like a probe-point, at the tip (Plate XL,
fig. 28). The spinous region quickly shortens in the succeeding forms, which show a
most interesting series of gradations from the first appearance of the secondary process,
the shortening of the probe-point and its gradual modification into a claw and a knob-
like tip with an oblique edge between it and the secondary process (Plate XL, fig. 29).
The spinous region in these is comparatively broad and short. Then, as the spinous
region diminishes inferiorly, the secondary process shortens and disappears, the bristles

50

384 HALOSYDNA GELATINOSA.

at the ventral edge, with short spinous regions having a marked dorsal curve, presenting
only a short knob, smoothly rounded, at the tip (Plate XL, fig. 30).

The dorsal cirri are stout, shorter than in M. castanea, brownish, and, in spirit, taper
from base to tip, a few short clavate cilia occurring on the surface. The ventral cirri
are slender and tapering, and only reach a little beyond the bases of the nearest bristles.
A few short clavate cilia are present.

Habits. — This species is probably commensalistic on another form, probably an
echinoderm, but hitherto it has occurred at St. Andrews only in the free condition, viz.
as thrown on the beach after storms.

n i

Genus XIX. — Halosydna, Kinberg, 1857.

Body linear-oblong ; head continuous anteriorly with the bases of the median and
lateral tentacles. Eyes large. Palpi smooth. Xuchal collar with a prominent flap.
Segmental eminences distinct and the papillae long. Proboscis with twenty-two frilled
papillae along each border. Only two short, wide, and glandular gastric caeca pass
forwards into the peripharyngeal space. Scales eighteen pairs, large, soft, and with a
frilled outer border, not covering the dorsum. Dorsal division of the foot minute, with
slender, simple, and finely spinous bristles. Ventral division rather long, with numerous
somewhat slender bristles with tips of varying breadths.

Halosydna gelatinosa, M. Sars i 1860. Plate XXV, fig. 5.

Specific Characters. — Head ovoid, pinkish, running into the bases of the lateral
tentacles. Transverse diameter greatest. Eyes large, furnished with lenses, close
together, the larger anterior pair on the lateral prominence, and the posterior close
behind. The median tentacle is long, smooth, and tapering, with a filiform tip, and the
lateral tentacles are also long. The tentacular cirri agree with the median tentacle in
length and structure. The palpi are of moderate length, and smooth. Most of these
organs are tinted pale madder-brown. Body elongated, bristled segments forty-
three, barred transversely with pale greyish-brown belts between the pedicles for
the scales. The nuchal collar has a prominent flap, which covers the posterior part of
the head. Segmental eminence distinct, and the papilla long. The proboscis is charac-
terised by about twenty-two frilled papillae along each border in extrusion, and a
muscular fold at each side. Scales eighteen pairs, not covering the dorsum completely,
large, soft, and rounded, with a folded or frilled outer border in spirit. The outer region
is studded with minute trifid papillae, but the margin is quite smooth throughout. The
dorsal division of the foot is minute, with slender simple and finely spinous bristles. The

1 Kinberg, in his description of the genus, gives little to discriminate it from Lepidonotus except
the numerous elytra and the elongated body.

HALOSYDNA GELATINOSA. 385

ventral division is rather long, and has a long cone for the spine superiorly. The bristles
are somewhat slender, with the spinous region of varying breadth, the superior simple
and long, the middle and inferior broad and bifid. Dorsal cirri rather long, slender,
and smooth, with a slight swelling (and a dark belt) below the filiform tip. The ventral
cirrus does not reach the adjoining tip of the fleshy part of the foot.

Synonyms.

1820. Polynoe foliosa, Savigny. Syst. des Ann., p. 23.
1826. Polinoe „ Risso. L'Europ. Merid., 414.

1834. Polynoe „ Audouin and Edwards. AnneL, 89.

1835. „ gelatinosa, Sars. Beskr. og Iagtt, p. 63, Tab. 9, fig. 25.
1851. „ foliosa, Grube. Fam. d. Ann., 37.

1853. Aphrodita cirrosa, Dalyell. Powers Creator, ii, 164, pi. xxiv, f. 1 and 2.

1858. Halosydna gelatinosa, Kinberg. Freg. Eugen. Eesa, Zool., p. 19, Tab. 5, fig. 26.

1860. Polynoe gelatinosa, Sars. Christ. Yid. Selsk. Forhandl., 1860, p. 58.

1865. „ „ De Quatrefages. Ann., i, p. 249.

„ Lepidonotus imbricatus, Baird. Johnston's Cat. Brit. Mns., p. 340.
„ Alentia gelatinosa, Mahngren. Nord. Hafs.- Annul., 81.

1866. Halosydna {Alentia) Jeffreysii, Ray Lankester. Trans. Linn. Soc, 25, p. 377, pi. Ii, figs. 12, 19

—21, 26, 27.

1867. Alentia gelatinosa. Mahngren. Ann. Polychset., 14.

1869. Halosydna gelatinosa, Mcintosh. Trans. R. S. E., 25, p. 408, pi. xv, f. 6.

1870. Polynoe foliosa, Grrube. Archiv f. Naturges., 1870, p. 288.

1875. Alentia gelatinosa, Ehlers. Annel. 'Porcup./ op. cit., p. 34.

„ Halosydna gelatinosa, Mcintosh. Invert, and Fishes St. A., p. 117.

1876. „ „ idem. Trans. Z. S., ix, p. 388.
1879. „ „ Tanber. Ann. Danic, 82.
1883. Alentia gelatinosa, Levinsen. Nord. Annul., 196.

1886. „ „ Langerhans. Zeit. f. w. Zool., 40, p. 251, Taf. 15, f. 6.

1888. Halosydna gelatinosa, De St. -Joseph. Ann. d. sc. nat. (7), v, p. 154, pi. vi, f. 6 — 21.

1891. ,, {Alentia) gelatinosa, Hornell. Op. cit., 237.

Habitat. — Everywhere distributed round British shores — from Shetland in the north
to the Channel Islands in the south. It is generally found under stones and in crevices,
in laminarian roots between tide-marks, or in the valves of old shells in deeper water. It
ranges also to the Scandinavian coasts, as well as to Madeira. A small specimen, 13'5
mm. long, is described by Ehlers (1875) from the 'Porcupine ' Expedition of 1869, from
the great depth of 1366 fathoms, on a bottom of fine mud. This, so far as my experience
goes, is a rare habitat, and no station or date is given — only the position of 54° 54' N".
and 10° 59' W.

Head (Plate XXVIII, fig. 11) with the transverse exceeding the antero-posterior
diameter, so that it forms an ovoid. Anteriorly the prominent base of the median
tentacle takes origin between the lateral lobes, and thus well within the anterior margin,
while the bases of the lateral tentacles are continuations of the head, no peaks being
present. The eyes are proportionally large and prominent, and have lenses. The larger
anterior pair occupy the projecting lateral region ; the smaller posterior pair are situated

386 HALOSYDNA GELATINOSA.

immediately behind, and, from the rapid narrowing of the region, are thus nearer each
other. In an example of medium size from Bressay Sound in July the eyes were larger
than usual, and those of each side in contact. The lenses were also larger. The
reproductive elements did not appear to be much developed. Sir J. Dalyell mentions
that a large one, also from Shetland, had apparently only two eyes, but it is possible
they were confluent. In life the head is pinkish, and in the preparations iridescent.
In the majority the head is prettily mapped out by a pale median belt running forward
to the tentacle. The prominent lateral regions on which the eyes are situated are pale ;
while in front are two pinkish or brownish-pink smoothly rounded areas — indicating
the brain. The median tentacle is somewhat long and tapering, with a filiform tip. In
spirit it is slightly enlarged below the latter. The surface is quite smooth. The lateral
tentacles, the bases of which are lower than the median, are remarkable for their length,
being only a little shorter than the median, 1 and this is confirmatory of the nomenclature
adopted in the group. They are tinted brownish. The tentacular cirri agree with the
median tentacle in structure and length. The palpi are of moderate length, brownish,
tapering, and quite smooth.

Body upwards of 2 inches, sometimes 3 J inches in length, slightly tapered in
front and much more distinctly tapered posteriorly ; rounded dorsally and flattened
ventrally. The dorsum is somewhat regularly banded across with greyish or greyish-
brown granular belts, and occasionally touches of white occur in the middle line of the
coloured belts. These alternate with a pale belt between the pedicles of the scales,
Between each pale belt there is thus a central band bordered with pale lines — extending
between the papilla on the cirrigerous foot, and a belt in front and behind, each of
which extends to the neighbouring segment. The segment-junction is, therefore, in the
middle of each band. No pale band occurs opposite the first pair of pedicles, but the
second and third, as well as the succeeding, have them. In some the pale belts are
more or less invaded by pigment. The caudal region behind the scales is continuously
grey. In front the nuchal collar forms a free flap, which is sometimes bifid and crenate,
but it is not papillose. The under surface is pale or slightly yellowish and iridescent.
The proboscis in many tints, the anterior region to about the sixth foot, the rest being
iridescent bluish with a red line in the centre. The segmental eminence is distinct,
and the papilla is evident at the fifth bristled foot, and continues to the posterior end
of the body. It is longest and largest about the beginning of the posterior third.

The proboscis and digestive system differ from that of the previous genera in having
a muscular fold at each side in extrusion immediately below the angle, and divisible into
two halves, so that a doubly papillose aspect is given in certain views. Moreover all the
papillas at the tip of the extruded organ are more or less lobed. The six central are
large, and when viewed from the exterior do not differ very much from the ordinary
forms, except that they are more slender. On their inner aspect, however, they send off a
frill sloping towards the aperture. The succeeding eight on each side form a diminishing
series with frilled and lobed ends. The total number of papillas is thus twenty-two.
Each of the upper sharp horny jaws bites to the left of the corresponding ventral in

1 Kinberg states that the antennaa are only about half the length of the palpi, and that the scales
are rugose.

HALOSYDNA GELATINOSA. 387

extrusion. Both are sharply ridged. A comparatively large number of the preparations,
including those from the stomachs of fishes, have the proboscis extruded.

Only two gastric casca, proportionally short and wide, pass forwards into the peri-
pharyngeal space. They are glandular throughout. The tip seems to end in a series of
diverging fibrous processes which fix it in the dorso-lateral space. The glandular lining
of the stomach is arranged in a series of trumpet-like processes projecting inwards.

The alimentary canal is generally empty, but in some it contains muddy debris with
spicules of sponges and other organic materials, together with the skin, bristles, and
hooks of Terebella3. De St. -Joseph says that it feeds on minute crustaceans. Sir J.
Daly ell gave a graphic account of its rapacious habits.

The scales (Plate XXXIII, fig. 12) are eighteen pairs, large — but do not quite cover
the dorsum, have a translucent greyish or slightly purplish aspect, or translucent
brownish anteriorly, and dull bluish or greyish posteriorly. A pale spot in each marks
the scar for attachment. In some of the anterior scales a few dark specks occur along
the posterior border. They are soft and more or less irregularly rounded, the first pair
having a marked fold at their outer and anterior border. They are devoid of cilia at
the margin, and present a minutely cellular aspect by transmitted light. Moreover
along the inner and anterior border the surface is densely studded with minute papillae,
which are short and broad — generally with a trifid end, two stronger spines laterally,
and a smaller and more acute median, as Langerhans pointed out. They extend over a
considerable surface of the region indicated. Most of the scales have frills or folds on
their outer margin. The finely branched nerves radiate from the scar of attachment
throughout the entire scale. They are highly sensitive, so much so that when irritated
the annelid will sometimes turn on its back to avoid interference. The scales are fixed
to the following bristled feet: 1, 3, 4, 6, 8, and so on to 22, 25, 28, 31, 34, 37, and 38.

A curious solid rounded body of minute size occurred in the substance of one scale.
Its contents were granular. In some also minute opaque white specks were present
behind the scar.

Feet. — The first foot has a spine, the tip of which projects beyond the region in a
sheath of cuticle and granular epiderm. No trace of a bristle was present in any
specimen examined, and in this it agrees with Malmgrenia.

In the second foot the dorsal division is marked chiefly by the tip of the spine,
which projects a short distance from the dorsum of the long ventral division. Behind
the spine is a small tuft of straight, slender, tapering bristles — finely spinous. The
combined lobe is comparatively long, nearly cylindrical, and ends in a long conical
process for the spine. The ventral bristles are short and slender, extending beyond the
conical process at the tip of the foot. They are slightly enlarged at the commencement
of the spinous region, and then taper to a hair-like extremity. Their tips, which have a
dorsal curve, diminish in length from above downward. The ventral cirrus of this foot
extends as far as the tips of the bristles.

In the third foot the dorsal division is marked only by the projection of the long
process enclosing the tip of the spine, and a tuft of similar bristles to those in the
previous foot, only the tips are less acute. The ventral division is proportionally
shorter and thicker, and the upper border of the process for the spine is now continuous

388 HALOSYDNA GELATINOSA.

with the dorsal margin of the foot. The bristles are somewhat stronger and the spinous
regions shorter. The tips of the upper series are less acute, and those of the stouter
inferior show traces of the expansion. The backward curve of the lower series is
marked. The ventral cirrus reaches only to the tip of the adjoining fleshy part of the
foot.

In the next (fourth) foot a slight elevation internal to the long sheath for the spine
exists, as an indication of the dorsal division. The tips of the ventral bristles are now
shorter and wider.

In the typical foot (Plate XXX, fig. 11) the dorsal division forms a small process
on the upper and anterior border, and is best seen in the elytrophorous feet, since the
enlarged base of the dorsal cirrus in the others obscures it. The inconspicuous tuft
consists of slender translucent bristles with faint serrations, and tapering to a fine point
(Plate XLI, fig. 1). The bristles of the ventral series are comparatively slender and
translucent, while the tips are flattened out in varying degrees. The superior series
(Plate XLI, fig. 2) have elongated spinous regions, and simple, slightly blunt tips ; while
the broader middle series (Plate XLI, fig. 3) have a strong curved hook at the tip, with
a secondary process immediately beneath. The flattened spinous region is marked by
the oblique lines from the rows of spines as in Malmgrenia. Langerhans specially refers
to the changes in the bifid tip of the ventral bristles. The ventral cirrus does not now
reach the tip of the fleshy part of the foot.

Posteriorly the bristles become extremely slender, with a capillary tip. The ventral,
however, retain a trace of the enlargement at the base of the spinous region. The
bristles on the last foot are stout. The ventral cirrus extends almost as far as the tip of
the inferior division of the foot.

Parasites on the bristles are rare.

The dorsal cirri are rather long, slender organs, with a filiform tip, and a dark belt
at the slight swelling immediately below. Their surface is smooth. A considerable
nerve passes up the centre of the organ nearly to the tip, giving off' branches as it
proceeds.

Habits. — This species is fond of clinging to the under surface of stones in pools near
low-water mark, from the north of Scotland to the Channel Islands, and no marked
difference in size is found on contrasting specimens from the extremes. In Bressay
Sound it is very abundant in old bivalve shells, in crevices in the "roots" of the
tangles, and on the surfaces of the valves of the " horse "-mussels bound together by
the foregoing " roots." Sir J. Dalyell many years previously procured it from this
region — under shells. He gives an interesting account of the rapacity of one in con-
finement which devoured its fellow, the teeth of the proboscis audibly striking the
glass as it darted it out to conclude its meal. In various Polynoida3 this tapping on
the glass occurs occasionally in confinement. Fishes seem to find it out readily in
deep water, for large examples are frequent at St. Andrews in the stomach of the cod
and haddock.

It is active and irritable, and frequently ruptures its body if held by the middle, or
when put in spirit. Sir J. Dalyell found it timid and nocturnal, yet watchful for prey.

Reproduction. — Eipe males occur at St. Andrews towards the end of November. A

POLYXOE SCOLOPEXDPINA. 389

large female, again, was laden with fairly advanced ova in July in Herm, so that the
period of deposition could not be distant, probably in autumn. They were somewhat
less advanced in a large female during the same month in Shetland, no free ova occurring
in either case. In August the condition of the ova was very similar to the last (e. g. in
one sent from Arran by Dr. Howden). Sir J. Dalyell, again, found that in the middle of
July a large specimen produced in confinement about 10,000 eggs, the mass having a
reddish hue as it lay on the bottom of the vessel. De St. -Joseph found ripe specimens
with the ova and sperms within a membrane, as shown by Claparede in Hermadion
fragile, but he omits to mention the date.

De Quatrefages adds nothing new to the account of the species. Grrube found that
this author's Polynoe foliosa in the museum at Paris agreed with the present species, as
Malmgren had hinted.

Ehlers gives a somewhat detailed description of the small specimen (13*5 mm.) from
the ' Porcupine. 5

Genus XX. — Polynoe, Savigny, 1820.

Head elongated antero-posteriorly, with a somewhat indistinct peak on each side,
beneath which is the lateral tentacle. Anterior and posterior eyes widely separated.
Palpi with dense rows of clavate papillae. Body linear-elongate, with many segments
(over 100). Segmental papilla long. Scales fifteen pairs, with a broad belt of papillae
on the surface anteriorly. Dorsal division of the foot minute, with small spinous bristles
truncated at the tip. Ventral division has a single strong hastate bristle superiorly, and,
below, strong bifid bristles with short spinous regions.

The median area for the nerve-cords is comparatively wide, the oblique muscles
having a broad attachment to its exterior border. A firm membrane bounds the area
internally, and a symmetrical longitudinal muscle runs above it on each side of the
median line.

Commensalistic.

Polynoe scolopendeina, Savigny, 1820. Plate XXV, f[g. 7.

Specific Characters. — Head pinkish, with the long diameter antero-posterior ; a
somewhat indistinct peak on each side. A pair of distinct eyes in front of the nuchal
collar, and a larger pair with lenses, scarcely visible from the dorsum, near the anterior
part of the peak, and directed forward and slightly outward. Median tentacle larger
than the palpi in spirit, with a slight enlargement below the filiform tip, and densely
clothed with short clavate cilia having truncated tips. Lateral tentacles of moderate
length. Palpi have dense rows of minute clavate papillae. Body linear-elongate, 2 to 5
inches long, bristled segments from 80 to over 100, reddish brown, or brown on the
dorsum, the pigment being arranged in various touches and bars. The posterior ventral
region has also brown bars and touches, especially on the segmental eminences. The

390 POLYNOE SCOLOPENDRINA.

papilla of the latter are long. Scales fifteen pairs, dappled brownish, with a dark patch
over the scar and surrounded by a pale ring. The larger examples have a dark metallic
lustre from the pigment. They have a series of minute papilla? as a broad belt along the
anterior region, and the outer edge shows very minute rounded or short clavate papilla?
— the homologues of the cilia. The dorsal division of the foot is minute, and bears
rather small, spinous, truncated bristles. Ventrally, as a rule, a single strong hastate
bristle occurs superiorly, and, below, a series of strong bifid bristles with short spinous
regions. The dorsal cirri are somewhat short, with a filiform tip, and with the same
short cilia as on the median tentacle. The ventral cirri are short, ciliated, and tapering
— not reaching the tip of the fleshy part of the foot in spirit. Commensalistic.

Synonyms.

1820. Polynoe scolopendrina, Savigny. Syst. des Aim., p. 350.
1834. „ „ Aud. and Ed. Annel., p. 92, pi. i, f. 17 and 19.

1840. „ „ Johnston. Ann. Nat. Hist., v, 307, Tab. 5, f. 1—9.

1851. „ „ Grrube. Fam. Annel., 37.

Sars. Yid. Selsk. Forh., 1860, p. 62.

„ ,, variegata, Grrube. Annel. Oersted., 49.

1860. „ scolopendrina, Sars. Yidensk. Selskab. Christ., 1860, p. 62.
1865. „ „ Johnston. Cat. B. M., 119, pi. xi, f. 1—9.

Malmgren. Nord. Hafs-Ann., 82, Tab. 10, f. 11 ; and Ann. Polycli., 15.

„ Lepidonotus scolopendrinus , De Quatrefages. Ann., i, p. 263.

1873. Polynoe scolopendrina, Marenzeller. Sitzb. der k. Akad., vol. lxix, p. 419.
„ „ crassipalpa, idem. Ibid., p. 412, pi. xi, f. 1.

„ „ Johnstoni, idem. Ibid., p. 420.

1874. „ „ idem. Sitzb. d. k. Akad., 69 (sep. Ahd.), p. 14.

1875. „ scolopendrina, Grrube. Jalirb. Schles. Gresells. {fide De St.-Joseph), 1875 Breslau,

.1876, pp. 53 and 68.

1876. „ „ Mcintosh. Trans. Z. S., ix, 389.
1879. „ „ Tauber. Ann. Danic., 82.
1883. „ „ Levinsen. Nord. Annul., 196.

1888. „ „ De St.-Joseph. Ann. d. so. nat. (7), v, p. 183.

1891. „ „ Hornell. Op. cit., p. 235.

Habitat. — In the tubes of Terebella nebulosa, between tide-marks, Herm, and
between the chinks of rocks (gneiss) in muddy sand in the burrows of Lysidice. The
large race abounds on the eastern shores of North Uist in the tubes of Terebella nebulosa
and other Terebellids attached to the under surfaces of stones close to low- water mark.
Not uncommon also off the east and west coast of Ireland (Prof. Haddon and Dr. Scharff)
and in the English Channel (Hornell). It extends to the shores of France and to the
Mediterranean.

The pinkish head (Plate XXYIII, fig. 9) forms an ovoid, the long diameter being
antero-posterior. A central groove terminates in front at the basal process (ceratophore)
of the median tentacle, and on each side is a somewhat indistinct peak, which, however,

POLYNOE SCOLOPENDEINA. 391

is differentiated from the lateral tentacle beneath. A pair of distinct eyes lie in front of
the nuchal collar. The considerably larger anterior pair are scarcely visible from the
dorsum, and are situated in the smaller southern forms on the outer border near the end
of the peaks, a long interval thus existing between the anterior and the posterior pairs.
In the large Hebridean examples these large eyes occupy almost the whole peak, and their
direction is more forward than outward. The anterior eyes, as a rule, look forward and
slightly outward, and are very conspicuous from the front. A minute lens, less distinct
in the larger than in the smaller forms, appears in the centre of the anterior pair. In
the preparations the brownish median tentacle is longer than the palpi, tapering from
the base to the slight swelling below the filiform tip. It is somewhat densely clothed
with short cilia, most of which have truncated tips, only a few being clavate — the
latter shape being due to an ovate rather than a rounded tip. The lateral tentacles
are of moderate length, and have a slight enlargement below the filiform tip. They
are similarly clothed with truncated cilia. The palpi are of average length, and have
rows of closely arranged clavate papillae with truncated tips. These are longest and
most cylindrical inferiorly, shorter and more markedly clavate superiorly. Smaller
forms extend on the tapering extremity of the organ. The tentacular cirri resemble
the median tentacle in regard to form and cilia.

Body linear-elongate, If to 4| inches or even more long, and having from 80 to
102 bristled segments. The iridescent dorsum in the larger examples is mottled with
reddish brown, which, behind the proboscidean region, is somewhat regularly arranged
between the lateral eminences, and by-and-by divided into two by the dark median
tubercle which commences about the twentieth bristled foot. These three distinct sets
of elevations (two lateral and a median) continue to the posterior end of the body. A
ridge-like fold forms a buttress in front and another behind the lateral tubercle, and
each bounds the seal-like brownish mottling. Brownish pigment also occurs on the
elevations at the bases of the dorsal cirri. The median tubercle is situated towards the
anterior part of the segment, the lateral towards the posterior part. In the large forms
the reddish hue of the dorsum shines through the scales, and tints the entire region,
relieved by the steel-like glitter of the darkly pigmented portions of the scales. In the
smaller forms the dull yellowish back is minutely flecked with brown. Every alternate
foot in front has a dark brown patch on the eminence for the cirrus. The ventral
surface is iridescent, pale brown or dull yellowish in the specimens from the Channel
Islands, and posteriorly marked by touches of brown — chiefly on the segmental
eminences, which are distinct, and have a large and somewhat clavate papilla which
projects backwards between the feet. The segmental papillas become distinct on the
sixth foot, and continue nearly to the posterior end. In the centre of the body is a
reddish streak. In the large Hebridean forms the under surface and feet are of a dark
orange — with the reddish central line — and iridescent. Moreover, in some, broad bars
of pigment occur on the ventral surface throughout in more than the posterior half. Two
races thus occur, the smaller yellowish-brown southern form, and the reddish-brown
Hebridean.

The body terminates posteriorly in two anal styles, which, like the cirri, are pro-
portionally short.

51

392 POLYXOE SCOLOPENDRINA.

Digestive System. — The pre-gastric caeca are two in number, both being short and
broadly club-shaped. The third is transverse. All go deeply into the intermuscular
spaces at the feet. The movements of muscles and feet must thus have considerable
effect on the contents of the caeca.

Scales (Plate XXXIII, figs. 13 — small example, and 14 — larger example) fifteen
pairs. They vary in appearance according to the condition of the specimen; thus the
smaller race from the Channel Islands has the anterior scales dappled brownish with a
dark patch over the scar for attachment, and surrounded by a broad pale ring. In the
larger specimens from the Hebrides they have a dark metallic (steel-like) iridescence.
When removed they are translucent, pale anteriorly, marked by a dark belt, speckled
with translucent spots round the inner three-fourths of their circumference, and with a
pale centre. They vary, in any given specimen, in size according to state of development,
covering the dorsum in some nearly as far as they extend, whereas in others with
developing scales an interval occurs between the pairs. In the large forms the first five
pairs cover the dorsum completely, the rest leave a space in the centre. The first pair
are rounded — with minute cilia anteriorly, the succeeding somewhat reniform, and the rest
more or less rounded ovoid. Though smooth under a lens they have a dense series of
minute papillae as a broad belt along the anterior region, and at the outer border a series
of minute papillae or cilia, and one or two also occur along the posterior border. Finely
branched nerves ramify throughout the scale from the scar for the pedicle. In the small
specimens from the Channel Islands the papillae (spines) on the scales are proportionally
large, as observed in the sketch.

Feet. — The dorsal division of the first foot presents a single small dorsal bristle with
about seven or eight spinous rows, and a smooth tip.

The second foot has dorsally a group of somewhat tapered, short, slightly curved
bristles with well-marked spinous rows. Ventrally the slender bristles are also short,
the spinous regions being bent dorsally, and tapering to smooth bifid tips. The rows of
spines are well marked, — that is, are at moderate distances from each other. The tips of
one or two of the inferior bristles are simple. The large ventral cirrus of this foot has
numerous cilia with clavate and truncate tips. The bristles in the succeeding feet gradu-
ally approach the typical form, which is found about the twelfth or thirteenth bristled
foot. In shape the typical foot (Plate XXIX, fig. 17) presents dorsally the comparatively
short tapering cirrus, then the eminence for the dorsal bristles. The inferior division is
bifid, with a long anterior process and a shorter posterior cushion, the margin beneath
having an inward slope from above downward. Even in the second foot the upper
ventral bristle is stouter than the others, with strongly bifid tip and spinous rows. It
increases in strength and becomes more boldly bifid as we proceed backward, attaining a
large size in the fifth and sixth, and then becoming simple about the tenth bristled foot,
while only traces of the spines remain. As a rule only one strong bristle occurs in the
typical foot, which has below the dorsal cirrus thQ tuft of short, slightly curved, and little
tapered dorsal bristles arising from a small eminence (Plate XLI, fig. 4). These bristles
taper a little from the base, but end in a blunt tip (Plate XLI, fig. 7), which is curiously
wrinkled so as to appear bifid. The spinous rows are somewhat close, yet fairly distinct.
A careful examination of various specimens, however, shows that this condition of the

POLYXOE SCOLOPENDRINA. 393

dorsal bristles is due to external influences, probably connected with commensalism.
The tip has evidently been destroyed, so that only the lower part of the bristle remains.
The stump, indeed, is often slightly thickened, and presents a furrow, giving the bifid
appearance already adverted to. In an example dredged off the Hebrides by Dr. Grwyn
Jeffreys the dorsal bristles are unusually complete, yet even in these the extreme tip
has disappeared and a probe-pointed end remains. This specimen shows, however,
that the normal condition of such a bristle is finely tapered to a filiform tip. The causes
of this remarkable change, which does not seem to affect the ventral bristles, may be the
secretions of the Terebella, or the friction of the bristles against the walls of the tube or
tunnel, the latter the most probable.

The ventral division forms a short triangular lobe, the process for the spine being at
the apex. In some a bar of dark pigment occurs near the tip of the foot, just at the
base of the bifid region. Above the spine is the single large bristle, the shaft of which
is about twice the diameter of those adjoining. The tip is hastate and the point acute
(Plate XLI, fig. 5). As already indicated, this bristle is formed from a bifid one, the
secondary process disappearing as we proceed backward, and likewise the spines— traces
of which, however, even far backwards, are left in the faint transverse lines on the
dilated region. The relative proportion of the two kinds of bristles is shown in figs. 4
and 5. All the bristles which follow are bifid, with short spinous regions and moderately
stout shafts (Plate XLI, fig. 6, representing one of the superior forms with a longer tip,
and fig. 8 showing the tip more highly magnified).

Toward the posterior end the large superior ventral bristle disappears. Before this
change takes place, however, two strong bristles occur on the foot as in front. The
dorsal tuft becomes minute posteriorly, and in the terminal feet the bristles are slender
and tapered — with very distinct spinous rows.

The dorsal cirri, which in life are pale, extend in the preparations only a little beyond
the bristles, and gently taper to the slight swelling below the filiform tip. They have
numerous short cilia with clavate and truncate ends, and a few small cilia occur on the
base of the slender extremity. The ventral cirri are small and tapering, wdth similar
cilia. They do not reach the tip of the fleshy part of the foot. In some they are
rendered conspicuous by dark pigment at the base.

Nerve-cords. — In transverse section nucleated cells are found externally and beneath
the cords, especially at the ganglionic enlargements. Strands from the exterior fibrous
layer also passed into the area of the cord in section, and in many such fibres seemed to
join the nucleated cells lying in the median line below the cords.

Reproduction.— Females with well-developed eggs were found in the tube of Terebella
nebulosa at Herm in July, 1868, and in the chinks of rocks in tracks of Lysidice. In the
latter case males well advanced were also observed.

Habits.— This form is more sluggish in its habits than Harmothoe imbricata or other
ordinary type. It is difficult also to conceive how the Terebella can be comfortable with
such a large commensalistic form in its tube, especially when the powerful bristles
of the Polynoe are considered. So far as observed it keeps its head towards the anterior
end of the tube, and thus its ordinary position is in agreement with that of the Terebella.
It is brittle, breaking into several pieces in lifting from one vessel to another.

3W POLYNOE SCOLOPEKDBJNA.

Its food consists of cellular substances with a few sponge-spicules.

It is phosphorescent. On placing it in spirit luminous flashes were emitted from
the bases of the feet, and the same emissions were caused by irritating the posterior end
with the forceps.

The remarkable difference in size and coloration between the southern forms from
the Channel Islands and those from the Outer Hebrides is an interesting feature, but one
in the British Museum from Falmouth is between four and five inches, so that caution in
making deductions is necessary.

The most marked variety is that from the chinks of the hard gneiss of the Channel
Islands— in the burrows of Lysidice. It is less than two inches, very narrow-, and there
is a tendency to have dark brown pigment at the mouth and anterior ventral region.
It thus approached in bulk Lysidice itself, and the tints anteriorly were not dissimilar.

The species was first found by d'Orbigny at La Eochelle, and by Savigny and
Audouin and Edwards on the shores of the Channel. The latter authors state that
it lodges in tubes of sand and shells agglutinated by a secretion, though they also
found it in company with Terebella. No instance of its occurrence in an independent
tube, manufactured by itself, is known in this country, and it is possible Audouin and
Milne Edwards may have met with it only in an empty tube of Terebella.

De Quatrefages points out that Johnston's species diverges from that of Audouin
and Milne Edwards, since it has 110 segments. Further, that the appendages of the
head differ in proportion. Thus the median antenna is much longer than the inferior
tentacles, which are very large and conical; the lateral, again, are proportionally
small. The superior tentacles differ correspondingly. For the rest, the figures of the
bristles given by the respective authors diverge much. This author, however, laboured
under a misapprehension on the subject.

Marenzeller in 1874 distinguished this from Savigny's species by the fact that the
tentacle was longer than the palpi, while the tentacular cirri were shorter than the palpi :
there are three rows of wart-like papillas on the dorsum of the segments. Moreover, his
P. crassipalpa comes near it, and may be the same form. I have not seen any reason to
think that Savigny's species differed from Johnston's, though two varieties exist ; and it is
satisfactory to find that Baron de St.- Joseph agrees with me in this respect as well as with
regard to the P. crassipalpa of the able naturalist of Vienna. Mr. Hornell observes that
nearly every haul of the dredge off Anglesey brought up at least one specimen, and in
one case it emerged from the tube of Thelepus cineinnatus.

In the recent remarks 1 of Dr. H. F. Johnson on the Pacific annelids it would have
been very interesting to find the results of a comparison of such commensalistic
species as Polynoe reticulata, sp. nov., or P. gigas, sp. nov., with the well-known
P. scolopendrina.

1 ' Annel. of the Pacific Coast/ Californ. Acad. Ser., 3rd ser., i, No. 5, 1896, p. 170 et seq.

ENIPO KINBEEGI. 395

Genus XXI. — Enipo, Malmgren, 1865.

Head subcircular without peaks. Eyes rather small, two posterior, two median and
lateral. Palpi thick, subulate, and, like all the tentacles, smooth.

Body much elongated (about 100 segments). Segmental papillae distinct. All the
segments have dorsal cirri.

Scales fifteen pairs, minute, smooth, subcircular, only occurring anteriorly.

Feet well marked and long. Dorsally is a minute tuft of slender serrate bristles.
Ventrally are slender bristles with tapering, boldly spinous regions and a capillary tip,
while a few are bifid.

Enipo Kinbekgi, Malmgren, 1865.

Specific Characters. — Head subcircular without peaks. Eyes rather small, two at
the posterior border and two on the lateral prominence. Lateral tentacles subulate,
short. Palpi thick, subulate. Tentacular cirri of moderate length. All these appendages
are smooth. Body much elongated, of about 100 segments. Segmental papilla distinct.
Scales fifteen pairs, minute, subcircular, pellucid, smooth, leaving the posterior part of
the body uncovered. Eeet well marked, long, bearing dorsally a minute tuft of slender
serrate bristles. Ventral bristles mostly slender, with a tapering boldly spinous region
ending in a capillary extremity, but a few have hooked tips with a secondary process. All
the segments have dorsal cirri, which reach only a little further than the bristles. Ventral
cirrus subulate, of moderate length — sparsely ciliated (fide Malmgren).

Synonyms.

1865. Enipo Kinhergi, Malmgren. Xord. Hafs-Ann., p. 83, Tab. x, f. 12.

1867. „ „ idem. Ann. Polych., p. 15 (sep. copy).

1873. „ „ Kupffer. Jahresb. der Com. der Deutsch. Meere, 1871, p. 150.

1875. „ „ Mcintosh. Invert, and Fishes, St. A., p. 116.

1876. „ „ idem. Trans. Z. S., ix, 388, pi. lxx, f. 7—10.
1879. „ „ Tauber. Ann. Danic, 83.

1883. „ „ Levinsen. Xord. Annul., 196.

Habitat. — Deep water off St. Andrews Bay. The examples have occurred in the
stomachs of cod and haddock (E. M.). It ranges to Christiania Fjord, to Drobak, to
Bahusia (Malmgren), and the Baltic.

Head. — The only British specimens hitherto procured have been from the stomachs
of cod and haddock caught off St. Andrews Bay, and the head has been so injured that
no description could be made. Malmgren observes that the head is subcircular, without
anterior peaks, the base of the median tentacle occupying the region. Eyes apparently

396 ACHOLOE.

rather small ; two in front of the nuchal collar, and two on the lateral cephalic promi-
nence. Lateral tentacles short, subulate. Palpi thick, subulate. Tentacular cirri about
the length of the palpi in spirit. All these cephalic appendages are smooth.

Body. — Much elongated, narrow, of about 100 segments, and about 65 mm. long.
Segmental papilla distinct (fide Malmgren).

Scales. — Fifteen pairs, minute, subcircular, pellucid, smooth, leaving the posterior
region of the body uncovered (fide Malmgren).

Feet. — The feet are rather elongate, carrying dorsally a minute tuft of slender hairs,
finely serrated (Plate XLI, fig. 9). The ventral bristles are described by Malmgren as
of one kind only, but this form shows two kinds, viz. (1) that indicated by Malmgren, and
represented in Plate XLI, fig. 10, the rows of spines in all the examples being apparently
less numerous and wider apart than in Malmgren's figure ; and (2) a few with character-
istically hooked tips and a secondary process beneath (Plate XLI, fig. 11). The rows of
spines in the first series are not opposite, but alternate — as observed in end views
(Plate XLI, fig. 12). A slight enlargement occurs at the commencement of the spinous
region.

All the bristled segments, as in P. scolopendrina, are furnished with smooth dorsal
cirri, which extend only a little further than the bristles. Ventral cirrus subulate, of
moderate length, sparsely ciliated.

The appearance of this northern form only in the stomachs of fishes shows how
readily such may escape the various instruments of capture used by naturalists.

I have placed this under Malmgren's genus and species, supposing that he had
overlooked the ventral bristles with the bifid tip. Should they be absent in his form,
then the species from St. Andrews should bear the specific name of Elisabeths, from its
discoverer.

Kupffer (1873) shows that the antennas are three-ringed in examples from the
Baltic, but this and other features require careful re-investigation in connection with
possible specific differences.

Genus XXII. — Acholoe, 1 Claparede, 1870.

Head elongated from before backward, and running into the bases of the tentacles.
Xo peaks. Four large equidistant eyes. Palpi smooth and short. Body sublinear,
flattened, segments numerous. A segmental eminence but no distinct papilla. Cirri on
every foot, and a T-shaped branchial process. Scales numerous. Feet short. Dorsal
division minute ; bristles few and small, with a minute hook at the tip. Ventral division
bearing bristles with long and strong shafts and short spinous regions having a sharp
hook at the tip. The nerve-cords seem to be comparatively large, ovoid in section, and
have the cuticle and granular epiderm externally. Internally a firm membrane separates
them from a well-marked layer of longitudinal muscular fibres in the median line, as in
Po lyn oe scolopen drin a .

1 One of the Harpies.

ACHOLOE ASTEKICOLA. 397

Aoholoe astericola, Belle Ghiaje, 1823.

Specific Characters. — Head somewhat elongated from before backward, with four-
large equidistant eyes — all visible from the dorsum. Anterior peaks absent or indistinct,
the head running into the bases of the lateral tentacles. Median tentacle of moderate
length, subulate, with a very few clavate cilia. Similar cilia occur on the short lateral
tentacles. The palpi are smooth and rather short. Body sublinear, flattened, segments
numerous and of a peach-blossom or flesh colour. Segmental eminence, but no distinct
papilla. Scales forty-five pairs, rounded or ovate, smooth, with a very few minute
papillaB at the anterior and inner border. The feet are somewhat short, the dorsal
division minute, the ventral bevelled from below upward. In the cirrigerous feet a
T-shaped branchial process. Dorsal bristles few and small, gently curved and tapered,
with a minute hook at the tip, which is smooth. The spinous region is short, and the
spines minute. The ventral bristles have long and strong shafts, a slight dilatation at
the commencement of the very short spinous region, and a smooth boldly marked sharp
terminal hook. Dorsal cirri somewhat short, tapered, and with a very few clavate cilia.
The ventral cirri are short, smooth, subulate, and tapering.

Synonyms.

1823. Nereis squamosa, Delle Chiaje. Mem. s. g. Anim., ii, pp. 368, 400, 425, Tav. 19, f. 7.

1841. Polynoe astericola, idem. Descriz. e notom., v, 57, 106, Tav. 129, f. 7.

1855. „ malleata, Grube. Arcliiv f. Xaturges., Bd. xxi, 81, Taf. iii, f. 1.

1857. „ astericola, Sars. Bid. til Kundsk. om Middelhav. Littoral-Fauna, Christiania,

p. 104.
1865. „ asterinse, Carrington. Proceed. Lit. and Phil. Soc., Manchester, iv, p. 176.
1870. Acholoe astericola, Claparede. Suppl. Ann. Chet., Nap. (sep. copy), 18, pi. ii, f. 1.

1875. „ ,, Marenzeller. Sitzb. der k. Akad. (sep. Ahd.), 14.

„ „ ,, Panceri. Atti Accad. Sc. Napoli, vol. vii, p. 13, Tav. 3, f. 3 — 5.

1876. „ ,, Mcintosh. Trans. Z. Soc, ix, 389, pi. lxx, f. 11, 12.
1884. „ ,, Y. Cams. Fauna Medit., 202.

1891. „ „ Hornell. Op. cit., p. 236.

Habitat. — It was first procured in Britain by the late Dr. Carrington on Southport
Sands clinging to Astropecten irregularis ; while Prof. Percival Wright got it at Galway.
Abroad it ranges to the Mediterranean, occurring on various star-fishes of the genus
mentioned.

The head (Plate XXVII, fig. 17) is somewhat elongated from before backward in
the softened preparations, of a roseate hue from the ganglia, and Claparede shows four
large equidistant eyes — all visible from the dorsum. 1 The anterior peaks are absent or
very indistinct, the head on each side running into the base of the lateral tentacles, while
in front of the median groove is the base of the median tentacle. The trilobed processes

1 The eyes had disappeared from the rather softened preparations which represented the species.
For these I am indebted to the late Dr. Carrington, of Eccles, near Manchester.

398 ACHOLOE ASTERICQLA.

alluded to by Claparede seem to be the result of compression. The tentacle is subulate,
with a few clavate cilia. The lateral tentacles are short and subulate, with a fewclavate
cilia. The palpi are smooth, rather short (in spirit). The tentacular cirri are also short,
with a few clavate cilia. Claparede observed that the dorsal base of the latter has a
band of vibratile cilia on its upper surface, and he considered that such was the homo-
logue of the branchial processes of the feet.

Body sublinear, flattened, and ranging from one to two inches in length. It is
very little tapered in front — much more so posteriorly. The segmental eminence is
distinct, but no papilla is visible in the preparations. Dr. Carrington describes the
general aspect as peach-blossom or flesh coloured.

Scales (Plate XXXIII, fig. 15) are forty-five pairs (Claparede 1 ), rounded or
in front somewhat reniform, apparently smooth, but having a series of minute papilla
along the anterior and inner border, and with finely branched nerves. They are rather
thin and translucent, and have a blackish belt round the border, the centre being pale.
In some the blackish belt is confined to the posterior and outer border. On the
anterior aspect of the pedicles for the scales is a band of vibratile cilia.

Feet. — Only the spine remains in the preparations in the first foot, and Claparede
shows the same condition from life. In this respect it agrees with Malmgrenia
and Halosydna.

The second foot has a few short dorsal bristles — slightly curved, and with a short
spinous region, which does not quite reach the smooth and pointed tip. The ventral
series are stout and somewhat short, with a distinct curvature at the upper part of the
shaft, and a short spinous region tapering from a basal enlargement to an acute and
slightly hooked tip. The spinous rows are well marked. The chief change in the third
foot is the increase in the strength of the ventral bristles, and in the more distinct hook
of the smooth extremity. A ciliated band occurs along the upper and anterior edge in
this and other anterior feet without elytra.

The typical foot (Plate XXXI, fig. 4) in those segments which have cirri has a
T-shaped branchial process on the dorsum — figured by Claparede from life, and showing
a diverticulum of the intestine, and ova from the perivisceral chamber in the interior. Its
inferior surface is richly ciliated. Such a process may fairly be called branchial. The
foot is comparatively short, with a small dorsal papilla for the bristles, and a short ventral
division sloping from below upward to the spine, and slightly bifid at the tip — when viewed
from above. The dorsal bristles (Plate XLI, fig. 13) are short, somewhat curved, and little
tapered distally below the bare region at the tip, which is acute and slightly hooked.
The spinous rows are minute. The ventral bristles have long and strong shafts— the
middle exceeding the superior and inferior in bulk, a slight dilatation at the commence-
ment of the very short spinous region, and a smooth boldly marked sharp hook at the tip
(Plate XLI, fig. 14). The spinous rows gradually diminish from below upwards, and no
larger process occurs at the base of the hook, as in several forms.

The dorsal cirri are comparatively short, apparently devoid of an enlargement below
the tips, and with a few clavate cilia. The ventral cirri are smooth, subulate, and

1 Carrington says twenty pairs or more of white scales.

ACOETID^. 399

tapering. They do not reach the tip of the fleshy part of the foot (in spirit), though
they pass beyond the bases of the bristles.

Nerve-cords. — At the sides of the nerve-cords are numerous nucleated cells, and the
cords themselves in section present a peculiar areolar or reticulated appearance — a
feature, however, which may be due to the mode of preparation. The granular epidermic
layer seems to be largely developed.

Habits. — The late Dr. Carrington, of Eccles, found it at Southport in considerable
numbers, along with Harmothoe lunulata, in the ambulacra! grooves of Astropecten
irregularis, his attention having been attracted by the bluish phosphorescence when the
star-fishes were put in fresh water. It is very fragile, so that it is difficult to obtain an
entire example ; yet it is sluggish. Claparede, again, procured it on the same star-fish at
Naples in company with another Annelid, Stephania flexuosa.

Reproduction. — Dr. Carrington observed ova in his examples from Southport, but no
date is given, though it is probable that they were obtained during the storms of winter.
The ova were seen through the dorsal papillse. M. Claparede procured those with ova
and sperms at Naples in the winter of 1868-9.

Delle Chiaje first found this species in the ambulacral grooves of Astropecten
cmrantiacus and pent acanthus at Naples, and gives a figure of the entire animal, which is
of a pinkish hue.

Amongst allied forms, Webster, in his 'Annelids of the Virginian Coast,' describes
Lepidametria commensalis with scales extending the whole length of the body, viz. 50 to
90 mm., and living in the tube of Amphitrite ornata (Verrill). Moreover he also describes
a minute form (which he terms " Antinoe parasitica") parasitic under the scales. It
has peculiar hooks in the posterior segments (ventral series) — he thinks for holding on.

This species is apparently that referred to and figured by Delle Chiaje in 1823, but
his reference to the previous note by Baster could not be verified. It has been thought
best to retain the specific name astericola subsequently given to it by the Italian author,
the original one of squamosa being less characteristic.

Family IV.— ACOETIDiE, Kinberg, 1857.

Head without a facial tubercle. Median tentacle arising from the middle of the
anterior lobe ; bases of the lateral tentacles covered by the ocular peduncles. Palpi long
and tapering. Body elongate, flattened. Scales numerous. Scale-bearing segments
alternating with those bearing cirri. Pharynx exsertile, with numerous papilke on the
margin — the median dorsal and ventral tentaculiform. No pregastric caeca. A
segmental eminence, but no distinct papilla.

52

400 PANTHALIS (EKSTEDI.

Genus XXIII. — Panthalis, Kinberg, 1857.

Scales smooth, flat, rounded or campanulate, covering the dorsum anteriorly,
leaving the rest bare in the middle. Foot with a dorsal process in front, a trilobed
median and a small ventral process. Dorsal bristles brush-like at the tip ; median
spinous on one side, and terminating in a long whip ; ventral with curved tips — finely
tapered and spinous.

Panthalis (Erstedi, Kinberg, 1857.

Specific Characters. — Head somewhat urn -shaped, with the rounded bosses for the
eyes (ommatophores) in front, and the small median tentacle extending a little beyond
them. Posteriorly it runs smoothly into the body. The median tentacle springs some-
what behind the bases of the pinkish ocular peduncles. The lateral tentacles arise close
together beneath the ocular peduncles, and their tips extend further than the median.
The palpi are long, flesh-coloured, and tapering, with minute conical papillse towards
the extremity. The tentacular cirri are longer and stronger than the median tentacle, and
like the latter, smooth. Body 3J inches in length, tapering in front and more distinctly
posteriorly. The pearly white or flesh-coloured dorsum is finely streaked transversely.
Inferiorly a ventral ridge and then a median groove stretch from the shield-shaped area
behind the mouth. Proboscis cylindrical, with y§ papillse, the upper median forming a
long smooth tentacle, the inferior a shorter process with an arch at its base. Maxillse
elongate, with a strong dorsal ridge ending in a hook, and a flattened blade with six to
eight teeth. Scales thirty-nine pairs, rounded or campanulate, smooth, the first three
pairs covering the dorsum, the rest leaving the centre bare. Foot with a dorsal process
in front, a trilobed median and a small ventral process, and with two spines. The
dorsal bristles are long and brush-like at the tip (bipennato-penicillate, Kinberg), the
median stout and with short broad tips, spinous on one side, and terminating in a long
whip (aristate, Kbg.), and ventrally a series of long slender bristles with curved tips
finely tapered and spinous (subulato-serrate, Kbg.).

Synonyms.

1857. Panthalis (Erstedi, Kinberg. Eugen. Resa. Zool., ii, p. 25, Tab. vii, f. 34.

1859. „ „ D.anielssen. Norske Vid. Selsk., Skrifter., Bd. iv, Hefte 2, p. 115.

1861. ,, „ Sars. Nyt. Mag. Natur., xi, Hefte 3, p. 253.

„ „ „ idem. Forh. Skand. Naturf., viii, Kjobenh., 1860, p. 625.

1863. „ „ „ Nyt. Mag. Naturv., xii, Hefte 3, p. 628.

1863. „ „ „ Geol. og Zool. Reise, 1862, p. 46.

1865. „ „ Malmgren. Nord. Hafs.-Ann., p. 85.

1867. „ „ idem. Ann. Poly., 16.

1869. „ „ Mcintosh. Rept. Brit. Assoc., 1868, p. 338.

PANTHALIS CERSTBDL 401

1876. Panthalis (Erstedi, Mcintosh. Trans. Z. S., ix, 389, 404, 405.

1879. „ „ Tanber. Ann. Danic,, 83.

1883. „ i9 Levinsen. Xord. Annul.,, 197.

1893. „ }} Marenzeller. Polychaet. des Grnndes, 5, Taf. 1, f. 2.

1895. „ i} Arnold Watson. Trans. Liverpool Biol. Soc, ix, 1895, 169, pis. ix and x.

), ,', Lacaziij Pruvot and Eacovitza. Arch. Zool. exper., 3rd serie, hi, 428, pi. xix, f.

84—104.

» „ Marenzellerij idem. Ibid., 442, pi. xix, f. 105 ; and pi. xx, f. 106 — 110.

Habitat. — First dredged in Britain by Dr. Gwyn Jeffreys in 78 fathoms, 35 miles off
Skerries, Shetland, in 1868 ; 358 fathoms in the Atlantic (station 6) during the ' Por-
cupine ' Expedition of 1870 ; by the ' Triton ' at 516 fathoms ; and recently in the Irish
Sea by Prof. Herdman and Mr. Arnold Watson. It stretches to the Mediterranean,
medium-sized examples having been procured in 40 to 80 fathoms off Jigeli in Algiers,
in 40 to 80 fathoms during the 6 Porcupine ' Expedition of 1870 ; while Marenzeller
obtained it in the eastern part of that sea off Jaffa. It is also found off Norway and
Sweden. Its distribution is thus extensive.

The head (Plate XXVIII, fig. 16) is somewhat urn-shaped in the preparations, with
the rounded bosses for the eyes in front laterally, and the median tentacle in the centre,
then dilating from the bases of the former, and again narrowing posteriorly where it
runs into the body, without a nuchal collar. The smoothly rounded tips of the ocular
peduncles retain no pigment in the preparations; but in life, besides the ocular pigment,
the peduncles are reddish. Pruvot and Eacovitza consider them the homologues of the
lateral peaks of the Polynoidse — a supposition to which little objection can be taken,
since they occupy the whole anterior region of the head. The median tentacle arises
rather behind the bases of the ocular peduncles, and thus is behind the middle of the
entire region (including the ocular peduncles). Its basal region is short, while the
tentacle itself is a somewhat small subulate organ with a tapered tip which extends
beyond the peduncles. The lateral tentacles arise close together beneath the ocular
peduncles ; and their tips reach a little further than that of the median. The palpi are
flesh-coloured, long, tapering, and with minute conical papillse which commence a little
above the base and extend almost to the extremity, only a very short portion of the
filamentous tip being bare. They are very minute inferiorly, but increase in size distally.
Moreover each papilla has a basal granular region, and a clear tip like a jewel in its
setting. The tentacular cirri are longer and stronger than the median tentacle, though
much shorter than the palpi.

Body 3J inches and upwards in length, tapering a little in front and more distinctly
posteriorly. The dorsum is pearly white in front, flesh-coloured posteriorly, the red dorsal
blood-vessel enlivening the general hue of the finely and transversely striated integument.
Inferiorly is the deep median furrow and two lateral ridges, the former terminating in
front in a broad shield-shaped area. The ventral surface in life is opalescent, with a
golden shade towards the sides. Mr. Arnold Watson describes the posterior extremity as
being broadly forked from the backward direction of the last pair of feet. No complete
specimen, perhaps, has been seen. Anteriorly the segmental eminence is very slightly
indicated, but it forms a distinct elevation posteriorly. No papilla is visible.

402 PANTHALIS CERSTEDI.

Proboscis (Plate XXXIV, figs. 3 and 4) exsertile, cylindrical, famished with yf
papillge, the median — upper and lower— longer, especially the superior. In the British
forms it projects from the mouth like a tentacle, and tapers from base to apex. The
inferior, also conical, is much less, and has an inwardly projecting hood or arch at its
base. The four maxillse are elongate, brownish, strongly hooked at the tip, which is
a process of the dorsal rib, and with an attached flat blade, the inner edge of which has
six teeth (Kinberg says six to eight teeth, while Marenzeller shows in his figure traces
of six).

Xo pregastric cseca occur, and in this respect it agrees with the Sigalionidae. The
stomach is very muscular. The lateral intestinal ca3ca are large and elliptical, with a
narrow neck.

Scales (Plate XXXIV, fig. 5) thirty-nine pairs (Kinberg), in life pearly white,
semi-transparent, the first three pairs flattened, covering the dorsum, the rest cam-
panulate, and about one-third the breadth of the dorsum on either side, the centre being
bare. "During life these do not rest upon the body, but in front are tilted up, so as to
meet at an angle above the prostomium, the last few pairs of elytra also assuming a
similar position. ... A constant rising and falling of the elytra, as though to
facilitate the passage of water for the purpose of respiration, was observable"
(Arnold Watson). They are perfectly smooth, of fair thickness, and richly supplied
with nerve-trunks, as in the PolynoidaB. Mr. Watson found in a living example that
posteriorly the last two pairs only met in the middle line, whereas in a specimen
in the British Museum, dredged by Sir John Murray in 44 fathoms, the last six pairs
did so.

Feet. — The dorsal division of the first foot bears two long and rather slender spines,
and several long slender bristles, finely spinous from a short distance above the base to
their capillary extremities. The slenderness of these hairs is in contrast with the con-
dition of the homologous organs in the Polynoidas. The spines are minute.

The second foot is somewhat complex, and is specially interesting in connection with
the action of the parts in the formation of the tube, as recently described by Mr. Arnold
Watson. The foot is essentially bilobed, — that is, divided primarily into a dorsal and a
ventral division, the latter, it is true, being again subdivided. The dorsal division has
its spine, and forms an even ridge dorsally — terminating in a projecting globular or clavate
knob — directed downwards and projecting as far as the ventral division. Beneath are a
series of stiff bristles of the type seen in the foregoing process, but much stronger. They
taper from the base to the slender apex, and the short but distinct spines begin a little
above the former. The main ventral division is trilobed. The uppermost bristles
spring rather above the lobe, point upwards and outwards, and have stout shafts ending
in a spinous region dilated at the base and tapering to a slender tip. As we proceed
downwards the spinous region becomes more slender and tapering — the enlargement at
the base of the region gradually diminishing, and the shafts also becoming more slender.
An accessory lobe (bract) occurs at the ventral border, and in it the bristles, though
retaining the same type, become more slender, and the tapering spinous region shorter.
The ventral cirrus of this foot agrees with the homologous organ in the Polynoidse, and
is apparently used in the same manner. Though certain modifications exist in this foot,

PAXTHALIS (ERSTEDI. 403

yet the general structure is so much in accordance with the type in others that the
functions in all probability are not very diverse.

The third foot presents a short subulate dorsal cirrus and a somewhat clavate dorsal
lobe, with finely serrated slender bristles. The ventral is a massive but short lobe, having
superiorly a small group of bristles of the type of those in the foregoing foot, the spinous
regions being slightly bent backward and downward, with prominent rows of spines at
the commencement. Beneath are about six short and strong bristles with spear-shaped
tips, slightly curved and quite smooth, —in short, the extremities of the typical series
without the spines, but with the accessory process at the tip. These cover the main or
central region of the foot, with intervals between. From the accessory process at
the ventral edge of the foregoing lobe springs a group of more slender bristles of the
type of the superior series, with well-marked alternate spines. In ordinary views (in
spirit) the serrated concavity of these bristles is directed upward. As Pruvot and
Racovitza observe, one or two simple serrated bristles occur at the inferior border of this
foot. They probably indicate the original condition of the series. The subulate ventral
cirrus extends beyond the fleshy part of the foot.

The fourth foot has the short slender dorsal bristles under the lobe superiorly. In
the ventral division the bristles preserve the same type superiorly, but they have spread
a little downward. The tips of the great spines beneath are somewhat longer, and the
upper has a slender terminal whip. In like manner the slender inferior bristles have
spread upward beyond the two lower strong bristles, their structure, however, showing
no marked difference from those of the previous foot. The accessory ventral lobe is a
mere notch.

The fifth foot exhibits no noteworthy change except the increased distinctness of the
dorsal lobe, and the same may be said of the sixth.

The seventh foot agrees with the foregoing, and shows very well the series of
globular warts or papillse along the dorsal ridge above the cirrus. There are about eight
of the stout median bristles with the enlarged tips and terminal brush.

The eighth foot inaugurates a change, for, superiorly, the slender bristles with the
brush-like tips have appeared in the ventral lobe, while the dorsal division is only
indicated by an elevation without bristles. There are about five of the strong median
bristles with the filiform brush, while the inferior group of slender forms with the curved
spinous tips remain as before. The glandular apparatus for the peculiar secretion
commences in this foot,

The foot and bristles gradually assume the typical condition (Plate XXX, fig. 8),
but the foot varies little from the foregoing, presenting a smaller superior and a larger
inferior spine projecting in the centre of the median lobe, a rounded process superiorly,
then the more prominent median lobe, and an inferior process. The brush-shaped bristles
superiorly have very long slender shafts, which slightly dilate distally, then taper and
terminate in a point. From the sides of the tapering part a series of long hair-like
spines project like a brush (bipennato-penicillatge, Kinberg), the enlarged region at the
base having a shorter series (Plate XLI, fig. 15). The hair-like bristles either form a
hair-pencil, as in the figure, or a broader brush at the tip. These bristles occupy about
the upper third of the median lobe. Eight or nine strong aristate bristles then follow,

404 PANTHALIS (ERSTEDL

and their shafts are considerably longer than in front ; they dilate in a spindle-shaped
manner (in antero-posterior view, Plate XLI, fig. 16) at the end of the shaft, and then
taper to a blunt point. In lateral view (Plate XLI, fig. 17) this region is somewhat
spathulate in outline, the tip being broad, with the end of the shaft slightly bent
backward, and finely striated. More than the distal half is covered with fine spines
(pinnately arranged), which increase in length distally, and finally terminate at the dorsal
edge in a long slender whip of such fibres, one being stronger and far longer than the
others, and projecting from the midst of a basal series of large hairs (Plate XLI, fig. 17).
These bristles (Grannenborsten of Marenzeller) appear to be capable of repair, one
presenting two fractures below the tip, yet quite stiff and useful, the callus in each case
being more coarsely striated than the normal bristle. In one small example from 358
fathoms in the Atlantic these bristles have shorter and more acutely tapered tips, and
the spines cover three-fourths of the edge, commencing just above the base. Another
and larger example from 516 fathoms (H.M.8. 'Triton') shows similar features. The
terminal whip is a true prolongation of the shaft, with lateral spines.

At the ventral edge is a group of bristles with long shafts as slender as the superior
brush-like forms, a well-marked shoulder or curve as in the anterior bristles (which
they closely resemble), and a long tapering curved finely spinous tip (subulato-serratse,
Kinberg). The spinous rows of the curved region are prominent, while the long tapering
region beyond has its spines somewhat longer just after its commencement, and thus in
antero-posterior views presents a broader feathered arrangement (Plate XLI, fig. 18).

The feet retain the fundamentally bifid condition to the posterior end, both spines
being present, though the dorsal lobe is only marked by a slight eminence (devoid of
bristles), to which the spine goes. All the bristles are greatly elongated, but they
preserve for the most part the characters seen in front. Thus the smoothly rounded
ventral division has superiorly long slender bristles with the dilated and serrated tip.
The shafts of the strong median series are also much elongated, and the dilated tips are
densely spinous like a stiff brush, and closely akin to the condition of certain forms in
Aphrodita aculeata. No filamentous terminal brush occurs, but the sides of the club-
shaped tip are densely bristled, and the spikes project beyond the tip. The filamentous
tip may have been abraded in these cases, but this is uncertain.

An elaborate account of the golden yellow fibrous secretion by the spinning glands in
Polyodontes maxillosus, with numerous finely drawn figures, is given by Dr. Hugo Eisig
in his beautiful ' Monograph on the Capitellidae,' l and the arrangement seems to be very
similar to that in Panthalis. Dr. Eisig considers the fibrous golden yellow secretion as
homologous with the spines and bristles of these and other Annelids, and enters into a
lengthened description of the structure of similar glandular secretions in both Vertebrates
and Invertebrates. The glands are long tubular structures which pour their secretion
externally by an aperture in the dorsal papilla above the foot. The threads thus
secreted are mingled with mud in the formation of the tube, and Mr. Arnold Watson has
described the mode by which the animal ruptures the anterior end, and, throwing it
outward, adds layer upon layer in this manner, so as to constitute a massive tube.

1 Naples, 1887.

PANTHALIS (BRSTBDL 405

Pruvot and Racovitza's view that the spinning glands are the homologues of the
dorsal setigerons glands, forming the material for the tube instead of the dorsal bristles,
is not free from doubt.

Reproduction, — In a large example procured in July, 1868, in Shetland (78 fathoms),
a series of large ova occurred in the perivisceral chamber. The embryo must therefore
attain considerable size before extrusion.

Habits. — Like others of the group, Panthalis (Erstedi inhabits a tube of mud, about
3J inches long and about 1 J inches in diameter, with loose extensions of mucus at either
end, thus concealing the entrances, and for an example of which I am indebted to Prof.
Herdman. The estimated internal diameter of the tube is usually about f inch, while the
thickness of the walls in the centre is about J inch. The tube is composed of a number
of layers of the thread-like secretion interspersed with mud, these layers not being
parallel, but curving outwards, since, according to Mr. Arnold Watson, from whose
interesting account 1 of the habits of the species the foregoing is taken, rupture of the
anterior end frequently takes place.

The animal crawls along the surface of the ground, can reverse itself in its tube,
and can also swim a little in the water.

The American Euarche tubifex described by Ehlers 3 makes a curiously ringed
arrangement of the anterior end of the tube, probably due to the gradual narrowing of
the calibre of the tube at this end.

Kinberg in 1857 described the species thus : — The cephalic lobe with the peduncles
equal to a fifth part of the length of the palpi, tentacular cirri longer than the tentacle ;
bristles of three kinds, subulate, serrulate, bipennato-penicillate, bearded. No f oot papillse.

Marenzeller published an excellent account, with good figures, of the bristles, as
well as pointed out the functions of the spinning-glands in regard to tube-making.

Pruvot and Racovitza give two new species of this genus from Banyuls with accurate
and beautiful figures, viz. P. Lacazii and P. MarenzellerL After a careful study of the
first-mentioned as given in the authors' descriptions and figures, I have doubts as to the
need for specific distinction from the British form, though there are a few minor varia-
tions, such as the increased size of the so-called branchial papilla in front. It would
also appear to be a question whether the French authors have not placed too much
weight on the mere curvature of the ventral bristles of their Panthalis Marenzelleri*
especially as Kinberg's artist may have had a mounted preparation for illustration, and
thus the curves would disappear. The twist or double curvature referred to is present in
all the examples of Panthalis CEJrstedi that have come under my observation, and the
bristles of the first pair of feet (bearing the tentacular cirri) are likewise present. The
comparative length of the tentacle is not always to be relied on, though in young
specimens the tentacle seems to be about the same length. The same remarks apply to
the median papilla of the proboscis. The absence of the ceratophore is remarkable.
The bristles in young forms show the finer characters better than in the adults — in which

1 Ibid., pp. 170, 183, &c.

2 < Annelida of the " Blake," ' &c, p. 54, Taf. 12, 13.

3 " Faune des Annel. de Banyuls," 'Arch. Zool. exper./ 1895, p. 451.

406 SIGALIONID^].

they have been subjected to considerable friction. The most noteworthy distinctions of
the French authors are the condition of the second foot, which is exceptionally large,
and the commencement of the spinning-glands on the ninth foot instead of the eighth.
Further investigation will probably clear up the doubtful points. It has to be stated,
however, that considerable variations exist in the common form (P. (Erstedi) in regard to
the length and bulk of the second pair of feet. Thus, for instance, these were unusually
large in an example procured by the ' Triton ' in 516 fathoms ; arid the median tentacle
was also much longer than usual, though it did not reach the tips of the lateral tentacles.
The spinning-glands began in this on the eighth foot as in the ordinary examples.

Family V.— SIGALIONID^E, 1 Kinberg, 18

-3/

Cephalic lobe rounded, often with a nuchal collar posteriorly. No facial tubercle.
Median tentacle, when present, generally long, produced from the median part of the
cephalic lobe, and with ctenidia at the sides of the base (ceratophore). Lateral
tentacles fused with the base of the tentacular cirri, the tips only emerging. Eyes four,
occasionally only two, or absent. Palpi long, attenuate and smooth, with buccal ctenidia
at the bases.

Body long and narrow. Pharynx with f yy f§ papillae and teeth. No pregastric
caeca. First pair of feet carried in front of the head. Dorsal bristles spinous and
tapering. Ventral bristles compound, the terminal region being often long, multi-
articulate and bifid. Ctenidia on all the feet. Elytra and cirriform gills alternate in
the anterior segments up to the twenty-sixth ; those of the middle and posterior part
furnished with both elytra and cirriform gills.

Granular epidermic area for the nerve-cords deep, the upper arch being covered by
the insertions of the vertical and oblique muscles, the latter attached below the former
— on each side of the nerve-area — without intermingling to any extent.

The genus Sigalion was established by Audouinand M. Edwards in 1834, and adopted
by Cuvier in the second edition of the ' Kegne animal' 3 for two species, remarkable
for their genera] organisation amongst the Aphroditaceans by the simultaneous presence
of superior cirri and elytra on all the feet. They did not see eyes in the spirit-preparations,
and no branchiae, the fringes of the elytra perhaps taking the place of the latter. They
grouped the Sigalionidas under the Aphrodisiens vermiformes along w4th the Acoetes.

Kinberg's fifth family of the Aphroditea is the Sigalioninze, which he characterises
as having a long narrow body, the cephalic lobe rounded, occasionally elongate. No
facial tubercle. Tentacle not always present, and generally long, the basal region

1 Dr. George Johnston observes, (C Perhaps formed from atyaXoeig, curiously or anomalously
made ; but Sigalion is a name of Harpocrates, the companion of Esculapius and Hygeia, by whom
physicians were obliged to swear that they would observe a religious silence in their profession."

2 T. iii, p. 207.

STHENELAIS. 407

produced from the median part of the cephalic lobe. Antenna?, when present, arising
from the anterior part of the cephalic lobe. Eyes four, occasionally only two or none.
Near the base of the tentacle are two fossa? covered with a membrane — perhaps auditory
organs. Palpi long, attenuate, smooth. Tentacular cirri furnished with many bristles.
Pharynx with f yy yf papilla? and teeth. Dorsal bristles serrate, ventral compound.
Elytra and cirri alternate in anterior segments, those of the middle and posterior part
furnished with both elytra and cirri. He gives four genera — Sthenelais, n., Sigalion,
Aud. and Ed., Leanira, n., Psammolyce, n. He split up the old genus Sigalion into
Sthenelais and Sigalion. Sthenelais he characterised as having a cephalic lobe rounded
anteriorly, except where marked by the base of the tentacle. Antenna? short, affixed to
the base of the tentacle. Dorsal and ventral divisions of the foot of equal length, the
former with serrate bristles, the latter with superior subulate-serrate — occasionally
bidentate, and with inferior articulate, the apex being bidentate or serrate. Elytra
covering the entire dorsum.

Ehlers, in his ' Annelids of the United States Expeditions ' (1887), follows Grube's
arrangement of this group with Kinberg's subdivisions.

Genus XXIV, — Sthenelais, Kinberg.

Cephalic lobe rounded anteriorly, with a nuchal collar posteriorly in the prepara-
tions. A pair of ctenidia at the base (ceratophore) of the median tentacle ; lateral
tentacles fused with the first foot; tentacular cirri more or less separate. Palpi long,
subulate and smooth, springing from the ventral surface of the cephalic lobe, but fusing
with the first foot. A pair of scoop-shaped ctenidia at the base. Scales covering the
dorsum ; fringed. Dorsal and ventral divisions of the foot of equal length. Segmental
eminence at the base of each foot, and a ciliated funnel-shaped process on the foot. A
well-developed branchial process on every foot, and on the dorsal ridge beneath it three
T-shaped ciliated organs. Dorsal bristles long, stiff, finely tapered, and spinous. Upper
ventral bristles with simple tips — boldly spinous ; next come compound bristles, at first
with a terminal region of three segments, then with one joint, and at the ventral edge
with one to four joints — all with bidentate tips. They are arranged in the foot after
the outline of a horseshoe (in transverse section). Nerve-cords in a deep area, almond-
shaped in section.

Grube, in the ' Annulata Semperiana,' defines the genus thus : — Body vermiform,
segments more or less numerous, alternating. Elytra on the second, fourth, fifth, and
on the unequal segments to the twenty-third, the intermediate segments bearing branchia?,
the rest furnished with both organs. Scales covering the entire dorsum. One frontal
(median) tentacle. Two palpi (his subtentacula). First foot thrown forward with the
cephalic lobe, and furnished with bristles and short tentacles. Other feet biramous and
with ventral cirri. Superior bristles simple, barbed ; inferior bristles compound or with
other kinds. Elytra thin. 1

1 Vide previous brief description in c Sitz. Schles. Gesell./ 1874.

53

408 STHENBLAIS BOA.

Carus in the ( Prodromus Faunae Mediterraneae ' gives the following : — Head with
median antenna and generally two short lobulated or fleshy-membranous organs at the
base; maxillae well developed; feet all furnished with branchiae, and the anterior pair
turned to the front, so that the cirri and setae form part of the cephalic apparatus ; body
elongate, vermiform.

Pruvot and Racovitza l have recently drawn attention to the homologies of the head-
processes in Sthenelais, and express the view that the subulate cirrus which springs from
the inner side of the bristle-tuft of the first foot (bearing the tentacular cirri) is the lateral
tentacle (their lateral antenna). In their descriptions they further point out the exact
arrangement of the dorsal and the ventral bristles, the latter forming a horseshoe with
the opening in front, and with a spur dorsally and ventrally, each of the latter being
characterised by a change in the structure of the bristles.

1. Sthenelais boa, Johnston, 1833. Plate XXVI, figs. 7 and 8.

Specific Characters. — Head crimson, broadly ovate, bounded by a nuchal collar
posteriorly. Median ridge passes in front into the broad basal process (ceratophore) of
the median tentacle, which is short and subulate. Two flap-like organs (ctenidia) occur
at each side, the anterior the longer. Two eyes on each side, the larger anterior pair
almost hidden by the latter processes, and looking forward and outward. The posterior
pair are on the dorsum, a little behind the bases of the processes. A truncated papilla
at each side of the head posteriorly. Head fixed to the massive parts of the first feet,
which bear externally two tapering cirri of considerable length. Behind the dorsal
tentacle is a T-shaped ciliated process. To the inner side of the ventral cirrus of the
first pair of feet is a sickle-shaped blunt appendage (ctenidium) — richly ciliated, and
probably branchial. A short subulate cirrus (lateral tentacle) exists superiorly on the
inner side of the bristle-tuft. The long, subulate, and smooth palpus springs from this
basal process, which also carries a double series of long, tapering, slender, spinous
bristles. Body elongated, little tapered anteriorly, but gradually diminishing posteriorly.
It is rounded dorsally and flattened ventrally, greyish or bluish-green, the latter or flesh-
colour characterising the iridescent ventral surface. Segmental eminence at the base of
each foot, and a ciliated funnel-shaped process on the foot. Scales 156 pairs ; mostly
reniform, and with numerous minute papillae, while the outer border bears a series of
large simple papillae. A well-developed branchial process exists above each foot, and on
the dorsal ridge beneath it are three T-shaped ciliated organs (ctenidia). The dorsal
bristles are long, somewhat stiff and finely tapered, as well as spinous from a little above
the base to the apex. Upper set of the ventral bristles with simple tips, boldly spinous;
then some with a tapering bifid appendage of about three joints, the end of the shaft
having a few rows of spines. The main part of the division, forming the large
curve of the horseshoe, has stout bristles with bevelled ends to the shafts, and a short
beaked appendage of one segment. Inferiorly the lower spur of the horseshoe has

1 ' Archives Zoolog. exper./ p. 452, 3 e Serv, iii, 1895.

STHENELAIS BOA. 409

more slender forms, with a few rows of spines on the slightly expanded distal regions of
the shafts, and an appendage of one to four joints. A group of long papillae occurs at
the anterior edge of the superior division, one above the spine of the ventral division,
and a ciliated or papillose frill is found at the base of the lowest slender bristles. Yentral
cirrus elongate, subulate, barely reaching the tip of the fleshy part of the foot.

Synonyms.

1833. Sigalion boa, Johnston. Loud. Mag. Nat. Hist., vi, 322, f. 42.

1839. „ „ idem. Ann. Nat. Hist., ii, 439, pi. xxiii, f. 6—15.

1843. „ Idunse, H. Rathke. Nova Act. Acad. Cassar. Nat. Cur., xx, 150, Tab. ix, f. 1—8.

1851. „ „ G-ruhe. Fam. Annel., p. 38.

n ,, boa, Williams. Eept. Brit. Assoc, 1851, p. 201.

„ „ Idunse, Sars. Nyt. Mag. Natur., xi, 3, p. 254.

1861. „ „ idem. Yid. Selsk. Forh., 1861, p. 1 (sep. copy).

1865. „ „ Malmgren. Nord. Hafs.-Ann., p. 86.

„ Sthenelais I.dunse and S. boa, De Quatrefages. Ann., i, p. 276.

„ Sigalion boa, Johnston. Cat. Brit. Mus., 124, pi. 13, f. 6.
1867. „ „ Parfitt. Annel. Devon., p. 19.

1873. Sthenelais Idunse, Sars. Bid. Christ. Faun., p. 14.

„ Sigalion Idunse, Willemoes-Suhm. Zeit. f . w. Zool., xxii, p. 347.

1875. „ „ Ehlers. Annel. ' Porcupine/! 869, op. cit., p. 18.

1876. Sthenelais boa, Mcintosh. Trans. Zool. Soc, ix, p. 390.

1879. Sigalion Idunse, Tauber. Ann.. Danic, 83.

1880. Sthenelais Idunse, Langerhans. Zeit. f. w. Zool., xxxiii, p. 276, Taf. xiv, f. 6.

1883. Sigalion Idunse, Levinsen. Nord. Annul., 199.

1884. „ „ V. Cams. Faun. Medit., p. 205.

1888. Sthenelais Idunse, De St. -Joseph. Ann. d. Sc. Nat. (7), v, p. 187, pi. viii, £. 55.

1890. Sigalion Idunse, Malaquin. Ann. Boulon., p. 23.

1891. „ boa, Hornell. Op. cit., p. 238.

Habitat. — Eanges from Shetland to the Channel Islands, being procured by the
dredge in the former and between tide-marks in the latter. It appears to be a generally
distributed species, frequenting both shores of Ireland, and extending to Norway and
Sweden. Ehlers gives a depth of 60 to 80 fathoms in the Minch during the ' Porcupine '
Expedition of 1869. Shores of France.

Read (Plate XXIX, fig. 1) of a fine crimson hue, somewhat broadly ovate, and
separated from the body by a nuchal collar. Anteriorly a median ridge terminates in
the broad basal process for the pale median tentacle, which is comparatively short. At
the sides of the basal process (ceratophore) are a pair of flap-like organs (ctenidia), the
anterior being the longer, the posterior the shorter. Both are rather flattened, and move
freely with the broad tentacular base (first foot) on the head— which is fixed. At their
base externally on each side are the eyes, the anterior and larger pair of which are almost
hidden under these processes— when viewed from the dorsum, especially as they are also
placed on the front of the head so as to look forward and outward. The posterior pair
lie a little behind their bases on the dorsum. At the posterior border on each side is a
well-marked process or papilla with a truncated tip (one of the " organes nucaux "). The

410 STHENELAIS BOA.

head is fixed inferiorly to the massive bases of the tentacular cirri (first foot). Externally
two tapering cirri of considerable length occur, one, the longer, from the dorsal edge, and
another a little shorter from the ventral edge above the palpi. Behind the dorsal tentacle
is a T-shaped ciliated process (ctenidium). To the inner side of the ventral cirrus is a
sickle-shaped blunt appendage (cuilleron cephalique, or buccal ctenidium), richly ciliated
and continuous with a well-marked ridge on the inner surface of the basal process. On
the inner side of the bristle-tuft superiorly, again, is another short subulate cirrus, the
homologue of the lateral tentacle (Pruvot and Racovitza). The long subulate and smooth
palpus springs from the middle of the ventral face of the basal process, and has a scoop-
shaped flap (ctenidium) ensheathing it internally. The first foot carries a double series
of bristles directed forward and inward, and consisting of somewhat stiff, long, slender,
curved bristles closely spinous towards the tip. The upper series of bristles had many-
stalked infusoria like Bhabdostyla.

Body four to eight inches in length ; elongate, not much tapered anteriorly, but
gradually diminishing posteriorly ; rounded dorsally and flattened ventrally, composed
of 170 to 200 or more bristled segments. The dorsal surface is marked anteriorly by
transverse lines, and the median region forms an elevated ridge, and sometimes has a
bluish-green or ochreous colour. The post-cephalic ridge has at its sides small
ciliated processes, which at first sight resemble parasites. The ventral surface
is of an iridescent flesh-colour, or bluish-green (from eggs) marked by the red
central vessel, and with a raised transverse line between the centres of the feet.
Posteriorly the body terminates in two tapering caudal styles. 1 A segmental eminence
occurs at the base of each foot, and on the adjoining region is a little funnel-shaped
papilla, but apparently unconnected with the former. In the anterior feet the funnel is
in the centre of the base of the foot, but it by-and-by moves forward, so that through-
out the rest of the body it is near the anterior edge of the foot. The funnel-shaped
papillse are continued from the fourth bristled feet to near the top of the tail; about
twenty segments, however, are devoid of them. In the males this funnel-shaped
organ does not appear to be present, but the segmental eminences are very prominent,
and have an oblique crest. In spirit the sperms sometimes form masses of a rope-like
character externally, as if they had issued (on immersion in spirit) from a minute pore.
The cuticle is considerably thickened over the nerve-area.

The proboscis has eleven papilla on each half (dorsally and ventrally). These are
large and somewhat conical, slightly constricted at the base, and with a projection
internally about the middle. The teeth do not seem to bite alternately, as in the
Polynoidas, but the lower pass in front of the upper pair (Plate XXVI a, fig. 21).

No pregastric ca3ca are present, but short intestinal caeca occur laterally.

Scales (Plate XXXIII, fig. 16), 156 pairs, covering the entire dorsum, of a greyish
hue, the anterior especially showing numerous dark brown grains ; all however possess
them, unless when recently reproduced. Towards the posterior end they have a
broad band of dark grey or brownish grey along the posterior border internally, and
occasionally externally a dappled condition or grains of the same dark grey colour. A

1 Body terminated by four small cirri (De Saint-Joseph).

STHENBLAIS BOA. 411

very pretty and symmetrically tinted arrangement thus pertains in this region. The
first pair of scales are somewhat ovate, the rest more or less reniform, and they are
tough, — adhering, moreover, with considerable tenacity to the tubercles. Their surface is
entirely covered with minute papillae, which are often tinted brownish, and along the
anterior and neighbouring part of the outer border are a series of massive and simple
cilia, somewhat constricted at the base, then dilating and tapering to a blunt tip. Some
show a branch or knob near the base. In the other scales a portion of the outer and
anterior region is devoid of the minute papillae, whilst amongst the cilia of the outer
border are numerous minute globular papillae. Both extend here and there within the
border. The inner or rounded anterior lobe of the scale has the papillae up to its border.
The scales occur on the first bristled foot, third, fourth, sixth, eighth to twenty-fourth,
twenty-sixth, twenty-seventh, &c, to the end of the body. In the terminal region the
small scales retain the same characters, though a patch devoid of papillae occurs on the
anterior part of the ronnded inner lobe, and the cilia along the external border are
smaller, more slender in proportion, and more numerous. The minute papillae on the
surface occasionally project beyond the border posteriorly in all the scales, and the gradual
elongation of the cilia externally shows that they are modifications of the same organs.
The largest papillae occur on the exposed part of the scale. Finely branched nerves
supply the entire organ. The scales often have a blackish or brownish deposit of
granular matter on their surfaces. It seems to be the same as that which occurs in the
Polynoidae. In young examples the papillae on the scales are proportionally large, and
occasionally sand-grains adhere. The cilia also are somewhat larger.

Feet. — The second foot shows dorsally two ciliated lobes. It has two powerful
spines. The dorsal division is indicated by the nature of the bristles and by the issue of
the spine beneath them. The dorsal bristles consist of the same slender, elongated,
finely spinose forms seen in the previous process, the tips of most being broken. They
have the infusorial parasites formerly indicated and fine filaments of an alga. The
ventral division bears much stronger bristles, the stout shafts of which merge into the
distal spinous region without evident change. The slightly alternate rows of spines are
prominent, and this region ends in a long, tapering, articulated appendage terminating
in a bifid tip, the dorsal process being hooked. Just above the lower edge of the ventral
region a series of large, lobate papillae project, and below the bristles are a curved series
of minute papillae. The ventral cirrus, as in the previous groups, is large and
tapering, its tip extending considerably beyond the fleshy part of the foot. The funnel-
shaped papilla external to this organ is absent.

The third foot shows a T-shaped ctenidium dorsally. The dorsal division of the
foot is more clearly indicated by a fissure, and its lower border is furnished with large
clavate papillae. The bristles curve upward and inward, their convexity being thus
external. The bristles of the ventral division have already a tendency to form groups.
Thus the upper six of the ventral series consist of moderately stout shafts with simple
tapering tips which are coarsely spinous inferiorly and finely spinous distally, the tip
being smooth. Then follow the stouter forms, as in the second foot, with spinous
regions distally and a jointed, tapering, bifid appendage. Amongst these, however, are
some with an unjointed terminal appendage, a powerfully hooked tip, and a strong

412 STHENELAIS BOA.

secondary process, so that the end is like a bird's beak. Those at the ventral edge
again are much more slender, have a slightly dilated spinous region, and a five-jointed
bifid appendage. Two groups of clavate papillae occur on the edge of the foot above
them and a row of smaller papillae at their bases ventrally. The ventral cirrus extends
fully as far as the fleshy part of the foot, and internal to it is a funnel-like, ciliated
process.

The typical foot (Plate XXXI, fig. 5) presents dorsally a richly ciliated branchial
process somewhat subulate in outline, and which is evident at the fourth foot. Beneath,
on the dorsal curve, are three T-shaped ctenidia. Another ctenidium lies on the posterior
border of the foot deeply placed in the interpedal fissure. In the middle of the body others
occur on the dorsum of the foot and behind the elytrophore. Pruvot and Racovitza observe
that they all have the same structure, viz. conical epidermic cells with their bases at the
broad end and their points at the pedicle of the organ; they are richly ciliated distally.
The dorsal lobe is considerably less than the ventral; the bristles (Plate XLI, fig. 19) are
long, slender, and boldly spinous from a little above the base to the tip. Viewed from the
side they form a beautiful fan directed laterally with the digit-like papillae at the anterior
border of the base. Though slender they are somewhat stiff and curved — with the
concavity upward (Plate XLI, fig. 20). The spines form even transverse rows,
thus differing from the alternate series so common in this group and in the
Polynoidae. The transverse arrangement, however, may render them more brittle. The
dorsal lobe is separated from the ventral by a deep fissure, and has a series of long
papillae at its tip. The ventral lobe is massive, with a truncated extremity which bears
the spine in its middle, and usually a single digit-like papilla. A fold of skin,
arising at the upper part of the lobe, passes down posteriorly and curves to the front,
separating the more slender ventral bristles from the stout series. In the posterior part
of the body the commencement of this ridge of skin forms a free flap. The upper group
of the ventral series consists of the moderately stout forms with the simple spinous tips
(Plate XLI, fig. 21), and also a few with about three joints to the tapering, bifid
appendage, the tip of the shaft having a few rows of spines. The centre of the foot is
occupied by a series of stout shafts bevelled at the tip, and with a short appendage of one
segment (Plate XLI, fig. 22), and the beak-like, bifid extremity. In some, transverse
striae indicate traces of spinous rows. Those of the upper series have longer and more
slender appendages. Ventrally is a group of slender bristles forming a row directed
forwards and outwards, then downwards and outwards, each with a few rows of spines
on the expanded distal end of the shaft and appendages of one to four joints (Plate XLI,
fig. 23). These spring from a notch at the ventral edge and within the tip of the foot,
the inner border of the notch being marked by a series of small papillae. In the region of
the spine, which pierces the middle of the foot, is a long and somewhat fusiform papilla.
So far as the structure of the bristles is concerned, the type remains the same in the
terminal feet, though the bristles diminish in size. The median stout bristles have longer
and more slender hooked appendages, and the backward curve of the end of the shaft
is more marked. The papillae are fewer. The ventral cirrus is subulate and tapering,
extending to about the tip of the fleshy part of the foot. A spur or process occurs at
its inner border above the basal segment. A well-marked ctenidium is placed a little

STHENELAIS BOA. 413

internal and anterior to the cirrus. The dorsal hairs are prone to deposits of mud, algoid
growths, and stalked Infusoria, their colour thus being often brownish or greyish ; while
Loxosomse abound on the feet, under the scales, and other parts.

Pruvot and Racovitza give a good figure, 1 showing the arrangement of the lobes,
papillas, and bristles in a typical foot. Dorsally the papilise (stylodes) occupy the
anterior region and pass beneath the division. Yentral]y, the flaps, which they call
parapodial bracts, envelop the foot above and beneath, as well as superiorly in front, the
anteroinferior being scolloped, each process being terminated by a sensitive palpocil.

Reproduction. — Large specimens in the Channel Islands were laden in August with
fairly advanced greenish eggs in enormous numbers. The males had also the peri-
visceral chamber charged with sperms, which issued, after immersion in spirit, from the
bases of the feet.

Development. — Larval forms occurred, towards the end of October, with bluish
anterior digestive organs in the metatroch stage, with two kinds of bristles, — viz.
long, curved, serrated bristles, somewhat like those of Polynoe, and shorter com-
pound forms. Viewed from the front superiorly (Plate XXVIa, fig. 22), two eyes lie on
each side. An oblique lateral view shows the feet of one side and the caudal cirri.
The first and last bristle-tufts are simple and short, and about seven pairs of feet are
visible. Only a single, long, and comparatively strong, swimming bristle occurs in each
foot, the curved tip being distinctly and somewhat widely serrated on the convex edge.
Such, therefore, would appear to be a modification of the serrate dorsal bristle, which is
functional at this stage. The ventral bristles had dilated ends to the shafts — with the
usual oblique termination and a distal region of two segments. It is uncertain whether
these young forms pertained to this genus or to Sigalion.

Habits. — They are somewhat sluggish animals, usually lurking under stones between
tide-marks, where they rest on a sandy bottom, though occasionally small examples are
found in pure sand as at Southport. They form curious coils in vessels in confinement,
and appear to be nocturnal. So far as my specimens go, the finest are those between
tide-marks, Herm, while next to these, and only a very little less, are those between
tide-marks at St. Andrews. Those procured in the sand of the Zetlandic voes are much
smaller, and the same may be said of those from the outer Hebrides. Average examples
come from the west coast of Ireland.

This species was introduced, in 1833, to science by Dr. Gr. Johnston, who found it in
Berwick Bay under stones near low-water mark. He says it is somewhat sluggish, but
burrows in sand with rapidity. Moreover that it is the Goliath of its race, and preys on
its fellow-worms. It is probable that Rathke, in 1843, referred to this species under
the title mentioned (S. Idunae). The Sthenelais Edwardsii of De Quatrefages (1865)
seems to be this species. He placed next the foregoing the Sigalion Mathilda of
Audouin and Edwards, for he thought they included two species under that title. He
entered the Sthenelais Idunx of Rathke, and the S. boa of: Johnston as separate species.

Langerhans (1880) describes Sthenelais Idunse, from a depth of twenty fathoms off
Madeira. He points out the reddish colour of the head (from the brain), the ringed base of

1 Op. cit., p. 463, fig. 12.

414 STHENELAIS ZETLANDICA.

the tentacle (his antenna), the ciliated rosette of the first segment, chitinous papillae, and
fringed scales (simple papillae), and gives the detailed structure of the feet with figures.
De Saint-Joseph (1888) observed many examples of Hal acarus olivaceus on the body of
one. He agrees with me in thinking that S. Idunse, Rathke, is identical with S. boa, Johnst.

2. Sthenelais zetlandioa, Mcintosh, 1876.

Specific Characters. — Head absent, and body fragmentary. Scales somewhat reni-
form or irregularly rounded, densely covered with low flat papillae, the outer and part of
the posterior edge bearing a closely arranged series of minute clavate cilia. Dorsal lobe
of the foot rather long, and obliquely bevelled from above downward, bearing a tuft of
bristles that are longer and more finely spinous than in S. boa. Beneath the spine are
a series of small papillae with truncated and papillose extremities. Central lobe irregu-
larly conical, with numerous papillae, at the tip of which are the secondary processes or
warts. Along the ventral border is a series of globose warts. The upper ventral
bristles have proportionally strong shafts, with four or five rows of spines at the distal
end, and a terminal bifid appendage usually of two joints. The slender inferior ventral
have three divisions in the terminal appendage — all with a hooked tip, and a secondary
process beneath.

Habitat. — Dredged off the Shetland Islands by Dr. Grwyn Jeffreys in 1867. Mr.
Harvey-Gibson found a fragment (?) off Port Erin, Isle of Man.

Synonyms.

1876. Sthenelais zetlandica, Mcintosh. Trans. Zool. Soc, ix, 390, pi. lxx, f. 15 — 17.
1886. „ „ Harvey- Gribson. Verm. Liver., p. 151.

1891. ■„ „ Hornell. Op. cit., p. 238.

Head. — Absent in the preparation, and the anterior end injured. The proboscis
appears to have the ordinary structure — with the horny teeth, and the dorsal and ventral
rows of conical papillae.

Body. — About the size of Sthenelais limicola, and having a similar appearance.
The scales are somewhat reniform, or irregularly rounded anteriorly, and the entire
surface densely covered with flat papillae, which on the folded edge of the scale form
low, smooth warts, larger in proportion than in Sthenelais boa. The outer and part of
the posterior edge again bear a somewhat closely arranged series of minute clavate cilia
(Plate XLI, fig. 24), almost globular at the commencement on the posterior border, and
with minute processes or palpocils on the summit, those on the outer edge being more
slender than those on the posterior border.

The feet (Plate XXX, fig. 14) have dorsally a branchial process and three ciliated
T-shaped organs beneath. The superior lobe is rather long, and obliquely bevelled
from above downward. It bears beneath the spine a series of rather small papillae,
which have truncated extremities provided with several accessory papillae. The
superior bristles are longer in proportion than in $. boa, but do not form so broad a fan

STHBNBLAIS ATLANTICA. 415

when viewed laterally. They taper to a delicate extremity, and their rows of spines are
distinctly marked, though somewhat finer than in 8. boa. The inferior lobe is irregularly
conical, and has numerous papilla?, the largest near the spine, and just above the
inferior group of bristles. Moreover below the latter is a cluster of smaller papilla? fur-
nished with the secondary processes or warts at the tip. After a brief interval, a series of
globular warts occurs along the ventral border of the foot. The superior ventral bristles
(Plate XLI, fig. 25, in calcium chloride, and with the basal part of the terminal
appendage slightly folded) have comparatively strong shafts, with four or five rows of
spines at the dilated distal extremity, visible, however, only at its edge. The terminal
appendage as a rule consists of only two divisions, occasionally a more slender form at
the upper border having three. In those with two the terminal portion is only one
fifth the length of the basal. The claw at the tip is distinct and much curved, and the
secondary process projects at a slight angle, the point often touching the tip of the claw.
The basal joint is finely striated, as is likewise the dilated end of the shaft. As we
proceed downwards these bristles become less robust, and the shaft has a more distinct
dorsal curvature, but the terminal region of two divisions remains. Bach of the more
slender ventral groups, again, has three divisions in the terminal appendage (Plate XLI,
iig. 26, which represents one of the larger forms). The bristles throughout are tinted
of a light brownish hue.

The ventral cirrus is subulate, and it reaches nearly as far as the fleshy part of the
foot (in spirit). The terminal region, which is elongate-ovoid, is marked from the
succeeding by a distinct shoulder, as if articulated.

The absence of the head makes the generic relations of this species doubtful, but in
the meantime it may be placed under Sthenelais until a more complete example is
obtained. It is one of the many rare forms which the persevering explorations of
Dr. Gwyn Jeffreys in the Zetlandic seas brought within our knowledge.

The 8. fuliginosa of Claparede l is an allied form, and the S. minor of Pruvot and
Racovitza 2 also approaches this species in regard to the structure of the papilla? and
the general character of the bristles, but differs in the presence of papilla? along the
ventral edge of the foot, and it has fewer papilla? on the scales than in the British form.

3. Sthenelais atlantioa, Mcintosh, 1876.

Specific Characters. — Head somewhat ovoid. Median tentacle subulate, and with
a terminal joint. A pair of eyes close together on each side,, near the base of the
tentacle. Palpi long, smooth, and tapering. Tentacular cirri show a terminal differen-
tiation, as in the median tentacle. Body as in S. zetlandica. Scales thin, rounded or
ovoid in front, reniform throughout the rest of the fragment, covered with sparsely
distributed but clavate cilia, and having a fringe of short clavate cilia, which are only
absent from a portion of the inner and adjoining anterior margin. They are longer and
more numerous than in S. zetlandica. Dorsal lobe of the feet bevelled from above

1 < Ann. Chet. Naples/ p. 94, pi. iv, f. 2.

2 ( Archives Zool. exper./ 1895,, p. 465.

54

416 STHENELAIS ATLANTICA.

downward, and witli papillae like those of 8. zetlandica. The bristles are similar, though
in the smaller specimen finer. The ventral lobe is conical, with long papillae,— as
in 8. zetlandica. The upper ventral bristles have about six rows of spines at the
dilated distal region of the shaft, and the terminal process has three or four segments.
The tips are slender a!nd bifid, the secondary process having no appreciable angle to the
axis. The next lower forms are stouter, with a beak-like extremity. The inferior
bristles are slender, with about two rows of spines at the end of the shaft, and a
tapering terminal appendage of two or four divisions, bifid at the tip. Apparently three
ciliated pads (ctenidia) occur on the dorsum. The ventral cirrus is slender, and rather
shorter than the fleshy part of the foot.

Synonym.
1876. Sthenelais atlantica, Mcintosh. Trans. Zool. Soc, vol. ix, p. 405, pi. lxxii, f. 16, 17.

Habitat.— Prom the entrance of the British Channel to the Atlantic deeps in
305 fathoms, Station 2, < Porcupine,' 1870. The ground seemed to have been sandy.

Head (Plate XXIX, fig. 2) somewhat ovoid in outline, a small subulate tentacle with
a terminal joint in front, and a pair of eyes situated close together on each side near
the base (ceratophore) of the organ. The palpi are very long, smooth, and tapering,
and the tips of the tentacular cirri show a terminal joint as in the median tentacle.

Body.— Slightly narrowed in front, but as the feet project laterally and forward, the
outline anteriorly is really broad and gently curved laterally. Only about 10 mm. of
the body remain, and the specimen appears to be small.

Scales (Plate XLI, fig. 27) somewhat thin, rounded or ovoid in front, reniform
throughout the rest of the fragment, covered with sparsely distributed but distinct
clavate papillae, and having a fringe of short clavate cilia, which are only absent from
a portion of the inner and adjoining anterior margin. Four of the longest, indeed, occur
on the outer lobe of the anterior margin near the hilus. They are longest anteriorly.
The cilia (Plate XLI, fig. 27) are decidedly longer and more slender, as well as more
numerous than in 8. zetlandica, and palpocils occur frequently on the extremity.

Feet (Plate XXX, fig. 14) resemble those of 8. zetlandica, having a projecting
dorsal lobe bevelled from above downward, and the papillae have similar dimensions.
The bristles are somewhat finer, and the serrations or spinous rows closer, but the
specimen is smaller. The ventral lobe is conical, with similar long papillae, the secondary
processes or warts at the tip of these being distinct. The superior ventral bristles
(Plate XLI, fig. 28) have about six rows of spines at the dilated distal regions of the
shaft; and the terminal appendage has three or four segments, the basal, in the case of
those possessing four, being about as long as the three distal. The tips are slender and
bifid, the secondary process having no appreciable angle to the axis, so that it lies
close to the hook. The next lower series are stouter, with about four distinct spinous
rows on the distal end of the shaft, and a terminal appendage of one or two segments,
the tip resembling the beak of an eagle. The inferior are slender, with about two
rows of spines at the dilated end of the shaft, a terminal appendage of two or four
divisions, and a tapering, delicately bifid tip. The shafts of all the ventral bristles have
a backward curve distally. A branchial process, and apparently three ciliated pads

STHBNBLAIS LIMICOLA. 417

(ctenidia) occur on the dorsum, but the specimen is not in a condition to speak decisively.
The ventral cirrus is slender and rather short, the tip being shorter than the fleshy
part of the foot. The preparations would seem to indicate a joint at the tip. In a
former note it was mentioned that some minute warts appear along the ventral margin of
the foot, but the preparations are now doubtful on this point.

The species comes near 8. zetlanclica, yet the points indicated seem to warrant
separation. Both species approach Olaparede's S. ctenolepis from the Mediterranean, 1
but yet are sufficiently distinct.

4. Sthenelais limicola, Ehlers, 1864.

Specific Characters. — Head ovoid with the long diameter transverse, and median
space bounded by a brownish crescentic line on each side. Anterior and larger pair of
eyes often scarcely visible from the dorsum, being situated under the lateral processes
at the base (ceratophore) of the median tentacle, and looking forward. Posterior pair of
considerable size, a little behind the base of the median tentacle. Both occasionally show
a pale speck in the centre, as if from a lens. Base of the median tentacle (ceratophore)
somewhat conical (in spirit) ; tentacle of moderate length, the filiform tip being slightly
enlarged, and with a trace of an articulation. On each side of the base is a flattened
spathulate process (ctenidium). First foot fused ventrally with the base of the long,
tapering, smooth palpus, with a small sheath-like process (ctenidium) at its base
internally; then come a filiform ventral tentacular cirrus and a much larger dorsal
cirrus, and internally to the ventral cirrus a broad scimitar-shaped process (ctenidium).
Above and behind the latter is the small terminal region of the lateral tentacle.
Proboscis trumpet-shaped in extension, with eleven conical papillas dorsally and ventrally.
The teeth appear to bite as in 8. boa. Body elongate, more than two inches long, and
having about 128 segments. It tapers to a slender tail with two styles. Segmental
eminence opposite each foot ; no papilla. Scales smooth, translucent, brownish, covering
the back. First pair rounded, each with short clavate cilia along its outer border, and
a few larger digit-like forms at its anterior and outer margin. The others are more or
less reniform, the outer margin thin, folded, and having irregular processes — simple, bifid,
or irregularly divided. In the posterior scales the outer margin is bilobed, and in those
near the caudal region a belt of large round vesicles occurs in the hypoderm in front
of the scar: a touch of brown is present in some scales. Feet with branchial process,
and three ciliated pads (ctenidia) dorsally. The dorsal lobe is prominent, somewhat
clavate in outline, and has anteriorly four or hve long papillaa (stylodes) from its upper
end, the spine projecting inferiorly. The bristles are long, slender, tapering, and finely
spinous. The ventral lobe is shorter and broader, somewhat conical at the tip, and
bears one leaf-like lobe above the spine, and a smaller lobule at the ventral edge, a long
papilla (stylode ?) likewise being attached to the former region. Upper bristles like
those in S. boa, with simple spinous tips, only more slender. The next have slender
shafts with long tapering tips of twelve or more segments, and minutely bifid. A

1 ' Ann. Chet. Naples/ 88.

418 STHBNBLAIS LIMICOLA.

stouter series follows, also with long bifid tips, the shafts becoming more slender in the
lower group. Lastly is a more delicate group at the ventral edge, having bifid distal
appendages of from five to ten articulations. The ventral cirrus is long and tapering,
with a terminal segment.

Synonyms.

1853. Aphrodita arcta, Dalyell. P.ow. Creat., ii, p. 170, pi. xxiv, f. 14.

1864. Sthejielais limicola, Ehlers. Borstenw., 120, Taf. iv, f. 4—7 ; Taf. v.

1868. „ leiolepis, Claparede. Ann. Chet. Nap., 96, pi. iv, f. 3, and pi. vi, f. 1.

1869. „ limicola, Mcintosh. Trans. R. S. E., xxv, 410.

1875. „ „ idem. Invert, and Fish., St. A., p. 118,

1876. „ „ idem. Trans. Z. S., ix, pp. 390, 476, pi. lxx, f. 13.
1884. „ „ Y. Carus. Faun. Medit., p. 205.

1891. „ „ Hornell. Op. cit., p. 238.

1895. „ „ Pruvot and Racovitza. Arch. Zool. exper., p. 473, pi. xx, f. 122, 123.

Habitat. — First dredged by Dr. Grwyn Jeffreys in sixty to eighty fathoms off
North Unst, in St. Magnus Bay, and the Outer Haaf, Skerries, in Shetland in 1867. It
is very abundant off St. Andrews, probably in sand, and is tossed on shore in large
numbers during certain storms. While it is not common in the dredge, the cod seems
to find it readily, and so do the flounder and other fishes. In the 'Porcupine'
Expedition of 1869 it occurred in 30 to 370 fathoms off the Irish coast, and a small
eyeless variety in 420 fathoms. It ranges from Shetland to Cornwall, and has been
found chiefly in water of some depth, — never, at any rate, between tide-marks. It
extends likewise to Norway (Canon Norman), to Quarnero in the Adriatic, to the shores
of Canada, and the United States of America.

Head (Plate XXIX, fig. 3) somewhat ovoid in outline, with the long diameter
transverse. The median space is marked by a brownish crescentic line on each side,
which Ehlers compares to an H. The posterior pair of eyes are situated a little behind
the base of the median tentacle, are of considerable size, and sometimes show a pale
speck in the centre, as if from a corneal lens. The anterior pair are scarcely visible
from the dorsum (indeed, they escaped Ehlers), being situated under the lateral
processes (ctenidia) at the base (ceratophore) of the median tentacle, and looking
straight forward. They are somewhat larger than the posterior, and also occasionally
show a pale speck in the centre. The ceratophore of the median tentacle is somewhat
conical in spirit, but Ehlers says cylindrical in life, and from it a tapering tentacle
(ceratostyle) of moderate length passes. The filiform region shows a slight enlargement
at the tip in spirit, and a trace of an articulation at its commencement. On each side of
the base is a flattened spatulate process (ctenidium) with long cilia. The frontal lobe
carries ventrally the very long, tapering, smooth palpus, with a small sheath or
membranous collar at its base internally, then a slender filiform ventral cirrus ; dorsally a
much larger dorsal cirrus, and internally to the ventral cirrus a broad scimitar-shaped
process (cuilleron cephalique, Pruvot and Racovitza). A little behind and internal to the
dorsal cirrus is the tip of the lateral tentacle, which Racovitza followed backward to its
attachment to the head. In his description Ehlers omits the scoop-like process at the

STHBNELAIS LIMICOLA. 419

base of the palpus. His view of the parts, indeed (Taf. iv 9 fig. 6), differs from what
our specimens would have exhibited in life.

Proboscis. — The teeth are similar to those of 8. boa, the lower pair apparently
biting in front of the upper ; but preparations are not always to be relied on in this respect.
The number of conical papillas dorsally and ventrally is eleven. It is curious that few or
none eject the organ in spirit, whereas the gastric juice of the stomach of the cod causes
full protrusion, the broad end of the trumpet-shaped structure being distal. Moreover
a pair of oblique elevated ridges occur dorsally and ventrally near the tip.

Body. — The body is elongate, some of the imperfect specimens being nearly two
inches in length. Ehlers gives 57 mm. and 128 segments, and one of the contracted
perfect forms is larger than Ehlers' examples. Pruvot and Racovitza record a length
of 95 mm., and a total breadth of 4 mm. It tapers posteriorly to a delicate tail which
has two caudal cirri. The body is rounded dorsally, flattened ventrally. A segmental
eminence occurs at the base of each foot, but no distinct papilla.

Scales numerous, translucent, rounded in front, irregularly ovate or somewhat
reniform posteriorly, and cover the back. They are of a light brownish hue, best marked
in front. They occur on segments 2, 4, 5, 7 — 25, and on all the others (Ehlers). The
first scale, which has the scar for attachment behind its centre, presents a series of short
clavate cilia along its outer border, and a few longer digit-like cilia at its anterior and
outer margin. 1 Finely branched nerves pass from the scar throughout the scale. Its
surface is quite smooth. The typical anterior scale (Plate XXXIY, fig. 9) has externally
a thin folded margin possessing irregular processes, either simple, bifid, or divided into
several flaps. The surface is smooth, though often crossed in the preparations by fine
lines, probably creases or folds. Between the elytrophore and the anterior curvature, and
stretching on each side beyond, is a series of rounded bodies like papillse (the " petit
tubercles irreguliers " of Pruvot and Racovitza), but they do not project from either
surface of the scale, and appear therefore to be internal, as a reference to the condition
in the posterior scales more clearly demonstrates.

The posterior scales (Plate XXXIV, fig. 10) are bilobed externally, each lobe
forming a rounded process, which either approaches its fellow over the T-shaped fissure
or slightly overlaps. Moreover in the terminal scale a belt of large vesicular organs
occurs in the granular layer of the epiderm, in front of the scar for attachment. They
consist of a large pale central region surrounded by a ring of smaller bodies. Each of the
large circular vesicles has a cell with a nucleus and nucleolus, and the ring seems to be
composed of nucleated granular cells. They indeed resemble papillas with a ring of
secondary processes at the tip, but they appear to be internal, and a further development
of the structures noted anteriorly. These continue to the posterior extremity.

At Banyuls Pruvot and Pacovitza found a large transverse touch of brown on the
posterior region of the scales, and between the elytrophore and the anterior curve a
few minute tubercles, best marked and most numerous in the posterior scales. As
above mentioned, however, these structures project neither from the dorsal nor the
ventral surface of the scale ; they are vesicular and internal rather than tubercular and
external.

1 Neither Ehlers nor Pruvot and Racovitza mention these.

420 STHENELAIS LIMICOLA.

Feet. — The tufts of bristles from the first foot project somewhat outward dorsally,
obliquely inward ventrally. They all appear to belong to the dorsal type, consisting of
long, slender, finely serrated bristles with hair-like tips. The dorsal tuft is the longer ;
the inferior, moreover, presenting proportionally more of the smooth shaft at the base.
The dorsal group alone possess a spine, and Pruvot and Kacovitza consider the inferior
group as representing the ventral tuft.

The second foot has its dorsal lobe only slightly separated from the ventral, but the
spine has a special free papilla. Just above it three extremely elongated papillse
(stylodes) project from a common base. A long tuft of dorsal bristles — very finely
serrated — arises from this lobe. They are directed forwards and inwards. The ventral
division of the foot is bluntly rounded, extends further outward than the dorsal, and
bears three groups of long papillaB, each springing from a basal process. The first arises
from the tip above the spine, and has attached to it about four long papillas, the longest of
which projects further than the dorsal, since its basal process is carried out by the ventral
division. The knob enclosing the spine gives attachment also to four, while a prominent
peduncle just above the group of slender inferior bristles carries three. All these
processes are somewhat translucent, granular, and with a cellulo-granular and almost
transparent tip. The upper ventral bristles have tips of great length and tenuity, no less
than about twenty divisions occurring in the terminal appendage, which is capillary at the
tip, and, if fringed with minute algas, it resembles a pinnate hair.

The feet gradually merge into the typical form (Plate XXXI, fig. 6), which dorsally
below the branchial process shows three ciliated pads (ctenidia), and a few small clavate
papillas on the region between them and the bristles. This (dorsal) lobe is somewhat clavate
in outline, and has four or five long papillae (stylodes) from its upper terminal region, the
spine passing out inferiorly. The bristles are slender, long, finely spinous, tapered, and
form a fan directed upwards, outwards, and slightly backwards. Parasitic structures
are common on these bristles.

The ventral division is shorter and broader than the superior, and is somewhat
conical at the tip, while a large flattened and leaf-like lobule projects above the spine,
and a smaller lobule at the ventral edge. The large lobule shows a process or papilla
extending beyond it, and in its granular epidermic layer are large glandular areolaa. The
superior ventral bristles have the type of those in S. boa, only more slender. They are
simple tapering bristles with bold spinous rows at first, and then more minutely spinous
towards the slender tip (Plate XLII, fig. 1). Beneath are a few with stout curved shafts,
a short bifid terminal region, and a single articulation (Plate XLII, fig. 1), an arrange-
ment which also occurs in the following three or four segments. The next are rather
slender bristles with long tapering tips, having from nine to nearly twenty segments
(Plate XLII, fig. 2) with a minute hook at the tip. Then follows a stouter series
(Plate XLII, fig. 3) with long tips, also bifid, the shafts becoming more slender as we
proceed downward; and lastly, a much more delicate group at the ventral edge, with
from five to ten segments in the terminal region, which also has a minutely bifid tip.

The ventral cirrus is long and tapered, with a terminal segment (Plate XLII, fig. 4),
or occasionally two.

In the posterior feet little change takes place in the dorsal lobe, except a diminution

STHENELAIS JEFFKEYSIL 421

in the number of papillae. In the inferior division the flattened lobe becomes a much
smaller rounded process above the spine, and has a large papilla attached to it superiorly.
The simple spinous bristles of this lobe disappear, and a few bearing three rows of spines
on the end of the shaft and a terminal division of three segments occur, the tip being-
simple. Others show four articulations. Above and below the spine are stouter bristles
having tips of a single joint, or of two or three, strongly bifid, and this condition holds
in front in certain Norwegian examples.

The slender inferior series show very delicate tips, with about four joints and simple
hair-like extremities. The lobe beneath the spine in these feet does not diminish so
much in proportion as that above it,

Reproduction. — Specimens tossed on the beach at St. Andrews in February had
numerous large ova, so that the reproductive period would seem to be early spring,

Habits. — It appears to be a dweller in sand, and is somewhat sluggish in confinement.
The finest specimens in my collection came from St. Andrews, but those from Shetland
and Polperro are also of considerable size.

Dalyell's A.phroclita arcta may be this species, or an allied form. Unfortunately no
distinctive feature is given.

The Sthenelais leiolepis of Claparede, from Naples, approaches this species in the
form of the scale, as pointed out by the Swiss author. It is distinguished from it by the
occurrence of only one ciliated mammilla below the branchia. Pruvot and Eacovitza unite
them, and consider the distinctions rested on imperfect observations.

5. Sthenelais Jeffreysii, Mcintosh, 1876.

Specific Characters. — Head broadly ovate, with a median riclge running forward to
the base of the tentacle, which is longer and thicker than in S. limicola. Lateral regions
of the head form smooth ovoid lobes. No eyes in the preparation. The massive first
foot carries dorsally the lateral tentacle internal to the bristle-bundle, and the dorsal
cirrus externally, both extending beyond the tips of the bristles. Just beneath the
former is the lamellar process, while ventrally is the shorter and more slender ventral
cirrus. The long, smooth, tapering palpus, with its sheath-like lamella at the base —
superiorly and internally — arises below the process. Body narrow, probably about two
inches in length, and with numerous segments. Scales smooth, translucent, and devoid
of pigment in the preparation ; first pair probably rounded, rest reniform. On the
external border are long, slightly tapered papillae, perhaps more numerous in the anterior
scales, and they may disappear posteriorly. The foot has a branchial process, and three
ciliated pads (ctenidia) along the upper edge. The dorsal lobe is somewhat clavate,
bevelled at the tip, with three long papillae, and a tuft of tapering bristles bearing fine and
rather closely set rows of spines. The ventral division has a conical tip, with one or two
papillae at the apex, and one on each of the lobes. The upper ventral bristles have four
rows of spines on the distal end of the shaft, and a most delicate tapering terminal
process of twelve to fifteen articulations ending in a hair-like tip. Below the spine the

422 STHENELAIS JEFFREYS!!.

shafts are stronger, and with less delicate and less numerously jointed tips. Some have
one or two rows of spines at the distal end of the shaft, and a bifid tip, with a claw and
secondary process, the joints ranging from one to three, the latter kind being inferior.
The delicate inferior group (below the lobule) have also one or two rows of spines on
the distal end of the shaft, a long tapering terminal process of six or seven joints, and a
bifid tip. The subulate ventral cirrus extends nearly to the tip of the fleshy part of the
foot.

Synonym.

1876. Sthenelais Jeffreysii, Mcintosh. Trans. Zool. Soc, vol. ix, p. 406, pi. lxxii, f. 18, 19; pi. lxxiii,

f. 1 and 2.

Habitat. — Dredged in the Atlantic off the west coast of Ireland (Galway) in lat.
53° 16' X., and long. 12° 42' W., in 165 fathoms, Station 9, < Porcupine,' 1869.

Head (Plate XXIX, fig. 4) broadly ovate, with a median ridge running forward to the
base of the tentacle, which is a tapering organ, longer and thicker than that of S. limi-
cola. The lateral regions of the head form smooth ovoid lobes. No eyes are visible in
the preparation. The first foot carries the following organs : dorsally the lateral
tentacle (of some length) inside the bristle-bundle, and externally the dorsal cirrus, both
reaching in spirit beyond the tips of the bristles. Just beneath the former is the
lamellar process, while ventrally is the shorter and more slender ventral cirrus. The
long, smooth, tapering palpus with the sheath-like lamella at the base — superiorly and
internally — arises from the ventral aspect of the process. The structure of the parts
thus diverges from that in S. limicola, and leans to those with more numerous processes,
as for instance S. boa.

Body narrow, probably about two inches long, and with numerous segments.
Scales (Plate XXXIV, fig. 13). — First pair rounded, the rest more or less reniform,
and all smooth, translucent, and devoid of pigment in the preparations. On the external
border are long, slightly tapered papillae, perhaps more numerous in the anterior scales,
becoming fewer as we proceed backwards, and finally in the posterior scales disappearing
altogether, — that is to say, if the first form agrees with the closely allied Norwegian
species. The number of cilia (Plate XLII, fig. 5) on the external border thus varies, ten
being a usual number anteriorly, and their great length is in contrast with those of S.
boa (Plate XXXIII, fig. 16). In the latter the edge of the scale has been doubled, so
as to show (somewhat out of focus) the smaller papillse on the surface.

Feet (Plate XXXI, fig. 7). — Superiorly is the long branchial process, and three
ciliated pads (ctenidia) along the dorsal edge. The dorsal lobe is somewhat clavate,
bevelled at the tip superiorly, and bears three papilla?, one at its tip, one projecting
from the upper bevelled region, and another near it — the two latter springing from the
anterior aspect of the foot, and a tuft of the usual slender bristles with fine and rather
closely set rows of spines. The ventral division is somewhat conical at the tip, and has
one or two mammillary papillse at the apex to which the spine goes, and one on each of
the lobes — superiorly and inferiorly. These are generally constricted at the base. The
superior ventral bristles are somewhat slender, have four rows of spines on the enlarged

STHENELAIS JEFFKEYSIL 423

distal ends of the shafts, and a most delicate tapering terminal process with a hair-like
tip, and comprising fourteen or fifteen segments, the basal being much longer than the
others (Plate XLII, fig. 6, which shows a bristle somewhat compressed by others). Below
the spine are some with shafts considerably stronger, and with less delicate and less
numerously jointed tips, which, however, end in a fine point. From the spine to the
inferior lobule of the foot somewhat strong shafts of similar character are distributed,
the inferior especially showing one or two rows of spines on the dilated distal region,
while the terminal processes are of several kinds. The upper have a single terminal
division consisting of a segment with a well-marked claw and secondary process which
fills up the concavity. Just above the inferior lobule of the division are some with three
segments in the terminal region, viz. a basal two thirds the entire length, and two short
distal articulations, the last with a claw. A few have only two segments (Plate XLII,
fig. 7). The inferior series, arising below the lower lobule, are delicate translucent
bristles, having one or two rows of spines on the distal part of the shaft, and a long
terminal process of six or seven articulations (Plate XLII, fig. 8), the secondary process
filling up the hollow. Ventrally three series thus occur : (1) the stronger superior with
tapering filiform tips ; (2) the stout shafts with the short tips, of one, two, or three
articulations ; and (3) the slender inferior with long, tapering, bifid tips. All are very
delicate and translucent, and the basal region of the terminal process is often wrinkled.

The necessity for carefully regarding the nature of the bristles is well illustrated in
a very closely allied form brought by Canon Norman from Norway, and which in almost
all its characters corresponded with the present species. A glance at the bristles of the
Norwegian form showed that the dorsal series were much denser; the upper ventral
series had nine or ten rows of spines at the distal end of the shaft, and a short acutely
pointed terminal region of nearly a dozen segments. A series with more numerously
jointed (eighteen to twenty articulations) and finely tapered tips followed, the shafts having
two or three articulations in the upper examples, the rest being smooth; then a
group with a single distal segment ending in a well-marked claw and secondary process ;
below was a group with similar shafts, but with tapering jointed terminal pieces ending
in a hair-like tip ; finally, the ventral series consisted of delicate bristles, with a slender
terminal region of about five articulations and a bifid tip. The differentiation thus at
every step was made clear.

The ventral cirrus is subulate, with an articulation at the tip, and extends nearly as
far as the end of the fleshy part of the foot.

Habits.— Only a single example has been obtained, so that it would not seem to
stretch to shallower waters. Where so much difficulty exists in capture and examination
it is unsafe, however, to make statements on this head. The species is sufficiently
defined, and should easily be identified. It probably frequents a sandy bottom.

This form clearly leads through Eusthenelais to Leanira.

55

424 STHENELAIS.

6. Sthenelais ? sp.

Specific Characters. — Head unknown. Body long and narrow. Scales on every
foot in the posterior region, large, covering the dorsum, reniform in outline, with a notch
externally as well as at the hilus, perfectly smooth on surface and border. Branchial
process unusually long and straight ; three ciliated pads (ctenidia) beneath it along the
dorsal curve. The dorsal lobe of the foot is clavate, with a long, slender papilla stretch-
ing from the apex. Dorsal bristles boldly spinous, and rather long. Ventral lobe forms
an irregular spear-head, the longer slope being inferior. Above the spine is a prominent
hump bearing a papilla. Inferiorly behind the lower bristles is another elevation. The
superior bristles are slender, ends of shafts with eight or nine whorls of spikes and
apparently simple tips, with many articulations. Some with stronger shafts and shorter
tips occur below, others being slender with a minutely bifid tip. Ventrally are bristles
with a few spines at the tip of the shaft, and long six- to eight-jointed tips. Ventral
cirrus long and subulate, and reaching as far as the apex of the foot.

Synonyms.
1896. Sthenelais, n. s., Mcintosh. Sc. Proceed. K. Dub. Soc, vol. viii, n. s., p. 403.

Habitat. — South-west Ireland, long. 45°, 325 fathoms (E. I. A. Exped., 1886).

Read absent.

Body seems to be long and narrow, with prominent feet. Only the posterior region
remains.

Scales (Plate XXXIV, fig. 12) on every foot in the posterior fragment, large,
covering the dorsum of the narrow body, and reniform in outline. They are perfectly
smooth on surface and border, and thus differ from the other British species. A shallow
notch occurs at the external margin, and a more acute one at the hilus. The distribution
of the nerves is well seen.

Feet (Plate XXXI, fig. 8) with an unusually long straight branchial process dorsally,
and three ciliated pads (ctenidia) beneath it on the dorsal curve. The dorsal lobe is
clavate (narrower at the base), much bevelled dorsally at the tip, and with a long
slender papilla stretching from the apex. The dorsal bristles form a long tuft of rather
boldly serrated bristles superiorly, and they diminish towards the ventral edge. The
ventral lobe resembles an irregular spear-head, the longer slope being inferior, and the
apex from which the spine projects is prominent, and bears a papilla. Above the spine
is a prominent hump, which also possesses a papilla. Inferiorly is another prominence
behind the lower group of bristles. The superior ventral bristles are slender, the distal
ends of the shafts having eight or nine whorls of spikes, the end apparently simple — in
the form of a tapering acicular process, with a needle-like tip — a condition probably due
to repair, since others show a many-jointed needle-like tip. Bristles with stronger
shafts follow — with shorter simple tips, numerously jointed. Some of the more slender
shafts at the ventral border of the stout series present many-jointed tips, with a minute

EUSTHENELAIS HIBERNICA. 425

claw. Then follow a large fan-shaped group of most slender bristles, with a few spines
at the tips of the slightly curved shafts, and long (six- to eight-) jointed hair-like tips.

The ventral cirrus is long and subulate, and its tapering tip reaches to the apex
of the ventral lobe.

As far as can be observed this is the nearest approach to Leanira, only the more
slender forms of the stouter series of bristles in the ventral division showing very finely
bifid extremities. Unfortunately the condition of the single specimen leaves much to be
desired.

Genus XXV.— Eusthenelais, Mcintosh, 1876.

Characters as in 8the?ielais, but the scales are unknown. Dorsal bristles slender,
elongate, and finely spinous, tapering to a hair-like point. Upper ventral bristles
slender, the distal end of the shaft having seven to nine whorls of spikes, and a terminal
tapering process of fifteen to eighteen articulations ending in a hair-like tip. The
stouter bristles at and below the spine have shorter divisions to the terminal whip. The
slender group at the inferior edge has long (fifteen- to eighteen-) jointed terminal processes
with a minutely bifid tip.

EUSTHENELAIS HIBERNICA, Mcintosh, 1876.

Specific Characters. — Head broadly ovate, with a median region and two well-marked
ovoid lobes at the sides. Eyeless in the preparations. Median tentacle similar to that of
Sthenelais Jeffrey sii, with a filiform tip. At its base are two small processes (ctenidia)
which do not reach the tip of the ceratophore. The appendages springing from the
first foot agree with those of the species mentioned. The dorsal cirrus proper (external)
is slightly shorter than the median tentacle, and the tip is filiform. The shorter and
more slender lateral tentacle lies to the inner border dorsally. Beneath are two pro-
cesses ; an inner, short, broad, and blunt (cuilleron) ; and an outer, slender, tapering,
ventral cirrus, less than half the length of the dorsal. Palpi long, smooth, and tapering,
with a scoop-shaped lamella at the base superiorly and internally. Body about two
inches long, with numerous segments. Scales absent. Dorsal lobe of the foot some-
what clavate, and bevelled at the tip. It has three or four long papilke. The bristles
are slender, elongate, finely spinous, and tapered to a hair-like point. Ventral lobe
conical, with a rounded lobule in front bearing a long papilla — stretching beyond the tip
of the dorsal lobe. At the spine is another somewhat fusiform papilla, and on the
inferior lobule a longer and more slender papilla. Upper ventral bristles with slender
shafts, the distal region having from seven to nine whorls of spikes, while the terminal
tapering process has from fifteen to eighteen articulations. Near the spine are bristles
with stouter shafts, devoid of spines distally, and with shorter divisions in the terminal
process. The slender inferior series have in some a few spines on the end of the shaft,
and a numerously jointed (fifteen to eighteen) bifid tip. Ventral cirrus long and subu-
late, almost reaching the tip of the fleshy part of the foot.

426 EUSTHEXELAIS HIBERNICA.

Synonym.
1876. Eusthenelais hibemica, n.g. and s., Mcintosh. Trans.. Zool. Soc., vol. ix, p. 407, pi. Ixxiii, ff. 4 and 5.

Habitat— Dredged in the 'Porcupine' Expedition of 1869 at Station 8, off the
west coast of Ireland (Galway) in 106 fathoms, and again in the Expedition of 1870 off
Cape Sagres in the Mediterranean, in 45 fathoms.

Head (Plate XXIX, fig. 5) as in Sthenelais Jeffreysii, broadly ovate, with a median
region and two well-marked ovoid lobes at the sides. Eyeless in the preparations. The
median tentacle is similar to that of the species mentioned, and has a filiform tip. At its
base are two small processes (ctenidia), which do not reach to the tip of the basal
segment. The appendages springing from the first foot agree with those of Sth.
Jeffreysii. The external process, corresponding to the dorsal cirrus, is slightly shorter
than the median tentacle, and has a filiform tip. The shorter and more slender lateral
tentacle lies to the inner border dorsally. Beneath are two processes,— an inner, short,
broad, and blunt (cuilleron), with the flattened tip extending beyond the peduncle; and
an outer, slender, tapering, ventral cirrus, which is not half the length of the dorsal.
The palpi are long, smooth, and tapering, with a scoop-shaped lamella at the base
superiorly and internally.

Body probably about two inches in length, and having numerous segments. It is
rounded dorsally and flattened ventrally.

Scales absent. In all probability the scales will be found to present long papillae on
their external border — if we may judge from its close resemblance to Sthenelais Jeffreysii
and its Norwegian ally, as well as to Sth. fuliginosa, Claparede. 1

Feet. — The second foot bears a scale and a long ventral cirrus, and the third a
dorsal cirrus of considerable length, while the fourth has a branchial process arisinp
from the homologous base. The bristles in the second foot have already mapped them-
selves out more or less as they occur behind, though of course less distinctly. The
branchia is short anteriorly, but gradually increases in length till about the middle of the
body. A single large ciliated pad (ctenidium) exists on the dorsal edge of the foot, and
three smaller in the curve below the cirrus. In the typical foot (Plate XXXI, fig. 9) the
dorsal lobe is somewhat clavate, and bevelled at the tip superiorly. It bears three or
four long papillse. The dorsal bristles are slender, elongate, and tapered to a fine hair-
like point. All are finely spinous, one series more distinctly, and another less distinctly
so. The ventral lobe is conical at the tip, has a rounded lobule in front, bearing
a very long papilla, which extends beyond the tip of the dorsal lobe. At the spine is
another somewhat fusiform papilla, and on the inferior lobule a longer and more slender
one. The superior ventral bristles have rather slender shafts, and the distal ends are
furnished with from seven to nine whorls of spikes. The distal region is a slender
tapering process of from fifteen to eighteen joints, ending in a capillary tip. These
resemble the bristles of Sthenelais and Sigalion rather than those of Leanira, since the
necklace-like canaliculi are absent. The figure (Plate XLII, fig. 9) represents a bristle
adjoining the superior lobe, the spinous rows on the tip of the shaft being more numerous,
while the jointed terminal region is shorter.

1 ' Ann. Cha3t. Nap./ p. 94, pi. iv, ^g. 2.

SIGALION MATHILDA. 427

At and below the spine are a series of bristles having stouter shafts, without spines at
the distal ends, and bearing shorter divisions to the terminal whip. The slender group at
the ventral border of the foot, that is below the inferior lobule, have in some cases one
or two spines at the end of the shaft, and long tapering and numerously (fifteen to
sixteen) jointed terminal regions, with a minute claw and secondary process at the tip
(Plate XLII, fig. 10). These bristles also lean to the two forms mentioned, and not to
Leanira.

The ventral cirrus is long and subulate, reaching almost to the tip of the fleshy part
of the foot. It has an articulation at the extremity.

Habits. — This species would seem to frequent sand in deep water, but it may yet be
found inshore.

It is closely allied to Claparede's Sthenelais fuliginosa.

Genus XXVI. — Sigalion, Audouin and M. Edwards, 1830.

Head elongate from before backwards. Median tentacle absent ; lateral tentacles
short and papilliform, fixed to the anterior part of the cephalic lobe. Scales covering
the dorsum ; processes pinnate or with long papillae from the axis. The feet resemble
those of Sthenelais. Dorsal division clavate and furnished with a papilla. Bristles as
in Sthenelais. Ventral division somewhat truncate, and with a papilla internal to the
bristles, which are all bifid. Branchiae on every foot 1 (in S. Mathildse). Nerve-trunks
rounded in section, and the granular epidermic area enclosing them is expanded
inferiorly. Segmental eminence placed ventrally at the base of each foot.

1. Sigalion Mathilda, Audouin and M. Edwards, 1834.

Specific Characters. — Head elongate from before backward, with a truncate an-
terior border, having a slight peak at each side. Behind the border are two pairs of
small black eyes, the anterior nearer each other and larger. Body elongate (three
to five inches), little tapered anteriorly, and gradually diminishing posteriorly.
Segmental eminence situated ventrally at the base of each foot. Colour greyish-
brown or pinkish. Proboscis with seventeen to eighteen papillae on each lip.
Scales about sixty-four pairs, adherent, somewhat quadrate or rhomboidal, with
the posterior and inner corner rounded off. The processes along the external
border are pinnate, with long cylindrical papillae (about twenty on each side),
and at the base these are irregularly distributed around the central axis. Foot
with a club-shaped dorsal division, which has a single subulate papilla, and simple,
tapering, finely spinous bristles— increasing in strength externally. The inferior

1 Cams, in his ' Prod. Fauna Metfit./ states that in the anterior part of the body the branchiae
alternate with the elytra, but that posteriorly they occur on all the feet.

428 SIGALION MATHILDA.

division is somewhat truncate, sloped from above downwards and inwards, and has a
small papilla internal to the bristles, which superiorly are whorled, then finely serrate at
the tip. Next follow larger bristles, with the distal end of the shaft curved and spinous.
The distal region is segmented, and tapers to a fine bifid point (minute beak). Ventrally
is a dense group of similar forms, but without spines at the end of the shafts. They
are most slender at the ventral ed^e.

Synonyms.

1 830. Sigalion Mathildx, Cuvier. Eegne An., 207.

,, „ „ Bathke. Fauna Norweg., 151.

1834. „ „ Aud. and Edwards. AnneL, 105, pi. ii, ff. 1—10.

1840. > „ „ Grnbe. Actin., Echinod., &c, p. 84.

1848. ,, „ Gervais. D'Orbignv, Diet. d'Hist. Nat., xi, p. 601.

1851. „ „ Grube. Earn. d. Ann., p. 38.

•1855. „ „ Peters. Archiv f. Naturges., vol. xxi, p. 38.

1857. „ „ Kinberg. Eugen. Eesa, vol. ii, Zool., 2, p. 9.

1865. „ „ De Quatrefages. Ann., i, p. 279.

» „ Carringtonii, Carrington. Proceed. Lit. and Philos. Soc. Manchest., vol. iv, p. 179.

1875. „ Mathildse, Mcintosh. Invert, and Fishes St. Andr., p. 118.

1876. „ „ idem. Trans. Zool. Soc., vol. ix, p. 408.
1884. „ „ Y. Cams. Faun. Medit., p. 204.

1898. „ squamaium, De St.-Joseph. Ann. d. sc. nat., Zool. (8), vol. v, pp. 239, 241, pi. xiii, ff.

22—29.

Habitat. — Generally distributed round the British shores. Of large size and rather
abundant between tide-marks in the Channel Islands and at St. Andrews, while those
from Shetland are small. Most of the specimens are littoral, generally under stones,
though occasionally small pale specimens occur in sand, as at Southport. It extends,
however, to some depth in the adjoining waters. It is' occasionally tossed on the sands
at St. Andrews in considerable numbers after storms, and is also not uncommon in the
stomachs of cod, haddock, and flounders (E. M.). The first two are especially partial
to this species. They sometimes take refuge in tubes of Terebella when cast ashore.
The species seems to be partial to sand. In the ' Porcupine ' it occurred off the Algerian
coast between Capes Falcon and Tenez.

Head (Plate XXIX, fig. 6) somewhat elongated from before backwards, oblong or
cylindrical in outline. The anterior border is truncate, with a slight peak at each end.
Behind the latter are two pairs of small black eyes, situated close together on each side
of the middle line, the pairs not far apart, The anterior pair are nearer each other and
also somewhat larger than the posterior pair. From the elevation of the head they look
forward and upward. Posteriorly only a transverse furrow marks the boundary of the
head. The head bends downward in front, and has soldered to it the two short and
somewhat conical lobes of the first feet, which bear externally (dorsally and ventrally)
short subulate cirri, and dorsally and internally a tuft of very slender, tapering bristles,
most minutely serrated and directed upwards and inwards. Beneath is the long
tapering palpus, the tough cuticle of which is usually thrown into fine transverse

SIGALIOX MATHILDAS. 429

wrinkles, so that it resembles in a decayed specimen the trachea of an insect. The head
is thus raised above these processes, and is confined to the dorsal region. A short
conical process on the inner and upper border of the foot may indicate the lateral
tentacle.

Body three to five inches in length, and with segments about as numerous as in
Sthenelais boa (180), elongate, very little tapered anteriorly, so that in the preparations
it is almost truncate, and gradually diminishing posteriorly, though terminating in by no
means a slender tail, from the tip of which a remarkably long and slender caudal style or
cirrus extends. The functions of this delicate appendage would appear to be sensory,
and it is curious that the tail is often specially modified in sand-dwellers, such as
Nephthys, certain Opheliidx, and Spionidae. The dorsal arch is slightly rounded laterally,
flattened in the middle when the scales are present, the prominent papillae for the latter
occupying most of the surface, and in the ripe forms showing ova through the walls.
The ventral surface is in life flattened, and in the preparations usually presents the
aspect of a long riband with rounded edges. A segmental eminence occurs at the base
of each foot, but no distinct papillae could be made out.

In life the colour of the dorsum is greyish-brown with a central bluish-green mark,
or dull greyish-white or pale pinkish in front with a dark grey stripe from the proboscis.
In some a pale brown streak marks the inner margin of each scale. The ventral surface
is iridescent, bluish-green or pale pinkish, with the red central vessel.

Proboscis. — The number of papilla on each side (dorsally and ventrally) is about
sixteen, though from overlapping one or two more may occur, e.g. seventeen or eighteen.
The teeth bite alternately, the lower passing to the right of the upper.

Scales (Plate XXXIV, fig. 14) about sixty-four pairs, and of the hue already
mentioned. They are firmly adherent. The surface is smooth, and they have the
external border supplied with a series of pinnate processes. The first pair are smaller
and somewhat triangular in outline, with a series of the typical pinnate processes along
the outer border. Moreover at the bases of these are a few simple papillae, and in some,
instead of the pinnate process, a group of long, simple papilla? occur at one end of the
series. The scales gradually assume a quadrate outline, the outer border bearing the
papillae being nearly straight, while the posterior and inner corner is rounded off. The
processes 1 (Fig. 83) have a stout central stem, which tapers to a slender point. The
axis is granular. The pinnae or papillae arise on the scale even below the process, and
are continued on it somewhat irregularly at the base, but by-and-by assume a more
uniformly pinnate arrangement. The papillae are pale throughout. Occasionally one of
the processes is bifid. In vertical section the dorsal and ventral coverings of the scale
are joined by a close series of fibrous strands. Both surfaces are remarkably smooth.
As we proceed backwards the scales become more elongated transversely, but again
diminish posteriorly.

The first scale shows no branchial process, but all the rest are provided with such,
the organ being long and sickle-shaped as well as richly ciliated.

Feet. — The second foot is bifid, presenting an elevated dorsal lobe with a tuft of
simple, tapering, finely serrated bristles as in the first foot, the spine issuing at their

1 < Ann. Nat. Hist./ August, 1898, pi. ii, f. 14.

430 SIGALION MATHILDA.

lower border, and dorsally having a papilla for the scale. The inferior lobe is irregularly
conical, the upper slope ending in a rounded process pierced by the spine, and the margin
trending downward and slightly backward from this to the abrupt angle inferiorly. The
upper bristles have slender shafts with slightly dilated extremities and an oblique edge, and
present only traces of rough processes, the long tapering distal region having numerous

Fig. 33.— Papilla of scale of Sigalion MatJdldce, Aud. and Ed. x 280.

segments, the tip apparently being simple. The group below the spine is similar, except
that the ventral series show a distinct spine or two towards the end of the shaft. The
ventral cirrus is subulate, and projects beyond the tip of the fleshy part of the foot.

The third foot has a similar form, but papillas occur dorsally (a conical upper and
two boss-like inferior), and ventrally is a rounded papilla internal to the cirrus. The
ventral bristles now present a claw at the tip,

A typical foot (Plate XXXI, fig. 10) shows dorsally three ciliated pads (ctenidia)
along the dorsal edge. The dorsal lobe projects somewhat further outward than the
ventral, is club-shaped, with a sinus superiorly, and a long, slightly tapered papilla a
little below the dorsal bristles. The normal extremity of the division seems to be tilted
upwards, the bristles standing obliquely upwards and backwards from the dorsal edge.
Internally they commence as extremely fine hair-like forms, with very minute spikes.
Towards the outer edge these simple bristles become much stronger, show slight longi-
tudinal striations of the shaft, and a curvature where the spinous distal region commences.
They then taper to a very slender extremity. De Saint- Joseph 1 states that these bristles
end in a bifid point, an error of interpretation probably due to imperfect specimens. The
minute spikes project upwards and outwards when the bristle is in position (Plate XLII,
figs. 11 and 12). Fine transverse or oblique lines indicate the rows of spines. The spine
passes out below the bristles and above the papilla.

The inferior division of the foot presents superiorly a small papilla internal to the
bristles, a somewhat short series of bristles (Plate XLII, fig. 13), the tips having
proximally a series of longer spines in whorls, five or six of which are very distinct, and

1 Op. cit., 1898, p. 239.

SIGALION BUSKII. 431

a tapering tip with finer spines. The bristles below these have larger shafts — dilated
and slightly curved towards the extremity (Plate XLII, fig. 14), with numerous rows of
spines on the convex edge. The distal region is jointed, and terminates in a beak or
claw. Then follows a dense tuft of similar compound bristles without spines at the tip
of the shaft, and a long, jointed, tapering, terminal region with a minute claw (Plate
XLII, fig. 15). They become more slender at the ventral margin (as in Plate XLII,
fig. 16).

Besides these slender forms there are, below the spine, one or two somewhat stouter
bristles, which show a single segment in the terminal bifid region (Plate XLII, fig. 17).
The ventral cirrus is long and slender — reaching to the tip of the fleshy part of the foot
— in spirit.

The foot at the posterior end retains much of the foregoing structure, both as
regards the fleshy parts and the bristles. Those at and above the spine of the ventral
division are somewhat stronger, and the ventral cirrus is proportionally longer than in
front.

The branchiae commence on the fourth foot, and occur apparently on every foot.
The statement of Carus, therefore, that in Sigalion the branchiae alternate with the
scales anteriorly is not borne out in this species.

Reproduction. — A fragmentary specimen procured off the Algerian coast had well-
developed ova on August 27th (' Porcupine,' 1870).

Habits. — In confinement its habits are similar to those of Sthenelais boa.

This Annelid, which formed the type of the family, was found by Audouin and
Milne Edwards at the islands of Chausey, and measured about five inches long by three
or four lines broad, and with 180 segments.

The scales of this species and the general characters approach very closely the
Sigalion squamatum of Delle Chiaje, even closer than Claparede — the able interpreter of
some of the more doubtful Annelids described by the Italian naturalist — supposed,
especially since the difference in regard to the eyes has now been removed. Neither
Delle Chiaje nor Claparede, however, show the ciliated mammillae on the dorsal edge of
the foot, so that this doubtful distinction alone remains. In his supplemental volume
Claparede specially refers to the distribution of the nerves in the papillae for the scales,
and also in the periphery of these organs, thus giving them extreme sensibility. He
mentions the occurrence of certain spicules or rod-like bodies on the pinnae of the
papillae, probably of a parasitic nature ; but such have not occurred in the British
species. The males of S. squamatum were whitish, the females of a fine rosy hue — due
to the eggs. Leidy's 1 8. Mathildas from New Jersey is a Sthenelais.

2. Sigalion Buskii, Mcintosh, 1869.

Specific Characters. — Head somewhat pear-shaped, broad in front and narrow
posteriorly. Body stouter than in S. Mathildas, pale greyish in spirit. Scales more or

1 ' Marine Invert./ &c, p. 148, pi. ii, f. 53.

56

432 SIGALION BUSKH.

less quadrate anteriorly, with the inner edge rounded. The straight external edge has a
series of about sixteen pinnate processes, besides a few simple papillae posteriorly. Each
process has a granular stem, to which is attached a series of lanceolate granular lamellae
with a narrow papillary tip. In the typical foot the inferior lobe is shorter than in 8.
Mathildas, broadly clavate, and with a single papilla on its tip. The bristles are attached
to the whole tip (i. e. superiorly, terminally, and inferiorly), and are long and minutely
spinous. The ventral lobe is devoid of the superior papilla seen in S. Mathildas, and has
a more prominent process for the spine. The compound bristles (with jointed tips
t ^-below the forms with the short tapering spike) have the terminal portion of the shaft
covered with whorls of somewhat sparse spikes, which are more numerous than in
8. Mathildas, while the stouter bristles beneath have close rows of minute spines on the
same portion of the shaft. One or two below the spine-papilla have short tips of one or
two segments. The slender ventral series have only about two whorls of spikes on the
.end of, the shaft, and the long jointed tips have, as in the upper series, minute beaks.
The ventral cirrus is long, slender, and subulate, extending beyond the fleshy part of
the foot.

Synonyms.

1869. Sthenelais dendrolepis ? Mcintosh. Trans. E. S. E., vol. xxv, p. 409, pi. xii, f . 12; pi. xv,

f. 4, 5.
1876. Sigalion BusMi, Mcintosh. Trans. Zool. Soc, vol. ix, p. 391, pi. ]xx, f. 14.

Habitat.— Dredged in 90 fathoms off North Unst, Shetland, by Dr. Gwyn Jeffreys, in
1867.

Head (Plate XXIX, fig. 7) somewhat pear-shaped in outline, broad in front and
narrow posteriorly, where it is only separated from the nuchal collar by a slight furrow.
The anterior border is smoothly rounded, and abuts on the conical firstfoot. It is quite
pale, no eye-specks being visible in the preparation. In the example a small and some-
what clavate papilla (tentacle) projected from the centre of the anterior border of the
head— a remarkable condition in Sigalion. 1

Body somewhat stouter than in Sigalion Mathildas, and in this, respect resembling
Sthenelais boa, but having the same general shape both dorsally and ventrally. The
example is incomplete, but it would seem to reach a similar length to the species first
mentioned. The segmental eminences exist at the bases of the feet posteriorly, but no
distinct papilla is visible. The general colour of the dorsum is pale greyish.

Scales. — The first pair of scales are almost ovoid, only the outer border is broader
than the inner, and has eight pinnate processes besides a smooth projection in front,
and some isolated papillaa posteriorly. Except at the latter part the margin is quite
smooth. At the bases of the processes, however, isolated papillas occur as in 8.
Mathildas, one or two of which are attached to the central axis as in the species just

1 The head had been injured, so that the question might be raised as to whether this was not
one of the lateral tentacles pushed out of position. After careful examination, however, the view
above mentioned was held.

SIGALIOU BTJSKII. 433

mentioned. No branchial process occurs on the papilla supporting this scale, — indeed, it
only appears on the papilla of the fourth scale on each side.

Passing backward the scales assume a somewhat quadrate outline, which in a typical
example anteriorly has the inner end rounded, while the external border has about
sixteen beautifully pinnate processes, besides a few simple papilla? at one end (Plate
XXXI Y, fig. 15). The pinnate processes have a tree-like figure — more robust than in
8. Mathildas, and the pinna?, instead of being hyaline cylindrical processes, are lanceolate
granular lamella? with a narrowed papillary tip (Plate XXYI, fig. 9). The posterior
scales are longer transversely, and have fewer pinnate processes, bnt the structure is
essentially the same. The granular epiderm (hypoderm) of the curve between the pro-
cesses is denser, more opaque, and in this condition extends up the stalk to the first pinna?.

Feet. — The first pair project forward just beneath the head, and are bluntly conical
with a single spine, which passes forward and upward near the dorsal cirrus— a slender
and subulate organ, with a slightly dilated tip in the preparation. It springs from the
outer and upper lateral region of the foot. The ventral cirrus arises from the lower
part of the foot, somewhat internal to a vertical line from the former, to which it is
similar in outline. The tuft of slender simple bristles is directed forwards and upwards ;
they are minutely spiked. The palpus is fused with the base of the foot inferiorly, and
is shorter proportionally in the specimen than in S. Mathildas. It is finely tapered
and quite smooth. Only a blunt papilla occurs in the region of the lateral tentacle.

The second foot agreed with that in the former species, presenting a well-marked
though short dorsal lobe, with a spine and a tuft of simple bristles, and a somewhat
trifoliate ventral lobe with a series of bristles. These have long slender shafts with a
few spikes on the dilated distal end, and a long jointed and finely tapered distal region,
with a slight trace of a bifid (beak-like) tip.

In the typical foot (Plate XXXI, fig. 11) the superior lobe is shorter than in
S. Mathildas, and broadly clavate, the expansion of the tip in lateral view being greater
than in the former species. It bears a brush of simple minutely spinous bristles, which
spring from the superior and inferior borders as well as the tip. The stronger and
longer forms arise above the spine ; the more slender occur for the most part below it, and
especially the inner group. The papilla at the tip of the superior lobe appears a short
distance behind the front. Three ciliated pads (ctenidia) exist on the superior border
between the branchial process and the tip. A stalked infusorian is present on the bristles.

The ventral division is devoid of the superior papilla observed in S. Mathildas, and
has a more prominent papilla for the spine. The ventral bristles are characterised, in
contrast with S. Mathildas, by having rather longer terminal processes with bifid tips.
The superior series which adjoin the forms with the short tapering spinous region have
the terminal portion of the shaft covered with whorls of somewhat sparse spikes
(Plate XLII, fig. 18), which are much more numerous than in S. Mathildas, while the
stouter bristles below have close rows of minute spikes on the same portion of the shaft
(Plate XLII, fig. 19, and in some the rows are closer than in the example figured). One
or two below the spine-papilla have only a single articulation in the terminal region,
while others have two, and the ends of the shafts in these are smooth, or in the former
with only traces of spinous rows. The slender ventral bristles have generally about two

434 LEANIRA HYSTRICIS.

distinct whorls of spikes at the end of the shaft, and the basal division of the terminal
process is broader than the latter.

The ventral cirrus is slender and subulate, and the tip extends beyond the fleshy
part of the foot.

Reproduction. — The species is a female with a few ova, the majority probably
having escaped either before or after capture in July.

This seems to be a deep-water species. It is named in honour of the late Prof. G.
Busk, whose patient and accurate investigations in various groups are so well known,
and whose genial interest in many a young worker in science will long be gratefully
remembered;

An allied form is the Sigalion Edwardsi of Kinberg 1 (1858), from the Atlantic — off
La Plata.

Genus XXVII. — Leanira, Kinberg, 1857.

Anterior border of the head fixed to the bases of the feet above the palpi, furnished
with a somewhat slender median tentacle, usually with a short process at each side of
the base ; lateral tentacles minute ; tentacular cirri unequal.

Scales not covering the dorsum anteriorly, smooth. Ciliated pads (ctenidia) along
the dorsal arch above the feet. Dorsal lobe of the foot rather more prominent than the
ventral, papillose, and with long spinous bristles. Inferior division has compound
subulate bristles, with the tapering terminal region pectinate-canaliculate. Branchiae
commence in front, and are attached to the scale-peduncle on the dorsum of each foot.
Nerve-cords somewhat flattened or ovoid in section, and the area covered by the oblique
and vertical muscles. Segmental organs as in the group.

The genus was established by Kinberg in 1857, while Grube produced additional
features in his ' Annulata Semperiana.'

V. Carus, again, 2 includes Leanira under Sthenelais, but does not allude to the
condition of the bristles.

Leanira hystricis, fflilers, 1874.

Specific Characters. — Head smoothly rounded. The awl-shaped median tentacle is
remarkably short and small. No eyes in the preparations. The anterior border of the
head is fixed to the bases of the feet above the palpi, which are long, tapering, and
smooth, with scoop-shaped lamellae at their bases. The foot above the palpus bears
three processes : superiorly a tentacular cirrus about a fifth the length of the palpus ;
inferiorly a minute organ of the same nature, extending only a short distance beyond the
peduncles ; and a minute awl-shaped process (lateral tentacle) attached to the base of
the peduncle superiorly. The body is small and slender, and between two and three
inches in length. Scales rounded, translucent, and perfectly smooth in outline and

1 Op. cit., p. 30, Taf. 9, f. 41, &o.

2 f Fauna Mediterranean (1884).

LEANIRA HYSTRICIS. 435

surface. The foot presents a superior lobe rather more prominent than the ventral, and
bears long, simple spinous bristles and two papilhe. The inferior division of the foot
has two somewhat larger papillae, and the characteristic bristles with the tapering
extremities and moniliform markings. The branchial process begins as a minute organ
on the twenty-fourth foot.

Synonyms.

1874. Leanira hystricis, Ehlers. Ann. Nat. Hist. (4), vol. xiii, p. 292.

1875. „ „ idem. Annel. c Porcup./ 1869, Zeitschr. f. wiss. Zool., xxv, p. 35, Taf. 2,

f. 5—11.

1876. „ „ Mcintosh. Trans. Zool. Soc, vol. ix, p. 408, pi. lxxiii, f. 6—8.
1885. „ „ idem. Ann. f Chall./ vol. xii, p. 155, pi. xxiii, f. 9.

Habitat. — Dredged in the 'Porcupine' Expedition of 1889, off the south-west of
Ireland (Station 2), at a depth of 808 fathoms on a bottom of soft sticky mud;
also at Stations 23, 87, and 20, the last at a depth of 1443 fathoms, generally on mud
and Grlobigerina ooze. In the c Challenger ' it was got off the Azores at depths of 900
and 1000 fathoms, and in the 6 Knight-errant ' in 530 fathoms. Yerrill enters it in his
list from Cape Cod to the St. Lawrence.

Head (Plate XXVIII, fig. 17) smoothly rounded, little elevated, and slightly dusky
from the presence of pigment along the anterior border. The median tentacle is
remarkably short and small, shaped like the handle of an awl — narrow at the base,
dilated in the middle, and tapering to a blunt tip. No eyes are visible in the prepara-
tions. The anterior border is fixed to the bases of the feet above the palpi, which arise
close together on each side of the middle line inferiorly. They are long, tapering, and
smooth, and at the base of each, towards the inner and ventral surface, is the scoop-
shaped lamella. Immediately above and soldered with the base of the palpus is the first
foot, bearing three processes, viz. superiorly a tentacular cirrus about a fifth the length
of the palpus ; inferiorly a minute organ of the same nature, and extending only a short
distance beyond the peduncle ; and a minute awl-shaped process (lateral tentacle)
attached to the base of the peduncle superiorly. The latter is similar in form to the
median tentacle near which it is placed. Inferiorly the oral aperture has prominent
rugose lips, with a blunt papilla on each side of the median fissure in front. None
showed traces of the bristles usually present in allied forms on the bases of the
tentacular cirri.

Body comparatively small and slender, none of the incomplete specimens from the
'Porcupine' measuring more than an inch; but an example from the ' Challenger '
reached about two inches, and was also incomplete. The external appearance of the
body agrees with that in Sthenelais.

Scales. — The first and second scales are small and rounded. The rest are also
more or less rounded, translucent, and perfectly smooth in outline and surface.

Feet. — The second foot is directed forward, its dorsal division being represented by
a rounded process, which bears about a dozen digit-like papilla? and a series of fine
bristles resembling those of the ordinary foot (Plate XLII, fig. 20). The ventral lobe
has superiorly and inferiorly a papilla larger than those of the dorsal lobe. The bristles

436 PHOLOE.

have whorls of spikes on the distal region of the shaft and tapering extremities (Plate
XLII, fig. 21), the stoutest being next the spine, and the more slender forms with spines
at the distal end of the shaft occurring at the ventral border.

The superior lobe gradually increases in size until, in the typical foot (Plate XXXI,
fig. 12), it projects about as far as the ventral, and the bristles become stronger and
longer. A few smooth hairs occur in each bundle. The digit-like papillas, however,
diminish in number as a rule, only two occurring in each division of the foot in the
middle of the body, those of the inferior lobe being the larger. Posteriorly (Plate
XXXI, fig. 13) the superior division has three or four papillse above the bristle-bundle,
the inferior frequently only a single large, pedunculated, clavate process. The ventral
cirrus is also reduced in size. The inferior bristles of the ventral series of the same
region have a more distinct enlargement at the distal end of the shaft (Plate XLII,
fig. 22).

]STo branchial process occurs until the twenty-fourth foot, and then it is minute.
Posteriorly it gradually elongates, so as to extend outward as far as the tip of the foot.

The bristles are somewhat shorter and proportionally stouter than in Leanira
Yhleni, Mgrn.; no ciliated pad exists on the dorsal edge of the foot, and the papillas of
the latter do not show the disparity in size characteristic of L. Yhleni. The ventral
cirrus is also shorter, and in the preparation shows no process at the base.

The species diverges from L. tetragona in regard to the tentacle, bristles, and other
parts.

Habits. — A deep-water species.

At the anterior end of a fragmentary specimen a crustacean parasite was fixed in
the dorsal muscles.

Bhlers gives a detailed description l with figures of this species, his largest example
being only 19 mm. long. He shows the tentacle of three segments, or at least it is thus
figured and described. Eyes absent. He adds nothing novel to the description in the
* Transactions of the Zoological Society.' He observes that the species approaches the
L. Quatrefagesi of Kinberg from the Atlantic, off the La Plata. 2 A more minute
investigation of Kinberg' s specimen, indeed, is necessary before all doubts as to the
specific separation are removed. They closely agree in regard to the structure of the
scales, bristles, and general condition of the head and its processes, and seem to differ
chiefly in the structure of the feet and in the absence of eyes in the British form.

Genus XXVIII. — Pholoe, 3 Johnston, 1839.

Head furnished with a single short median tentacle. Two pairs of eyes, more or
less connate. Body linear-oblong. Scales on alternate segments in the anterior part of

lf Zeitsch. f. w. Zool./ 1875.

2 ' Freg. Eugen. Resa/ p. 30.

3 Carus, in the c Prodromus Faunas Mediterraneaa/ includes this genus under the sub-family
SigalioninsD, Pholoe being characterised by having a linear-oblong body, elytra on alternate segments

PHOLOE MINUTA. 437

the body; posteriorly a pair on each segment. First foot with two tentacular cirri;
without bristles. Dorsal lobe of the foot with slender, tapering, spinous bristles.
Inferior division with stout, falcate, compound bristles. Nerve-cords forming triangular
flattened areas in section on each side of the median line, the oblique muscles passing to
their insertion above them.

De Quatrefages (1865) gave the group three "antennae," a facial tubercle, and two
tentacles, the upper of which was bifid. Alternate scales anteriorly, while posteriorly
all the segments had scales. No dorsal cirri. He thus had an imperfect acquaintance
with the genus. Grrube's account 1 was more accurate.

Pholoe minuta, 0. Fabricius, 1780.

Specific Characters. — Head somewhat rounded, bearing a short subulate median
tentacle, with a few papilhe on its surface. Two eyes on each side — connate, the
anterior being the larger. There are two short tentacular cirri, also with a few small
papillas on their surface. Two prominent papillae project behind the eyes, and sometimes
overlap them. The palpi are somewhat massive, short, tapering organs, with a smooth
surface. Body small, forty-five to seventy segments, and about three-quarters of an inch
long as a maximum. It -has two slender styles posteriorly. Scales ovate or reniform,
with a series of cilia having moniiiform tips along the outer border, and more sparsely
along the posterior border; while the surface, especially at the inner region, is areolated.
Dorsal lobe of the foot forms a prominent process with a convex margin externally for
the dense tuft of bristles, which are slender, tapering, and spinous. Yentral lobe an
oblique cone with numerous papillas over the surface. The shafts of the bristles are
stout, the distal convexity having numerous spikes. The terminal piece is short and
falcate, and the edge is generally spinous. The ventral cirrus is short and tapering.

Synonyms.

1776. Aphrodita long a, 0. F. Miiller. Prod. Zool. Dan., p. 218, n. 2646.

1780. „, „ 0. Fabricius. Fauna Gronl., p. 313, n. 293.

„ „ minuta, idem. Ibid., p. 314, n. 294.

1820. Polynoe minuta, Savigny. Syst. des An., p. 26.

1828. Palmyra ocellata, Johnston. Zool. Journ., vol. iii, p. 329.

1834. „ minuta, Aud. and Ed. Ann., p. 95.

1839. Pholoe inornata, Johnston. Ann. Nat. Hist., vol. ii, pp. 437, 438, Tab. xxiii, f. 1 — 5.

1843. „ minuta, Oersted. Gronl. Armal. Dorsib. (Danske vid. Selsk., Afh. x), p. 169, Tab. i,

f. 3, 4, 8, 9, 16.

,, „ baltica, idem. Annul. Dame. Consp., p. 14, f. 21, 34 — 36, 40.

in the anterior part of the body; posteriorly a pair on each segment; rami of the feet connate;
superior bristles capillary, inferior falcate. He includes also Sigalion, Psammolyce, and Sthenelais, in
the order given, under the Sigalioninge.
1 ' Sitz. d. Schleisch. Gesell./ 1874.

438 PHOLOE MINUTA.

1844. Pholoe assimilis, Oersted. Kroyer Nat. Tidskr. Anden Baekke, Bd. i, p. 404.

1851. „ minuta, G-rube. Earn. d. An., 38.

1865. „ inomata, Johnston. Cat. B. M., p. 121, pi. xiii, f. 1 — 5; and Baird, p. 340.

„ „ minuta, Malmgren. Nord. Hafs-Ann., p. 89, Tab. xi, f. 13, and An. Poly., 17.

„ „ „ De Quatrefages. Hist. Nat. Anneles, vol. i, p. 188.

,, „ inomata and baltica, idem. Ibid., p. 190.

1873. „ minuta, Mobius. Jahresb. Com., 1871, p. 112.
„ „ „ Metzger. Ibid., p. 175.

„ „ „ Ehlers. Sitzungsb. Phys.-med. Soc, Erlangen, vol. v, p. 8.

1874. „ „ Malm. G-oteborgs Kongl. vet. o Yitt. Samhalles Handl. Ny Tidsfoljd,

Haftet xiv, p. 76.

1875. „ „ Mobius. Jahresb. Com., 1872, p. 167.

1878. „ „ Lenz. Jabresb. Com. Anhang, 1874, p. 12.

1879. „ ,, Theel. Annel. Nov. Zemb. Kongl. Sv. vet. Akad. Handl., vol. xvi, p. 24.
„ „ „ Tauber. Ann. Danic, p. 84.

1883. „ inomata, Levinsen. Nord. Annul., p. 199.

1884. „ „ Webster and Benedict. Ann. Mass., p. 701.
1886. „ „ Marenzeller. Porif., &c., Jan Meyen, p. 12.
1891. „ ,, Hornell. Op. cit., p. 239.

1896. „ eximia, Micbaelsen. Polych. Fauna, p. 12, pi. i, f. 2.

1897. „ minuta, Mcintosh. Ann. Nat. Hist., ser. 6, vol. xx, p. 169.

1898. „ „ Michaelsen. Gronland. Ann., p. 122.

Habitat. — Everywhere on the shores of Britain from Shetland to the Channel
Islands, where it attains the maximum size, and from the tidal region to the coralline
ground. It lurks under stones between tide-marks, especially in pools. It is some-
what difficult to find, and perhaps is best obtained by placing suitable stones in vessels
of sea water overnight, and then examining the water-line next day.

The variety inomata is more common on the eastern coast, whilst the variety eximia
is more frequent in the west and south.

The species extends to Jan Meyen and to the shores of America (Verrill). Lenz
found it in the grass-wrack region of the Baltic, and Michaelsen in the tubes of
Sabellaria sjpinulosa at Heligoland.

Head (Plate XXIX, fig. 8) somewhat rounded and comparatively small, having
anteriorly a rather short median tentacle with a few papillse on the surface. The eyes
are four in number ; but as the pigment of the pairs touches on each side, they are
connate. 1 Lateral tentacles absent. Two prominent papillse project immediately
behind the eyes. The palpi are somewhat massive, short, tapering organs, with a smooth
surface. Two short, tapering, tentacular cirri occur on each side, with small papillse
sparsely distributed on the surface.

Body of forty-five to seventy segments, and about three-quarters of an inch as a
maximum length, small, elongated (almost linear), but slightly more diminished pos-
teriorly than anteriorly in young specimens. The dorsum is convex, the ventral surface
flattened, with a median groove in the preparations. In life the dorsum is of a pale
pinkish colour, slightly grained with brownish on some of the scales. A reddish mark
occurs in front, with a dark greyish patch a little behind. Some, as Dr. Johnston

1 Malmgren observed that in P. minuta the eyes were four, and approximated.

PHOLOE MINUTA. 439

observes, are of a yellowish-brown colour, dusky along the sides. The centre of the
dorsum is uncovered by the scales. Posteriorly are two slender, subulate styles.

Proboscis 1 forms a short muscular organ with teeth, as in the Sigalionidae (biting to
the left), and nine short but distinct papillae along each arch. Moreover in extrusion,
just behind the lateral furrow separating the dorsal and ventral arches, are two papillae.
A median and two lateral bosses (elevations) are also present in the basal segment when
viewed from the dorsum.

Scales (Plate XXXIY, fig. 17). — The first pair in the variety inomata are somewhat
rounded, as if an isosceles triangle had its angles smoothly removed. The scar for
attachment is situated nearer the posterior than the anterior border. The latter has
numerous short clavate cilia along its edge — to the number of about fifteen, while the
posterior border has about nine longer cilia, somewhat moniliform in outline from
constrictions. The surface of the scale anteriorly has also a row of cilia internal to that
along the border, and a few are scattered in the area in front of the scar, All the cilia
have traces of palpocils at the tip. Only the inner border of the scale is smooth.

In contrast with the first scale of Pholoe minuta, Fabr., from Greenland, the fore-
going has fewer cilia. Thus there are upwards of forty along the anterior border of the
arctic form, and twelve, proportionally shorter than in P. inomata, along the posterior
edge. Moreover these organs are more numerous on the surface in front of the
scar.

The shape in the second pair becomes transversely elongated with an anterior
incurvation. The moniliform cilia along the posterior border are more numerous, while
the smaller along the outer edge are fewer ; and the same may be said of those on the
surface.

The succeeding scales are irregularly rounded, with a somewhat even external
border, which has moniliform cilia, while those along the posterior edge are few and
widely separated. The scale is areolated, especially towards its inner border, which,
along with the anterior, is smooth.

The posterior scales are more elongated transversely and have fewer cilia (about six
of the large moniliform kind), both on the abbreviated external border and on the
posterior edge. The scar is nearer the anterior than the posterior edge.

In comparing the larger broad anterior scales with those from Greenland and
Canada (Plate XXXIY, fig. 18) comparatively little difference is observable, both having
from eighteen to twenty-three moniliform cilia externally.

On the whole, therefore, the view that P. minuta and P. inomata are the same
forms is borne out by the scales.

In the variety eximia the first pair of scales have a similar shape to those of the
type, being irregularly rounded, and, from the size of the examples, smaller than in the
latter. The outer border has a series of longer cilia, fewer in number but similar in
structure. They encroach somewhat on the anterior border, or rather some of the
isolated cilia scattered over the surface near the edge project beyond it, but none of the
smaller clavate cilia so characteristic of the type minuta are present on this edge.
These large isolated cilia occur both external and posterior to the scar for attachment.

1 Described from an arctic example.

57

440 PHOLOfi MINUTA.

The second pair are elongated transversely as in the other form, but are dis-
tinguished by the great length of the cilia on the outer border. A few cilia also occur
along the posterior edge.

The succeeding scales in the anterior third do not differ much in shape from those
of P. minuta, but the cilia are much longer and stand stiffly out. They occur on the
external margin and the outer half of the posterior edge. They are less numerous than
in the large examples of the typical form, but agree with the smaller in this respect.

The posterior scales have about the same number of cilia, but they stand stiffly out
like rays.

This form is distinguished from the preceding in spirit by an olive spot on the scale,
with a pale centre at the scar for attachment. Occasionally, as in an example of this
variety from Lochmaddy, the cilia on the scales are fewer and longer, being filiform
tapering processes without the terminal enlargement.

In both varieties the areolse, especially towards the inner edge, are well marked.

Feet. — The second foot is bifid, with two well-developed spines. The dorsal lobe
forms a rounded eminence with a smooth surface. The bristles are comparatively short,
slender, and taper to a very fine hair-like tip, minutely spinous. The inner forms taper
more abruptly than the outer, and the tips are often bent nearly at right angles to the
base. The ventral lobe is longer than the former, bluntly conical, and with numerous
small papillae on its surface. The ventral bristles have stout shafts dilated at the tip,
and with numerous rows of spines on the convex margin, thus differing from those of
the arctic examples (P. minuta, Pabr., typical), which have few. The distal region
forms an elongated process — hooked at the tip, and with a series of spikes along the
ventral edge. The tip in the arctic forms is more robust and proportionally shorter.

In the typical foot (Plate XXXI, fig. 14) the dorsal lobe presents a prominent
process and a broad, slightly convex margin externally for the bristles, the spine
piercing the apex of the convexity. The bristles (Plate XLII, fig. 23) form a dense
tuft, directed outwards and downwards, and are slender and tapering, with well-marked
spinous rows.

The ventral lobe has the shape of an oblique cone, with the spine issuing from the
apex and with numerous papilla over the surface. The shafts of the bristles (Plate XLII,
fig. 24) are somewhat shorter and stouter than in the northern form (P. minuta, typical),
and the convex edge of the tip has more numerous spikes. The falcate tip is shorter,
more curved (the hook more pronounced), and the spines along the edge are often absent.
Indications of these, however, are seen.

Posteriorly, the dorsal bristles have finer spikes, and the ventral have fewer rows of
spines at the convex distal region of the shaft, while the terminal portion is more
slender and proportionally longer. The papillae on the ventral division are fewer and
somewhat longer. The ventral cirrus is short and tapered.

In examples from Lochmaddy only three or four spikes occur on the distal end of
the shafts of the ventral bristles. Some from Bressay Sound show more numerous
though finer spikes at the distal end of the shafts, as in those from St. Andrews. In a
small one from St. Magnus Bay they were too indistinct for determination, though
they appeared to be smooth. In a small example from Herm no spikes were present

PHOLOE MINUTA. 441

on the distal pieces of the ventral bristles. In specimens from the St. Lawrence,
Canada, the bristles agreed more with the arctic forms, and one had a translucent
granular deposit on them. The spikes at the end of the shaft were indistinct, but this
also occasionally occurs in the British examples.

In considering the two varieties (inornata and eximia) it is clear that no reliable
distinction can be drawn from the structure of the bristles, and this is probably more
important than the condition of the cilia on the scales. It is true the end of the shaft is
most spinous — that is, has a longer series of spines from above downwards — in var.
inornata, and that the dorsal bristles are more distinctly spinous, and the ventral warts
or papillae more conspicuous, but the characters are not new, and only vary in degree,
and are probably due to the surroundings, with which we are, perhaps, not fully
acquainted. The steps from var. eximia to var. inornata, and thence to the typical
minuta from the arctic seas, are easy, both as regards scales and bristles.

After a careful consideration of all the facts it has been thought best to unite the
varieties into a single species.

Reproduction. — Male elements as minute granules occurred in August at Lochmaddy.
At St. Andrews ripe males are common at the end of June, so that the breeding period
would seem to be in July. The spermatozoa have globular heads and long tails, and
resemble those of Lepidonotns squamatus, a species with the same breeding period. The
examples of P. minuta from Greenland present ova in July.

Development. — Trochospheres, apparently of this species, occur near the bottom on
September 12th, presenting only four crenations posteriorly to indicate the seg-
ments.

On September 10th the youngest stage has a bluntly conical prostomial region,
which appears to occupy about half the bulk of the animal. Two small black specks
are situated posteriorly. At each side are the short tentacular cirri. Four bristled feet
follow, with stout bristles having the typical structure. Two spines occur on the dilated
end of the shaft, the most prominent being inferior. Besides these stout bristles each
foot has one or two much more slender forms, with a long, tapering, diaphanous terminal
piece, evidently a larval ventral bristle. The dorsal bristles have the adult structure,
being slender, tapering, hair-like, serrated bristles. The dorsal cirrus is much enlarged
at the base, but with a slightly tapered tip.

What appear to be three pairs of scales have the aspect of globular organs on a
pedicle, each with several large truncate papillas projecting from the edge. These
papilla are out of all proportion to the condition in the adult. Viewed from below the
ventral cirrus shows a basal swollen region (ceratophore) and a two-jointed cirrus, while
some distance inwards is a globular papilla. The body, which is marked by transverse
lines at this stage, terminates posteriorly in a broad pygidium.

In the next stage (which, however, is of the same date— September 10th, 1888) the
snout is truncated, and the caudal process has a dimple in the middle. Then the head
becomes denned as an oblong mass, rounded in front and flanked by the two tentacular
cirri, which now are directed forward on each side. There are still only four bristled
feet. The cilia on the scales are longer, and two short, broad, caudal cirri have
appeared under the pygidium.

442 PHOLOE MINUTA.

The bottom tow-net procured, on February 4tli in four to five fathoms, a young
example of Pholoe having thirteen feet on each side. The head bears in front two
closely approximated median eyes, and two more widely separated towards the posterior
border. Both pairs of eyes have a rounded, lens-like differentiation. The median
tentacle is well developed and subulate, while three smaller and the stump of a fourth
(making two tentacular cirri) project in front. These are minutely but sparsely
papillose. The palpi have enlarged basal regions, and taper to a blunt point. The body
is flattened and nearly of uniform diameter, though tapered a little in front and more
distinctly posteriorly, where it terminates in a rounded " boss " on each side of the vent,
the globular cirri of the last pair of feet generally projecting on each side a little in
front. The proboscis is furnished with well-marked teeth.

The feet show dorsal and ventral divisions, each with a spine and the characteristic
bristles, besides the cirri. At least four pairs of scales were present, most with five cilia
on the outer border, though one (probably the first) had seven. Lines radiating from
the centre of the scale to the base of the cilia probably indicated nerve-strands.

Habits. — They break, when lifted with the forceps, like the Polynoidas, but are
sluggish, lurking under stones between tide-marks. The best way to obtain them,
indeed, is to chip fragments from the under surfaces of stones covered with various
growths in pools near low water mark, and immerse them in sea water for a day or
two. The Annelids either occur at the water-line of the vessel or are found by examining
the debris at the bottom.

P. minuta was first found by Otlio Fabricius on the shores of Greenland, and
afterwards by Oersted in the same region.

Dr. Thomas Williams 1 (1858) credited this species with the only vascular system he
had been able to see in the Aphroditaceans in the form of a vessel, carrying a colourless
fluid in contact with the nerve-cord, and slowly undulating with pulsations.

The Pholoe synophthalmia of Claparede seems to be a closely allied if not identical
form. 2 Marion and Bobretzky, 3 who agreed with this author as to the specific distinction
of P. synophthalmia, point out the proper interpretation of the cephalic appendages, and
that the first segment is fused with the cephalic, so that the first scale is borne on the
second segment. De Saint-Joseph likewise follows Claparede in separating P. synoph-
thalmia.

Marenzeller 4 (1893) describes a new species from the African shores of the Mediter-
ranean (Santorin), in which the dorsum of the body is covered with papilla?, and the
scales have more numerous cilia. The bristles seem to approach closely those of the
common form.

Dr. Michaelsen lately (1897) regarded the variety eximia as a distinct species, but
he relied chiefly on the divergence of the scales and other points already alluded to. It
is sufficient to consider eximia a variety, and chiefly a smaller variety. He kindly sent
me slides for examination, so that no dubiety might exist.

1 'Philos. Trans./ 1858, p. 135.

2 ( Annal. Chet. Naples/ 79, pi. iii, f. 1.

3 ' Ann. Sc. Nat./ 1875, pp. 8, 9.

4 ' Polychaten des Grundes/ 1893, p. 6, pi. i, f. 3.

PI. XXIV

Figs. 1,2, Nat. & von Graff, *,5,J.M.A.
Fig. 3 R.M.&A.T.M.
Fms. 6-9 E.B.

PLATE XXIV.

Fig.
1. Dorsal surface of Spinther miniaceus, Gr. Enlarged.

*2. Ventral surface of the same. Enlarged more highly.

3. View of a reddish example of Euphrosyne foliosa, Aud. and Ed., showing both dorsal

and ventral surfaces. Enlarged.

4. Aphrodita aculeata, L., from the dorsum. Slightly enlarged.

5. Ventral view of the same. No median furrow occurs in this specimen.

6. Head and anterior region of the foregoing. Enlarged.

7. Dorsal view of a dark variety of Hermione hystrix, Aud. and Ed. Enlarged.

8. Cephalic region of the same. Enlarged.

9. Dorsal view of Lsetmatonice filicornis, Kbg. Enlarged.

PL XXV.

5.

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ith.Werr

PLATE XXV,

Fig.

1. Lepidonotus squamatus, L., from the dorsum. Enlarged.

2. Head of the foregoing in life. Enlarged.

3. A large specimen of Gattyana (Nychia) cirrosa, Pall., from the dorsum. Enlarged.

4. Harmothoe longisetls, from the dorsum. Enlarged.

5. Dorsum of a living example of Halosydna gelatinosa, Sars. Enlarged.

6. Head of Harmothoe imbricata, L. Enlarged.

7. Anterior end of a large example of Polynoe scolopendrina, Sav., from North TJist.

Enlarged.

PL XXVI

Figs . 1, 2 , 5 & 7 & 8 ( partim ) A .W
M 6,7&8 R.M.

» 3 E.L. 4- J.M.A.

Lith.Wei

PLATE XXVI.

Fig.

1. Lepidonotus clava, Mont., from the dorsum. Enlarged.

2. Lagisca floccosa, Sav. (propinqua, Mgrn.), from the dorsum. Enlarged.

3. Harmothoe imbricata, L., from the dorsum. Enlarged.

4. Posterior end of an example of the same, from which three pairs of scales have been

removed to show the ova on the dorsum. Enlarged.

5. Evarne impar, Johnst., from the dorsum. Enlarged. A very dark example.

6. Scalisetosus pellucidus, Ehlers, with regenerating scales; Channel Island*.

Enlarged.

7. Anterior end of Sthenelais boa, Johnst. Enlarged.

8. Posterior end of the same. Enlarged.

9. Process from the edge of a scale of Sthenelais Buskii.

PI. XXVI A

PLATE XXVI a.

Fig.

1. Spermatozoa of Harmothoe imbricata. X Zeiss, 1 oc, D obj.

2. Spermatozoa of Lepidonotus squamatus. X idem.

3. Trochospheres of H. imbricata assuming a greenish hue. Magnified.

4. View of a more advanced trochosphere of March 28th, with the projecting mouth to

the left. X 116.
5 and 6. Lateral and dorsal views of other examples. X 52.

7. More advanced larva with traces of feet, but without bristles, or with such only

indicated.

8. A further stage in which the feet have minute bristles. (June 26th.) X 82.

9. Larval Polyno'e with well-marked feet and bristles, minute scales, and anal cirri.

X 52.

10. Nectochaete stage of Harmothoe. (April 20th.) x 52.

11. Nectochaete stage of a Polynoid (June 28th), with median and lateral tentacles and

anal cirri. X 52.

12. Nectochaete stage of Harmothoe, sp. (April 27th.) X 50.

13. Nectochaete stage of Polyno'e, sp. (July 18th.) -^ inch long, x 50.

14. More advanced nectochaete conditions in a form approaching Lagisca or Evarne.

(October 25th.) x about 40.

15. Dorsal view (in spirit) of nectochaete stage of a form approaching Lepidonotus

squamatus. (June.) x 40.

16. Ventral view of the foregoing. Similarly magnified.

17. A slightly later stage. Similarly enlarged.

18. Stage with developing palpi, and with additional eye-spots. Similarly magnified.

19. Stage of the same form with distinct palpi, median and lateral tentacles, x about 35.

20. Dorsal view of Panthalis (Erstedi, from a figure kindly sent by Mr. Arnold Watson.

21. Proboscis and jaws of Sthenelais boa. Enlarged under a lens.

22. Metatroch stage of Sthenelais or Sigalion seen from above. (October 22nd.) X 50.

23. Post-larval Sigalion, pelagic condition. (October 28th.) x 50.

24. Dorsal view of Pholo'e minuta. Enlarged.

25. Spermatozoa of Pholo'e minuta. (June 24th.) X 350.

PI. XXVII

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PLATE XXVII.

Fig.

1. Oral region of Aphrodita aculeata, L., in contraction and from the ventral surface.

Enlarged.

2. Head of Lxtmatonice filicornis, Kbg. Enlarged.

3. Head of Lagisca Elisabethse, n. s. Enlarged.

4. Head of Lepidonotus clava, Mont. Enlarged.

5. Head of Gattyana cirrosa, Fabr. Enlarged.

6. Head of Acanthicolepis asperrima. Enlarged.

7. Head of Evarne Johnstoni, Mel. (large example). Enlarged.

8. Head of Harmothoe lunulata, D. Chiaji. Enlarged.

9. Head of Eunoa nodosa, Sars. Enlarged.

10. Head of Lagisca Jeffrey sii. Enlarged.

11. Head of Harmothoe marphysse, Mel. Enlarged.

12. Head of S calls etosus pellucidus, Ehlers. Enlarged.

13. Head of Evarne impar, Johnst. Enlarged.

14. Head of Lagisca floccosa, Sav. Enlarged.

15. Head of Harmothoe areolata, Grube. Enlarged.

16. Anterior end of Eurythoe borealis, Sars, dorsal view. Enlarged.

17. Head of Acholoe astericola, D. Chiaji. Enlarged.
-18. Head of Phyllantinoe mollis, Mel. Enlarged.

PI. XXVIII.

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F.Huth.LiflrT Edm 1

PLATE XXVIII.

Fig.

1. Head of Harmothoe zetlandica, Mel. Enlarged.

2. Head of Lagisca extenuata, Grube. Enlarged.

3. Head of Harmothoe spinifera, Ehlers (olim Sibbaldii). Enlarged.

4. Head of Harmothoe Ljungmani (=H. Macleocli). Enlarged.

5. Head of Lxnilla setosissima, Sav. Enlarged.

6. Head of Hvarne Hubrechti, Mel. Enlarged.

- 7. Head of Harmothoe Fraser-Thomsoni, Mel. Enlarged.

8. Head of Malmgrenia andreapolis, Mel. The median tentacle is imperfect. Enlarged

9. Head of Polynoe scolopendrina, Sav. Enlarged.

10. Head of Antinoe Sarsii, Mgrn. Enlarged.

11. Head of Halosydna gelatinosa, Sars. Enlarged.

12. Head of small example of Evarne Johnstoni, Mel. Enlarged.

13. Head of Harmothoe antilopis, Mel. Enlarged.

14. Head of Antinoe mollis. Mel. Enlarged.

15. Head of Malmgrenia eastanea, Mel. Enlarged.

16. Head of Panthalis (Erstedi, Kbg. Enlarged.

17. Head of Leanira hystrieis, Ehlers. Enlarged. The specimen was somewhat im-

perfect.

21

Pi. XXIX.

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PLATE XXIX.

Fig.

1. Head of Sthenelais boa, Johnst. Enlarged.

2. Head of Sthenelais atlantica, Mel. Enlarged.

3. Head of Sthenelais limicola, Ehlers. Enlarged.

4. Head of Sthenelais Jeffrey sii, Mel. Enlarged.

5. Head of Eusthenelais hibemica, Mel. Enlarged.

6. Head of Sigalion Mathildas, Aud. and Ed. Enlarged.

7. Head of Sigalion Bushii, McL Enlarged.

8. Head of Pholoe minuta, Fabr. Enlarged.

9. Anterior foot (about twelfth) of Lagisca extenuata, Grube. X 24.

10. Anterior foot of Lagisca floccosa, Sav. ( = propingua, Mgm.). x 24.

11. Twelfth foot of Lagisca Elisabeths, McL X 40.

12. Tenth foot of Harmothoe spinifera, Ehlers. x 24.

13. Anterior foot of Harmothoe Ljungmani, Mgrn. X 28.

14. Anterior foot of Harmothoe antilopis, McL X 24.

15. Anterior foot of Harmothoe Fraser-Thomsoni. X 26.

16. Anterior foot of Harmothoe marphysse, McL X 40.

17. Anterior foot of a large example of Polynoe scolopendrina, Sav., from North

Uist. x 15.

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PLATE XXX.

Fig.

1. Foot (about twelfth left) of Harmothoe imbricata, L. X 12.

2. Anterior foot of Harmothoe zetlandica, Mel. X 24.

3. Anterior foot of Harmothoe areolata, Grube. X 24.

4. Seventh foot of Harmothoe lunulata, D. Chiaje. X 40.

5. Anterior foot of Malmgrenia castanea, Mel. X 24.

6. Anterior foot of Evarne Johnstoni, Mel. X 20.

7. Anterior foot of Evarne impar, Johnst. X 30.

8. Foot of Panthalis GErstedi, Kbg. X 15.

9. Foot of Sealisetosus pellucidiis, Ehlers. X 40.
(l0y' ; Anterior foot of Evarne Hubrechti, Mel. X 20.

11. Anterior foot of Halosydna gelatinosa, Sars. x 18.

12. Anterior foot (about eleventh) of Loenilla setosissima, Sav.

13. Anterior foot of Antinoe finmarchica, Mgrn. X 40.

14. Foot of Sthenelais zetlandica, Mel. X 32.

15. Anterior foot of Sealisetosus assimilis, Mel. X 40.

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PLATE XXXI.

Pig.

1. Scale of Oattyana cirrosa, Pall., from St. Andrews. X 12.

2. Anterior foot of Antinoe Sarsi, Mgrn., " Knight- Errant," 558 fms. X 20.

3. Foot of Malmgrenia andreapolis, McL, St. Andrews. X 24.

4. Foot of Acholoe astericola, D. Oh. X 28.

5. Anterior foot of Sthenelais boa, Johnst. X about 15.

6. Anterior foot of Sthenelais limieola, Ehlers. x 30.

7. Anterior foot of Sthenelais Jeffreysii, Mel. X 24.

8. Foot of Sthenelais, Z. X 30.

9. Foot of Eusthenelais hibemica, McL (Imperfectly preserved.) x 24.

10. Anterior foot of Sigalion Mathildm, Aud. and Ed. x 24.

11. Anterior foot of Sigalion Buskii, McL X 24.

12. Anterior foot of Leanira hystricis, Ehlers. x 40.

13. Posterior foot of the same. X 40.

14. Foot of Pholoe minuta, Fabr., from St. Andrews. X 90.

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LiSKfEaW

PLATE XXXII.

Fig.

1. Anterior scale of Lepidonotus squamatus, L. x 12.

2. Anterior scale of Lepidonotus clava, Mont, (spirit), x 12.

3. Scale of Eunoa nodosa, Sars (dried), British Museum. X 12.

4. Anterior scale of Acanthicolepis asperrima, Sars. X 12.

5. Eighth scale of Lagisca floccosa, Sav., from Baltic. The spines are rather distinct,

X 20.

6. Anterior scale of Lagisca Elisabeths, Mel. x 40.

7. Anterior scale of Lagisca Jeffrey sii 9 Mel. X 20.

8. Anterior scale of Lagisca extenuata, from 680 fathoms in the " Porcupine." X 12.

9. Scale of Lagisca floccosa, var. "Porcupine," No. 42 (p. 302). x 40.

10. Anterior scale of Harmothoe imbricata, L. x 12.

11. Anterior scale of Harmothoe Fraser-Thomsoni, Mel. X 24.

12. Anterior scale of Harmothoe lunulata, D. Ch. X 24.

13. Anterior scale of Harmothoe marphysse, Mel. x 44.

14. Anterior scale of Harmothoe Ljungmani, Mp-rn. X 30.

15. Anterior scale of Harmothoe zetlandica, Mel. X 24.

16. Anterior scale of Harmothoe antilopis, Mel. X 28.

17. Anterior scale of Harmothoe areolata, Grube. X 12.

18. Anterior scale of Evarne impar, Johnst. x 20.

19. Anterior scale of Evarne Johnstoni, McL, from a Norwegian example. X 28.

3

-

PI. XXXIII.

.

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F.Hutli,Litli T Edin r

PLATE XXXIII.

Fie.

1. Anterior scale of Evarne Hubrechti, Mel. X 20.

2. Portion of the scale of a small example of Halosydna gelatinosa, Sars. X 210.

3. Portion of the inner edge of the first scale of Harmothoe areolata, Gr. X 90.

4. Anterior scale of Harmothoe spinifera, Ehlers. X 20.

5. Anterior scale of Loenilla setosissima, Sav. X 20.

6. Anterior scale of Scalisetosus assimilis, Mel. X 38.

7. Anterior scale of Scalisetosus pellueidus, Ehlers. X 45.

8. Anterior scale of Antinoe finmarchica, Mgrn. X 30.

9. Anterior scale of Antinoe mollis, Mel. X 18.

10. Anterior scale of Malmgrenia castanea, Mel. X 40.

11. Anterior scale of Malmgrenia andreapolis, Mel. X 24.

12. Scale of a small example of Halosydna gelatinosa, Sars. X 15.

13. Scale of small example of Polynoe scolopendrina, Sav. (from Herm). x 40.

14. Scale of the large form of Polynoe scolopendri?ia 9 Sav. (from North Uist). X 24.

15. Scale of Acholoe astericola, D. Chiaje. X 28.

16. Tenth scale of Sthenelais boa, Johnst., from St. Andrews. X about 30.

17. Anterior scale of Antinoe Sarsi, Kbg. X 20.

') Y

PI. XXXIV.

.

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18.

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12

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F.HuOi.lithTEdin:

PLATE XXXIV.

Fig.

1. Proboscis of Scalisetosus pellucidus. X under a lens.

2. Proboscis of Lagisca floccosa. X under a lens.

3. Lateral view of the proboscis of Panthalis (Erstedi. X under a lens.

4. Antero-posterior view of the same. X under a lens.

5. Anterior scale of the same form (" Triton," 1882). X 24.

6. Anterior scale of Sthenelais boa (adult). X 17.

7. Scale of Sthenelais zetlandica. X 32.

8. Anterior scale of Sthenelais atlantica. X 55.

9. Scale from the anterior third of Sthenelais limicola. X 12.

10. Posterior scale of the foregoing. x 30.

11. Proboscis of Sthenelais limicola. X under a lens.

12. Scale of Sthenelais, Z. X 18.

13. Anterior scale of Sthenelais Jeffrey sii ( u Porcupine," off Ireland). X 17.

14. Anterior scale of Sigalion Mathilda with branchial process. Enlarged.

15. Anterior scale of Sigalion Bushii. Enlarged.

16. Dorsal view of the anterior end of Pholoe minuta. X under a lens.

17. Anterior scale of Pholoe minuta, var. inornata (Lochmaddy). X 90.

18. Anterior scale of Pholoe minuta, var. (St. Lawrence, Canada). X 40.

3

PL. XXXV.

.

22 a

WCI.ainai. del.

IfizrtewiBros.i

PLATE XXXV.

Fig.

1. Dorsal cirrus of Euphro syne folios -a, x 310.

2. Branchial processes of Euphrosyne armadillo.

3. Tips of branchial processes of Euphrosyne folios a. X 236.

4. Tips of branchiae of Euphrosyne Bobertsoni, Frith of Clyde. X 90.

5. Bristles of Spinther miniaceus.

a. Yentral bristle. X 90.

b. Tip of a somewhat shorter and stouter form. x 350.

c. Hook from the ventral region. X 350.

6. Upper bristle of Euphrosyne Bobertsoni, n. s. (Clyde), with shorter tip. X 210.

7. Lower bristle of Euphrosyne Bobertsoni, n. s. (Clyde), with longer tip. X 210.

8. Bristle of Euphrosyne armadillo. Oc. 1, obj. D, Zeiss.

9. Dorsal bristle of Euphrosyne armadillo (Norway). x 350.

10. One of the slender bristles of Euphrosyne armadillo (Norway). x 350.

11. The same. X 350.

12. The same. X 350.

13. Bristle of Euphrosyne armadillo. X 350.

14. Bristle of Euphrosyne armadillo. Zeiss, oc. 1, obj. D.

15. Dorsal bristle of Euphrosyne folios a. x 236.

16. Bifid bristles of Euphrosyne folios a (seen on edge). Zeiss, oc. 1, obj. D.

17. Slender serrated bifid bristle (dors.). Zeiss, oc. 1, obj. D,

18. Profile view of curved bifid bristle. Zeiss, oc. 1, obj. D.

19. Bristles of Paramphinome pulchella, a, from the inferior lobe; b, c, from the superior

lobe. X 700.

20. Dorsal bristles of Eurythoe borealis from a posterior foot. X 350.

21. Bifid dorsal bristles of Eurythoe borealis (Channel Islands). X 350.

22. Slender bristles of Eurythoe borealis (Channel Islands). X 350.

23. Ventral bristle of Eurythoe borealis (Channel Islands). X 350.

24. Bristle of first ventral series of a typical foot of Aphrodita aculeata. X 55.

25. Bristle of the middle of the second row of the ventral series. X 55.

26. Tip of posterior hair (lower series) near last foot (Aphrodita aeuleata). X 360.

27. Tip of dorsal hair forming felt (Aphrodita aculeata). X 350.

28. Bifid (serrated) dorsal bristle of Euphrosyne Bobertsoni. X 300.

3

PL

W.C.l/L adrvat del

Mintem Bros. sc. sr, invf

PLATE XXXYI.

Fig.

1. Palpus of Lsetmatonice filicoruis, a little beyond the middle. X 90.

2. Strong bristle from the upper (third) ventral series in the foot of Aphrodita aculeata.

X 55.

3. Bristles from a posterior foot of Aphrodita aculeata. X 360.

4. Tip of a dorsal spine of a British Lsetmatonice producta . x 55.

5. Ventral bristle of Lsetmatonice producta, var. x 40.

6. Tip of dorsal spine of Lsetmatonice filicoruis with four hooks on one side. X 55.

7. Tip of ventral bristle of Lsetmatonice filicoruis. X 55.

8. Lateral view of a dorsal spine of Lsetmatonice filicoruis. x 55.

9. Tip of bristle on the dorsum of an anterior foot of Rermione hystrix. X 90.

10. Tenth foot (about) of Aphrodita aculeata, viewed from the ventral surface. Enlarged

under a lens.

11. Tip of one of the attenuate spines on the dorsum of an anterior foot of Rermione

hystrix. x 210.

12. Tip of dorsal spine of Lsetmatonice filicoruis . X 90.

13. Foot of Rermione hystrix. Enlarged under a lens.

14. Tip of dorsal spine of Rermione hystrix with shield. X 90.

15. Tip of spine of Rermione hystrix with shield removed, lateral view in balsam. X 90.

16. Foot of Enrythoe borealis, Channel Islands. X 90.

17. Pinnate ventral bristle of the second foot of Aphrodita aculeata. X 90.

18. Stout smooth bristles of Aphrodnta aculeata. X 90.

19. Bristle (spinous) from the posterior end of A. aculeata. X 55.

20 a front, and 20 b lateral view of a bristle from the same region of A. aculeata.
X 55.

21. Tip of the palpus of Lsetmatonice filicoruis. X 55.

22. Anterior foot of Aphrodita aculeata, from the posterior face. X under a lens.

23. Papillge of the facia! tubercle of Aphrodita, aculeata. X 90.

24. Outline of three papillas of the facial tubercle of Lsetmatonice filicoruis. x 90.

PL.XXXv

W.C.M.^dxia^dei

Mxzvterm Bros, i

PLATE XXXVII.

Fig.

1. Papillae of the tip of the proboscis in Aphrodita aculeata. X 55.

2. Papillae of the proboscis of Lsetmatonice filicomis. X 55.

3. Papillae of the proboscis of Lsetmatonice producta. X 40.

4. Papillas of the proboscis of Hermione hystrix. X 55.

5. Tip of tentacle of Hermione hystrix. X 55.

6. Second bristle of the second foot of Hermione hystrix. x 90.

7. Ventral bristle (at ventral edge) of the same foot of Hermione hystrix. X 90.

8. Smaller bristle from the posterior foot of Hermione hystrix. X 90.

9. Ventral bristle of Lepidonotus squamatus. X 80.

10. Ventral bristle from the middle of the twelfth foot of Lepidonotus squamatus . X 210.

11. Ventral bristle from the middle of the twelfth foot of Lepidonotus clava. X 210.

12. One of the short dorsal bristles of Lepidonotus squamatus. X 180.

13. One of the longer bristles of Lepidonotus squamatus. X 280.

14. Group of bristles at the base of the tentacular cirri (first foot) of Lepidonotus clava.

X 210.

15. Dorsal bristle of Lepidonotus clava. X 210.

16. Superior ventral bristle of Gattyana cirrosa. X 258.

17. Ventral bristle from the middle of the foot of Gattyana cirrosa. X 258.

18. Dorsal bristle of the middle of the foot of Gattyana cirrosa. X 210.

19. Bristle from the base of the tentacular cirri of Gattyana cirrosa. X 210.

20. Tip of upper bristle of the first foot of Eunoa nodosa. X 210.

21. Tip of the upper bristle of the first foot of Eunoa (Erstedi. X 210.

22. Ventral bristle of the first foot of E. nodosa. X 210.

23. One of the smaller ventral bristles of the first foot of Eunoa (Erstedi. X210.

24. Tip of a dorsal bristle (average example) from an anterior foot of Eunoa nodosa.

X 350.

25. Tip of a dorsal bristle (average example) from an anterior foot of Eunoa (Erstedi.

X 350.

26. Dorsal bristle of Eunoa nodosa (coast of Durham). X 210.

27. Ventral bristle of Eunoa nodosa (coast of Durham). X 210.

28. Tip of a dorsal bristle of Acanthicolepsis asperrima. X 210.

29. Ventral bristle of Acanthicolepsis asperrima. X 210.

30. Tip of another example in Acanthicolepsis asperrima, showing minute process. X 350.

31. Tip of one of the longer dorsal bristles of Lagisca floccosa. X 350.

32. Tip of a median ventral bristle of the same. X 350.

33. Tip of a bifid dorsal bristle of Emphrosyne liohertsoni. X 350.

J> /

p L. XXXVIII

-

Mm.tera.1: -

PLATE XXXVIII.

Fig.

1. Superior ventral bristle of Lagisca floccosa. x 350.

2. Dorsal bristle of variety of Lagisca floccosa. X 350.

3. Dorsal bristle of Lagisca floccosa, var. (p. 302). x 350.

4. One of the longer dorsal bristles of Lagisca Elisabeths. X 350.

5. Superior ventral bristle of Lagisca Elisabeths. X 350.

6. Inferior ventral bristle of Lagisca Elisabeths. X 350.

7. Dorsal bristle of Lagisca Jeffreysii. X 350.

8. Superior ventral bristle of Lagisca Jeffreysii. X 350.

9. Bristle from the middle of the ventral series of Lagisca Jeffreysii. x 350.

10. Dorsal bristle of Lagisca e,vtenuata, Gr. x 90.

11. Tip of the same bristle. X 350.

12. One of the longer (not longest) ventral bristles. X 210.

13. Tip of another ventral bristle, showing the secondary process. X 350,,

14. Tip of a dorsal bristle from a large specimen of Harmothoe imbricata. x 350.

15. Superior ventral bristle of Harmothoe imbricata. x 350.

16. Median ventral bristle of Harmothoe imbricata. X 350.

17. Dorsal bristle of Harmothoe spinifera, Ehlers. X 350.

18. Tip of superior ventral bristle of Harmothoe spinifera, Ehlers. X 350.

19. Inferior ventral bristle of Harmothoe spinifera, Ehlers. X 350.

20. Front view of a dorsal bristle of Harmothoe Zetlandica. X 210.
20 a. Lateral view of the same. X 350.

21. Superior ventral bristle of Harmothoe Zetlandica. X 350.

22. Dorsal bristle of Harmothoe Ljungmani, Mgrn. (olim H. Macleodi). x 350.

23. Superior ventral bristle of Harmothoe Ljungmani. X 350.

24. One of the larger dorsal bristles of Harmothoe antilopis, Mel. X 350.

25. Superior ventral bristle of Harmothoe antilopis. X 350.

26. One of the stouter ventral bristles from the neighbourhood of the spine, x 350,

27. Dorsal bristle of Harmothoe halia'eti, Mel. X 350.

3?

PL. XXXIX.

"W! C .M . a.d na.t del

MirL-be-rrx Bros- s=-

PLATE XXXIX.

Fig.

1. Superior ventral bristle of Harmothoe haliaeti viewed antero-posteriorly. X 350.

2. Another of the same form seen laterally, x 350.

3. Inferior ventral bristle, x 350.

4. Dorsal bristle of Harmothoe Fraser-Thomsoni. x 350.

5. Ventral bristle of Harmothoe Fraser-Thomsoni. X 350.

6. Dorsal bristle of Harmothoe Fraser-Thomsoni with parasitic Loxosoma. X 210.

7. One of the longer dorsal bristles of Harmothoe marphysse. x 350.

8. Superior ventral bristle of Harmothoe marphysse.

9. Median ventral bristle (in profile) of Harmothoe marphysse,. X 350.

10. Antero posterior view of one of the same. X 350.

11. Inferior ventral bristle. X 350.

12. Profile view of one of the longer dorsal bristles of Harmothoe lunulata, D. Ch. X 350,

13. Anteroposterior view of another dorsal bristle. X 350.

14. One of the shorter dorsal bristles. X 350.

15. Superior ventral bristle. X 350.

16. Median ventral bristle, x 350.

17. Tip of a dorsal bristle of Harmothoe areolata, Grube. X 350.

18. Superior ventral bristle. X 350.

19. One of the most characteristic median ventral bristles. X 350.

20. One of the stouter dorsal bristles of a large Evarne impar, from Herm. X 350.

21. Superior ventral bristle. X 350.

22. Lower median ventral bristle. X 350.

23. One of the stronger dorsal bristles of Evarne Johnstoni. X 350.

24. Tip of a dorsal bristle. X 700.

25. Superior ventral bristle. X 350.

2Q. Bristle from the middle of the ventral group. X 350.

27. Tip of a superior ventral bristle, x 700.

28. Tip of a bristle from the middle of the ventral series. X 700.

v<

J

PL. XL.

W.C.]vr.adtia.t.del

Mm-ber-a Br OS . sc. et imp .

PLATE XL.

Fig.

1. Dorsal bristle of Evarne Hubrechti. X 60.

2. The same, x 60.

3. Tip of dorsal bristle, x 210.

4. Tip of ventral bristle. X 350.

5. Dorsal bristle (average) of Loenilla setosissima, Sav. x 350.

6. Ventral bristle (average) of Loenilla setosissima, Sav. x 350.

7. Tip of dorsal bristle of Antinoe Sarsi. X 200.

8. Tip of ventral bristle of Antinoe Sarsi. x 200.

9. A nearly straight dorsal bristle of Antinoe finmarchica. X 210.

10. An outer slightly curved bristle from the same group. X 210.

11. Median ventral bristle (South-west Ireland). X 210.

12. Dorsal bristle from posterior region of Antinoe mollis. X 210.

13. Median ventral bristle of Antinoe mollis. X 210.

14. The same. X TOO.

15. One of the largest dorsal bristles of Phyllantinoe mollis. X 350.

16. Ventral bristle. X 350.

17. Dorsal bristle of Scalisetosus pellueidus. x 700.

18. Ventral bristle. X 700.

19. Distal region of a dorsal cirrus, x 90.

20. One of the larger dorsal bristles of Scalisetosus assimilis. x 350.

21. One of the shorter dorsal bristles. X 350.

22. Ventral bristle. X 350.

23. Dorsal bristle of Malmgrenia castanea. X 350.

24. Superior ventral bristle. X 350.

25. Inferior ventral bristle. X 350.

2(u Tip of a ventral bristle from Valencia, showing secondary process. X 350.

27. One of the larger dorsal bristles of Malmgrenia andreapolis. x 700

28. Superior ventral bristle. X 350.

29. Median ventral bristle. X 700.

30. Inferior ventral bristle. X 700.

PL.XLI

I'f.C.M.a.dna.t.del.

Mffibem 3r-os.sc.et imp •

PLATE XLI.

Fig.

1. Dorsal bristle of Halosydna gelatinosa, Sars. X 180.

2. Superior ventral bristles, x 180.

3. Median ventral bristle. X 180.

4. Dorsal bristle of Polynoe scolopendrina. X 350.

5. Large bristle above the spine of the ventral division of the foot. X 90.

6. Bristle of the other series adjoining the spine. X 90.

7. Tip of a dorsal bristle. X 350.

8. Tip of one of the series represented in fig. 6. X 350.

9. Dorsal bristle of Enipo Kinbergi. X 350.

10. Ventral bristle with simple tip. X 350.

11. Bifid ventral bristle, x 350.

12. Antero-posterior view of a large ventral bristle with a simple tip. X 350.

13. Dorsal bristle of Acholoe astericola. X 350.

14. Ventral bristle. X 350.

15. Brush-shaped bristle from the superior border of the foot of Panthalis (Erstedi,

X 350.

16. Antero-posterior view of a bristle from the upper third of the median lobe of the foot.

X 210.

17. Lateral view of the same bristle. X 210.

18. Bristle from the ventral edge of the foot. X 210.

19. Dorsal bristle of Sthenelais boa. X 90.

20. Fragment of the former, showing spinous rows. X 350.

21. Superior ventral bristle with simple spinous tip X 350.

22. Stout bristle from the centre of the ventral division with a single appendage, x 350.

23. Bristle from the inferior series. X 350.

24. Portion of the edge of a scale of Sthenelais Zetlandica. x 350.

25. Superior ventral bristle in calcium chloride- X 350.

26. Inferior ventral bristle. X 350.

27. Edge of a scale of Sthenelais atlantica. X 350.

28. Superior ventral bristle. X 350.

PL.XUI

-^

. : M.a.ina.t.del. Fid . Ze , J.M.A-

Brod.sc]et

PLATE XLIL

Fig;

1. Second series of ventral bristles with single terminal joint in foot of Sthenelais

limieola. X 350.

2. Slender bristle beneath the former. X 210.

3. One of the stouter bristles following the preceding. X 210.

4. Tip of a ventral cirrus. X 210.

5. Papillae on the edge of a scale of Sthenelais Jeffreysii. X 350.

6. Superior ventral bristle. X 350.

7. Bristle from the inferior lobule, with two divisions in the terminal process. X 350.

8. Bristle from the ventral edge of the group. X 350.

9. Superior ventral bristle of Eusthenelais hibernica X 350.

10. One of the slender bristles at the ventral border of the foot. X 350.

11, 12. Portions of a dorsal bristle of Siyalion Mathildas. X 280.

13. Superior ventral bristle. X 280.

14. One of the series below the former. X 280.

15. One of the next series without spines at the end of the shaft. X 280.

16. A more slender form at the ventral margin of the foot. X 280.

17. One of the stronger bristles of the ventral lobe, with a single terminal process.

X 280.

18. Portion of the end of the shaft (with its whorls of spikes) of one of the superior

bristles of Sigalion Buskii. X 350.

19. Similar region of one of the next series. X 350.

20. Dorsal bristle of Leanira hystricis. X 350.

21. Anterior ventral bristle. X 350.

22. Ventral bristle from the posterior region. X 350.

23. Dorsal bristle of Pholoe minuta. X 280.

24. Ventral bristle. X 280.

25. Foot of Lepidonotus squamatus. X 20.

26. Tenth foot of Lepidonotus clava. X 12.

27. Tenth foot of Gattyana cirrosa. X 40.

28. Foot of Eunoa nodosa. X 18.

29. Foot of Acanthicolepis asperrima—hehmd the middle of the body. X 24.

30. Anterior foot of Lagisca Jeffreysii. X 24.

t^