THE DANISH

INGOLF-EXPEDITION

■

VOL. V, PART 1.

CONTENTS

HECTOR F. E. JUNGERSEN: PENNATULIDA.

" QJ

m

" O BY

PUBLISHED AT THE COST OF THE GOVERNMENT

THE DIRECTION OK THE ZOOLOGICAL MUSEUM OF THE UNIVERSITY.

•< r *&2MS£*^"'

COPENHAGEN.

H. HAGERUP.

PRINTED BY BIANCO LUNO.
1904.

THE DANISH INGOLF-EXPEDITION.

VOLUME V.

1.

PENNATULIDA.

BY

HECTOR F. E. JUNGERSEN.

WITH 3 PLATES, i CHART AND 3 FIGURES IN THE TEXT.

->H3«f-

COPENHAGEN.

PRINTED BY BIANCO LUNO.
1904.

Published, July the 13 th 1904.

CONTENTS.

Pennatulida.

Introductory remarks .

Pag

I

Remarks on the systematic classification of the Pennatulids 4.

List of the northern Pennatulids 10,

Fam. Pennatulidse K611 11

Pennatula Lam 11

Pennatula aculeata Kor. Dan 11

— phosphorea L.var. Candida Marsh.&Fowl. 14.

— grandis Ehrb 16

— prolifera n. sp 1

Fam. Virgularids K611., emend 24

Virgularia Lam 24

Virgularia mirabilis (O. F. M.) 25

cladiscus nom. nov 33.

Stylatula Yerrill 37

Stylatula (Diibenia) elegans Kor. Dan 38

Fam. Pavonaridse Jungersen 39.

Pavonaria K611 39

Pavonaria finmarchica (M. Sars) 39

Halipteris Koll 44,

Halipteris christii (Kor. & Dan.) 45

Fam. Funiculinidae Koll., emend 49.

Funiculina Lam 49

Pag.

Funiculina quadrangularis (Pall.J 49.

Fam. Protoptilidse Koll 51.

Protoptilum Koll 51.

Protoptiluni carpenteri K611 51.

thomsoni K611 '. 55.

— denticulatuin n. sp 59.

Distichoptilum Verrill 62.

Distichoptilum gracile Verr 62.

Fam. Anthoptilidse Koll 65.

Anthoptilum Koll 65.

Anthoptilnm grandiflorum (Verr.) 66.

— murrayi Koll 67.

Fam. Kophobelemnonidse Koll 68.

Kophobelemuon Asbjorns 68.

Kophobelemnon stelliferum (O. F. M.) 68.

Fam. Umbellulidae Koll 74.

Umbellula Cuv 74-

Umbellula lindahlii Koll 75.

— encrinus (L.) 79-

General review of the occurrence and distribution of the

northern Pennatulids 87.

Literature 93.

4isnr

Pennatulida.

By

Hector F. E. Jungersen.

In the present treatise an account is given, not only of the Pennatulids brought home by the Ingolf ,
but also of material in later collections from within the territory of the Ingolf-Expedition and from
the adjoining regions which were investigated afterwards. Thus it comprehends the results from
the surveying cruises of the Diana at Iceland and the Faeroe Islands, from the expedition to East-
Greenland by Messrs. Amdrup and Hartz, from the participation of cand. mag. Ad. Jensen in the
investigation of the «Michael Sars in the North Atlantic during the summer of 1902 under the
direction of Dr. Hjort, and from the cruise of the «Thor», the steamer for the international investi-
gations of the sea, during the summer of 1903 under the direction of Dr. J oh. Schmidt; finally it
includes also the material sent to our museum from Mr. Jonsson, district physician on the Yestman-
Islauds, and by Mr. Miiller, agent in Thorshavu.

Our knowledge of the Pennatulids rests in several respects on a rather slight footing; this
holds good also for the apparently so well examined Scandinavian forms, in spite of the numerous and
beautiful treatises which have appeared from Norwegian scientists. I have tried personally therefore
to examine all the forms that have hitherto been described from the coast-regions of Scandinavia,
as well as the forms from the territory adjoining the one which was investigated by the Ingolf .
For this purpose I have examined the collections in Christiania, Bergen and Stockholm; from the
museums in these towns I have received later all that I wished to examine more closely and to
compare with our own material here in Copenhagen; for this great liberality I beg to offer my best
thanks to Professors R. Collett and Hj. Theel, and to Conservators Dr. A. Appellof and Mr.
J. Grieg. I have further had the opportunity at the British Museum of bringing into the comparison
part of the material of the Challenger-Expedition; on the other hand, I have searched in vain in
several English museums for the Pennatulids from the expeditious of the Porcupine , the Triton ,
and the Knight-Errant*. The result of my endeavours has thus been a revision of the Pennatulid-
fauna of a large sea-territory, viz. the Polar Sea between Europe and Greenland, the sea to the
West of Greenland, and the northern part of the Atlantic down to 55 Lat. N. and to the meridian
of Cape Farewell. This revision, as will be seen, has led to a somewhat new conception of several
species hitherto established; further, several forms have been added, of whose occurrence within
territory so far north we have hitherto known nothing, and could know nothing; of undescribed
species only two have been added.

The Ingolf-Expedition. V. I. j

PKNXATUUDA.

I have followed K6 Hiker in regard to the terminology; some of his terms have been used
from much older times, and are based on the external [resemblance to a feather, especially conspicuous
in forms like Pennatula, Ptcrocidcs etc. The stem of the colony is accordingly divided into the
shaft, rhachis, carrying all the individuals, and the peduncle, which is naked and under natural
conditions sunk into the bottom of the sea; polyp is the name used only for the individuals posses-
sing the entire equipment of an octocoral, tentacles etc., zooid for the dwarf individuals wanting
sexual organs and tentacles (only in Umbellula do they carry one tentacle). The individual is said to
be provided with a calyx, when the basal part of its body 1 ) is stiff so as not to be retractile, but
the upper part of the polyp with mouth and tentacles can be retracted and hidden in it. The upper

edge of the calyx may be more or less marked, in the former case it is
V provided with lobes, up to eight in number; the axial (dorsal) side of the

individuals is turned towards the stem, the ab axial (ventral) away from
the stem. Wings, alse or pinnse, is the name of the oblique series of
polyps placed transversely on the stem, where the polyps are mutually
coalesced.

I have differed only in one point from Kolliker; what he de-
scribes as the ventral side of the colony, I call the dorsal, and
vice-versa (except in Renillcfl see below); this I shall try to explain more
closely.

In the great majority of Pennatulids the colony, as is well known,
is constructed bilaterally. The plane that divides the stem longitudinally
into two corresponding parts, has quite naturally always been interpreted
as a dorso-ventral one {V~D in the annexed figure); but whether the
side of the stem in the figure denoted by D, is to be called the dorsal (po-
sterior), or the ventral (anterior), has always been decided quite arbitrarily.
The earlier authors -- with the exception of Lamarck — have generally
called it the dorsal, or posterior side. Kolliker, in his monograph (p. 5),
declares it to be the ventral for no other reason than that it is am
zweckmassigsten >. Later, all authors have adopted the decision of Kolliker
although some of them (e. g. Verrill, Koren and Danielssen) had earlier used the contrary desig-
nation. The feature distinguishing this side of the colony, is first and foremost that new polyps
develop on either side, away from the median line; thus, the polyps placed nearest to this line are
the oldest in each transverse series; further, the median line itself is always «sterile» in so far as
polyps are never formed in it; only rarely is it covered with zooids (for instance Kop/iobclaunoii);
generally along the median line there is a more or less narrow naked streak, increasing in breadth
down towards the peduncle.

The opposite surface of the stem ( V in the figure) is characterized by the fact that the new
polyps are developed from both sides in the direction towards the median line; accordingly, the

Fig. 1.

Diagram, part of a Penna-
tulid-rhachis. V—D indicates
the direction of the middle
plane, i — 8 polyps in one
transverse series; the figures
give the age of these polyps,
no. 1 being the eldest one;
the arrows indicate that the
polyps accordingly appear
and are developed to either
side away from the naked
streak D-D.

1 ) By the body of the polyp is understood the part of the polyp projecting from the surface of the stem; in reality
some portion of each individual issuing from the stem is inclosed in the latter.

PENNATUUDA.

youngest polyps in each transverse series are those nearest to this
line; the median line and its more immediate surroundings is often
also sterile of polyps; when this is the ease the naked streak, how-
ever, is narrower here than on the opposite surface, with the exception
of the lower part towards the peduncle (Renilla, however, is a remark-
able exception). Thus the two median surfaces of the stem are gener-
ally easily distinguished r ). Making the median line our starting-point
we might say that the surface D shows a centrifugal development
of the polyps, the surface Vs. « centripetal one; so far we might well
abandon the terms ventral and dorsal side, and instead of them use
centrifugal and centripetal side; these latter names, however, are
somewhat clumsy in descriptions, as is also the case, I think, with
the terms proposed by Bourne 2 ): prorhachis (for D) and metarhachis
(for V); dorsal and ventral side are, and will always be, the most
handy names, if only a rational choice were agreed upon. In m y
choice I have started from the following consideration: the stem of
the sea-pens, rhachis -\- peduncle, is the direct product of the individual
developed from the egg (the oozooite of bacaze Duthiers); this
primary individual is for some time a solitary polyp of the typical
octocorallian structure ; accordingly, it is bilaterally symmetrical with
respect to a plane naturally called a dorso-ventral plane; the mouth
and pharynx (stomodaeum) are oval with the longer axis in the plane
of symmetry; this plane bisects two median chambers, mutually diffe-
rent and differing from all the others: only one of them contains re-
tractor muscles, and this one is on the same side as the ciliated groove
(sulcus, siphonoglyphe3)) of the pharynx; it has long ago been agreed
on calling this csuicar side the ventral side in all polyps of octo-
corals. If we will keep this designation here, we must accordingly
be consistent, and also use it for the corresponding, homologous side
of the stem of the sea-pens. Several years ago by following the
development of Pennatula phosphorca, I have shown that the < sulcar
side of the primary polyp is really the side V, the «centripetal side»;
only this side, then, can justly be called the ventral side.

Whichever appellation, however, is preferred it is obvious that
in all Pennatulids the same name is to be used for the same (homo-
logous) side. This, Kolliker (and with him the later authors) has

Fig. 2.

Fig- 3-

Young specimens of Renilla reni-
formis (Pall.) and Pcnnatula phos-
phorea L. , seen from the dorsal-
side; about 2 1 2 times the natural
size. P terminal or primary polyp,
p, p' lateral polyps, p' younger than
p\ ^terminal zooid, .r dorsal zooids;
s' (only in Renilla) zooids placed on
the polyps; n naked dorsal streak.
The arrows indicate that the polyps
grow in the direction of the unseen
ventral side of the colony.

M Even in such a form as Distichoptilum (PI. I, Figs. 12—14) or in young stages for instance of 1 ' irgtilaria (PI. II
Fig. 25). The polyps placed in a single series on either side of the rhachis will always incline somewhat towards the side V.

2 ) Anthozoa in Ray Lankester: Treatise on Zoology. Part II. 1900. p. 31.

3) The ciliated groove, as is well known, is not developed in the older polyps of the sea-pens, but yen," highlv
developed in the zooids and the quite young polyps (Hicksou, Phil. Tr. 18S3).

PENNATULIDA.

not done, in so far as lie has made a mistake in the comparison with regard to one form, Rcnilla.
Kolliker decided, without further explanation, that the most naked surface in all bilateral Pennatulids
was to be termed the ventral side; now, in all pennate and spicate forms the most naked side is really
homologous, being the centrifugal side>; it is easily seen however that the naked «underside» of the
reniform disc is the -centripetal side», new polyps budding only at the margin of the disc, and the
direction of their development is seen to be away from the centre of the polyp-bearing overside*.
Tli is may be seen in every specimen of Renilla; but it has been further proved by the examination
of its development by Wilson. From his examination it is also seen that the naked side of the
Renilla corresponds to the ventral or sulcar side of the primary polyp; but Wilson has not seen either
that it corresponds to Kolliker' s dorsal side» in the other Pennatulids; I have, however, proved this
fact in my treatise, quoted above, on Pennalula phosphorea. Nevertheless the matter does not seem to
be understood yet with regard to Renilla; Delage and Herouard (Traite de Zoologie concrete, T. 2,
2, 1901) take the same erroneous view as Kolliker, so does Moroff; whether it is also the view of
Bourne is not distinctly seen, but from his endeavours to derive Renilla from an Llmbelhila-Yike
original form I am inclined to think so. To prevent any misconception in future I figure here a Renilla
and a Pennatula, seen from the same -- homologous -- side, the side I designate as the dorsal one.

Remarks on the systematic classification of the Pennatulids.

Recent authors have generally adopted the system set forth by Kolliker in the Challenger
Report vol. I, 1880. According to this, the order Pennatulida is divided into four groups (sections), in
the first three of which the colony has a bilateral formation; when the sub-sections are omitted this
system is as follows:

I. Pennatulece.

Fam. Pterocididw.

Pterocidcs Herkl.

Godeffroya K611.

SarcopJiylluin K611.
Fam. Pennatulida.

Pennatula Lam.

Leioptilum Verr.

Ptilosarcus Gray.

Halisceptrum Herkl.

Fam. Virgularidce.

Virgularia Lam.
Scytalium Herkl.
Pavonaria K611.

Fam. StylatulidcB.

Stylatula Verr.
Diibenia Kor. & Dan.

u I caiitlwptilum K611.

II. Spicatce.

Fam. Funiculinidce.

Funiculina Lam.

Halipteris K611.
Fam. StaehyptilidcB.

Staehyptihtm Koll.
Fam. Anthoptilidm.

Anthoptilum Koll.
Fam. Kophobelemnonida.

Kophobelemnon Asbjorns.

PENNATULIDA.

Sclerobelemnon Koll.

Bathyptilitm Koll.
Fam. Umbellulidae.

Umbellula Lam.
Fain. Protocaulida.

Protocaulon Koll.

Cladisais Kor. Dan.
Fam. Protoptilidee.

Protoptilum Koll.

Lygomorplia Kor. Dan

Microptilum Koll.

Trichoptilum Koll.

l.iptoptiliiiu Koll.

Sclerqptilum Koll.

III. Renillece.

Fam. Renillida.

Renilla Lam.

IV. VeretillecB.

Fam. Caver nularidce.

Cavernularia Val.

Stylobcle union Koll.
Fam. LituaridiC.

Lituaria Val.

Vcrctillum Cuv.

Policclla Gray.

Clavella Grav.

Studer (Versuch eines Syst. der Alcyonaria. Arch. Naturgesch. 53. Jahrg. 1. 1887, p. 22, and
Challenger Report vol.31, p. XXVII) has followed Koren & Danielssen 1 ), and like these authors
added a fifth to the four groups of Kolliker: Gondulece with fam. Gaud/tlidcs, genus GondztlKor. Dan.
but has otherwise restricted himself to the inserting of the later established genera into the families
of Kolliker {Sciophyllum Verr. in fam. Pennatulidce, Svava Kor. Dan. in Stylatulidcp, Gunneria Kor. Dan.
and Distichoptihnu Verr. in Protoptilidee). Later (Note prelim, etc. Camp, de l'Hirondelle 1890) he
has inserted into the fam. Pfcrocididce the new genus Gyrophyliinii Stud. Kiikenthal (Diagnosen
neuer Alcyonarien aus der Ausb. d. Deutschen Tiefseeexpedition. Zool. Anz. 25. Bd., 1902, p. 302) has
added still another (the sixth) group: Verticilladece: fam. Chuuncllida;, genera C/mnella Kiikenth.
and Ampliiantlms Kkth. Bourne (Anthozoa in the treatise on Zoology edited by Ray Lankester,
part II, 1900) almost without any alteration — at all events, without any essential alteration — has
followed Kolliker; it has to be remarked, however, that he regards tiie two families of Kolliker Vir-
gularida: and Stylatulidcz as subfamilies of one family ] r irgidarida'\ Pcnnatulida: and Pterocidida Koll.
are regarded also as subfamilies; and the family Gbndulidce is placed beside, and in the same group
with these families; finally also Funiculi nida and Bathyptilidce are degraded to subfamilies of one
family, as are also the two families of Veretillinse. The system has been more highly modified in
Delage & Herouard (Traite de Zoologie concrete T. 2, Partie 2, 1902); here there are five groups:
i) Frondina (Renilla), 2) Umbellina (Umbellula), 3) Juncina (Spicatce -\-Vcretillctr of Kolliker), 4) Pennina
(= Pennatulece of Kolliker), ^Acaulina (Gonditl). The extent of the families remains otherwise almost
unaltered; the family Protocaitlidm of Kolliker, however, has been placed under the family Kop/10-
belemnonincB D. & H., and the genus Deutocaulon Marsh. & Fowler has been included in the same
family; the genus Cladiscus Kor. Dan. has been removed to the family Protoptilina; the genus Svava
Kor. Dan. to the family Virgularince\ several genera of the same family, as Lygus Herkl., Radicipes
Stearns, as also some of the genera established by Gray in the family Peintatulince are also enumerated.

M Nye Alcyonider etc. 18S4.

PENNATULIHA.

I cannot regard the grouping of Delage & Herouard as an improvement on the system of Kolliker,
and most of the genera, found in these authors bnt not in those before mentioned, will have to be
abandoned.

Nor can I, for my part, regard the system of Kolliker as satisfactory ; the characters, by which
the division into groups has been made, hardly always make the line of separation in a natural way;
when, for instance the first group Pennatulea is made to contain the forms in which the individuals
of the polyp-series are coalesced to form wings, this feature is seen not to be of absolute validity: in
the genus Virgularia, species are found in which the wings are partly dissolved into free individuals,
or, more properly, in which only some individuals are coalesced, and only to quite a slight extent
[Virgul. bromleyi K and Virg. cladiscus mihi); the same is found in the genus Stylatula (in the sub-
genus Dubcnia). On the other hand, forms are found in the group Spicatw, in which a coalescing of
some individuals in an oblique series takes place, in which, accordingly, there is as distinct a beginning
of wings as in the mentioned Virgulari<F\ this is seen in Anthoptiliim grandiflorum Yerr., while in
another species of the same genus, A. vmrrayi K. the polyps are quite separate, and partly even not
arranged in regular oblique series. A one-sided prominence, given to this single character, will thus
easily separate closely related forms into two groups; but when other structures are also taken into
consideration, I think that the four groups of Kolliker may be kept --at least provisionally — but,
to be sure, with altered contents. If we knew more of the development of the colony in a larger
number of Pennatulids than is the case now, the mutual relations of the forms might certainly be
seen more clearly, and then, perhaps, the grouping might be different. For the present, it is exceedingly
difficult' to reach a sure classification ; difficulties occur on all points, especially in limiting off the species.
Most of the features that have generally been used as specific characters prove to be more or less
useless: the number, grouping, size of the polyps and zooids often change considerably during growth,
as also all the relative sizes of the parts of the polyps and the stem, which are moreover exceedingly
influenced by the degree of contraction. Nor does the anatomical structure give many holds for the
classification, as it is upon the whole too uniform ; the spicules also give but slight help : species of
the same genus may quite lack spicules or be abundantly provided with them (for instance, within
the genus Umbellula); nor is the form and size of the spicules often of much assistance; in specimens
of the same species both the number and the size may be very varying (for instance, in Funiculina
quadrangiilaris, Kophobelemnon stcllifcrum). Amongst the better characters, I reckon the presence or
absence of a calyx on the polyps, and the form of the calyx. For the delimitation of genera and
families, these features along with the grouping of the polyps and zooids in the developed colony will
for the present be of most value.

I am not in a position to undertake a complete reformation of the system of the Pennatulids
on the basis of the material I have hitherto had for examination, but on several points 1 shall be able
to revise, with greater certainty than has before been possible, certain parts of the system of Kolliker
and the later additions to, and alterations of, this system; on other points I shall have to be content
with giving suggestions more or less positive. The more partictdar reasons for the position of the
forms dealt with in the following section, will be given there; in this place I shall only give some
conclusions with regard to these forms.

PENNATULIDA.

In the group Pennatulece, the genus Halisceptrum Herkl. will have to be removed from the
family Pennatulidm to the family Virgularidm\ it is quite obvious that this genus is closely related
to the genus Virgularia itself; perhaps it might even be embodied into it (see later under Virgularia)\
further the genera of the family Stylatulidm show so close a resemblance to Virgularia, that they can
scarcely be regarded as forming a separate family ; they are certainly more closely related to the genus
Virgularia than this genus is to the genera Scytalium Herkl. and Parovaria K611. ; on the other hand,
I should be inclined to separate these latter genera into a separate family (or, at all events, a sub-
family) characterized -- in contradistinction to Virgularia and the Stylatulids (with the exception of
Acanthoptilum) — among other things by the fact that the sexual organs are only developed in the
fully formed polyps of the older wings, and that new polyps, at an}' rate in the younger forms, bud
out on the upper, older part of the rhachis; in this family Pavonaridm mihi must further be included a
member of the group Spicatm, viz. the genus Halipteris Koll., although its polyps are not united into
wings; but in most other features it shows the greatest resemblance to Pavonaria Koll., much more
than to Funiculina, with which it has been placed by Kolliker. The genus .SVv?r'« Kor. Dan., which
Studer has referred to the Stylatulids, Delage & Herouard to their Virgularincr, will have to be
dropped as a genus; it is only a species of the genus Virgularia; Lygus, which in Del. & Her. is
given as a separate genus, is synonymous with Virgularia; and Radicipes Stearns (Proc. U. S. Nat. Mus.
Vol.6, 1883, p. 96, PI. MI) is no sea-pen at all, but a Gorgonid (a species of Strophogorgia Perc. Wr.);
the genera of Gray: Ptilella , PJiosphorclla, Crispclla (the family Pennatulinm Del. & Her.), and
Argentella (of their Pteroeideidinm) have already, and certainly rightly, been condemned by Kolliker
in his monograph. Further, the genus Stachyptilum Koll., in my opinion, is to be included in the
family Pennatulidm.

In the group Spicatir of Kolliker (== Umbellina -\- Juncina — the family Veretillinm in Del. &.
Her.) the greatest alterations will have to be made. In the first place the whole family Protocaulid,c
must be done away with: two of its genera, Protocaulon Koll. and Dctttocaulou Marsh. & Fowl., because
they are young stages of Virgularia-spedes, Cla discus Kor. Dan., because it is only a (wrongly inter-
preted) species of Virgularia, in reality identical with Svava Kor. Dan. (for further particulars see
under Virgularia mirabilis and V. cladiscus mihi). Consequently, all inquiry may be omitted with
regard to the question whether Cladiscus > is to be included in Protoptilida- or Protocaulidir, and
whether Protocaulon and Deutocaulon, as has been done by Del. &. Her., may be placed under the
family Kophobelemnoninm.

Again, most members of the family Proloptilida: must be dropped as separate genera. Lygo-
morpha Kor. Dan. is a young form of Halipteris (for further particulars see under H. cAristii); Microp-
liluui Koll. of Pavonaria; Leptoptilum Koll. and Trichoptilum Koll. are young forms of Funiculina (see
under this latter); Gunneria Kor. Dan., which was inserted not only by the authors of the genus, but
also by Studer and Delage & Her., is a wrongly determined Kophobelemnon (see under K. stelli-
ferum); further, the genus Scleroptilum Koll. must be separated from this family; its polyps, in spite
of their dense provision of spicules, want a calyx, and it can only be by an oversight of Kolliker that
it has been placed in a family especially characterized by him as provided with a calyx. Finally we
have only left Protoptilum Koll. and the later added Distichoptilum Verrill. These two genera, at all

S PENNATULIDA.

events, have the common feature that their polyps are provided with a well developed calyx, the
abaxial part of which is fully formed, often with calyx-teeth, while its axial side is more or less con-
nected with the stem; young stages of Protoptiluiii, or such simple species as Prof, carpenteri (if it is
not only a young stage also) resemble Distichoptilum very closely; the arrangement of the zooids,
however, is quite different, as Protoptihtm is provided with numerous dorsal zooids, and moreover with
zooids everywhere on the rhachis between the polyps, while Disticlioptilum has only lateral zooids, two
for each polyp.

The other forms of the group Spicatcr in which the polyp has developed a calyx: Funiculina,
Haliptcris, and Stacliyptilitin, seem to me to show no close relation, so that the)' certainly cannot —
as has been done by Bourne -- be placed in one family. On the contrary, the genus Stachyptilum
K611. seems to me to be altogether a stranger in this group; as far as I can see from the description
and figures of Kolliker, I think it must be referred to the first group, Pennatulecr, and be placed there
in the family Pcnnatiilidcr; the whole form of the colony and the arrangement of polyps and zooids
is as in Pennatula: the polyps are placed quite regularly in oblique series, but the members of one
series are not mutually coalesced; the dorsal surface of the stem is covered with zooids, as are also
the lateral and ventral intervals between the polyps; apart from a slighter development of the points
of the calyx it looks like a Pennatula with free polyps.

The genus Funiculina occupies a quite isolated position by wanting real zooids; only the quite
young polyps appear temporarily as zooids; by and by they increase in size, get fully developed ten-
tacles, form sexual organs, and become perfect polyps; the arrangement and growth of the individuals
is not regular, although they, as it were, tend towards a regular arrangement in transverse series.
From all these facts, Funiculina seems to me to occupy an especially primitive place below all known
sea-pens, and, at all events, a sufficiently peculiar place for it to form a separate family in which no
other genus can be included for the present. Haliptcris, which has been placed as its nearest ally,
since Kolliker in his monograph had made this arrangement, is in reality far removed from it; the
resemblances found are common to most long and slender sea-pens. In Haliptcris, there is as strong a
contrast between zooids and polyps as in any other sea-pen; in the calyx of its polyps the abaxial side
is far more developed than the axial one where teeth are quite wanting in the calyx. In reality, I
think that the original reference of the typical species H. christii to « Virgularia has more nearly
approached the correct thing; it is, as stated above, a Pavonarid; as in Pavonaria the calyx of the
polyp is provided with two abaxial teeth; the only essential difference is that the polyps do not
coalesce to form real wings.

With regard to the genera with polyps without calyx, Scleroptilum K611. shows the
simplest form of colony; as it shows no immediate relation to the other known genera it must, I
presume, form a separate family Scleroptilidm 1 ).

The genus Anthoptilum must still form the type of a family Aiithoptilidcc; but in this family

■) The genus was established by Kolliker (Chall. Rep. Vol. I, p. 30, PI. VII, fig. 29) for the species 5. grandiflorum
and dwissimim, both from considerable depths in the Pacific off Japan; Verrill has later found another species S. gracile,
of a length up to 1 foot, in the depths of the Atlantic to the east of North America (Cape Hatteras) (Rep. Coram. Fish and
Fisheries for 1S83 (1885) p. 510 [8], PL III, fig. 6, and Am. Journ. Sc. Vol.28, 1SS4, p. 219); the description in the latter place
speaks of c,ventral» zooids, and such zooids, according to Kolliker, ought to be wanting.

PENNATULIDA.

the genus Bcnthoptilum 1 ) of Verrill may probably also be included; further, I think that the two
families Kophobelemnonida and Umbellulidce. are to be kept unaltered. Umbellula I regard — contrary
to Delage and Herouard — as a particularly specialized form, related to Kophobelemnon; an Umbellula
might be thought to be derived from a Kophobelemnon-like form in which the polyps in the greater
part of the rhachis were checked so as to be kept in the zooid-stage; the zooids in Umbellula are
somewhat more developed than ordinary zooids, in so far as they are generally provided with one
tentacle; and a certain resemblance to Kophobelemnon is undeniably seen in such Umbellula-species
as U giintheri K. in which the polyp-bearing part of the rhachis is lengthened in a club-shaped
manner. The radial structure seen in certain Umbelhila-species (for instance U. encrinus), is, at all
events, a secondary development, having plainly arisen from an original bilateral formation, as is
sufficiently shown by the younger stages. To Umbclhclidce , the Chunnellida Kkth. are no doubt
related; in these latter also the radial arrangement of the polyps is plainly a secondary one; the
bilateral formation may unmistakably be traced in the arrangement of the zooids, but may also be
seen in that of the polyps. I think it superfluous to form a separate main group for this family.
According to this, the section Spicatm will have to be divided into two subsections:

A. The polyps with calyx.

Fain. Funiculidce {Fttuiculina Lam.).

» Protoptilidcr {Protoptilum, Distichoptilum).

B. The polyps without calyx.

Fam. Scleroptilidce [Sclcroptilum).

» Anthoptilida [Anthoptilum, Benthoptilum).

» Kophobelemnonidce [Kophobelemnon, Sclerobelemnon, Ba/hyptilum).

» Umbclhdida> {Umbellula).

» Cliunellidce [Chunella, Amphianthus).

The sections Renille<r and Veretillea: must probably still remain. Even if Renilla in its
development, shows a distinct connection with a common primitive Pennatulid, it is so peculiar
when fully grown, that it does not agree with other groups. This holds good to a smaller degree
with regard to Veretillece ; the radial arrangement of polyps and zooids in these forms, however, is a
peculiarity justifying a separation, at least provisionally. I think it probable that this radial arrange-
ment will meantime prove to be of secondary origin, since a bilateral formation may be traced in the
inner structure of the stem in Veretillum (comp. Kolliker Mouogr. p. 430). It is rather remarkable
that no young stage of this group is as yet known, although it is represented, in the Mediterranean
for instance, by two rather common genera: Veretillum and Stylobelcvinon. Both these groups: ReniUem
and Verctillea>, I must regard as very specialized and anything but primitive Pennatulids.

With regard, finally, to the group Gondulece (Acaulina Del. & Her.), it must be quite omitted;
it has been established only on the & genus » Gondtil, which is nothing but a mistaken fragment of
Pavonaria jinmarchica (see under this species).

i) B. serium Verrill, from the depths of the Atlantic to the east of North America (Cape Hatteras). Rep. Conim. Fish
etc. 1883, p. 510 [8], PI. II, fig. 4, and Am. Journ. Sc. (3), Vol. 29, 1885, p. 149, note.

The Ingolf-Expedition. V. l. 2

IO

PENNATULIDA.

List of the northern Pennatulids.

By northern Pennatulids are here meant the species known from the Davis Straits and Baffins
Bay, the Greenland Sea, and the part of the Atlantic north of 55 N. Lat. and east of the meridian
of Cape Farewell.

The reduction made in the previous section of formerly established genera, will appear clearly
in a list of the Pennatulids from this territory, which has been the special object of my investigations;
to this is further to be added that I have found it necessary to unite together several of the earlier
established species. Notwithstanding the fact that several forms have now been added from within
this geographical territory, which were not hitherto known to occur there, and among these two
new species, the total number of species will be seen to be diminished to about two thirds of the
number which might be enumerated from previous publications.

The following list A contains the earlier established species, the list B the species as I now
think they should be. In the latter list, the species marked with an asterisk have been brought home
by the Ingolf-Expedition itself.

A.

Pennatula phosphorea L. |

2. distorta Kor. Dan. I

3. aculcata Kor. Dan.
grandis Ehrenberg

5. Virgularia affinis Kor. Dan.

6. mirabilis (L.)

7. Cladiscus gracilis Kor. Dan.

8. loveni

9. kollikeri

10. Svava glacialis

1 1. Deutocaulon hystricis Marsh. & Fowl

12. Diibenia abyssicola Kor. Dan.

13. elegans

14. borealis

15. Pavonaria finmarchica (M. Sars

16. Gondii/, mirabilis Kor. Dan.

17. Hahpteris christii (Kor. Dan.)

18. Lygomorpha sarsii

19. Stichoptiluni arcticum Grieg

20. Protoptilum tor turn

I -

B.

1. Pennatula phosphorea L.

2. » aculcata Kor. Dan.

3. grandis Ehb.

4. * prolifera n. sp.

5. Virgularia a/finis Kor. Dan.

6. » wira bills (L.)

7. * Virgularia cladiscus nom. n.

= 8. Stylatula (Diibenia) elegans Kor. Dan.

9. * Pavonaria finmarchica (M. Sars)

= 10. Hahpteris christii (Kor. Dan.)

PENNATULIDA.

II

26
27
28.
29
30

3 1
32
33

21. Protoptilum lofotcnse Kor. Dan.

22. » mohni

23. • carinatuni

24. armatum

25. Ftmiculina quadrangularis (Pall.'

Kophobclcmnon stelliferum (O. F. M.)
abyssorum Kor.Dan.
moebii » »

leuckartii Koll.

Gunncria mirabilis Kor. Dan.

Bathyptilurn carpenteri Koll.

Umbellula encrinus (L.)
gracilis Marsh.

n. Protoptilum thomsoni Koll.

12

J 3

14.

16.

17

*Protoptilum carpenteri Koll.

denticulatum n. sp.
*Distichoptilum gracile Verr.
*Funiculina quadrangularis (Pall.)
* Anthoptilum grandiflorum (Verr.)
* Anthoptilum murrayi Koll.

18. *Kophobelemnon stelliferum (O. F. M.|

19. Bathyptilum carpenteri Koll.

20. * Umbellula encrinus (L.)

21. * » lindahli Koll.

Fam. Pennatulidce Koll.

Pennatula Lam.
Pennatula aculeata Kor. & Dan.

PL I, fig. I.
Pennatula aculeata Kor. Dan. 1858. Forhandl. i Vidensk. Selsk. i Chnstiania 1858, p. 25.

Kor. Dan. Fauna littor. Norvegise. III. 1877, p. 86. Tab. XI, figs. 8—9.
» distorta var. aculeata Kor. Dan. Nye Alcyon. Gorgon, etc. 1883, p. 24. Tab. XI, figs. 5—10.

phosphorea var. aculeata Koll. Monogr. p. 134 and 366.

Marshall. Penn. * Triton , 1883, p. 123. Tab. XXI, figs. 4, 5, 7;
Tab. XXII.
» americana Moroff. Octoc. 1902, p. 381.

The Ingolf> has brought home 2 specimens from Station 25 in the Davis Straits; the steamer
sThor , in 1903, has taken 10 specimens at its Station 167, one specimen at St. 166, to the south of
the Vestman-Islands. Of the two Greenland specimens, the smaller one especially (Nr. 1) appears so
peculiar from its violet colour and its long spines (dorsal zooids), that at the first glance it might be
thought to be a new species; moreover, the wings of the pen are unfortunately highly contracted and
broken by the hauling-in, so that the form of the pen has been somewhat distorted (comp. fig. 1).
Only the calyx-teeth and the points of the zooids are red, and the peduncle is of a reddish violet
colour quite down to the terminal bulb. The other, larger specimen (Nr. 2), which is excellently pre-

12

PENNATULIDA.

served, agrees completely, on the other hand, with Penn. aculeata as to outer appearance, has the same
long, narrow wings etc.; but in this specimen also the red colour passes into the violet, even if the
red is more conspicuous.

By closer comparison with our Museum's type-specimens of Penn. aculeata Kor. & Dan. from
Christianssund , complete correspondauce is found in such features as the number of polyps in the
wings, the double row of lateral zooids at the base of each wing, the 4 — 6 rows of dorsal aculei; but
in the length of these latter there is a conspicuous difference. Whilst the spines of the Norwegian
specimens are 1 — 2" 11 " long (according to Koren and Danielssen they may grow to a length of 3'3' r "":
Fauna litt Norveg. 3, p. 87), in the two Greenland specimens they are fully 4'"'" long. American
specimens in our Museum (from the Atlantic: 36 55' — 58 N. Lat. 7 1° 13'— 14' W. Long., depths 207 — 214
fathoms, and 40 N. Lat. 70 W. Long., 260 fathoms), both by their strong colour, which is redder, how-
ever, and by their long and stout spines, resemble the Greenland specimens very closely; when to this
is added, that the Iceland specimens brought home by the «Thor», stand as to colour between the
two Greenland ones, and that specimens of Penn. aculeata are also enumerated from several other
Atlantic localities with spines of more than 3 and up to 4 ,ni " (Marshall, « Triton >>-Penn. p. 124, Kolliker,
Monogr., Appendix p. 366), there can be 110 doubt that the specimens from the Davis Straits are justly
referred to this species. Both the colour and the size of the spines also, to judge from several state-
ments in the literature of Penn. aculeata, would seem to vary according to the depth at which it is
found; at great depths the ventral calyx-tooth of some of the large zooids is developed to a powerful
spine, and the colour becomes more intense, deeply red or violet.

The following features refer to the two Ingolf specimens:

Total length

Length of peduncle

Number of pinnulae

Length of a well-developed pinnula
Number of polyps in such a pinnula
Length of the largest aculei (zooids) .

Nr. 1.

105 mm
50 •

19/17

8-9

4. mm

Nr. 2.

115 mm

4 S

22/22
24 mm — 25 mm

7—9

4 nun

The specimen Nr. 1 (PI. I, fig. 1) is provided with spicules differing somewhat from the form
common to the species, being more slender, more spatulate at the end, and very strongly coloured;
some of the polyp-spicules, moreover, reach a very considerable length; the largest ones measured are
3-i20 mm long and o-io8 mm broad, whilst the smallest measured (from the tentacles) o-o8o mm in length
and ca. o-oo8 ,nm in breadth. The specimen Nr. 2 shows, both in the form of the spicules and their
length, a tolerably close agreement with the American specimens of P. aculeata examined. The longest
measured spicules from one of the aculei were 2 - i6o mn) long and o-oo,6 mm broad.

In the specimen Nr. 1 the top of the rhachis is so much contracted and partly damaged, that
its more particular structure cannot well be made out; in Nr. 2 the rhachis is terminated above by a
rather large individual with three-toothed calyx; this may possibly be a somewhat uncommonly deve-
loped terminal zooid, possibly the transformed terminal polyp; in the latter case, the terminal zooid
must have disappeared.

PENNATULIDA.

The ii specimens brought home by the steamer «Thor» from the south of Iceland, are, as
already indicated, of an intense, deep-red to violet colour; size and other proportions are given below:

Nr. I.

Total length

Length of peduncle

Number of wings 2 3 24

2o Inm

3 »

8

2 mm

Nr. 2.

Nr. 3.

Nr. 4.

Nr. 5.

Nr. 6.

Xr. 7.

Nr. 8.

Hi min 1 105 mm I0 2 mm joomm 86 m
58 »

Length of a well-developed wing
Breadth of such a wing above the base
Number of polyps in such a wing . .
Length of the largest spines (zooids) .

53 •

46 >

45 »

3 S »

J9/20

»/ig

17/17

17/17

22 tnm

jo mm

24 Inm

24 mm

3 »

ca. 3 »

3 »

3 »

10

IO

9

9

2 mm

t mm

2 S mm

3 mm

Jjcrom 7c mm

38 » 40

21/21 I4 M

2i mm j a mm

2'5 » 2 - 2 •»

9 7-8

2inm T mm

65 mm
30 »

'4/15

2 1 mm
2 »

7-8

^ mm

Nr. 9. Nr. 10.

65mm
35 »
r 3 13

jc mm
2 I

7

2*2 mm

Nr. 11.

14 »
8 D

21 mm I 3 mm

2-6—3 » 1-5 »

9- II j 3—4

^ — - -c mm c_ j nun

None of the first ten specimens show any distinct terminal polyp; some (Nr. 1, 4, 5, and 7), on
the other hand, have a distinct terminal zooid, in Nr. 1 with a spiniform, large, but distinctly two-
pointed calyx; in some, rudimentary wings with reduced polyps are found at the apex; in some
distinct top-zooids. Nr. 10 is a torn-off pen, not quite complete below. In the specimen Nr. 11 (from
.St. 166), on the contrary, the terminal polyp and terminal zooid are well developed; in spite of its
slight size it may already be recognized quite distinctly as belonging to the species aculeata\ a
few of the zooids are not only larger than the others already, but have also the long calyx-tooth
distinctly marked; moreover, the specimen is of the same dark, violet-red colour as the ten specimens
from St. 167.

Pcnnatula aculeata is originally known from the western coast of Norway; it is here (see
Grieg 30, p. 9) said to go farther north than P. phosphorea, viz. to the Lofotens, and to greater depths
(400 fathoms); it seems not to have been found farther south than Askevold (Sondfjord). Later it has
been found by the English expeditions of the Porcupine .>, Triton*, and <Knight Errant* at the
following places in the Atlantic: west of Brittany (48 26' N. Lat. 9 44' W. Long., 358 fathoms); north-
west of Rona (the Hebrides) (59° 40' N. Lat. f 13' W. Long., 556 fathoms, and 59 37' N. Lat 7 19' W. Long.,
520 fathoms) ; further, by the Prince of Monaco near the Azores (38° 34' 30" N. Lat. 30 43' 30" W. Long.,
1287 M.); by the French expedition of the «Caudan» in the Bay of Gascony (46 28' N. Lat. 7 W.Long. 1 ),
1710M.); by the Travailleur», west of the Spanish peninsula (P. Fischer, 23, p. 38). To these loca-
lities are to be added as new ones : south of the Vestman-Islands, 63 5' N. Lat. 20 7' W. Long., 557 M.
(the ,,Thor», St. 167, l V? *<PZV and 52 57' N. Lat. 19 58' W. Long., depth 956 M. (the «Thor», St. 166,
^/■j 1903). The species is not known from the North Sea nor the adjoining seas, the Skager Rak,
Kattegat etc., nor from the Mediterranean. On the American side of the Atlantic it is very common,
and is hitherto known from the Gulf of St. Lawrence as its northernmost locality (Whiteaves), to off
Chesapeake Bay. To this occurrence on the American coast is to be added the new locality, St. 25 of
the Ingolf», Davis Straits, 63° 30' N. Lat. 54°25' W.Long., 582 fathoms, by which observation its northern
limit on the American side has been carried considerably farther north. According to the above, this
species is assuredly to be regarded as a purely North-Atlantic one; it is not known south of the
Azores, nor on the American side farther south than 33 N. Lat. (Agassiz: Three Cruises of the

'I Reckoned from the meridian of Paris.

14

I'ENNATUUDA.

Blake, vol. II, p. 142. Bull. Mus. Comp. Zool. vol.15); it seems to be limited essentially to the margins
of this part of the Atlantic and to the slope into great depths. As to its bathymetric range
the lowest depth is 30 fathoms, at Norway, but upon the whole it occurs in greater depths; on the
American side, where, as mentioned, it is very common, it is still found at 1255 fathoms (Verrill:
Rep. U. S. Fish. Comm. 18S3 (1885) p. 509), but it is most abundant between 150 and 300 fathoms.

Whilst the very closely allied Penn. phosphor ea has not yet been found in America, a Penn.
americana has been described by Moroff. According to the description, it is beyond all doubt,
however, P. aculeata. M.'s specimen is also from Massachusetts (see Zool. Anz. 1902 and Zool. Jb.
17. Bd., 1902).

Pennatula phosphorea L. var. Candida Marsh. & Fowl.

A. Milnes Marshall and G. H. Fowler: Report on the Pennatulida dredged by H. M. S. : Porcupine».
Trans. Roy. Soc. Edinb. vol.23, P ar t 2, 1887, p. 456, PI. XXXI; PL XXXII, Fig. 3.

This form has been founded by Marshall and Fowler 1. c. on three fragments taken by
the «Porcupine» in 1869 at 62 1' N. Lat. 5° 19' W. Long. Our Zoological Museum has obtained 17
specimens from the Vestman Islands, 7 from the physician Thorsteinn J 6ns son, who, according
to «Skyrsla um hid islenzka nattiirufrseflisfelag arid 1897 — 98', p. 12, has also given a specimen to the
Natural History collection at Reykjavik; 8 specimens from the cruise of the «Diana» in 1900 (A. C.
Johansen); 1 specimen from 1901 (R. Herring), and 1 from the collection of the «Thor» in 1903.
The species is peculiar by its being perfectly white in spirit, almost all the spicules being colourless;
the animal, when living, may be of a slight reddish colour (according to Horring), and in a single
specimen the red colour is still distinctly seen (see below). Otherwise it agrees in several features
with Penn. phosphorea var. angustifolia Koll. (Mouogr. p. 130). The pinnules are slender, and the
calices of the polyps are long and conspicuous, provided with 8 long teeth formed by spicules; the
number of pinnules 011 either side 17 — 24 (often differing somewhat on the two sides of the rhachis;
comp. below). The tentacles of the polyps carry spicules along the dorsal side of the stem. The
large spicules, as elsewhere in P. phosphorea, are fusiform, and thus only conspicuously triangular at
the ends; in section, they are somewhat round; the smaller ones, on the other hand, are altogether
triangular. The largest polyp-spicules measured are ro72 mm long, and o - 072 mm thick; the smallest are
0*04 — o-048 mm long, but the general length of the spicules of the tentacles is 0-096 — o-i28 ,nm .

The specimens in hand show the following particular features:

Nr.

1

2

3

4

5

6

7

8

9

10

II

12

13

14

15

16

17

Total length in mm.

37

65

65

65

66

72

72

75

75

80

SO

80

S3

85

90

7S

Length of stalk. . »

15

27

3°

3°

32

3°

35

32

34

35

35

40

40

41

42

36

Number of wings. . .

"/12

22/21

•9/i9

2 °/i8

i7/i 9

22/ 23

22/22

i3/i 7

19/18

!9/20

23/24

21/21

20/21

22/21

20/20

2 5/2 3

iS/tS

Number of polyps in

one of the large

wings

6

9 — 10

S— 10

9

11

9

10

9

8

8-9

10—12

10

9-11

8— 11

9

10

8—10

Length of a wing in

10

15

11

13

14

12

11

16

14

17

16

ii-S

19

12

t5

12

13

PENNATULIDA.

1 5

The specimen Nr. i is distinguished by having rather a strong red tinge, so that it approaches
somewhat the common phosphorca; both polyps and zooids contain red spicules in considerable num-
bers; the polyps are red, but with white calyx-teeth and white lateral edges; the edges of each wing
are therefore white, and the red surface is streaked with narrow white lines marking the boundaries
between the coalesced polyps. A quite similar distribution of the colours, only with a far deeper red,
is not uncommon in typical pJwsphorea specimens from the Kattegat. Further, this specimen has kept
both terminal polyp and terminal zooid, as is also the case with the specimens Nr. 4 and Nr. 9. In
the specimens Nr. 3, 7, 12, 15, 16, and 17 the quite distinct terminal zooid is preserved in its full
development, but the terminal polyp is transformed into a zooid, either of equal size with the terminal
zooid (Nr. 15), somewhat smaller than this (Nr. 12), not much smaller (Nr. 3) — or quite atrophied
(Nr. 7, 16, 17). Although the specimens in question, with the exception of Nr. 1, are large and must
be considered as fully developed, the features mentioned are quite uncommonly distinct. The other
specimens show much altered features at the upper end of the rhachis; some specimens (e. g. Nr. 6)
show very beautifully a number (4) of large ^top-zooids>, amongst which, probably, both the terminal
polyp and the terminal zooid are to be sought. In the specimens Nr. 2 and Nr. 7, the peculiarity is
found that one of the dorsal zooids near the middle of the edge of the rhachis is developed as a soli-
tary polyp, 7 and fully 4 inm long respectively. The specimen Nr. 14 approaches somewhat the «lanci-
foliate form x ). Most specimens contain ripe sexual organs, and some seem to contain larvae.

Occurrence. This light-coloured form has been taken by the « Porcupine*, as mentioned, on
62°i'N. Lat. 5 19' W. Long., accordingly E. of the Faeroe Isles, at a depth of 114 (English) fathoms.
All the specimens here mentioned are from the Vestman-Islands; with regard to Nr. 2, 5, 8, 10,
n, 13, and 14 there is no information as to the depth or the bottom; but to judge from the attached
particles it must have been black Basalt-mud; Nr. 9 has been taken at 50 — 60 fathoms, E. of the
Vestman-Islands, 2 1 / 2 miles E.N.E. of Storhofdi; the others have been taken (the -Diana -, 1900, St. 41)
on clay-bottom , at a depth of 68 — 70 fathoms; Nr. 17, the «Thor», 1903, St. 169, 63 24-5' N. Lat. 20° 1'
W. Long., depth 120 — 158 m., ooze. All the specimens have been taken during the summer months
(July, August).

This pale or white variety probably occurs on restricted localities within the wide extended
area of the species, but with regard to the more particular conditions under which it is produced and
thrives, we have no information which could render any certain statement possible. The question can
scarcely be of a casual albinism, as all the specimens known from the Vestman-Islands belong to the
same pale variety, and in its red forms Penn. phosphorca is altogether unknown hitherto from any
locality at or round Iceland or the Faeroe Isles. Other discoveries of white forms of Penn. phosphorea
are, so far as I know, only referred to by two authors. Kolliker describes a white specimen in his
Monograph, p. 133; it belongs to the Museum in Copenhagen, and I have thus been able to compare
it with the form from the Vestman-Islands; it is very different from this latter, and Kolliker with
good reason has determined it as a subvariety alba of Penn. ph. var. lancifolia. Unfortunately, we

') When Marshall and Fowler give 4-2 r ' n as the greatest length of a pinnula in their specimens, this is surelv due
to a misprint; such a length is scarcely found in any form of P. phosphorea, and the figures on Plates 31 and 32 give also
a length of only ca. 14 — I5 ram .

j5 pennatuuda.

have no information at all as to the place where the specimen was found. Esper's Pennatula alba
which Kolliker has referred to his var. angustifolia, closely resembles, to judge from the figure, 22,
pi. VI, the specimens in hand from the Vestman-Islands; their colour is said to pass almost impercep-
tibly into a reddish one; but we have no information as to the place where these specimens of Esper
were found.

Verrill mentions a white specimen of Pennatida aculeata (Amer. Journ. Sc, vol.23, P-3 IO > an( ^
Report... tBlake p. 3) taken by the Fish-Hawk > at st. 1025, depths 216 fathoms; he says of it: «This
is doubtless only an albino* ; it appears to have been taken together with numerous specimens of the
common red form of Penn. aculeata (comp. Am. J. Sc. 23, p. 315).

Remarks. According to what has hitherto been known, it must be regarded as hardly pro-
bable that Pennatula phosphor ca should be found outside the Atlantic. Studer, however, in his preli-
minary report of the &Albatross» Expedition (p. 55) has given Penn. phosphorea as from the Pacific, from
o° 19' N. Eat. 90 34' W. Long., 331 fathoms, and Pennatula distorta, Dan. Kor, var. pacifica, n. var.»,
from i° 7' N. Lat. 8i° 4' W. Long. Of the former, it is said that the (single) specimen was not to be
distinguished from the Atlantic (i. e. the typical) phosphorea L. As to Penn. distorta Kor. and Dan.,
this species cannot be maintained, in my opinion; it is nothing but Penn. phosphorea with unusually
narrow and (casually?) distorted wings; and the Penn. distorta var. aculeata described by the same
authors, is obviously only ^distorted* specimens of the species Penn. aculeata. And with regard to
Studer's «Penn. distorta, pacifica^ his remark that the colour is whitish yellow, and that the calyx-
spicules appear to be longer and more conspicuous, seems to me to indicate that another species must
be in question; from the Asiatic side of the Pacific several Pennatttla-species are known, among others
just such forms with narrow leaves in which yellow or reddish yellow is prevalent in the colour, and
where the type of spicules, moreover, is different from that of phosphorea; also the depth, 1740 fathoms,
must be regarded as somewhat great for the supposed species. Th. Moroff (Zool. Anzeiger 1902,
p. 579, and Zool. Jahrb. Abth. Syst. 17 Bd., 1902, p. 380) has established a P. phosphorea var. longispinosa
from Japan (1 specimen); unfortunately, his figures show only that the specimen is a Pennatula; but
some remarks in the description would indicate that phosphorea is not in question (for instance the
broad streak between the « ventral* zooids).

Pennatula grandis Ehrb.

Pennatula grandis Ehrenberg. Die Corallenthiere des rothen Meeres, 1834, S. 66.

borealis M. Sars. Fauna litt. Norvegias, I, 1846, p. 17, PI. II.
Ptilella borealis Gray. Catalogue of Sea- Pens, 1870, p. 21.

This splendid species has not been taken by the Ingolf- itself; but during one of our sojourns
at Thorshavn I received from Mr. Finseu, apothecary, a specimen taken on «the bank south west
of the Faeroe Isles at a depth of 90 fathoms »; later, 19 specimens from the Faeroe Isles have been
sent to our Museum by the agent Mr. Miiller, taken on the fishing bank east of Fuglo, ca. 150
fathoms. With the exception of one individual (Nr. 20) all the specimens are large and well-developed.

I can add nothing essentially new to the former descriptions (by Sars, Kolliker, Koren & Danielssen,

PENNATULIDA.

17

Verrill, a. o.) of this species which can scarcely be confounded with any other. I shall only make a
few observations on the spicules, as these have not been described in particular by the earlier authors.
The spicules are of a marked triangular type ; in the larger types especially the polyp-spicules
(i. e. of the calices, the upper part with the tentacles containing no spicules) the three ridges
which constitute the spicule show their free edges to be distinctly thickened, whilst the sides
are hollowed; in the red spicules, therefore, the colour is deeper at these thick edges. The polyp-
spicules measure 0.144 — o.8g7 mm long, and 0.016 — o.048' nm broad. The colourless part of the peduncle
is provided with short, coarse spicules 0.088 — o.i20 mm long, the thickest ones being o.o24 mm thick; these
spicules are of a very slight reddish colour; the deep-red spicules from the baud-shaped edge of the
bulb are 0.096— o.200 mm long, and 0.016— 0.032""" broad.

The specimens in hand from the Faeroe Isles show the following particular features:

Nr.

1.

2.

3-

4-

5- 6. 7 -

8.

9-

IO. II. [2.

13-

14-

15-

16.

17-

iS.

19-

20.

Total length in mm.

445

445

435 435 433 43° 43°

425

420

420

410 400

398

595

39°

380

370 315

210

107

Length of peduncle,
to the edge of the

bulb . ... in mm.

125

no

90

130 124 120

no

108

no

125

no

82

92

IOI

80

90

105

30

Diam. of bulb -

32

34

3°

35

3 2 3°

30

29

27

30

27

30

34

27

35

20

23

23

7

Number of polyps
in a well -deve-
loped, arge wing

83

73

70

77

74

79

70

73

6S

82

7<S

67

72

78

81

62

84

25

Number of wings.

39/40

42/44

44/41

37/37

44/43

39/39

37/35

37/38

3 6 /35

37/40

3939

41/41

42 4I

3939

37/37

34/34

39/39

(3 I 32)

i8/ I?

Length of a large
wing . ... in mm.

. . .

45

44

35

41

23

38

43 45

47

22

Greatest breadth of

do in mm.

23 3°

...

26

25

23

22

27 24

17

8

Length of polyp-
calyx in mm.

4—5

4-5

3-5

3-5

3-4

5

5

4-6

5

4

The specimens Nr. 18 and 19 are defective; in Nr. 18 the lowermost wings are wanting, the tissues
of the bulb and the peduncle having been rubbed away; Nr. 19 is broken off below the bulb; the rhachis,
which is complete, is ca. 70 mm long. Nr. 20 is the smallest specimen of this species I have ever seen amongst
the large number I have had the opportunity to examine in our Museum and in Bergen and Christiania;
Verrill mentions that he has had several young specimens and one very young, but he gives no
measurements nor any further information. For the sake of comparison, I may just mention that the two
specimens of M. Sars, upon which the species P. borealis was established, were of a respective length
of 420 (16") and ca. 810""" (31"}, with 37 and 57 pairs of wings; in other Norwegian specimens measured
by Koren and Danielssen, the length is given as 250—780""" with 30—70 pairs of wings and 48—100
polyps, whilst Verrill mentions a specimen of 530 mm in length with 36 pairs of wings, as one of the
largest American specimens.

The colour in some specimens is brownish or dark brownish-red (especially so in Nr. 20, partly
in Nr. 18 and 19), but most frequently of a beautiful red or reddish yellow.

Occurrence. The species was only known hitherto from the fjords and coast-regions of
Norway, the eastern coast of North America where it is common on the fishing banks off New

The Ingolf-E*pedition. V I. 3

jg PENNATULIDA.

Foundland, Nova Scotia, and the northernmost of the United states (as far as off Nantucket Island
and Martha's Vineyard). To these localities are now to be added the banks round the Faeroe Isles.
It has been taken at depths from 50—1255 fathoms 1 ).

Pennatula prolifera n. sp.

PI. II, figs. 15-24.

From two stations in Davis Straits, 24 and 36, we have obtained a / 3 V,</-w/«/^-speeies in very
large numbers, from the former station about 40 specimens, from the latter upwards of a hundred;
but all the specimens are young ones, several of them as small and apparently as young as the
youngest known stages of Pennatula phospkorea 2 ); even the largest and most developed specimen is
still little developed, as will be shown more particularly below, so far as the pen's provision of polyps
is concerned. It is a very difficult thing therefore to characterize the species; or, to speak more cor-
rectly, it is impossible to diagnose it briefly by such features, as are elsewhere used in distinguishing
species within the genus Pennatula. That the species is a new one not hitherto described, will be
obvious, I think, from the following description, in which I shall give an account of the peculiar mode
of propagation in this species which has led me to give it the specific name prolifera.

All the specimens were white, with the exception of two from St. 36, which were light-red;
but these also have now turned as white as the others in spirit.

The largest specimen (from St. 24) has to some degree the penniform appearance (figs. 15, 16).
The total length is 35"™, in which the rhachis accounts for i8 mm , measured from the terminal zooid
(see below) to the lowermost polyp-bud, the peduncle itf/z"""; the breadth of the pen is 4.5 mm (the
polyps being pressed in against the stem); the rhachis itself is ca. 1.5""" broad, the terminal bulb of
the peduncle is 2.5 mm long and ca. i.5 ram broad. On either side of the rhachis are eight distinct wings
and below these four rudimentary wings.

The calyx-mouth of the polyps is provided with 8 long and strong spines supported by long
spicules. The form of the calyx, as is ordinarily the case in the genus, is somewhat different accord-
ing to the degree of retraction; when the polyps are quite retracted, the calyx-teeth are closed and
the whole calyx has a lengthened oval form reminding one of a narrow grain of barley. The longest
calices are 5 mm long and ca. i mm broad. The tentacles are on the aboral side provided with longi-
tudinally arranged spicules, both on the stem and the lateral branches.

With regard to the arrangement of the polyps it has first to be pointed out that no terminal
polyp is present: the pen ends above in two polyps placed (almost) side by side and equally developed;
between their bases the upper end of the rhachis itself is found, formed by a large zooid (with two
calyx-points). Each of these two uppermost polyps forms a wing (I). This uppermost pair of wings
is separated from the next pair (II) by a somewhat long part of the rhachis, this latter pair again
by a somewhat shorter distance from the third pair (III), whilst the following pairs (IV — VIII) follow
one another more closely.

') Verrill: Rep. U S. Fish. Comin. 1SS3, p. 509.
2 ) Figured by Jungersen. 1888, PI. V, figs. 1—2.

PENNATULIDA. 19

Apart from the uppermost wing the other wings contain more than one polyp, as at all events
a rudiment of one or two polyps more is found in each wing to the ventral side of the oldest one.
In the uppermost pair of wings but one (II) the polyp of the left side (1) carries a small protuberance)*),
which I take to be the first trace of individual Nr. 2; the right wing has a distinct individual 2, but
still in the zooid-stage; in wing III, polyp 2 is large and well-developed on either side of the colony,
but nothing is seen of 3; in wing IV, polyp 2 is still larger, and 3 is present as a zooid; in wings
V— VII there are also 3 individuals, but 2 is smaller than in the preceding wings, and decreases in
development below, while 3 is still only a zooid in all of them; in wing VIII there are only two
polyps, of which Nr. 2 is only a zooid. The four rudimentary wings (IX— XII) consist, each of them
- more or less distinctly -- ot two individuals, both still zooids.

Of zooids, there are 2 longitudinal rows dorsally, one on either side, separated by a bare
streak (i. e. devoid of individuals), about o.5 mm broad below, narrowing above (a similar naked streak
runs along the rhachis between the polyps of the ventral side); in each row an indication of a
grouping into 3 — 4 zooids for each wing may be traced. The series is already- indicated below at the
smallest rudiments of wings. - - Between the uppermost wing and the next the series is very open,
and onlv one zooid is found at each of the uppermost polyps (I). The zooids look like small scales
supported by spicules. At the upper end of the rhachis the rows are completed by a median, con-
siderablv larger, single top-zooid, Z. provided with two distinct calyx-points 1 ). Of lateral zooids, one
seems to be begun immediately above each wing, with the exception of the two uppermost pairs of
wings. These rudiments, however, are very small, and difficult to determine with certainty.

The spiculation is copious and dense; the longest spicules are found on the polyps, the
next on the zooids and rhachis where they are mostly arranged longitudinally; there are also many
transverse, rather long and thin, spicules on the peduncle, intermingled lower down with longitudinal,
short, oval ones; towards the terminal bulb the spiculation diminishes to open, longitudinal rows of
short, oval spicules, and the bulb itself has no spicules, and is therefore transparent, so that the
interior longitudinal septum is seen (but no terminal pore; I think upon the whole that the existence
of such a pore in Pennatula normally is very doubtful 2 ).

The spicules (espeeiallv with regard to the long ones) are of quite a different type from that
predominant in Penn. phosphorca. The fact is that all the spicules (excepting the oval ones in the
peduncle) are distinctly and sharply triangular, recalling mostly those of Penn. grandis. The longest
measured are 0.784""" long, 0.04""" broad; the short and thick ones are respectively o.i6o mm and 0.04—
0.112""", and 0.032"""; the small oval spicules of the peduncle are 0.064— o.o88 mm long, and o.oi6 mm broad.

The calcareous axis terminates above, bent to the right, opposite to the third uppermost
wing, and below, also curved, just above the terminal bulb.

The sexual organs are fully developed; they may be seen shining through the po-
lyps without any further preparation, but still more distinctly by clearing with oil of cloves; in

■) In this specimen the top-zooid has by chance been wedged in between the two uppermost polyps, so that it is
only partly seen in the figure, but both from the dorsal and the ventral side.
2 ) Comp. Jungersen, 1SS8, p. 161.

3*

20

PENNATULIDA.

all the more developed polyps, from the uppermost to the two lowest, large eggs (or larvae) are
seen; in the rhaehis, on the other hand, no sexual products are found.

In the other specimens, about 150, that I have been able to examine, there is found, a so to
speak continuous series of stages, from the one just described down to a stage where the colony is
provided with only one pair of wings (I). Representatives of these stages may be found figured on
PI. II, figs. 17—24.

One specimen (St. 24) of a total length of 29""", is provided with six distinct pairs of wings, and
below these with tiny rudiments of two pairs more. Each wing of the uppermost pair (I) consist of
three individuals: one fully developed polyp, one somewhat smaller, and one still a zooid; the next
wing (II) shows the same development. The wings III — V still consist of three individuals, but two
are in the zooid-stage, and the most ventral, youngest one, is already very small in IV, in V both are
very small; VI consists of only two individuals, a small polyp, and a small zooid; both the rudiments
(VII, VIII) consist of a larger and a quite small zooid.

Another specimen (St. 24), of 25 mm , shows 5 pairs of wings and 2 pairs of rudiments; all the wings
consist of two individuals, the two uppermost of a developed polyp and one little developed, (wing I
on the right, however, has three individuals, being provided with a small zooid), the others of a polyp
and a zooid, both decreasing in size from above downwards; the rudiments consist of two zooids.
Other smaller specimens of 2i mm , also show 5 pairs of wings and 2 or 3 pairs of rudiments; others
again (25 — i6 mm ) 4 pairs of wings, 3 pairs of rudiments, others (17 — 13 1 "" 1 ) 3 pairs of wings, 3 pairs of
rudiments; others again 2 pairs of wings and 2 pairs of rudiments, PI. II, figs. 17, 18; in all these
specimens each wing consists of two individuals, a polyp and a zooid. Finally, we have several
specimens with only one pair of wings, and below one or two pairs of rudiments, and some specimens
with only one pairs of wings and no rudiments below (PI. II, tigs. 19, 20). Each wing in these latter,
however, consists also of two individuals, a developed polyp and a small zooid.

The proportion between the rhaehis and the peduncle as to length is somewhat varying; some-
times the former, sometimes the latter is the longer, but often they are about equal in length. Devel-
oped sexual organs (sometimes larvae) are seen in most polyps, at all events in the developed ones;
in quite small (io mm long) specimens indeed with ouly one pair of wings and 2 pairs of rudiments, I
have seen large and developed sexual organs in one or both of the large polyps. Even the smallest
specimen, of 6 mm , represented in figs. 19, 20, shows (by clearing with glycerine) the sexual organs, though
not yet ripe. All the specimens have the two rows of zooids (z), but their number in each row is
very varying, increasing with the number of wings; in the specimens with only one pair of wings
only one or two zooids are found in each row. The small lateral zooid above each wing is only
seen in the tolerably well developed colonies. A common feature in all the specimens is further, that
the stem contains a calcareous axis, and that the rhaehis terminates above in a large zooid (Z). This
series of stages therefore is evidently a developmental series, in which the last-mentioned specimens
with only one pair of wings are the youngest. That this developmental series belongs to a species
of the genus Pennahtla is undoubtedly a fact; but the rather obvious supposition that the young
stages of the pale variety of Penn. phospltorea var. Candida before mentioned might be in question, is
inadmissible on account of the difference in the type of the spicules, and a closer comparison with

PENNATUUDA. 21

the developmental stages known of P. phospJiorca renders it quite untenable. Penn. phosphorea
alwavs shows more numerous and more advanced polyps in the single wings, at stages where the
number of wings is as in specimens of the present new species, and the colony is upon the whole
more developed at corresponding sizes; on the other hand, the young colonies of P. phosphorea show
no ripe sexual organs. These features, however, might be thought to be influenced by the conditions
of life at such great depths as the one from which the new form has been obtained. The most con-
clusive thing to my mind, however, is the great difference in the construction of the upper end of
the rhachis. In young colonies of Penn. phosphorea the rhachis always terminates above in a terminal
polyp, and on the dorsal calyx-base of this there is a large terminal zooid which completes the rows
of dorsal zooids' )• Both terminal polyp and terminal zooid are found even in well-developed colonies
with complete pen-form (for instance, in specimens of more than 35'""', with 12 pairs of wings, the
largest of which are provided with 5 — 6 polyps; but more rarely in still larger specimens); in the new
species however, no stage not even the youngest, shows the least indication of a terminal polyp: the
rhachis ends as stated in a large zooid, and the two nearest polyps (or wings) are equally developed
and placed at almost the same height; thus the younger stages especially have a dichotomous appea-
rance, markedly different from that of Penn. phosphorea.

According to what has been known hitherto, the primary individual, developing directly from
the larva which arises from the egg, is in all Pennatulids a polyp with tentacles (see my treatise
pp. 162, 173); as this polyp grows greatly in length, its lowermost part below the calyx forms the
stem (rhachis ~|- peduncle) ; on this lengthened part the other individuals, polyps and zooids, appear
gradually (in some genera zooids are also formed on the polyps). During the growth of the colony
the terminal polyp may- later disappear; this is probably the case in most of the genera of sea-pens,
and it is the rule in Penn. phosphorea. Where the disappearance of the terminal polyp is not due to
a mere displacement (as in Kophobelemnon), it may be due either to an abortion or a transforma-
tion into a zooid. In the genus Pteroeides the latter seems to be the case; when the colony has
grown to a certain size, the end of the axis, at all events, carries a very large zooid instead of a ter-
minal polyp, (see Kolliker, Monograph, p. 356, my previous treatise, p. 172, and v. Koch 2, p. 398). In
Penn. phosphorea, the end of the rhachis of large colonies carries generally several very large zooids
top-zooids»; amongst these presumably, are both the original terminal zooid and — by a retrograde
transformation — the terminal polyp and some of the uppermost wing-polyps (comp. Jungersen p. 168,
and v. Koch). With these facts in view we return to our new Pe//nalula-species. It may be main-
tained with a probability, bordering on certainty, that this species like P. phosphorea, has also had a
stage not only with a terminal polyp, but also with a terminal zooid on its calyx-base. — I shall only
recall the fact that such a distant form as Renilla agrees completely in this respect with P. phosphorea.
How, then, has the terminal polyp disappeared here?

1) If a displacement had been the cause, one of the two uppermost polyps would have to be
regarded as the terminal polyp, and the large zooid of the end of the axis would represent the terminal
zooid. This supposition, however, is inadmissible, partly because this large zooid is not placed in the
same way as the real terminal zooid in P. phosphorea and Renilla, on the dorsal calyx-base of either

M Comp. my previous treatise p. 16S, and Marshall & Fowler, Penn. tPorcupine-, PL XXXII, figs. 5 and 7.

22 PENNATUUDA.

of the polyps in question, partly because both the upper polyps (I) always (or, at all events, with
very rare exceptions) distinctly appear as lateral polyps, i. e. wing-polyps, a polyp Nr. 2 sometimes even
a Nr. 3 being found at their base on the ventral side of the colony (whether these latter ever grow-
to their full size is of no consequence in this connection); in Penn. phosphorea (and Renilld) the term-
inal polyp is always single, and continues to be so.

2) The large zooid on the end of the rhachis might be interpreted as the terminal zooid, the
terminal polyp itself being aborted. This hypothesis would require so early an abortion of the terminal
polyp, that we know of no parallel in any other Pennatulid whatever.

3) The terminal polyp might be transformed into the large zooid, which ought, in this case,
to be denoted as the top-zooid . This interpretation would be met by a difficulty quite similar to
the one in supposition Nr. 2, and further by the fact that the original terminal zooid must have dis-
appeared at the same time — that is to say, at a very early stage. This seems little probable, when
it is remembered that v. Koch has in one case found the real terminal zooid to be preserved in a
specimen of P. phosphorea, in which the terminal polyp seemed to have disappeared, a full grown
specimen with 38 pairs of wings (comp. v. Koch, 2, p. 397, fig. Y).

Although, in spite of the difficulties indicated, the possibility is scarcely quite rejected that one
of the two latter suppositions, or both, may apply to this species, I believe I have observations which
show quite a different interpretation to be the correct one, and make the others superfluous.

The fact is that amongst the many specimens from St. 36 no less than 18 individuals, and 3
individuals amongst those from St. 24, show that this species is capable of transverse fission. By this
fission only the upper end of the rhachis with the uppermost pair of wings (I) is separated. The
separation always take place close above the second uppermost pair of wings (II), and before it is
quite accomplished the part of the rhachis situated between the two uppermost pairs of wings evidently
stretches longitudinallv ; by this means, the part prepared for separation becomes similar to the
the youngest known stages of development described above; compare figs. 19, 20 with the top of
figs. 21 and 22. A clearing with glycerine or oil of cloves shows, however, that all the specimens in
hand which are in the act of transverse fission, contain no calcareous axis in the top-part destined for
separation; this part contains only the continuation of the longitudinal partition which farther down
in the rhachis contains the calcareous axis. This axis itself terminates always immediately below the
point of separation, generally with a hook turned to the right but this may also assume a somewhat
different form (comp. fig. 22). This want of an interior calcareous axis is literally the only difference
that can be pointed out between the top and one of the youngest independent colonies. The specimens
showing the transverse fission belong to the large and middle-sized; the total length is between ca. 17
and 25" ln, ; the number of wings is 4 pairs with 2 or 3 pairs of rudiments, and up to 7 pairs and 3
pairs of rudiments; each wing is generally provided with two individuals, of which Nr. 2 is almost
always a zooid; yet the largest specimen, with 7 pairs of wings, shows three individuals in wings II
and III: a large polyp, a quite small polyp and a zooid. Developed sexual organs are found in some
of the specimens, but in several others, e. g. the one represented in fig. 21, I have not been able
to find them. The process of transverse fission itself is seen in somewhat different stages; where the
constriction of the rhachis is not deep, the top is least lengthened (comp. fig. 23); where the constriction

PENNATUL1DA. 23

is so deep, that the top is only connected with the other part of the colony by a quite thin string,
the top (on the dead specimen) is hanging down like a broken flower along the stem of the colony,
and then its rhachis is always prolonged stalk-fashion, most often bent into the shape of an S, so that
the top gets a still more independent appearance (comp. fig. 21). Unfortunately, I have had no oppor-
tunity of observing the separation itself, the further fate of the separated top, nor that of the
remaining part of the colony.

That I am not here guilty of a misinterpretation of chance mutilation is certain; to be sure,
casual injuries are not wanting in many of the specimens in hand, but they have quite a different
appearance; here the spicules on both sides of the constriction are placed quite regularly and in such
a way as to be evidently arranged for the purpose; broken spicules are never seen as in injuries;
moreover, the facts that the constriction always takes place at the same spot, that it is found in
different stages and in a somewhat large number of specimens, and further that the top grows in
length and emancipates itself, as it were, tend without a doubt to show that this species really is
able to separate off the top. Of course it may be doubted whether this is a normal process; but I
think it an obvious conclusion to take it to be normal, and to regard the phenomenon as a kind of
propagation. The whole individualized character of the top, ready for separation, seems to me to
indicate that this part will later live independently as a small colony, develop an interior calcareous
axis J ), continue to grow and increase its number of individuals. The rest of the colony, the mother-
colony , will then probably form the large zooid at the place of constriction. One single specimen
(fig. 24), without any constriction, seems to me of some interest in this connection ; in that place of the
rhachis where the constriction is seen in the other specimens, we have here a large, single and median
zooid (Z') projecting in a polyp-like manner and interrupting the two rows of smaller dorsal zooids;
it is provided with two lateral calyx-points in quite the same manner as the zooid which terminates
the rhachis above between the two wings I (Z). The interpretation of this feature might be that the
large top-zooid had been formed exceptionally before the constriction, this having not taken place or
having been delayed for some reason or other. The specimen in question is otherwise of a good size
(27 mm ), has 7 pairs of wings and 3 pairs of rudiments and contains developed sexual organs (even
larvte); the calcareous axis reaches only to the height of the pair of wings III, that is to say, to a
tolerablv great distance from the spot where the large zooid is formed.

My conclusion is accordingly, that all the mentioned features of the transverse fission observed,
tend to show that this species may justly be called Pennatula prolifrrn. We should then have found
a mode of propagation within the Octactinia somewhat reminding one of that known in the Poly-
actinia, for instance in Goniactinia prolifera (M. Sars), and the Fungias; but the question, with regard
to this sea-pen, is of a transverse fission of a colony.

Several facts will be more intelligible viewed in this light. In the first place, the circumstance
will now be readilv understood why the species was taken in great numbers at both places where it
was found, and that, in spite of this, it is only represented by colonies of a youthful character. It is
obvious that the colonies need only a slight degree of development to be capable of transverse fission

■) It might, however, be possible that an interior calcareous axis might come up, before the separation took place;
the contrary cannot be said to be proved by the specimens in hand.

24

PENNATULIUA.

(as in Goniactinia, it is only young individuals that divide); the new colonies separated off by the
division will soon get so far as also to divide; even the mother-colony itself will soon again be
able to throw off the top part, and so on. Secondly, the fact will be explained that in the whole
series of development found, no specimen, however small, is provided with terminal polyp and terminal
zooid; the smallest colonies ma}- possibly simply be tops of somewhat older colonies recentlv divided,
which older colonies had before divided — perhaps more than once. Accordingly, we do not vet know
the real primary polyp of this species, nor do we know the appearance of the species in its full-grown
state. Whether the finished Pennatu /«-shape is ever reached must remain doubtful; I think, however,
that the largest specimen in hand indicates that some individuals attain this form. This would not
be necessary for the sake of the sexual generation, as a great number of the imperfect colonies have
proved to be sexually ripe.

Occurrence. The Davis Straits; St. 24, 63 6 N. Lai. 56 W. Long., (ca. 40 specimens), and
St. 36, 6i c 50' N. Lat. 56 21' W. Long, (upwards of 100 specimens).

The depths are 1199 fathoms and 1435 fathoms; accordingly, the species seems to be a
marked deep-sea form; it has been found together with numerous other deep-sea forms; other
Pennatulids found there, were specimens of Distichoptilum gracile, a series of young stages of Koplio-
belcmnon stelliferum, and a young stage of Umbcllula lindahli.

Fam. Virgularidce Koll., emend.
Virgularia Lam.

Among the peculiar characters of this genus, I shall point out a few which have either not
been noticed or only imperfectly:

1) The polyps are provided with a calyx, the edge of which may be simply circular or
furnished with 8 more or less distinct small teeth without spicules; in the former case (for instance

Virg. viirabilis\ the edge of the calyx is not seen at all, when the polyps are most extended. I need
only mention that the calyx here, as elsewhere, means that the hinder part of the polyp-body is rather
stiff, so that the fore part with the tentacles may be retracted into it; thus the necessary stiffness is
found here in spite of complete want of spicules.

2) The shaft (the rhachis) may be divided into three regions; (a) that of the developed wings,
(b) that of the rudimentary wings, and (c) that of the stalk-zooids. (a) is the longest, upper part of
the shaft; the polyps are fully formed, with tentacles etc., but generally contain no sexual products 1 );
(b) may be rather long, and is on either side provided with close-set wings with undeveloped polyps
which still want tentacles, but contain sexual organs and ripe sexual products; these wings become
smaller below and gradually mere rudiments looking like low transverse ridges, and finally pass into
(c); this last region is provided on either side with a single row of small zooids, which I shall call
stalk-zooids. Close below these the peduncle or stalk begins; the boundary is generally indicated

An exception is found in Virg. a/finis Kor. & Dan. (see Kolliker, Monogr. p. 1981.

PHXXATULIDA. 25

by a slight constriction. Unless the examination is thorough, however, the peduncle seems to begin
where the rudimentary wings cease. The stalk-zooids are never developed to polyps, but keep the
zooid-form *).

3) New polyps are never seen in the act of formation at the ventral edge of the wings (as in
Pennatula and many other genera); this holds good not only of the whole region (a), but also far
down in (b); only in the smallest rudimentary wings are polyps seen to arise as elsewhere in the
Pennatulids; that is to sav in such a way, that the most dorsal polyp in each rudiment is the oldest,
the most ventral one the youngest. One gets the notion indeed, that all the polyps belonging to a
wing are begun contemporaneously 2 ); certainly every wing, as soon as it has the character of a wing,
whether its polyps are full}- formed or not, is provided with the number of polyps that it has
altogether. Nevertheless the polyp forming the dorsal edge of the wing continues always to be
the largest, the one terminating the ventral edge the smallest, even if the difference is often very slight.

4) The calcareous axis has always a broken end above; it generally projects even with a
more or less long naked end from the sarcosoma. This is a well-known and often mentioned fact;
but it has been interpreted as due to injury, whilst I think it is caused by the mode of growth of
the colony (for further particulars see under V. mirabilis).

5) All the species of the genus are provided with the regular radiate canals in the rhachis,
pointed out by Kolliker.

In all these features, and in several more, Halisceptrum agrees with Virgularia; it is quite
certain therefore, that the genus H. is to be referred to the same family as Virgularia; and it may
be doubted whether it ought to be maintained as a separate genus distinct from the latter.

Virgularia mirabilis |0. F. Midler).

Pennatula mirabilis O. F. Midler. Zool. Dan. Prodrom. 1776. 3074.

Zool. Danica. The Danish edition 1781 ( Straa-Sofja;reii »), p. 43; the

Latin edition. 1788, p. n, Tab. XI.
Virgularia mirabilis Lamarck. Anim. s. vertebres. 1816. T. II, p. 430.
Kolliker. Monogr. 1872, p. 190.
Ljungmanni K611. Monogr. p. 196.
multiflora Kner. Verhdl. k.-k. zool.-bot. Ges. Wien. 1858, p. 295.

i| Kolliker has first seen these facts in Halisceptrum and in Virgularia; but in the description of what he calls
lateraler Zooidstreifen , he does not distinguish between the zooids that become partly the wing-polyps, partly the lateral
zooids placed under the wings, and those that continue to be zooids and to be arranged in a long single series; these latter
are the stalk-zooids.

2| In this way the fact has been interpreted by Marshall; but everybody may, by careful examination of the most
rudimentary wings, see that this interpretation is not correct. M., however, was scarcely in possession of any specimen with
this part of the rhachis fully preserved. Otherwise he seems to be the only author, who has especially observed the unvarying
nature of the number of polyps in the wings. See: Report on the Oban Pennatulida, 1SS2, p. 63. This little treatise,
which amongst many well-known things, contains several new interesting facts, is not found in our libraries, so that, when
in 1S9S I communicated my conception of the mode of growth and development in Virg: mirabilis to the Natural History
Society here, I did not know that anyone had before observed the want of increase in the wings of Virgularia. The inter-
pretation of M. of the mode of growth, however, is essentially different from mine (see lateri.

The Ingolf-Expedition. V. I. 4

2 g PENNATUUDA.

Of this species, I have had the opportunity to examine a very large number 1 ), partly from
Danish seas, partly from various Scandinavian regions and from Iceland (the Vestman Islands). I
have been able to examine both well-developed and large specimens, to a length of 530 mra2 ), and also
voung stages down to a size of io mm , i. e. smaller than those hitherto mentioned in the literature.

In larger specimens I find the number of polvps in the developed wings very varying, from 12
to 4; specimens with three polyps in the wings or fewer are young stages which will be more particularly
mentioned later on. Kolliker, it is true, in his diagnosis of the species, states the number of polyps
to be 6 — 9, but in the more particular description, specimens occur with 10, with 11 — 12, and with 3;
O. F. Miiller gives the number as 8, Daly ell 8 — 10. The lateral zooids placed under each wing
occur, in a corresponding manner, in varying numbers; where they are most numerous they are
grouped in two transverse rows, in the young stages in a single row and in a number almost similar
to that of the polyps. The peduncle is always shorter than the pen, but otherwise of very varying
relative length. When Kolliker says: «Feder ungefahr 2 1 ,,—3 mal liiuger als der Stiel , his state-
ment is by no means always correct. The peduncle, here as in many other Pennatulids, may
obviously be stretched very much in the living state of the colony, and in a few preserved specimens
it may also be found much longer; in a specimen 350 mm long from the Kattegat I thus find it to be
250""" long (measured from the real boundary towards the rhachis at the cessation of the stalk-zooids).

Of the numerous young stages examined, most, and especially all the youngest ones, belong
to the museum in Stockholm. They form together a rather close series, elucidating, I ithink, the
essential facts of development and growth in the Virgularia. There is still, however, a regrettable
break between my youngest stage and the single primary polyp, the oozoitc, which we know
hitherto only from Daly ell (I.e. pp.188 — 189). In the month of June, Dalyell succeeded in hatching
some planulae> which swam about very actively (I.e. PI. XLVIII, fig. n); later, in July, he succeeded
in developing other plauuke into the hydra form, i. e. into a solitary, tentacled, lengthy polyp (figs.
12 — 14) without any interior calcareous axis. Apart from the fact that the tentacles developed several
lateral branches, these polyps did not change during the period of more than a month, in which it
was possible to keep them living. As to the size, unfortunately, no more particular information is
given s).

In conformity with what is known of [Renilla and) Pc una tit la phosphor ea, it is to be supposed
that this primary polyp grows in length, gains an interior calcareous axis, and develops on either side
a row of solitary polyp-buds following each other, as to age, from above downwards, so that a small
colony of the simplest penniform type is formed. Such a stage might be called the « Protocaulon ■■-
stage-t). My youngest stages are of a form approaching very near to this supposed one, the only
difference being that they are all wanting in a terminal polyp as the upper termination of the stem.

') Amongst these also, the specimens from our Museum mentioned by Kolliker in his Monograph.

2 ) Kolliker, in his diagnosis, says: bis zu 345 mm lang». Dalyell, (I.e. p. 182) however, had already stated a length
of 23 inches, or about 585mm.

3) Dalyell evidently — although with a cautious reservation — thinks these young polyps to be, not the beginning
of the stem which they really are, but a first wing-polyp. His words are: Indulging conjectures regarding the incidents of
futurity is perilous; however, the survivance of a nascent animal might shew it to be the first of an originating lobe, that
the flesh is carried up in a solid mass beyond the bone, and that in this manner both ends of Virgularia are fleshy».

4) After a genus established by Kolliker in Chall. Rep., but being really only a \ r irgu/aria\ see p. 30.

PENNATUUDA. 27

Together with some of the more developed stages they have been obtained by Loven in 1832, partly
at Kullen (Molle), partly at Bohuslan.

The yotmgest stages measure 10, 10-5, and ir5' mn . One specimen io-5 mm long, is figured on
PI. II, fig. 25, seen from the dorsal side. The upper, polyp-bearing part of the rhachis measures ca. 4 mm ,
the part with the zooids 3 - 5 mm , so that the length of the peduncle proper is 3™'". On either side of
the rhachis there is a row of five distinct polyps decreasing in size and stage of development from
above; the polyps of one side alternate with those of the opposite side; only in the row on the right is
there at the lowermost polyp but one, on the ventral side of the colony, a small rudiment of a winy-
polyp No. 2. Below each polyp one lateral zooid is found. In addition, the lower part of the shaft,
which is devoid of polyps, carries on either side a longitudinal row of four distinct and large stalk-
zooids. The uppermost is placed about i mm below the lowermost polyp-bud, and the zooids of the
two sides are somewhat alternating in position. At the first glance, the upper end of the stem seems
quite intact, and no naked part of the calcareous axis projects; but on a more thorough examination
the calcareous axis is seen to end quite abruptly, as if cut off, in the thin sarcosoma; accordingly, its
upper end does not taper to the fine thread, usually found in Pennatulids. Further, the top of the
stem is not occupied by a developed polyp which might be regarded as the terminal polyp, but there
are at the top two somewhat alternating and quite rudimentary polyps, without tentacles; below
each of them is a lateral zooid, so that they prove to be wing-polyps. Thus the features at the top
are already highlv changed; at all events, the terminal polyp and the upper end of the calcareous
axis have disappeared, and the two wing-polyps that are now the uppermost ones are about to atrophy,
or — what I think less probable — to regenerate. The specimen is excellently preserved, so that the
radiate canals in the rhachis are easily recognised, and there are no traces of generative organs.
The two other specimens agree in essential features with the one described, the only difference being
that all the wing-polyps are solitary. The specimen ny long') is provided with 6 6 solitary polyps,
each with a lateral zooid below, and four stalk-zooids. At the upper end of the axis a rudimentary
polvp is found only on the left side ; but this polyp has to an important degree a distinctly atrophied
appearance, and is very little similar to the young imperfect polyp-buds at the lower end of the shaft.

A number (8 — 9) of small colonies, n — 22 mm long, appear somewhat more developed, in that
some of the wings are provided with two polyps. A specimen is figured on PL II, fig. 26, seen from
the ventral side; it is 22 mm long, with a peduncle of 4\5 mm . Each of the wings of the upper part of
the rhachis consists of only one polyp (on the left side there are 4 such, on the right side only 2),
each of the following wings lower down have two polyps, likewise the distinct rudimentary wings
under these show two polyp-buds with perceptible difference in age. Under each of the wings with
two polyps and rudimentary wings two lateral zooids are found, under the upper solitary polyps only
one such zooid. The number of stalk-zooids is doubled (8 on the right side, 9 on the left), and the
lowermost indistinct rudimentary wings 2 ) almost reach the uppermost of them. To judge by the
size, each of the new stalk-zooids seems to have arisen in the interspace between two of the older

J ) In this specimen two parasites are present in the rhachis; on account of the transparent sarcosoma the lower one
may distinctly be recognized as a Distoma.

2 ) These are not given in the figure, as they are only seen under rather high magnifying power-..

4*

28

PENNATULIDA.

ones. At the top of the shaft a short portion of the calcareous axis projects nakedly, and the upper-
most wing-polyp is somewhat rudimentary. Quite similar features are found in the other specimens;
in some, the ventral polyp of the two-polyp wings is quite small, but a bud, and among the two-
polyp wings there may be a wing with only one polyp, but it, like the wings with two polyps,
is provided with two lateral zooids. At the apex of the stem one or two rudimentary polyps are found,
and above the uppermost there is generally a lateral zooid, which, together with the broken end of
the calcareous axis shows that the original apex must be wanting.

Other specimens again, up to a length of 4o ,nm , show two-polyp wings quite to the apex with
two lateral zooids, and likewise in the rudiment-region, which in the larger specimens is long, and
reaches to the numerous stalk-zooids ; a specimen of a length of 4o mm numbers on either side n — 12
wings with developed polyps.

A somewhat more advanced stage is represented by the specimen 45 mm in length (from Bohuslan)
figured on PL II, fig. 27. At the apex a naked end of the calcareous axis projects; uppermost are two
single, rudimentary polyps; then follow two-polyp wings, of which the two uppermost are rudimentary,
whilst the others (56) are provided with two fully developed polyps; next follow wings with three
polyps the most ventral of which is the smallest, and the farther we get down, the smaller it becomes;
but so long as the polyp-buds of the smallest wings in the rudiment-region may be discerned, the
number continues to be three. Of lateral zooids three are found under a 1 1 the wings. The stalk-
zooids are numerous (upwards of 16), and close-set, especially below. Similar features are seen in
several specimens, partly Swedish, parti} - from the Vestman Islands. Finally, we have a great number
of young colonies, 6o' 1 " 1 ' in length and others (Swedish, Icelandic specimens, one Danish fragment) in
which the developed wings and the rudiments, whose polyp-buds are distinct, contain three individuals;
at the apex, however, (if not damaged) a few two-polyp wings are regularly found, but with rudimentary
polyps. The number of lateral zooids is three or four; in the latter case one of them may be placed
in a second row. In the rudimentary wings where the most ventral polyp-bud is very small, the
corresponding lateral zooid is also indistinct. The rudiment zone is long and joins onto the line of
stalk-zooids; in a specimen (from Kristineberg) 6o mm long, with 15/15 fully developed wings I have
counted ca. 36 stalk-zooids in a closely packed series.

The larger colonies with four, five, and more polyps in the wings, which are generally found
in collections, although most frequently in fragments, connect directly with such stages as the last-
mentioned.

The young stages described accordingly show: 1) that the number of polyps in the wings is
gradually increased by one individual in the rudiment-zone, 2) that by the longitudinal growth of this
zone the new wings advance farther upward, as also the older wings above with fewer polyps, 3) that
contemporaneously with this growth, transformations take place at the apex, as the oldest wings, and
upon the whole the older parts of the colony, here atrophy and finally drop off; for, the fact that
all the well-preserved young stages, even the smallest, are provided at the apex with indisputable
rudimentary polyps, without tentacles and often much diminished, seems to me to be most naturally
interpreted as an atrophy. A new light is thrown on the condition found also in the older colonies
by this interpretation: it is evident that the same processes continue, as long as the Virgidaria is living.

PENNATULIDA. 29

It is well-known that most of the specimens found in collections are fragments; generally the
peduncle is wanting, often also the upper end. This, of course, is often owing to the fact that the long
and thin colonies with the fragile calcareous axis, are easily broken by the fishing gear, and especially
easily at the spot where they rise up from the bottom. But when Kolliker says that he has seen
no single specimen undamaged at the upper end, this statement is most likely due to a very common
misconception. The fact that a naked end of the calcareous axis, which moreover is abruptly
broken, projects from the sarcosoma, is generally taken to be due either to a contraction in spirit of
the soft parts of the colony, or to chance mutilation. That the former cannot hold good as a general
cause is easily seen: in many cases, the sarcosoma beneath the naked part of the axis is not contracted
at all or drawn away from its part of the axis, but wraps this part as firmly as in any other part
of the colony, and moreover specimens just caught and still living show the same feature. If the
condition was due to damage, the first thought occurring to one would be of such as might be caused
by the fishing gear. But the same kind of fishing gear may bring up other forms as long and fragile,
for instance Funiculina, in a quite undamaged condition, sometimes even together with Virgularise;
and again, there are several signs tending to show that Virgularia and allied forms, while they are
still growing undisturbed in their natural element, are already possessed of the naked end of the
axis; it is well-known that other organisms, for instance Serpulse, have been found attached to this
part 1 ); besides, certain Yirgularia; and Stylatuke have been observed in places where it was possible
to wade out at low water, and draw- them up with the hands, or where divers might easily see and
take them-); these forms showed the naked axis-stump. Here the mutilation cannot be due to the
fishing apparatus. Marshall^) has also been of opinion that the mnltilation might be due to the
fact that other sea-animals, especially fishes, had browsed on the tops. To support this supposition,
he refers to the fact that fragments, and especially often tops, of Virgularia have frequently been
found in the stomachs of haddock and cod. From the description Marshall gives of the fragments he
takes to be entire tops, it is clearly seen however that the calcareous axis was broken, and that it
even projected from the sarcosoma with a small naked part (Oban Penn. p. 56). Kolliker has perhaps
also had before him specimens with tops that were perfect:, in the signification in which Marshall
uses the word. In the Museum of Copenhagen however, we have a great number of fragments with
such perfect tops, and also a by no means small number of colonies, also with perfect i.e. normal
apices, from various collections after 1870. When the apex does not show some quite indisputable
mark of mutilation of the sarcosoma and the soft parts upon the whole, I regard it as normal; and
among the characters of a normal apex is an abruptly cut calcareous axis, whether it ends near the
soft parts or projects with a more or less long, naked part. On closer examination such a normal

■) Dalyell 1. c. p. 182; Koren and Danielssen, Fauna litt. Norv. Ill, p. 19. PI. IV, fig. 3d (Virg. affinis). In our
Museum are specimens of Virg. mirabilis from Danish seas (Samso Belt, Great Belt, collected by G. Whither; the Kattegat,
Dr. Joh. Petersen), where the top of the rhachis immediately above the soft parts continues in a long, round prolongation,
formed by closely entangled threads of alga- intermingled with bristles of Annelids, sponge-spicules. vegetable detritus, and particles
of mud; and the lower part of this enwraps the top of the calcareous axis of the Virgularia. One of these specimens is So mlI >
long, the extraneous prolongation 35 mm , a second is 72""", the prolongation 56^'", a third 67'""!, the prolongation 48"""; the
specimens mentioned are young stages with wings with three to four polyps.

-) Darwin, Journal of Researches p. 99 iStylalula Darwinii Kcill.i. Bell iThurston), Proc. Zool. Soc. Loud. 1890,
p. 463 ( Virgularia juncea Pall.).

5) Oban Penn. p. 59; Rep. Penn. -Triton . p. [33.

, PENNATUUDA.

apex will always show that the upper wings are quite rudimentary in a more or less large number; not
only are they smaller than those farther down, although, if their polyps can be counted, they may show
the same number as these; but their polyps are quite degenerated, without tentacles, or with reduced
tentacles. The degree of degeneration increases upwards; where a somewhat large number of reduced
wings occur, the polyps of the lower ones may still be counted, then follow some in which only a
single polyp seems to be preserved, and finally, instead of wings we find but low and narrow trans-
verse ridges just as indistinct as those far down in the rudiment-zone. All these features I interpret
as indicating atrophy. As the atrophied part gradually dies the upper part of the calcareous axis
is quite naturally denuded; and this denuded part decays and is broken off, like the top of the
shell of certain snails with . truncate shells - - or is perhaps thrown off, after the lapse of some
time, as a stag throws off its dead antler.

This process begins very early, as shown by the young stages described (in so small colonies,
that fishes cannot be suspected of having any share in the phenomenon), and it goes on -- normally
- hand in hand with the growth and increase of polyps in the rudiment region 1 ).

This interpretation of the mode of growth in the Virgulariae is certainly not quite new, as I
thought at the time when I gave my first communication upon it 2 ); it has in reality been set forth,
contemporaneously with the first real description of Virg. mirabilis, by O. F. Miiller. In the Danish
edition of Zoologia Danica (1781) it is said of Straa-Sofjasren p. 45: When it has grown to a certain
height, the outermost polyp-hooks by and by die and wither away from above downwards at the outer
end of the stalk, and only the shrunken skin remains some time. Finally this also drops away, and
the end of the stalk is denuded. Accordingly, the polyps perish at one end as they are generated at
the other, which shows the second resemblance to the mode of growth of the tape-worm; yet the
stalk (i.e. the calcareous axis) remains entire and undamaged . Marshall has also had the same idea
(Oban Penn. p. 58), but he immediately rejects it, as he does not think that the facts sufficiently support
it. I think, however, such facts have been advanced here; and especially by pointing out the features
in the small young stages, I think I have placed this interpretation on a better footing at all events
than has hitherto been done.

The young stages described are also of interest in another respect, as proving that two Penna-
tulid-genera (together with their family) will have to disappear from the system. One is the genus
Protocauhn of Kolliker (Rep. Challenger Penn. p. 26), with the species P. n/oltc, from a depth of 700
fathoms near New Zealand. My previous supposition, which was well-founded, that this species was
only a young Virgularia at a similar stage to that of V. mirabilis figured on PI. II, fig. 25, has been
fully corroborated by examination of the original specimen in the British Museum. The upper end
of the colony is not -- as might be thought from the figure I.e. PI. VII, fig. 23 -- provided with a
terminal polyp, but is imperfect after the manner of Virgularia; a small broken end of the cal-
careous axis projects distinctly. The polyps are solitary, in two alternating series as stated by Kolliker;

•) If this explanation was not correct, and the condition of the top was a result of violence, it would be a
strange thing if specimens were never found showing that the colon}- has managed to repair the damage. Daly ell has
proved that Virg. mirabilis is possessed of no slight power of regeneration. On a living specimen D. cut off both ends, and
hung up the middle part on a thread; in three weeks both ends were repaired by regeneration (I.e. p. 1S6).
) In the meeting of Naturhist. Forening on Nov. r i th 1S9S. See Vid. Medd. [898, p. 1.

PENNATULIDA. 31

but they are furnished with a clearly marked, although short calyx with distinct edge which may
even by longitudinal folds seem to be provided with 8 slight teeth ; under each polyp one lateral
zooid is found (seen by K. indeed, but he has not ventured to state it as a certainty); by clearing in
a suitable mixture of oil of cloves and alcohol it is seen quite distinctly, as also the long line of stalk-
zooids characteristic of Yirgularke ; here, these zooids reach almost to the lower end of the colony;
the peduncle proper is damaged. Thus, there is full conformity between this species and my youngest
specimens of V. miradilis; Protocaulon molle can only be a young stage of a Virgularia-sjiecies, and
probablv of Virg, gracillima K611. 1 ).

The other genus that must be rejected is Marshall and Fowler's Deutocaulon founded on
the species D. hystricis (Rep. Perm. Porcupine , p. 461, PI. XXXII, figs. 8, 9). I have not had the
opportunity of examining the type specimens, but the descriptions and the figures make it quite
certain that the five specimens (30— 48™'" long, badly preserved) are a Virgularia in stages corres-
ponding to my young stages on PI. II, figs. 26 and 27, and the ones slightly more advanced. Marshall
and Fowler have seen themselves and very correctly that the family Protocaulidcc of Kolliker, to which
they refer their Deutocaulon^ must be placed nearer to the Virgularia^ than in the system of Kolliker,
where it is far from these latter; they even hint as a possibility that Deutocaulon might prove to be
identical with Virgularia gracillima of Kolliker (from New Zealand), but they think that in this case
a new genus would have to be established for it. This is now superfluous; Deutocaulon is either
simply ]'. iiiirabilis or possibly V. cladiscus mihi. The locality and depth (south west of Ireland and
west of the Hebrides, 109 and no fathoms) might very well speak in favour of the former, but do
not, however, exclude the latter, which was taken at one of the places together with Deutocaulon ,
and is referred to by M. & F. as a Svava glacialis Kor. & Dan .

Among the great number of specimens of Virg. iiiirabilis I have examined, some show certain
differences from the normal which I think worthy of mention.

In a complete specimen from the Kattegat, 530 mm long, with wings with 12 polyps, small
wings with few (3 — 4) polyps are found, inserted here and there among the normal wings; in a few
places, a long solitary polyp is found between two normal wings, and under this a group of three
lateral zooids; in two places the uppermost, most dorsal zooid of a common group of lateral zooids
is transformed into a complete polyp (similar features, however, are known from Pennatula phosphorea\
comp. p. 15 under var. Candida). Further, this specimen is provided with two calcareous axes, of which
however, only one projects from the top with a naked end. In the middle of the rhachis both axes
are very thick; the one retains a considerable thickness, until it suddenly ends, but little rounded, in
the upper part of the peduncle, whilst the other, as is the normal case tapers downward and ends in
a bent hook, about 2 cm below the termination of the stalk-zooids. From the Great Belt and from
Samso Belt, we have further two other small specimens each with two calcareous axes. One is 50 ra:n
long, with wings with five polyps; both the calcareous axes project with naked ends over the top;
here also, only one of these forms the bent hook in the peduncle. The other specimen is a young

M This species has been found again at New Zealand, where it is apparently of frequent occurrence. Deudy:
Trans, and Proc. X. Zeal. Inst. 1S96, vol. 29 (N. S. vol. 121 p. 256. When Verrill (Am. J. Sc. 23, p. 312. notei thinks that Proto-
caulon may prove to be the young of Anlhoptilum , this supposition is no better founded than the one that Protoptiluiu
K611. is the young stage of Virgularia^.).

, 2 PENNATULIDA.

stage, 30""" long, with wings with 3 polyps, a qnite short pen with developed polyps, but a long rudi-
ment-region; here also two naked axial ends project from the sarcosoma at the top, one quite short
broken off, the other rather long (5— 6 mm ) and evidently in process of decay, being plainly divided
into a series of short, loosely connected pieces, like joints ')■

In a complete specimen, 423"™ long (the peduncle 8o mm ), with wings with six polyps, from the
Norwegian coast off Arendal, the wings of the left side are normal, whilst those of the right side,
even quite down in the rudiment-region are placed in an inverted position, so that the calyx-
openings and the tentacles of the polyps are turned towards the peduncle. It is well-known that
the wings have a considerable power of motion, and I have often seen that the}' may also be turned
in such a way as to make the polyps point downwards, but here the question is somewhat different;
in the present specimen, the group of lateral zooids belonging to the inverted wings is placed above
the wing instead of below it, and by serial sections I have convinced myself that both zooids and
polvps here turn the ventral side upward, pointing towards the top of the colony, whilst the
zooids and polyps of the correctly situated wings of the left side are placed normally with the dorsal
side turned upwards. The same abnormality is seen in two fragments of the upper part of the pen
in a specimen with four polyps in the wings, from Bohuslan, Flatholmen, and in another fragment
from the same locality, with three polvps in the wings, in which latter the abnormal side is disting-
uishable as the right one, the end of the stalk being preserved. The last-mentioned specimens belong
to the Museum in Stockholm. An abnormality, probably of a somewhat similar kind, is mentioned by
Kolliker in a specimen of Haliptcris cliristii (Mougr. p. 244); I have found no statement elsewhere of
such features having been observed before.

As to the synonymy of V. mirabilis, I have without any hesitation referred the V. Ljung-
manni of Kolliker to this species. He has established his species on a fragment (in the Museum in
Stockholm) from the Azores, and he himself expresses (Monogr. p. 197) a strong doubt as to its specific
difference from V. mirabilis; after all, I think that its geographical occurrence, so far distant from
what is otherwise known for mirabilis, has been the special reason of its being established as a separate
species. Later, the correctness of this species has hardly been carefully examined; it has, so far as I
know, only been mentioned later by Studer as having been obtained in fragments by <rHirondelle
in the Bay of Gascony, without, however, any further information as to its structure, and by
Whit eaves from the Gulf of St. Lawrence (Catal. Mar. In v. East. Canada, 1901, p. 33). So far as I
can discern from the careful description of this .species by Kolliker, there is no sufficient reason for
retaining it. The same can also be said for the V. mult, [flora Kner of the Adriatic, which Kolliker
himself seems to be most inclined to regard as a local variety. Only the fact that the specimen in
our Museum, in spite of its size, is provided with remarkably small polyps, could raise some doubt in
my mind; the polyps, however, are just as small in a large fragment of V. mirabilis from the Vestmau
Islands (Th. Jonsson); but this specimen has evidently been somewhat dried. No doubt also the
Virgularia sf>. mentioned by P. Fischer, from the west coast of France (Loire-Inferieure, at Croizic),

M A double calcareous axis in Virg. mirabilis has been observed before in a single specimen by Dalyell (1. c. p. 187)
PI. XUII, fig. 8. «.The bones were unequal, a longer and a shorter, the longer extending so regularly, that the shorter might
have been almost considered a fragment accidentally or supernaturally introduced along with it amidst the fleshy substance

PENNATULIDA. 33

is identical with V. mirabilis. It is said to resemble V. multiflora Kner, and to have its, < calicles aussi
profondement separes que cenx des V. mirabilis Mull. (Bull. Soe. Zool. de France, vol. 14, pp. 36, 37);
but a more particular description of it is wanting.

With regard to the distribution, I have to add here a new locality of no small interest,
viz. the Vestman Islands off Iceland. During the summer of 1900, about fifty large and small frag-
ments of specimens with wings with 4 — 9 polyps were taken here, also 14 young stages, most of them
with 3 polyps in the wings (the Diana , A. C. Johansen), and in the summer of 1901 the Museum has
received from Th. Jonsson, physician in the Vestman Islands, a large fragment (145™™ long) with (8 — 9)
polyps in the wings. Hitherto J', mirabilis was only mentioned from the coasts of Scandinavia, Den-
mark and Great Britain. On the west coast of Norway it reaches as far north as Lofoten (Grieg,
1. c. p. n), on the British coasts it does not seem to have been found farther north than Scotland; in
the Danish seas its southern boundary is given as the Great Belt and the Sound down to south
of Hveen (Levinsen, Hauchs Togter, p. 399); for England, in the Channel (Falmouth and Eddystone,
Marshall: Oban Penn. p. 75, W. P. Marshall (sen.): Jouru. Mar. Biol. Ass., vol. III. p. 335). The fact
of this species having now been found at the Vestman Islands not only removes its northern boundary
and considerably extends its western territory, but, when we consider that in recent years we have
been able, just at the Vestman Islands, to point out several Atlantic animals known as belonging to
the fauna, partly of the south-west of Europe, partly of the Azores, the probability that the « V. Ljung-
maiini ■ taken at the Azores and in the Bay of Gascony is identical with V. mirabilis rises almost to
a certainty. If my supposition that V. vntitiflora is also to be referred to mirabilis, be correct, the
Mediterranean will also belong to the territory of the species, the Adriatic at all events where V. multi-
flora has been taken in the Bay of Fiume (at the island of Veglia) 1 ); also at Naples (Nisida) has a
Virgularia been found (Kolliker, Monogr. p. 369), but of its more particular determination nothing is
as yet known; also at southern France, Cap Beam (Lacaze Duthiers, Comptes rend. Acad. Sc.
1891, and Corallaires du Golfe du Lion, p. 359). On the American side of the Atlantic the species
has not hitherto been found, according to existing statements; but according to what has been said
above of V. Lyungmanni K611. it no doubt occurs there also, since 15 specimens of V. <s.Ljungmanni
were taken already in 1872 at a depth of 200 fathoms in the Gulf of St. Lawrence north east of Cap
des Rosiers (Whiteaves I.e. p. 34).

The depths at which V. mirabilis has hitherto been found seem not to exceed 150 — 200 fathoms.
At the Vestman Islands it has been taken at 98 fathoms (the «Diana» St. 41) and at 49 fathoms (the
Diana St. 18).

Virgularia cladiscus noni. nov.

Cladiscus gracilis Kor. & Dan. Fauna litt. Norv. Ill, 1877, S. 101, Tab. IX, Fig. 13 — 15.

Loveni Berg. Mus. Nye Alcyonider etc. 1883, S. 23, Tab. XI, Fig. 1 — 4.

Virgtdaria tuberculata Marshall. Rep. Penn. H. M. S. < Triton 1883, S. 129, Tab. XXXI, Fig. 1 — 3.

Cladiscus Kollikeri Kor. & Dan. Norske Nordhavs Expedition, Pennat, 1884, S. 57, Tab. II, Fig. 8— 13.

Svava glacialis Kor. & Dan. Norske Nordh. Exp. 1884, S. 4, 5, Tab. I, Fig. 1 — 16.

>) From Trieste it is not known; at all events, it is not mentioned by Graeffe in Ubers. der Seethiere des Golfes
v. Triest, III. Coelenteraten. Arb. Zool. Inst. Wien, T. 5, 18S4.

Tlie Ingolf-Expedition. V. I. 5

34

PENNATULIDA.

The calyx of the polyp distinct, with 8 small papillae or wart-shaped calyx-teeth (sometimes,
however, they may be indistinct and seem to be wanting), often with 8 slight longitudinal ridges; the
wings formed of groups of 3 — 4, up to 6 polyps, somewhat coalesced only at the base; more frequently-
one of the outermost polyps is quite, or almost quite, free; below each wing a series sometimes double,
of 3 — 4 lateral zooids. The wings of the two sides are, in the younger part of the rhachis, placed
almost opposite to each other, farther up they are distinctly alternating, with a distance of up to 2 — 3"""
between each set of wings. The dorsal radiate canals on each set of wings grouped more or less
distinctly in a fan-shaped manner. Spicules are quite wanting (as in V. mirabilis). The colour of the
living animal yellowish white.

Of this beautiful Pennatulid the Ingolf has obtained four specimens at three different stations;
a fifth specimen (a fragment) has been taken by the < Diana » at the Vestman Islands. Only in one
specimen (Nr. 4) has the peduncle been preserved complete with its terminal bulb, in the three largest
(Nr.. 1 — 3) the lower part of the peduncle is wanting, but the corresponding end of the calcareous axis
with its fine, bent hook is preserved; as in other Virgularias, a denuded part of the axis projects above
from the sarcosoma.

Nr.

2.

Total length in mm.

The rudiment-region — the stalk-zooids . in mm.

Number of polyps in each developed wing

Length of a polyp in mm.

Number of lateral zooids under each wing

94
25
4

2

3—4

S5
20

3
2—2,5

3

71
20

4

2

3-4

32
10

3
2

3

40

In the upper part of the rhachis irregularities and variations are found with regard to the
wings and their number of polyps, which is probably, at least to some extent, connected with a normal
atrophy of the top part. At the top of the larger specimens, for instance, some of the upper wings
are quite wanting on one side, so that only something like a scan is left; towards the top there
may be wings with two polyps or even with one, or 3- to 4-polyp wings with one or two quite small,
probably atrophied, polyps. As mentioned above, the polyps are provided with a distinct calyx, about
j mm long, with eight small soft calyx-teeth; these teeth, however, may be very indistinct or even
imperceptible in some individuals of the same colony; the forepart of the polyp with extended ten-
tacles is generally of the same length as, or a little longer than, the calyx; whether the fully developed
polyps are able to retract this part completely into the calyx as in V. mirabilis, I must leave undecided;
in the specimens before me all the fully developed polyps, at all events, are stretched out. As in
other Virgulariae the rudiment-region contains the sexual organs, whilst the developed polyps are
sterile. The longest colony I have been able to measure myself, is io5 mm long (a specimen taken
north of Norway by the Barents-Expedition); Grieg (Norges Penn. p. 20) gives a size of even 486™
(a specimen from Fotlandsvaagen north of Bergen).

As is seen from the synonyms this form has been described before, but as constituting a separate
genus, or even two genera, and several species. The reason why I have given it a new specific
name within the genus Virgularia, and not retained its oldest specific name « gracilis •» is that the

PENNATULIDA. 35

name aVirgularia gracilis* has long ago been used by Gabb and Yerrill for another, Californian,
species 1 ). I have therefore used as specific name, Koren and Danielssen's oldest name for the genus
which, I think, is quite descriptive in itself.

Koren and Danielssen originally established the genus Cladiscus with the species CI. gracilis
on a single, fragmentary specimen (Fauna litt. III. p. 102); the genus was referred to the subfamily
Funiculinea of the family of the Virgularise in the (old) system of Kolliker; later, Kolliker (Chall. Rep.
p. 35) placed this genus far from the Yirgularice in his family Protocaulidcr 2 ). This erroneous posi-
tion - - far from the Virgularise - - is, I think, essentially due to the misinterpretation of certain
features in the structure of the colony by Koren and Danielssen, which misinterpretation, among other
things, led them to establish the genus Cladiscus. The fact is that in the original diagnosis of this genus,
only the following essential features are mentioned viz. that the cells are situated separately , and the
zooids ventral . (Lateral zooids are not mentioned). In the description of the single specimen of the species
CI. gracilis upon which the genus has been established, it is said of these zooids: «On the ventral

surface there appears a series of scattered zooids which, in the spaces between the groups of

cells are more agglomerated, become larger, and form strong ventral protuberances of a very peculiar,
as it were, crenulated appearance; the zooids, which are very much elongated, lying in the protuber-
ances nearly in fan-like arrangement (fig. 15 a). In each protuberance there are from 20 to 25 zooids .
Later (Bergens Mus. Nye Alcyonider etc. p. 23) with the help of a new species > CI. Loveni the generic
characters have been supplemented with a remark that the lower end of the axis is knob-shaped
(which is not correct); on the other hand, nothing whatever is said in the altered diagnosis as to
zooids. According to the description, the new species is not provided with the above mentioned
«ventral zooids -, but on the contrary with a single series of 3—4 lateral zooids below each wing. At
the same time, the authors protest against Kolliker's transferring the genus Cladiscus to the family
Protocaulida; in which the calyx of the polyps is said to be wanting'), justly drawing attention to
the fact that the polyps are here provided with distinct «cells ; but when they then place their genus
in the family Protoptilidw K61L, they are still farther in the wrong. Finally, the same authors again
find in a third species Cl.Kollikeri (Norwegian North-Atlantic Expedition p. 57) the ventral zooids ,
arranged upon the whole as in CI. gracilis, but bell-shaped and containing sexual organs (!) (comp. I.e.
PI. II, figs. 8—13); further, two transverse rows (each of three individuals) of lateral zooids below the
wings. From a close examination of the type-specimens of these three C la discus-species in the Bergen
Museum (where I have also examined the later added specimens of CI. Loveni, mentioned by Grieg
(1. c. p. 20)) it was easy for me to see: 1) that the specimen described as CI. gracilis has also lateral
zooids, 2) that in CI. Loveni the same ventral zooids- are found as in the two other «species , and 3) that
the lateral zooids in all three species may occur in a single row, or some may be placed in such a
way that the row seems double. As the number of polyps in the wings, three or four, which number
varies with growth, cannot be used here as a distinguishing specific character there can be no doubt
■1 This species is hardly caracterized in such a way, that one would recognize it from the original description only.
Coinp. KSlliker, Monogr. p. 215.

2) This family (under the section Spicalcr. subsection Jtinciformes) consists in Kolliker of the genera Protocaulon
Koll., and C/adisats Ivor. Dan.; as both the genera must be abandoned, the family must also be dropped. Marshall & Fowler
(Rep. Penn. <Porcupiue p. 4621 had seen already that its place was unnatural.

3) In iProtocau/on>, however, a calyx is found. See above p. 31.

5*

36

PENNATULIDA.

whatever that their three species will have to be united into one. Further from an examination of
the type-specimens of the genus Svava of Danielssen and Koren (North-Atlantic Exp. Penn. p. 45) —
the species Sv. glacialis with the var. alba — I have convinced myself that this genus agrees in every
respect with the ^Cladiscus-species* 1 ); here also « ventral-zooids of the same kind and the same
arrangement as in these species may easily be detected; they may always be seen even under a

magnifying glass — when the specimen is cleared in oil of cloves, or, what is often better, in bergamot.
But if the same method is used with regard to any tolerably well-preserved specimen of Virgiilaria
mirabilis, the very same ventral zooids will also be found there, arranged in the same way, only in
far greater numbers and in several layers. The fact is that these so-called ventral zooids , which in
some specimens of Cladiscus are seen so easily, in others only with difficulty unless in a clearing
medium, are not at all zooids, but on the contrary the eradiate canals > described by Kolliker
in Halisceptrum and Virgularia. There can be no question therefore of maintaining genera like
Cladiscus and Svava; they are one and the same species, and this species must belong to the old
genus Virgiilaria; it is even very closely allied to the species mira bills; indeed, young specimens of
this latter species with the same number of polyps in the wings can with difficulty be distinguished
from Virg. cladiscus mihi; on clearing, the young V. mirabilis also show radiate canals in smaller numbers
and in a single layer; the polyps, however — whose degree of coalesence in V. mirabilis is somewhat
varying, -- are never so slightly coalesced as in V. cladiscus, and distinct calyx-teeth, at all events,
are not found. That the species V. cladiscus may be provided with more than four polyps in the
wings is seen in a couple of specimens from the Dutch Barents-Expedition (1881) now before me,
which have five polyps in the wings (whilst others specimens from the same expedition have four
or three polyps in the wings), as also in a specimen from Finmark, Grolsund, and one from the Skager
Rak (both belonging to the Stockholm Museum): and Grieg (I.e. p. n, under Svava glacialis) men-
tions a specimen from Trondhjem Fjord with up to six polyps. A young stage, with two polyps in
the wings, two lateral zooids, 52 mm long, from the Skager Rak, is found in the Stockholm Museum.
This species may also vary in the way in which the polyps of the separate wings are connected, and
partly also in the degree of their coalescence (comp. Grieg 1. c. p. 19).

That I have also enumerated Virg. tuberculata Marshall among the synonyms will require no
long explanation. From the description in Rep. Penn. •Triton pp. 129 — 133 and from the figures,
every one will easily recognize its identity; and moreover, this specific name has been withdrawn in
the later work by Marshall and Fowler (Rep. Penn. Porcupine', p. 464), on account of its identity
with Cladiscus gracilis Kor. & Dan., and it is added that these authors find «the generic characters as
given by the Norwegian authors too slight to justify a separation of the form from Virgularia^ 2 ) one
must certainly allow that they are right here, although they pay no regard to the false «ventral zooids.:
whose nature was unknown to them. Strange to say, the same authors have retained Svava glacialis,

") The description bv Danielssen and Koren of the genus Svava, agrees so well with the one given earlier of the
genus Cladiscus, that it is astonishing they have not themselves seen the identity. And when it is stated for Svava that the
generative organs are only developed in the lower, imperfect polyps, whilst the upper, perfect ones are sterile, it is also strange
that their attention was not immediately drawn to the genus Virgiilaria, with regard to which Kolliker had long ago pointed
out the same fact.

2 ) Marshall figures no lateral zooids in his V. tubcrcu/ata ; from his remarks on p. 132 it is seen, however, that he
has noticed them, but does not venture to state them as certain.

PENNATULIDA. 37

although they had a specimen before them; but they have referred it to the right place, in the family
of the Virgulariae ').

Distribution. The IngolL has taken the species at the following localities: St. 6: 63 43'
N. Lat, 14 34' W. Long, 90 fathoms, bottom temp. + 7° (Nr. 4); St. 59, 65 N. Lat, n c 16' W. Long.,
310 fathoms, temp. -=-o.i° (Nr. 2), both these stations near Iceland; St. 138, 63 26' N. Lat, 7 56' W. Long.,
471 fathoms, temp, -f- o.6° (Nr. 1, 3), near the Faeroes to the north. The Diana has taken Nr. 5 at
the Vestman Islands at 68 fathoms. To judge from the hitherto known places, the species occurs in
different fjords along the Scandinavian coast from Finmark through the Skager Rak to Bohuslan; in
the deep channel of the Skager Rak it is evidently of frequent occurrence, to judge from the numerous
specimens from this place in the Stockholm Museum (taken by the Gunhild ) ; north of Norway it
has been taken (by the North-Atlantic Expedition and by «Barents ) up to 75 31' N. Lat. (between
Bear Island and Spitzbergen); north of Asia, west of Taimyr by the Vega -Expedition; by the English
expeditions ( Triton >, Porcupine ) it has been taken north of the Hebrides and southwest of Ireland
(51 51' N. L., 51° 51' W. L.), its most southerly occurrence hitherto known, while the Vestman Islands, about
20 W. Long., is the most westerly. It is remarkable that it may occur both in the warm and the cold
area; its proper territory, however, seems to be within the warm area; within the cold area, it has
only been taken at the two stations of the Ingolf^ enumerated above, and at four stations of the
North-Atlantic Expedition (18, 31, 124 and 251); but all these stations are near the boundaries to the
warm area, the Norwegian ones (according to the chart) even at the very border. At all events, it
has not hitherto been taken in the deep and cold polar basin, properly so called.

The bottom is commonly soft, clay or mud, sometimes sand (e. g. St. 6). The depth rather
varying, from 40 — 555 fathoms, most frequently, however, over 100 fathoms.

This species is not known from the American side of the Atlantic. On the other hand, a
Cladiscus -species is described from the Pacific, CI. Agassizii Studer (Note prelim, sur les Alcyon.
<Albatross > Exp.; Bull. Mus. Comp. Zool. Vol. 25, Nr. 5, 1894, p. 58) | under the family Protocaulidir\ s
of considerable size, 28o ram , with three polyps in the wings («Albatross >, St 3424, 20 15' N. Lat 106
23' W. Long., depth 676 fathoms). That Virgularia-species of « Cladiscus -form are also known from
other places in the Pacific, is seen from the Virg. bromlci K611. of the Challenger-Expedition, from
Japan (565 fathoms, temp. + 3°.3 C, provided with spicules), and V. gracillima K611. from New-Zealand
(at small depths, 10 fathoms or thereabout).

Stylatula Verrill.

In Fauna litt Norv. Ill, p. 92, Koren and Danielssen separated the genus Dilbenia from the genus
Stylatula for two Norwegian species, which at the same time they figured and described in detail, later a
third species was added. These three species cannot be maintained; they are only one species, and I think
the justification of the genus Diibcnia to be doubtful. To the best of my belief, Dubenia is related to
the other Stylatula-iorms (with St. gracilis Verr. as type) in quite the same manner as the 'genus
Cladiscus is to the typical Virgularia; here as there, the most conspicuous difference from the nearest

i) Danielssen and Koren had said nothing at all of the more particular relationships of Svava.

38

PENNATULIDA.

allies is that the polyps are almost free and only few in each wing; further, the polyps in Dubenia
are said to want calyces; generally, they are only contracted -- in very different degrees — and not
with the circle of tentacles fully retracted; but that this circle may be completely retracted, is seen from
Kor. S: Dau.'s fig. 3, I.e. PI. X 1 ), and it is also mentioned I.e. p. 95; when the circle of tentacles is re-
tracted the other part of the body of the polyp then forms a <calyx ; I find this feature to be quite
similar in Stylahda gracilis, in which, however, it is the rule that the individuals of the wing are
retracted. Finally, the lateral zooids are arranged in a group above each wing with numerous indi-
viduals, whilst in the other Stylatula-iorms the}' seem to be arranged in a single transverse row.
Altogether, these features seem to be too insignificant to justify a separation from Stylatula as an
independent genus; but, as my material of typical forms of Stylatula has been limited to one specimen
of St. gracilis, I think it better to leave the matter for future determination.

Stylatula (Dubenia) elegans Dan. Kor.

Virgularia elegans Dan. Videnskabsselsk. Forhdl. i Christiania 1859, S. 251.
Stylatula elegans Richiardi. Mouografia etc., 1869, S. 73.

K611. Monogr., 1872, S. 225, Tab. XVI, Fig. 137-38.
Dubenia elegans Dan. Kor. Fauna litt. Norv. Ill, 1877, S. 97, T. Ill, Fig. 1 — 7.

abyssicola => » Ibid. S. 94, Tab. X og XII, Fig. 1 — 3.

borealis » » N. Nordh. Exp. Penn., 1884, S. 97, T. Ill, Fig. 1—7.

abyssicola Marshall. Rep. Triton Exp., 1883, T. XXIII, Fig. 17—21.

From the Vestman-Islauds we have two fragments (17 and 20 mm , probably of the same colony)
of a '> Dubenia with four polyps in the wings, above each of which is a group of zooids consisting
of several rows; the large needles of the calcareous plate are 2.4™'" long, and at the thick end up
to o.240 mm broad; the smaller needles are 0.544— o.768"" n long, up to o.048 mm broad. The large needles
of the calcareous plate seem to grow in thickness at the proximal end in such a way that the original
long spicule is united with several of the smaller neighbouring spicules. The spicules along the aboral
side of the tentacle-stem are of a length of 0.064— o.i6o mm . The body of the polyps is without spicules,
or only provided with a few. The polyps measure up to 3"™ (but the size is here, as is so often the
case, very varying according to the degree of extension).

I have stated above that I cannot acknowledge the three established species as separate.
Neither from the descriptions, the figures, nor the specimens I have had the opportunity of examining,
can I discern any other difference than in the number of polyps in the wings and the number of
lateral zooids; and here where the number of polyps varies within the limits 2 — 6, this difference
means no more than in Virgularia, the mode of growth of the colony being the same as in that
genus (even Koren and Danielssen have given up laying absolute weight on the number of polyps
by making the form <s.smaragdinai> a variety of abyssicola). Only in appearance are the specimens of

') I find the feature to be the same in a great number of polyps in a specimen in our Museum; it is determined as
D. abyssicola by Koren and Danielssen, and has been sent from Bergen Museum.

PENNATULIDA. 39

«elegans» and of abyssicola> rather different; but this is due, partly to the colour, partly to the
different degree of retraction of the polyps and their tentacles, partly also, to the distance between
the consecutive pairs of wings. All such features are individually varying, e. g. compare Rolliker's
figures of St. elegans with those of Koren and Danielssen; both are true to life as I can certify
from the respective original specimens. As to D. borealis , its right as a separate species rests
on one single spicule! That a separate species cannot be maintained on such a character will
be admitted by all who have compared, for instance, a number of specimens of Funiculina qua-
drangular is.

Distribution. This form is known from the coast and fjord-regions of Norway, from Rofoten
and down through the Skager Rak, in the deep channel of which it seems to be common (found by
the German North-Sea Expedition of 1872 and by the Gunhild ); further, it is known from a locality
north west of the Hebrides (west of Rona) ( Triton-); to these localities must now be added the
Yestman Islands, and thus its territory to the west is considerably extended, and it is quite probable
that it will also be found elsewhere. Its bathymetric range is between 30 and 550 fathoms; at the
Yestman Islands it has been found at 68 fathoms together with Virg. mira bill's and V.cladiscus.

On the American side of the Atlantic, and only in the south, are as yet only Styla tula-species
having true wings with many polyps known (further particulars are wanting of the Sfylatula-species
taken by the Albatross » on the eastern coast of North America, off Chesapeake Bay, at a depth of
444 fathoms; Verrill: Am. J. Sc. Vol.29, p. 150, 1885).

Fam. PavonaridcE Jungersen.

Pavonaria K611.

Pavonaria finmarchica (M. Sarsi.

PI. II, Figs. 2S, 29; PI. Ill, Figs. 33— 36.

Virgularia finmarchica M. Sars. Nyt Mag. for Naturvidensk. 1851, S. 139; Fauna litt Norv., II, 1856,

S. 68, Tab. XL
Balticina finmarchica Gray. Catal. Sea-Pens, 1870, S. 13.
Pavonaria » Roll. Monogr., 1872, S. 243, Tab. XVII, Fig. 144.

Gondul mirabilis Ror. & Dan. Berg. Mus. Nye Ale. etc., 1883, S. 19, Tab. X.

This Pennatulid is said to have been taken at Iceland and brought to the Paris Museum by
Gaimard 1839 (Roll. Monogr. p. 243). According to its known distribution, its occurrence at Iceland
was verv probable, though up to the present we have had no other authority for it than the
above. From the results of the Ingolf , however, I am able to confirm the correctness of this
occurrence. At one of the stations south of Iceland (St. 47) the « Ingolf: has taken four specimens
of a Pennatulid which, after a thorough examination, I can state positively to be young stages of a
Pavonaria, and to belong without doubt to the species P. finmarchica.

4°

PENNATULIDA.

These four specimens show the following proportions:

Nr.

Total length in mm.

Length of the peduncle

Greatest thickness of the peduncle
Length of the polyp-calyx

167

140

20

20

2

2

1 — 2

1 — 2

136

37

15

I

The specimens Nr. 1 and 4 are unfortunately rather badly preserved; Nr. 3, which is figured on
PL III, fig. 33, twice the natural size, has been broken just above the peduncle (which has been added
after one of the other specimens); in the three specimens (2, 3, 4) the upper part of the rhachis is
bent with the concavity towards the ventral, polyp-bearing side; in the specimen figured in fig. 33,
moreover, a rather abrupt bending is found, ca. i5 mm from the point, possibly a healed up fracture.
The upper part of the peduncle is dilated in the form of an ellipse; the calcareous axis is also
enlarged here and reaches its greatest thickness; the calcareous axis is round, and ends both below and
above without being bent into a hook. There is no terminal polyp, but the rhachis ends as a naked
process containing the end of the calcareous axis. The polyps are placed along the ventral side of
the rhachis in two rows, each really belonging to one side; two polyps are often placed almost
opposite to each other, making, as it were, a pair; the}' then touch each other at the base, so that
no naked ventral streak is left between them; now and then, especially at the upper end of the
colony, groups of three polyps are found (comp. fig. 34); in this case one of these polyps is always
distinctly younger than the two others, and at its base is partly coalesced with the one belonging
to the same side as itself, and always placed obliquely immediately below; two such polyps form the
first beginning of a wing. In some part of the rhachis, especially in its lower half, large polyps
alternate tolerably regularly with smaller ones; several polyps (not only farthest down towards the
peduncle) have evidently appeared quite recently. The zooids are small and only found on the
ventral side, placed between the polyps, often but one or two above each polyp.

The polyps are provided with a very marked, conical calyx, the ventral edge of which projects
into two strong points; sometimes these points are very long. They are close together on the side turned
away from the stem, but on the side towards the rhachis they are separated bv a broader and much
deeper indentation, from which projects the fore part of the polyp with the circle of tentacles. The
largest calyces measure 2.5 — 3 mm to the end of the points. The calyx is abundantly provided with
spicules, especially dense in the long points; the other part of the polyps shows spicules only along
the aboral side of the tentacle-stem. The longest spicules of the calyx I have measured, are 1.024"""
long and fully 0.048 — 0.064""" broad ; the average size of the long calyx-spicules is 0.8 — 0.96""" in length
and 0.040 — 0.048"™ in breadth; the smallest calyx-spicules are 0.08 — 0.128""" long and 0.007 — 0.016""" broad ;
the tentacle-spicules have been measured to a length of 0.122 — o.i44 mm , and a breadth of ca. 0.0014""".

The whole stem is provided with lengthy parallel spicules, of a length of 0.096""", a few
rougher ones 0.144""" long and ca. 0.032""" broad. The lower end of the peduncle is sparingly provided,
and the spicules here become quite small, oval or round, 0.008 — 0.016""" long; here and there a few
longer ones, up to 0.064'""'. All the spicules — apart from the quite small ones — are of a marked

l'KXXATUIJDA.

41

triangular form being composed of three ridges. As to the colour, it can now (i. e. for spirit specimens)
only be said that the part of the polyps projecting outside the calyx, as also the stomach, is chocolate-
coloured, or deep reddish-brown.

These four specimens show so much correspondence with the Microptihim of Kolliker, estab-
lished ou a specimen of the species M. villemoesii from Japanese seas (Rep. Chall. Penn. p. 26, PL VII,
fig. 27) that the}- may undoubtedly be referred to this genus. This I have done originally; but on
a visit to the Bergen Museum in 1897 I had the opportunity of examining the type-specimens in that
collection of Lygomorpha Sarsii Kor. & Dan., and I very soon recognized this genus and species to be
young stages of Halipteris christii; now these Iugolf-Microptiles show no slight resemblance to
<s.Lygomorpha», as will also be seen from figs. 29 and 31 ou PL II. The arrangement of the polyps is
the same in both, as also the structure of the calyx etc. I therefore referred them to Halipteris, being
of opinion that both Microptilum Koll. and Lygomorpha Kor. and Dan. were to be referred to Halipteris
as young stages of this genus. Later, when I had the opportunity of studying the genus Pavonaria
more thoroughly, I came more and more to the conviction that the Ingolf-Microptiles would have to
be referred to Pavonaria; I wished therefore to test their resemblance to Lygomorpha by a direct com-
parison of the specimens. The Bergen Museum, with the greatest friendliness, met my wish and sent
me the type-specimens of Lygomorpha. With regard to Lygomorpha the result was the same; it is
decidedly a Halipteris, but it is as decidedly different from the Microptiles of the Ingolf . To men-
tion only a single case: the long calyx-spicules of the Litter are on an average twice the length of
those in Lygomorpha, whilst the spicules of this latter agree with those in Halipteris.

That I am right in referring these Ingolf- Peunatulids to the Microptilum of Kolliker will, I
think, be evident, when one compares Kolliker's figure of M. villemoesii with the fragment of the Ingolf-
specimen Nr. 2 shown in fig. 36, and also the descriptions. The most essential difference is that the
genus Microptilum is said to have only one ventral calyx-point. This one point, however, is
really two points fused together. At the British Museum I had the opportunity of examining the
type-specimen of M. villemoesii, and thereby to verify, not only the resemblance to especially the
smallest of the Ingolf-specimens (Nr. 4) already seen in the figure of Kolliker, but also just the
feature mentioned of the double nature of the calyx points. The Challenger-specimen, however, which
is only 65""" long, has a terminal polyp in which the calcareous axis ends; this polyp seems to be
provided with eight calyx-teeth, but in the others the calyx-teeth are arranged into two ventro-lateral
points with an indentation between them on the side turned towards the stem, where the circle
of tentacles of the polyps etc. may be seen; in most, the two teeth are joined close together ventrally
but the arrangement of the spicules distinctly shows the double nature. Except in the four upper-
most polyps a small, younger polyp is placed at each of the others, immediately above the larger
one. The form of the calyx, the arrangement of the polyps etc. leave no doubt that Micr. villemoesii
and our Ingolf-specimens belong to one genus; but it is beforehand little probable that they should
belong to one species; I shall, however, return to this question later.

Verrill has already said that the genus Microptilum of Kolliker is a young stage of Pavonaria;
he has set forth the opinion (Am. Journ. Sc. (3) Vol. 23, p. 311) that most, if not all, the members of the
families ProtocaulidcB and Protoptilidce of Kolliker are young stages of other Pennatulids; and though

The Ingoif-Expedition. V. I. 6

, 2 PENNATUUDA.

his supposition that Protocaulon belongs to Anthoptilum, and Protoptdum to V T irgularia, is quite errone-
ous, he is assuredly right with regard to Microptilum. He has had young forms of Pavonaria fin-
marchica in which he finds «the polyps isolated, thus forming two simple alternate rows, along the
rhaehiss as in Microptilum; and later (Bull. Mus. Comp. Zool., vol. XI, p. 4), he has described a young
specimen of Pavonaria finmarchica of a length of 70 mni , in the following manner: When most devel-
oped, there are two polyps in each oblique row, supported by two-lobed spiculose calicles; between
the wings there are four to six scattered zooids. Toward the upper part of the peduncle there is
but one well-developed polyp, with or without an additional young bud, in the oblique rows; and
here the calicles have the apex bilobed slightly, or not at all, and terminating in a single pointed
group of spicula. This part agrees essentially in structure with the genus Microptilum Kolliker .
The — somewhat imperfect — figure that he gives (1. c. PI. I, fig. 3) of part of a young Pav. fiiimarchica
is presumably of this described specimen. By a comparison of this figure with those on PI. II of the
« Ingolf >'s specimens, there will be scarcely any doubt left as to the identity of the form. That the
question is really of Pavonaria finmarchica - - this species being the only Pavonaria known from the
northern Atlantic --is made further certain for the American specimens, by the fact that Verrill has
evidently had sufficient material for comparison and sufficient intermediate stages. But without such
intermediate stages, I am equally certain with regard to the Ingolf specimens; a direct comparison
even with specimens of two feet or more will show the essential agreement in the features not
dependent on the growth. In the developed form of P. finmarchica the polyps are present in large
numbers, arranged in oblique rows and united in these by coalescing into wings; the single calyces
are completely coalesced almost to the very edge; the oblique transverse ridge thus formed show's,
however, in well-preserved specimens a series of long points supported by spicules on its abaxial sur-
face, apparently one such point for each polyp. A more thorough examination will show that to each
polyp-calyx belong really two long points, but the two adjoining each other in neighbouring calyces are
either coalesced, or one, the inner one, atrophied. Often, however, the oldest polyp of each wing, the
one at the dorsal edge of the wing, shows both its calyx-points; and when, as is frequentlv the case,
even in specimens of two feet or more, a solitary polyp is placed between the wings, the calyx of this
polyp has two ventral or ventro-lateral points. Here, as in the solitary young polyps at the lower border
of the rhachis towards the pedimcle — the edge of the calyx on the side turned towards the stem is
seen to be deeply hollowed; this may partly be seen also in the wings, when the attention is drawn
to it. The polyps have evidently some difficulty in retracting completely; at all events they are
generally somewhat expanded; their fore-part and tentacles (in spirit) are chocolate-coloured or deep
reddish brown. I have measured the largest of the spicules of the calyx to a length of i.28o mm and
a breadth of o.o8o mm ; on an average they are i.024 ,nm l° n & au d o.048 mm broad, (Kolliker, however,
gives their length to 1.8 — 2.i mm , their breadth to 0.06 — o.o8' mn ); these dimensions, as will be seen, agree
very well with those of the Ingolf specimens, and I may add that the form of the spicules is the
same. Both the form and especially the considerable size of the calyx-spicules exclude Halipteris
absolutely; this holds good also with regard to the long calyx-points; and the question could only
be of referring these young stages to the genera Halipteris and Pavonaria; of northern Pennatulids
they are the only ones in which the calyx is provided with two points.

PKNNATULIDA. 43

I have also included amongst the synonyms Gondii! mirabilis Koren & Danielssen, and I am
able to do so with perfect surety, as I have had the opportunity of examining the only existing
specimen of this form, the type-specimen in the Bergen Museum. It is not difficult to see that this
specimen is nothing but a mutilated fragment of Pavonaria finmarchica from which the calcareous
axis has been extracted; and thus originated the free edges of the four «longitudinal valvess. A
similar mutilation of fragments of other Pennatulids, for instance of Virgularia mirabilis, may also
give results which, on superficial examination, have a very striking semblance to being a distinct
organism. This « Gbnduh is either the upper end of a colony or a part near the top casually torn
from the axis, and attached to an Oadiua (Lopkohelia) by the secreted slime ; the basal surface or
« attaching discs mentioned and figured loc. cit, does not exist at all on the fragment, which on the
part in question, shows only the surface of a wound; the ^axial polyp is no terminal polyp at all, but
(by the mutilation perhaps?) an isolated polyp etc. When the long description of Koren and Danielssen
is read with the reservation, that in many things they have let themselves be deceived by preconceived
ideas, it will then be found that such features, as have not been quite disfigured by the mutilation,
agree very well with Pavonaria finmarchica, among other things for instance, the colour; but nobody,
who has not seen the original, would be able to suspect from the description the real state of the
case; now, however, every one who sees the specimen with the information given here, will certainly
recognize it Obviously, all the comments made by these authors and others on this Gondid mirabilis
with regard to its relation to other Pennatulids or Alcyonarise, its being an intermediate form between
Pennatulids and Alcyonids etc., are now quite superfluous. nGbndtih and the systematic group based
upon it will have to disappear and be forgotten as soon as possible!

Distribution. Pavonaria finmarchica has been found in the fjord-regions of Norway, from
Fiumark (Oxfjord) to the neighbourhood of Bergen; it seems also to extend farther south, into the
Skager Rak, to Bohuslan (at Koster, Grieg, I.e. p. n); it has further been found at Iceland (Gaimard),
and south of Iceland, St. 47 of the «Ingolf», 6i° 32' N. Lat, 13 40' W.Long., 950 fathoms; next it has
been found, frequently and in specimens more than one yard long 1 ), on the east coast of North
America, especially on the fishing banks. It has been taken at depths from 60 to 980 fathoms.
According to the localities where it has hitherto been found, its territory seems to be the whole
northern part of the Atlantic, and there is every possible reason to expect that it will be found again
in several places at Iceland (i. e. at the south coast and especially at the Vestman-Islands), and on
the fishing banks round the F^eroe Isles.

From the Atlantic another species has been described: P. africana Studer (Ubers. Anth. Ale.
Gazelle*. Berl. Monatsber. 1878, p. 672), 4 specimens of which were obtained on the west coast of Africa
at io° 12.9' N. Lat., 17 25.5' W. Long, depth 360 fathoms. The possibility is not excluded that we have
here really the same species which has so wide a distribution farther north; the very low wings and
the small number of polyps in these (4, or 5 to 6), specially pointed out by Studer, can scarcely -
the number of polyps at all events cannot — be regarded as reliable specific differences from P. fin-
marchica; at the top, the wings become higher and the polyp-calyx more distinct; the teeth of the
polyp-calyx are not very distinct, but this is often the case in northern specimens also, and here also

1) Iu the rich collection of P. finmarchica of the Christiania Museum is a specimen from Oxfjord, 5 feet long.

6*

j, PENNATULIDA.

the \vin°-s may be quite low. Whether the want of an enlargement of the calcareous axis at the
thickest part of the peduncle, is of any importance I must leave undecided, but I think it doubtful.
The colour, to be sure, is not in exact agreement with the statements of Sars with regard to living
P. finmarchica, but the difference is not so great as to make an identity of the species impossible.

The genus Pavonaria also occurs, but as other species, in the Pacific. In our Museum we
have representatives of two undescribed species from Japanese seas 1 ) and the Gulf of Korea. To
these must be added a young stage from the coast of Korea (depth 80 fathoms); it is 305 ,mn long, but
very slender. In spite of this considerable length it is still nearest to the Microptilum-stage; the
calyx-teeth are long as in the Ingolf specimens, and in many of the polyps also fused together so as
to appear as one tooth. The polyps are arranged in two rows, one belonging to either side, larger
(older) polyps and smaller (younger) alternating somewhat regularly, and where two polyps are
placed opposite to each other their bases touch; one zooid or only a few between each two polyps on
the same side, etc. Most probably it is the same species described at a still younger stage by K61-
liker as Microptilum willemoesii (Rep. Chall. Penn. PI. VII, fig. 27) taken south of Yedo, 34° f N. Lat.
138 E. Long., depth 565 fathoms.

In the Riksmuseum of Stockholm is a Pavonaria (1075, I4 /s 79) taken by the Vega-Expedition
at Bering-Island (depth 65 fathoms); and M or off (I.e. pp.390, 393) describes a Pavonaria dofieini
from Monterey in California, and a P. californica. Finally, we have several older notices of large
Pennatulids from the American side of the Pacific (originally by Gray called ■■■ Osieocella >, but he, to
be sure, established the genus on the naked calcareous axes); they have been described by Stearns
and Moss, and belong doubtless either to Pavonaria or Halipteris; the descriptions, however, are so
imperfect, that it cannot be decided with certainty which of these closely allied genera are in question
(comp. below under Haliptrris).

Halipteris Kolliker.

The genus Halipteris is very closely allied to Pavonaria K611. This may be seen, apart from
the anatomical structure of the colony, its polyps and zooids (with regard to which I may refer to
the monograph of Kolliker), especially in the polyp-calyx, the form of which is the same in
both genera, viz. a truncate, oblique cone whose abaxial side is the longer and whose abaxial edge
is provided with two calyx-points supported by spicules. The most essential difference between the
two genera is only that in Halipteris, the oblique polyp-rows are not coalesced to real wings; the
young stages therefore show so great a resemblance to one another as to be easily confused. To
separate these two genera into two different families, as done by Kolliker, must be regarded as
quite erroneous.

") The remarkable «genus» Echinoplilum described by Hubreckt (Proe. Zool. Soc. Loud. 1885, p. 512, Pis. 30, 31) is, I
think, one of the Japanese species, viz. a multilated fragment, from which, as is the case with Gondii/, the calcareous axis
has been torn out. As I have not, however, had the opportunity of seeing Echinoptilum itself, I can only advance this as a
supposition, and leave it to a more particular test.

PENNATULIDA.

45

Halipteris christii (Kor. & Dan.).

PI. II, Figs. 30, 31, 32.

Virgularia Christii Kor. Dan. Nyt Mag. for Naturvidensk. 5 Bd. 1848, S. 269; Faun. litt. Norv. II. 1856,

S.91, Tab. XII, Fig. 7— 12.
Norticina Christii Gray. Catal. Sea-Pens, 1870, S. 13.
Halipteris Kolliker. Monogr., S. 241, Fig. 146, 147.

Lygomorpha Sarsii Kor. Dan. Faun. litt. Norv. Ill, 1877, S. 99, Tab. IX, Fig. 7— 12.

Protoptiluni tortiim Grieg. Bergens Mus. Aarbog 1886, S. 13, Tab. VII, Fig. 19, 20, Tab. VIII, Fig. 1—3.
Stichoptilum arcticum Grieg. Ibid. S. 15, Tab. VIII, Fig. 4, 5, Tab. IX.

From a fishing bank ca. 20 miles east of FuglS, the Fceroe Isles, the Copenhagen Museum
received through Agent Miiller in 1899 a nue an( ^ complete specimen, io40 ,nm long, of which the
peduncle makes only 8o' r,m , and again in 1902 from the same donor an excellent, smaller specimen
from the Fseroe Isles, 4oo mm long (the peduncle 56""").

Young stages of this species differ so much from the grown form, that they have been
described as independent genera and referred to quite a different family. On some of them Kor en
and Danielssen have formed the genus Lygomorplia, on others Grieg has formed the genus Sticho-
ptiluni, whilst some specimens have been referred by the same author as a new species to the genus
Protoptihuii Koll. ; all these genera; were again gathered into the family Protoptilidce of Kolliker.
That I can now with perfect surety refer them to their right place as Halipteris christii, is due to
the circumstance that I have been able to examine the respective type-specimens, and to compare them
with one another, and with a large and good material of larger and full-grown specimens; upon the
whole I have had before me a tolerably entire series from a length of ca. 73""™ up to about four feet.

The most important aid to recognizing the younger stage as Halipteris christii is the form of
the calyx. The arrangement of the polyps will naturally vary; in the youngest specimens one
must expect a more or less distinct arrangement in two rows, in the somewhat older ones we must
expect to find new polyps on the inner side of the original rows (i. e. somewhat nearer to the ventral
median line).

In large specimens of Halipteris christii the polyp-calyx has a somewhat different appearance,
according as the polyps are stretched out or more or less retracted, and to this is to be added that
the two abaxial calyx-points may vary somewhat in size; generally they are short, one (the inner
one) is often quite indistinct, which may also hold good with regard to the outer one; this
however, will scarcely ever be found in all the polyps of one and the same specimen. When the
polyps are much expanded the mouth of the calyx is distended, and then the two teeth may be widely
separated and are easily overlooked; when, on the other hand, the polyps are quite retracted, the
mouth of the calyx is drawn together; then the calyx is very similar to a grain of wheat, and in
well-preserved specimens the calyx-points are always seen distinctly. They are supported by layers
of spicules continuing down the calyx. The longest of these spicules I have found to be of fairly
constant length - having measured them in several specimens, both large and small; on an average
they are some o.48o mm long, o.o32 mni broad ; the very longest I have found to be o.496 mni long, 0.040™™

4 6

PENNATULIDA.

broad (with these measurements those of Kolliker: 0.4 — 0.7, agree very well); the spicules are often,
as it were, divided longitudinally into, or composed of, several incomplete individuals; this holds good
also with regard to the shorter forms, as also for the tentacles.

Passing now to the hitherto wrongly understood young stages we shall begin with the
smallest specimen. It is only 73""" long (of which the peduncle makes ca. I5 mm ), and belongs to the
Bergen Museum, where it was found, together with two other larger specimens, in a glass labelled :
Protoptiluni tortum Grieg; Vadso, G. O. Sars-. It is figured on PI. II, fig. 30, three times the natural
size; if this figure is compared with fig. 32, which represents the top of a fully formed Halipteris
christii (viz. the above mentioned specimen from the Faeroe Isles of a length of 400""") magnified to
the same degree, the agreement in the form of the calyx will be so apparent that any further account
will be unnecessary; it may be remarked, however, that the calyx-points are upon the whole somewhat
more marked in the very young specimen.

It is evident that Grieg has founded the description of his Protoptihim tortum upon the largest
specimen in the same glass (143""" long), and this is the specimen figured 1. c. PI. VII, figs. 19 — 20,
PI. VIII, figs. 1—3; he does not even mention that he had two more specimens. In «Bidr. til de norske
Alcyonarier» (1886) p. 13 it is said of the calyx: «The edge of the cell is smooth; on the free side,
however, is found a small tooth very richly provided with spicules » J ). To this I shall only add that
in this as in the other specimens I find two teeth, one of which is in most polyp-individuals some-
what larger than the other.

The next smallest specimen I have seen is 86""" long (the peduncle only 7 mm ). It deviates
rom the former by the fact that the part of the rhachis where the polyps are developed, is much
longer, whilst on the other hand the lower part with the rudiments is extraordinarily short; the new,
inner polyps are upon the whole also larger. Together with a fragment i46 mm long 2 ) it was labelled:
iLygoviorplia Sarsii, Kristiausund, 80 — 100 fathoms. G. O. Sars coll.»

Next come two specimens, respectively 106 and i25 mm long, the peduncle respectively 20 and
25"™, labelled: zLygomorpha Sarsiz, Lofoten, Risvser. G. O. Sars coll.» These are the type-specimens of
Koren and Daniels sen of the genus and species stated on the label 3). Here all the polyps are
fairly well expanded, and the calyx consequently is distended; nevertheless, the calyx-points seem to
be comparatively long and are considerably more conspicuous than is commonly the case in adult
specimens with expanded polyps; they seem also to be somewhat richer in spicules, but an examina-
tion of the spicules shows the form and size characteristic of Halipteris. Nor have Koren and
Danielssen quite overlooked the resemblance; in Fauna litt. Ill, p. 100, they say: «The genus L. comes
nearest to the Halipteris, K611iker», but in the same breath they mention a series of features which

') p. 20: the free margin is furnished with an extremely minute papilla».

2 ) In this specimen the lower part of the rhachis together with the peduncle is wanting; in the upper part of tne
rhachis the appearance is already Halipteris-X\\at on account of the increase in the number of polyps, whilst the lower part
resembles more that of the small specimen ; on either side of the rhachis two polyp-individuals are inverted (comp. Virgul.
mirabilis above p. 32)'; here, the abnormal position is especially conspicuous, the long side of the calyx-cone and the two
points of the calyx-edge being turned towards the peduncle instead of towards the top, as in the other individuals.

3) That my measuresments are a little different from those in Fauna litt. Norv. p. loo, is owing to the fact that Koren
and Danielssen have paid no regard to the curves, and as to the peduncle, have not seen its real limit; the quite small
polyp-rudiments at the lower part of the rhachis have presumably been overlooked. In Halipteris the peduncle in reality is
always very short.

PENNATULIDA. 47

in their opinion, justifies the establishement of the new genus, which is placed between Haliptcris and
Funiculina. From a cursory examination of the large specimens of Haliptcris of a meter or more,
the resemblance, to be sure, is not very conspicuous; in most polyps of large specimens the edge
of the calyx seems often but little formed, as for instance is shown in the figure of Kolliker
(Monogr. PI. XVII, 146 — 147). A closer examination, however, of any toleiably well-preserved specimen
will always show a larger or smaller number of polyps, especially among the younger ones, for
instance in the lower end of the rhachis, in which the two calyx-teeth so conspicuous in Lygomorpha
are quite distinctly seen, and even rather long; and when once seen they may be traced in almost all
the polyp-calyxes. In the figures of Koren and Dauielssen of the adult Haliptcris in Fauna litt.
Norveg. 2 d part, PI. XII (figs. 7 — 10) they are also given very distinctly, and in the text it is said that
the calyces are of a «conical form and end upward in two points*. Kolliker says (Monogr. p. 244):
<Miindung ganzrandig mit einem kurzen Zahn und eiuer demselben entsprechenden schwachen Leiste
an der uuteren ventralen Seite, und swacher Andeutung eines ahnlichen Vorsprunges am gegeniiber-
liegenden Rande». Of the two ventral teeth also, the one which the polyp turns toward the naked
dorsal surface of the rhachis is generally the most developed. It is evidently this tooth that Grieg
has remarked upon in his description of ^Protoptilum tortum», whilst he has overlooked the other.
All other features, in which Lygomorpha Sarsii might seem to deviate from Halipteris christii, are only
such as are a consequence of its being a young stage: the arrangement and smaller number of the
polyps, the small size of the colony etc.

Next we come to some (3) specimens of lengths from 234 — 310""" '); they were labelled Sttcii-
optilum arcticums> Grieg. Vadso. 30 — 40 fathoms. G. O. Sars . Grieg has described them in Bergens
Mus. Aarb. 1886, p. 15 and p. 21, and given figures 1. c. PI. VIII and IX; of the calyx he says p. 16:
<The edge is provided with eight small, little prominent spines. The spicules are numerous in the cell,
being especially found in large numbers in one side of the spines, whilst the other side is without
spicules (PI. IX, fig. 6). The spicules never project over the edge, they are mostly placed longitudin-
ally, aud are narrow, pointed, o.2io mm long:. I must suppose that the statement here should be in
the spines of one (the ventral) side» ; and the fig. 6 quoted above shows the two ventral calyx-teeth
which I find in the polyps of all these specimens; as in the grown specimens, one of the calyx-teeth is
even the stronger one in most individuals. What has given Grieg the notion of the other six teeth
is only slight undulations ol the edge of the calyx; they may also often be seen in large Haliptcris,
and I take them to be the last traces of the equipment that must be said to be typical of the Penna-
tulids. Also with regard to the arrangement and number of polyps, these Stichoptilum*^ in accordance
with their greater size, are nearer to the •Haliptcris -character than is tLygomorpha*. For the rest,
Grieg has himself later observed the resemblance with the latter; in Ovs. Norg. Penn. (Berg. Mus.
Aarb. 1891) p. 22 he savs: When I enumerate these two species (viz. Protoptilum tortwm and Stichop-
tilum articum), it is not without some hesitation; they ought perhaps, to be enumerated as varieties
of Lygomorpha Sarsn\ which they resemble very closely*. (The succeeding observations on the varia-

») Between these and the preceeding, the above mentioned type-specimen of Protoptilum torium, 143™™ long, and
the «iji^<?wor^/;a»-fraguient, I36 mm long, from Kristianssund, fit in with regard to development and size.

4 8

PENNATULIDA.

tions of the calyx are partly iucorreet; the calyx, when it is correctly understood, will always give
good characters).

Stages such as the last mentioned are immediately followed by others as for instance, the
specimen 400" 11 " long from the Fseroe Isles. If we are first persuaded that the difference between
iLvgovwrphao and Haliptcris with regard to the teeth of the polyp-calyx, is only a difference in degree
dependent on the growth, and that all other features of the polyps are the same, it will easily be
seen that the other, apparently great, difference in the appearance of the colonies, the number and
grouping of the polyps and the zooids, is due only to age: at the same time that the colony — as is
commonly the case in the Pennatulids — grows in the lower part of the rhachis, at the boundary
towards the peduncle, and adds here new polyps and zooids, a constant increase in the number of the
polyps also takes place in Haliptcris in the older, upper part of the rhachis; as is indicated in the
young stages, new individuals grow up in the interspaces between the older ones. In this way
arise the short and little regular, oblique rows peculiar to Haliptcris. The irregular arrangement
seen in the younger stages is for the rest constant, solitary polyps or irregular little groups being
always found between the more regular oblique rows; and apart from the lower part of the rhachis,
the boundary line between the polyps of the two sides on the ventral side of the colony is, as in the
young stages, only seen with some difficulty; in some places towards the top it seems to be quite
wanting. The increase in the number of the zooids follows that of the polyps; Kolliker has carefully
traced the mode of development of both these kinds of individuals in the lowermost part of the
rhachis (Mongr. p. 245).

Distribution. Hitherto, the species has been known from various Norwegian fjords, from
the Varanger fjord (Hjailmo) to the Jader; also, the young stages were found within the same territory;
away from Norway it is only noted as having been found on the east coast of North England (Northumber-
land, Tynemouth; Alder: Ann. Mag. Nat. Hist. ; (3) vol. IX, p. 316). To these localities I may add the
Dogger Bank in the North Sea J ), and as a new locality also the Fseroe Isles, the fishing bank 20 miles east
of Fuglo (agent Miiller). Hitherto it has only been found at depths to about 200 fathoms. From
the American side of the Atlantic it is not hitherto known with full certainty. Verrill, to be sure,
(Proc. U. S. Nat. Mus. vol.2, 1879, p. 199, and later in Americ. Journ. Sc. and Arts vol. 23, 1882, p. 309)
mentions a Haliptcris (christii) as taken on the east coast of America on a fishing bank (Grand Bank),
but nothing more definite has appeared concerning it later; if the genus be correctly determined, it
is most probable that the species is also H. christii. It has probably a similar distribution to Pavonaria
finmarchica and Pennatula grandis etc. 2 ).

We have various indications that the genus Haliptcris is not restricted to the Atlantic. In

San Francisco Mining and Scientific Press* August 9 th 1873, and Proceed. Calif. Acad, of Sciences

1873, as also in Am. J. Sc. (3) vol.7, P- 68, Stearns refers a Pennatulid (up to 66 inches = ca. i68 cm

long) to the Pavonaria of Cuvier, as subgenus Vcrrillia, the species V. Blakci. In a note added (in

the last named place, p. 70) Verrill declares the form to be most closely allied to Haliptcris christii

(Kor. & Dan.), and accordingly Steams has later (Americ. Naturalist vol. 16, 1882, pp.55 — 56), called the

our specimens in the British Museum labelled: <■ Funiculi na quadrangularis. Dogger Bank. 94.6.24. 1 — 4».
1 It is possible, however, that this Haliptcris has later been tacitly withdrawn and placed b)' Verrill under Pavonaria
finmarchica.

PENXATTJLIDA. 49

form Halipteris blakei. The place of discovery was the Gulf of Georgia, Burrard's Inlet (eomp. also
Proc. Calif. Acad. Sc. Vol. V, 1873, pi. I, pp. 7— 12). From the description given I would not maintain
as a certainty, however, that this species is not a Pavonaria K61L, and at all events nothing more
particular can be said as to its specific difference from Halipteris christii. The description of the
calcareous axis with its lower enlargement agrees with both genera. The specimen from the same
locality, described and figured by Moss (Proc. Zool. Soc. Lond., 1873, p. 730), undeniably reminds one
most of a Pavonaria; but neither the description nor the figure yields any reliable basis for a determin-
ation. Naked calcareous axes of the same forms have been described by Gray (Catal. Sea-Pens,
p. 40; Ann. Mag. Nat. Hist. (4), IX, p. 405, X, pp.77, 406) as nOsteocella septentrionalis from the west
coast of America, and others similar as « Ostcoc. cliftoni , possibly from West-Australia. Thus the
genera Halipteris and Pavonaria are probably rather widely distributed in the Indo-Pacific Ocean.

Fam. FuniciiUnidce Koll., emend.

Funiculina Lam.

Funiculina quadrangularis (Pall).

Pennatula quadrangularis Pallas 1 ). Elenchus Zoophyt. 1766, p. 372.
Pavonaria antennina Cuvier. Regne animal. 1817, Vol. IV, p. 85.
Funiculina tetragona Lamarck. Anim. s. vert. Vol. II, 1816, p. 423.

quadrangularis Koll. Monogr. 1872, p. 256.
Leptoptilum gracile Koll. (Rep. Chall. Penn. 1880, p. 27, PL VII, Fig. 28).

var. norvegicum Kor. & Dan. Bergens Mus. Nye Alcyonider. 1883, p. 29, PI. XIII.
Funiculina arntala Verrill. Am. J. Sc. (3) Vol. 17, 1879, p. 240 and Bull. Mus. Comp. Zool. Vol. XI, 1883,

p. 6, PI. 1, Fig. 4.

South of Iceland the Ingolf» has obtained a young specimen of a length of 225 mm ; the length
of the peduncle cannot be decided with certainty, as the soft parts of the lower part of the rhachis
have been scraped off; otherwise the specimen is very well-preserved; the polyp-calyxes up to 3.5"""
long. In appearance it is strikingly similar to the Trichoptilum brunneum of Kolliker (Rep. Chall.
Penn. PI. VIII, fig. 31), and agrees also with the Leptoptilum gracile var. norvegicum of Koren and
Danielssen 1. c. PI. XIII. The polyp-calyxes are richly provided with spicules, the arrangement of
which quite agrees with fig. 3 of PI. XIII of the last-mentioned work; again, the tentacles are also

i) Bohadsch is stated by Kolliker to be the author who has given the earliest description of the species in: De
quisbusdam Animalibus Marinis, etc. Dresdae 1761. Here it is mentioned, chap. VI, p. 101 under Penna , as 110.3, P-rubescens
1 pinnis carens, tentaculis in corporis trunco positis ) and p. 112 as a species neglected by science, but known to fishermen
who called it Penna del pesce pavone >. His specimen which was incomplete, was of a length of 2 feet 10 inches ( niulto
vero longiorem earn fuisse uon dubito ), and has probably been from the Mediterranean (Naples? from which locality he has
several things'. However, I have found this species mentioned as early as 1655 in Museum Wormianum p. 235: Penuur
marina; nomine ad me transmissum ex Norvegia corpus gracile, oblongum, cartilagiueum, quadrangulum, interior substantia
albicante, quodammodo extremitatem penna; anserinae plumis denudata; referens, exterius membrana sublutea tectum. Tenuis
licet sit hsec penna, crassitie extremitatem pennae anserinae uon superans, longitudine tamen seqvat tres pedes Romauos.
Xullam video habere affinitatem cum Penna marina a Gesnero, Rondeletio, Aldrovando & aliis descripta, ut quo referam vix
habeam; hie interim locum obtineat. donee comodior detur .

The Ingolf-Expedition. V. r. 7

5 PENNATULIDA.

provided with spicules along the aboral side of the stem. The polyps contain well-developed sexual
products (eggs). The polyps are arranged in two alternating rows each corresponding to one edge
of the quadrangular calcareous axis ; between the developed polyps smaller ones are found, and between
these again numerous small ones more or less in the zooid stage; in the very smallest, however, ten-
tacles may be traced as small papillae, and their calyxes are provided with 8 points; true zooids
that remain as such, are not found. In all these features this specimen agrees with the «genus» Lep-
toptilum of Kolliker. At station 18, south west of Iceland, about midway between Greenland and Ice-
land, the Ingolf> has obtained one more specimen, also a Leptoptilum-stage, but only about 70™""
long. Unfortunately, it is little but a naked calcareous axis, the soft parts having been scraped
away except in a few places where some polyps are saved. They show the differences in size
characteristic for these young stages; although they are only badly preserved the determination is
quite certain; moreover, the axis is quadrangular. A very young and slender Leptoptilum stage, 28 mm
long (the top, however, is broken off, and the specimen is upon the whole very badly preserved) has
been taken by the steamer Thor in 1903, near the Vestman Islands (St. 167). It has been shown
by Grieg that Leptoptilum is only a young stage of Funiculina, and further that Leptoptilum gracile
— the Norwegian specimen (var. norvegicum Kor. & Dan.) as well as the pieces of the many specimens
of the Challenger Expedition (from the seas near New Zealand) which have been given to the Bergen
Museum — belongs to the species F. quadrangularis (Berg. Mus. Aarb. 1896, Nr. 9).

The genus Lrptoptilmn must accordingly be rejected, which assuredly, must also be the case
with regard to the genus Trichoptilitm of Kolliker. It has been established on a single specimen —
Tr. brunneum — obtained between Ceram and New Guinea; in all essential features, also, as mentioned
above, in outward appearance and very nearly in size '), it agrees exactly with the first named specimen
of Funiculina of the Ingolf . One difference, however, still remains : Trichoptilum brunneum is
provided with zooids, without spicules, which do not look like future polyps. As, however, the
specimen is somewhat damaged, their different appearance may possibly be due to maceration or the
like, or the question may be of a separate species of Funiculina. But I am quite convinced that the
genus» Trichoptilum is a young form of some Funiculina or other. The same conviction has
already been expressed by Verrill (Am. J. Sc. (3) vol.23, p. 312, note), and he proposes the name of
Funicul. brunnea*) for the species of Kolliker. The fact is that Verrill has observed its similarity to
young stages of his Funic, nrmata.

As to this species of Verrill, which is described in Am. J. Sc. (3) vol. 17, p. 240, and later
ibid, vol.23, p. 312 and Bull.Mus.Comp.Zool.vol.il, p. 6, PL I, fig. 4, it must, so far as I can
see, be regarded as identical with F. quadrangular is, since in the description I can indicate no
feature really separating these two species. The strong spiculation, which has evidently given rise
to the specific name armata, may very well be found in F. quadrangularis, which is very variable

') Kolliker gives 34.3™™ ; but the figure, which is said to be natural size, measures ca. 300 cnnl ; after having seen
the type-specimen in Brit. Museum I am able to decide that the mm. of the text must be a misprint for cm.

2 ) The opinion of Verrill is followed by Roule (Camp, du <Caudan» p. 307); but when he thinks his . Tric/raptilttm sp».
to be either a young Anthoptilum or a young Funiculina, he seems less clear about the matter {Anthoptilum, as is well
known, having no calyx and no spicules, apart from the microscopic small spicules of the peduncle, etc.); the question can
only be of Funicul. quadrangularis, which was before known from the coast off Arcachon.

PENNATUUDA.

5 1

with regard to spiculation; the want of spicules in the tentacles of this species, stated by Kolliker,
is for instance in no way a constant feature, and will rarely, I think, hold good for young colonies.

Distribution. The species has been found on the Norwegian coast, at least from Trondhjem
Fjord to into the Christiania- Fjord; also in the Skager Rak, in the deep channel of which it is evidently
numerous (many specimens, both large and small, have been taken in 1897 by Dr. J oh. Petersen;
comp. Vid. Medd. Nath. For. Kbhvn. 1898, p. 1 ; main- specimens have later been taken at the same
locality by the steamer Thor ); at Bohuslan; the west coast of Scotland, the Hebrides; midway
between the Hebrides and Sudero; the Fseroe Channel, in the warm area (among others by the «Michael
Sars , 1902); the Atlantic, west of Brittany (48 26' N. Lat. 9° 44' W. Long., K611. Monogr. p. 369), and
nearer the coast of Brittany, as also off Arcachon (P.Fischer, Bull. Soc. Zool. Fr., vol.14, P-34); a l so m
the northern part of the Bay of Gascony I Trichoptilum sp. Roule, Camp, du Caudan , p. 307); the
Mediterranean (Golfe du Lion, Naples, Adriatic Sea). To these localities is to be added the American
side of the Atlantic [P. armata Yerr.) from Sable Island to in the Carribbean Sea (Verrill).

The new localities, the stations of the Ingolf Nr. 40, 62 c N. Lat, 21 36' W. Long., and Nr. 18,
61° 44' N. Lat, 30° 29' W. Long., and the station of the Thor Nr. 167, 63° 5' N. Lat, 20 7' W. Long.,
accordingly extend the European distribution of the species in a way that can only serve to bear
out my supposition of the identity of armata with quadrangular is.

In the Atlantic the species is thus known so far, only north of the equator and inside
the territory of the positive bottom temperatures l ). The depths are given as from 10 — 30 fathoms
up to 769 fathoms; the above mentioned stations of the < Ingolf » exceed the greatest depth hitherto
known, one being 845 fathoms, the other 1135 fathoms. According to what is stated above, the
species, outside the Atlantic, may occur at New-Zealand, at a depth of 700 fathoms; but at all events
it is certain that the genus Funiculina belongs also to the Indo-Paeifie Ocean, the young stages of
Leptoptihuii and Trichoptilum having been taken there by the Challenger in two widely separated
localities, both south of the equator.

Fam. Protoptilidce K611.

Protoptilum Koll.
Protoptilum carpenteri Koll.

PI. I, Figs. 2, 3.

Protoptilum Carpenterii Koll. Monogr. 1872, p. 374, PI. XXIV, Figs. 223, 224.
aberrans Koll. Chall. Peun. 1880, p. 28, PI. VIII, Fig. 30.

In one locality, south of Iceland, the Ingolf has taken one complete and five more or

less incomplete specimens of a Protoptilum which is to be referred to the form P. carpenteri Koll.

') On PI. II, figs. 3, 3 a of the Chall. Report on the Alcyonaria (vol. 31, 1S891 Wright and Studer — among the
Strophogorgise, however — have figured a fragment of a « Trichoptilum , which, in their opinion, is perhaps a new species of
this genus . The very incomplete specimen has been taken at Station 143 of the Challenger'-, off the Cape of Good Hope.
The figure shows only three developed polyp-calyxes, and no incomplete polyps or zooids whatever, and just as little the form
of the calcareous axis: neither is anything said of the latter in the explanation. To judge from the figure the question may
be of a Funiculina, but it is impossible to decide whether it is F. quadrangularis.

7*

52

PENNATULIDA.

To the south of Iceland also (St. 164) the steamer < Thor has taken an incomplete specimen. As a rule
there is only one row of polyps on each side of the rhachis. The polyps are arranged, partly
alternating, parti y in pairs almost opposite to each other; the zooids are upon the whole also ar-
ranged in a ventral and a dorsal row on either side; the rhachis is narrow and thin, ca. 1 — 2 mm broad,
provided on the dorsal side only, throughout its whole length, with a narrrow streak bare of
individuals (fig. 3, n); as is generally the case this streak becomes broader down towards the peduncle - ,
on the ventral side also, there is a corresponding naked streak on the lower part of the rhachis, but
it soon disappears, and the opposite polyps of the two sides join, so that the separation is only marked
by the ventral zooids (fig. 2). The polyp-calyx is formed like a horn or (straight) cornucopia
abundantly provided with long, red spicules; the edge of the calyx has very slightly marked teeth
on the abaxial side, none of them distinct as a rule, and in no definite number anywhere; the axial side,
as is commonly the case in the genus Protoptilum, passes into the ccenenchyma; the length of the calyx
is 2 — 3 mm , the breadth 1 — i.i mm . The polyps are only provided with spicules along the aboral side of
the tentacle-stem. The zooids have a calyx of similar form and appearance as that of the polyps, only
much smaller. They are more numerous than the polyps; on the dorsal surface, there is generally 2 — 3
zooids opposite to each polyp on each side of the median streak, and one zooid between every two
polyps on the same side; on the ventral side there are considerably fewer, about half the number.

The colour of the peduncle was yellowish white (in spirit, colourless), the calyxes of the polyps
and the zooids were a splendid bright red on account of the coloured spicules (till now the colour
has remained unaltered in spirit); the whole rhachis gets the same splendid colour due to the
individuals being placed close together. The polyps themselves, like the zooids, were yellowish-white,
apart from the red streak on the tentacle-stem, due to the spicules; towards the point of the tentacle,
however, the red colour becomes faint or disappears, partly because the spicules decrease in size,
partly because they lose the colour themselves. The naked streak on the rhachis is yellowish white.

The calcareous axis is round, ca. o.4 mm in diameter.

In three of the specimens where the whole peduncle is preserved, it ends in a bulb; in a
fourth (Nr. 1 of the table below) there is no indication whatever of any enlargement, a proof that a term-
inal bulb is here, just as little as in other sea-pens, of no importance whatever as a specific character.

The spicules (triangular) are on the polyp-calyxes 0.240 — o.48o mm long, 0.016 — o.032 mm broad; on
the tentacle o.o64 mm long and up to o.oii2 mm broad.

Of the specimens in hand the six, as mentioned before, have all been taken at one locality:
St. 40, south of Iceland, 62° N. Lat. 21 36' W. Long, 845 fathoms, bottom temp. + 3°.3 C; the bottom
dark gray mud with numerous shells of Foraminifera and Pteropods [Limacina). The seventh specimen
has been taken (by the Thor) at 62 10.8' N. Lat. 19° 36' W. Long., r 3/ 7 1903, depth 2150 metres. The
measurements are as follows:

Nr.

I.

2.

3-

4-

5-

6.

128

100

75

"5

50

35

45

45

35

55

4

2,S

2,S

2

2

2

2,2

2

Total length in mm.

Length of peduncle »

Breadth of terminal bulb . »

Greatest breadth of rhachis

190

S7
4

2

PENNATULIDA. 53

Nr. i is entire and shows no terminal bnlb; Nr. 5 and 6 are fragments of therhaehis; in the latter,
the upper end of the rhachis is undamaged; half of Nr. 4 is a naked calcareous axis, and only in the
lower part of the pen are some polyps preserved; Nr. 7 is rather damaged, broken, and the uppermost
32 n,m only a naked calcareous axis. By clearing of Nr. 1 and 6 in oil of cloves, quite whole, undigested
shells of Limacina are observed partly in the oesophagus, and partly in the gastric sac of several
polyps; at the locality in question, Limacina was especially numerous in the plankton. This fact, I
suppose, indicates that these sea-pens, although bottom animals, get part of their food from the
animal life of the surface, individuals from this part sinking to the bottom; in a similar way I have
seen larvae of lamellibranchs forming a prominent part of the food of Virgularia mirabilis from the
Sound. Sexual organs were not seen on clearing; nor did an examination by serial sections show
any such organs. Closer examination, on the other hand, has shown the following interesting
features with regard to the inner structure. From the dorsal longitudinal canal of the rhachis, trans-
verse canals issue at quite regular and somewhat short interspaces towards the periphery, where they
open partly directly into the gastric sac of the dorsal zooids, parti y - in the interspaces between the
zooids — into a system of canals which is, otherwise, again connected with the zooids. These trans-
verse canals evidently correspond to the <radiate canals* of the Virgularia (and also of other Penna-
tulids); here they are characterized by being in the form of a long funnel, narrow at the
dorsal main canal and widening at the peripherial end; their epithelium stains very intensely, so
that they are seen with unusually sharp outlines. In each transverse section only one or two such
dorsal radiate canals are generally seen. Into the ventral principal canal open radiate canals situated
in the middle of the rhachis; they are much larger and wider than the dorsal ones, but run also
towards the periphery; their epithelium also stains very intensely. These wide collecting canals
connect the polyps and the other (ventral) zooids with the ventral main canal of the stem either
directly, or indirectly through a network of canals with less intensely staining epithelium. The
■collecting canals* are much less numerous, but much longer than the first-mentioned dorsal radiate
canals, and their course is a much less horizontal one; they go upwards or downwards, and only
rarely are more than two seen along-side each other. The small number of these collecting
canals, I suppose to be connected with the small number of the polyps; both features — to which may
also be added that no sexual organs were observed — might favour the belief that Prof, carpenteri
is only a young stage.

The features given above of the Ingolf >s specimens of this Protoptilum show the very greatest
correspondence with the Pr.aberrans of Kolliker; there is no doubt that these specimens might justly
be referred to this form (or the unnamed ones which Kolliker in the Chall. Report has added after-
wards as Nr. 2 and Nr. 3, which, however, no doubt belong to the same form). I have had the oppor-
tunity of seeing the originals in the British Museum, and I find no other differences between the specimens
of the Ingolf and the type-specimens of the Challenger- than the fact that the enlargement of the
lower part of the rhachis in which the sexual organs were found, mentioned by Kolliker as character-
istic of Pr. aberrans, is wanting (as in the unnamed Nr. 2), and further that the colour of our speci-
mens is much more intensely red. The first feature: the presence or absence of an enlargement is certainly
quite a casual one, and the presence or absence of distinct sexual organs in this place is certainly

ca PENNATUUDA.

quite special; as to the red colour, great stress is hardly to be put on it, as it probably is very widely
varying in intensity, for example in Pennatula phosphorea where the red colour may quite disappear
(comp. also Protoptiliiiu thomsoni) ; and the unnamed Challenger-specimen Nr. 2 is of the same intense
red colour as our specimens.

If I have, nevertheless, designated these specimens not as P. aberrans but as P. carpenteri, I
have done so because I think these two ^species to be synonymous, and the name carpenteri is
the older one. To be sure Kolliker says of Pr. carpenteri that the polyps are placed in two rows
011 either side of the pen; but it is very difficult to get a clear view of these double rows as
the ventral median line is not distinctly marked, and at all events they are not found through-
out the whole length of the rhachis. The figures given by Kolliker (Mouogr. PI. XXIV, 223, 224)
show an arrangement, essentially agreeing with that of some of the specimens of the Ingolf (comp.
the figured fragment, in which a two-rowed arrangement is quite distinctly beginning); and of
Pr. aberrans Kolliker himself says that in some places the polyps show a tendency towards the ar-
rangement seen in P. carpenteri. The whole difference in the arrangement of the polyps is certainly
only a difference in individual development, and cannot consequently be made the base of a separation
into different species of the forms aberrans and carpenteri, which agree so exactly in all other features.
But if it is a fact that the one-rowed arrangement of the polyps may here pass into a < two-rowed
one, the question arises, if Prof, carpenteri-aberrans is not upon the whole only a young stage of a
Protoptiluni with more longitudinal series of polyps and a correspondingly larger number of zooids.
This, perhaps, will turn out to be the case, and I think I may point out Pr. thomsoni as the further
developed form in question; there is really in the calyx-form, the potyps and zooids, the spicules etc.
of this species a tolerably close agreement with those of Pr. carpenteri, and all the differences are
more particularly such as might be due to differences in age and growth. As there is, however, a
recognizable difference in outward appearance from the least developed stages of the forms I refer to
Pr. thomsoni -- the whole colony of the latter is especially much more robust -- I have thought it
best for the present to keep the species Pr. carpenteri.

Occurrence. Pr. carpenteri has been found before in the Atlantic by the « Porcupine Ex-
pedition, at 48 31' N. Lat, io° 03' W. Long., 690 fathoms (the type-specimen of Kolliker); 48 50' N. Lat.,
11° 9' W. Long., 725 fathoms, 2 specimens (Marsh. & Fowl.); by the «. Challenger Expedition (Pr. aberrans)
south of New York, 37 25' N. Lat, 71 40' W. Long, 1700 fathoms; 31 34' N. Lat, 7 2° 10' W. Long., 1240
fathoms; 40 17' N. Lat, 66° 48' W. Long., 1350 fathoms. In addition, Verrill (Am. J. Sc. (3) vol. 23, p. 312,
note) states that he has found the same species (as aberrans) in shallower waters, and adds that in
some of the larger specimens the polyps are arranged in groups of two or three individuals. (When
Verrill on this account thinks the genus Protoptiluni to be ^evidently the young of Virgularia , he is
quite wrong, which need not be shown more particularly '))•

By the discoveries of the «Ingolf» and the «Thor», the territory of the species has accordingly
been considerably extended towards the north; probably it comprises the whole northern part of the
Atlantic inside the warm area, where the depth, bottom, etc. are suitable.

') Grieg has already and justly objected to this view (Norges Pennatulider pp. 15—16).

PENNATULIDA. 55

Protoptilum thomsoni Koll.

PI. I, Figs. 4— S.

Protoptilum Thomsoni Koll. Monogr. 1872, p. 373, PI. XXIV, Figs. 220 — 222.

lofotcnsc Dan. & Kor. N. North-Atlantic Exp. Penn. 1884, p. 61, PI. II, Figs. 14—20.

» mohni 63, » III, 1 — 7.

» carinatum » >■> 65, » III, > 8 — 11.

armatum 68, > IV, » 1 — 7.

Of a Protoptzlum-sj)ecies which I refer to P. thomsoni of Kolliker, I have had for examination
three considerable fragments taken by the Michael Sars in the summer of 1902 ; these fragments I
have been able to compare with a specimen from the Skager Rak belonging to the Riksmnsenm
in Stockholm, and with the material of the Bergen Museum amongst which were the type-specimens
of the above named four ■■species.* of the North-Atlantic Expedition.

The description of Kolliker is based on four considerable fragments of the rhachis; both the
upper and the lower end with the whole peduncle were wanting. Only the abaxial edge of the polyp-
calyx is free, and this edge is most frequently provided with four small points or teeth (to judge from
the figures 221 — 222, PI. XXIV, these teeth are as often wanting or slightly marked); the axial side
passes into the ccenenchyma, and downwards towards the peduncle the calyxes pass upon the whole
so very gradually into the rhachis, that their length cannot be determined exactly; the projecting
part itself, however, measures hardly more than 1.5 — 2 mm ; the calyx is richly provided with calcareous
spicules 0.36 — i.c>5 mm long and 0.027 — o.o66 mm broad. The zooids have an (incomplete, calyx supported
by calcareous needles which form a small point. The polyps are provided with a row of spicules
on the aboral side of the stem of the tentacles (o.n — o.ic/ 1 "" long, 0.01 — 0.04"™ broad). According to
Kolliker the arrangement of the polyps is difficult to define precisely; it may evidently be described
as two-rowed, i. e. two rows of polyps on either side of the median line of the colony (this line,
however, is only to be determined with difficulty on the ventral side); but K. prefers to describe the
arrangement as consisting of short oblique rows of three polyps on either side, being of opinion
that each of the dorsal polyps forms a group with the two lower polyps placed more ventrally; these
oblique rows would then have to be designated as ascending towards the dorsal side, whilst the
oblique rows (or the individuals of the wings) in the Pennatulids generally ascend towards the ventral
side. The zooids are found in large numbers, and everywhere on the rhachis between the polyp-
calyxes, with the exception of a streak along the dorsal surface of the rhachis. The colour is pink,
varying in intensity in the different specimens.

In the Scandinavian specimens before me the calyx of the polyps is rather varying with
regard to the degree to which it projects from the rhachis ; this holds good with regard to different
specimens as well as to different places in one specimen; but the axial side to the very mouth is
always incorporated into the ccenenchyma. Thus the measurement of the length and breadth of the
calyx may give very different results; it is especially difficult to determine the lower end of the calyx.
The edge of the calyx is also varying, being often quite without teeth, often with one large
and several smaller and indistinct; the most frequent number seems to be three, and these variations

56

PENNATULIDA.

may be seen in the same colon}-, even in neighbouring' polyps; the ealyx-edge of the zooids is just as
indistinctly furnished with projections; most frequently, it seems to me, without such. The tentacles
are provided with a distinct streak of spicules; otherwise the retractile parts of the polyps are naked,
whilst the calyx is furnished with long spicules, placed densely and numerously on the abaxial
side. The size of the spicules agrees upon the whole fairly well with the measurements given bv
Kolliker.

The arrangement of the polyps presents some variation, but the differences are evidently
connected with the different age and development of the colonies; also, the arrangement shown by
Kolliker for the type-specimens of 'the species Pr. thomsoni is represented here. A common feature
in all the specimens is, that along the dorsal median line of the pen runs a distinct, sometimes rather
broad streak devoid of polyps and zooids [n, figs. 4, 5, 6); it is broadest in the lower part of the pen
towards the peduncle; this streak is uncoloured (or yellowish white) like the peduncle. A similar
streak, more or less distinct, also runs along the opposite, ventral surface («' figs. 6, 7, 8); it begins
below at the peduncle where it is almost as broad as the dorsal one, but it very soon becomes
quite narrow, often only seen as a very fine line, in some specimens even interrupted in several
places by polyps placed quite mediauly. Otherwise the whole rhachis is covered with polyps and
zooids; the latter occur in very large numbers and fill all the interspaces between the polyps. On
the lowermost part of the pen (figs. 6, 8) undeveloped polyps together with zooids form a long, wedge-
shaped streak on either side, between the two naked streaks mentioned above.

In the most developed specimens the polyps may be said to be arranged in three or four
longitudinal rows on either side; but if we follow Kolliker, and take into consideration the oblique
rows ascending towards the dorsal surface, each of these contains generally four, five (or in a single place
six) polyps (comp. fig. 7); in reality, the arrangement on either side is nearest quincuncial. Less
developed specimens show two longitudinal rows on either side with two or three polyps in each
oblique rows; the least developed show something between one and two rows, i. e. they approach the
arrangement that may be found in P. carpenteri; in these specimens there seem to be transverse groups of
three polyps: one belongs to the right side, one to the left whilst the middle one is apparently placed
in the ventral median line ( Pr. lofotense ■). In places where the ventral median line is only indicated (it
may be very difficult or in some places quite impossible to see it) the real condition is that a polyp
has been added alternately on the right and the left side inside the one belonging to the original
first row (the dorsal polyps) nearer to, or, as it were, in the ventral median line; oblique rows
cannot be said to be marked here as in the others; or, if they are present, they must be growing
in the opposite direction and alternating in such a way, that opposite to an oblique row of two
polyps on one side there is a solitary polyp on the other side. In the table below, of all the speci-
mens which have hiherto been described and which I think must be referred to the same species,
some of these features have been indicated and some measurements given. The statement of the
total length means here only the length of the specimen in question, as no specimen is complete.
This species probably grows to as greath a length as such forms as Halipteris, Pavonaria or
Funiculina.

PENNATULIDA.

57

Nr.

I.
P. lofo-
tense

2.

-<P

mohiii

3-

4-
Pr. ar-
niatum

5-

Kolliker's

type of

P. thom-

soni

6.
*.P. cari-

iiatum

7-

8.

9-

to.

II.

P.
Gun-
hildce >

Total length ... in mm.

90

150

380

290

145

215

230

140

230

320

505

Length of peduncle

73

157

Breadth

2

5

Length of the spicules of the

calyxes in mm.

0.560
-0.S93

o.445
- 0.625

0.142
-0.S93

0.36

— 1.05

0.222
-1.023

0.992

0.192
—0.992

0.592
—0.960

Breadth

0.020

0.027

0.016

0.027

0.041

0.032

0.032

0.032

-0.035

-0.045

— 0.046

—0.066

-0.055

— 0.064

—0.056

— 0.048

Length of the spicules of the

tentacles in mm.

0.290
-0.3 1 S

0.26S

')

0.1 1
— 0.19

0.136

0.1 18

-0.128

0.192

0.12S
—0.192

Breadth

0.020
—0.027

0.027

')

0.01
—0.04

0.014

0.0080

0.016

0.016
—0.032

Length of the polyp-calyx

3-5

2-4

4-5

3

1-5—2

3—5

3-4

3—4

4(2-5)

2—4

2-4

Breadth

2

1

2

1.5—2

i.S— 2

i.S

Number of calyx-teeth
Number of longitudinal series

3

2—5

3

2-3

(most often) 4

3

3(0—2)

3

3(0-2)

3(o-5)

3(o-4)

of polyps

l(-2)

2

2

2

2

3

3

3

3

3

3(-4)

Number of polyps in each ob-

lique series . . .

I(-2) 2 )

2—3

2 2 )

2-3

3

3(-4)

3

3-4

4

4-5

5

The most complete specimen is Nr. n from the Skager Rak, in which the whole peduncle is
preserved; the upper end, however, is broken off 3). The four specimens Nr. 3, 7, 8, and 10 are of especial
interest because their upper end is intact; this shows that an}' supposition that the genus Protoptilum was
constructed like the Virgularise, is unwarranted ; when the top is so often wanting, and the calcareous
axis appears naked for a greater or less distance, this in Protoptihuu is with tolerable certainty only
due to mutilation, generally caused by the fishing apparatus. In the four mentioned specimens the
rhachis terminates above as a longer or shorter process (fig. 4 t\ bent towards the ventral side in a
more or less hook-like way; this process is covered with zooids to the point; the point itself may be
occupied by a zooid, which, however, does not appear to be larger or upon the whole built differently
from the others.

No stress can be put on the differences with regard to the size of the spicules; in the first
place, I have by no means always measured exactly the largest and the smallest specimens, and
secondly, the measurements of the specimens of the North-Atlantic Expedition and of the type-
specimen of the species P. tlioinsoni, have been made by Dauielssen and by Kolliker, and these
measurements I have not been able to control; besides, spicules are upon the whole often very
varying both in length and thickness in different individuals of the same species of Pennatulids.

As to the colour of polyps and zooids, and consequently of the rhachis as a whole, it is also

') Is said to be «as in the other species*.

2 ) The' oblique series are here most easily seen to be such, when viewed in a direction opposite to that in the other
specimens, viz. from the dorsal polyp upwards towards the ventral median hue.

3) This specimen was in the Riksmuseum in Stockholm labelled as *Pr. gunhildtz , but the name has never been
published (comp. Grieg. Ovs. Norges Penn. p. 21).

The Ingolf-Expedition. V. I. S

58

PENNATUUDA.

rather variable, but is almost always different shades of red; only Nr. 6 «Pr. carinatiuin is stated
by Danielssen to have been straw-coloured, the calyxes almost white. For several specimens the
colour of the calyxes of the polyps is stated to have been of a brick-colour; now, however, they are
almost quite bleached or only pink; exceptions in this respect are formed by the Swedish specimen
Nr. n and one of those taken by the Michael Sars» (Nr. 10), both being still of an intense brick-
red colour.

By a careful comparison of all the Scandinavian specimens I have not been able to find any
difference that might justify a division into several species. The four species established by Danielssen
on the material of the North-Atlantic Expedition: Pr. lofotense, armatum, carinatiiu/, and iiio/u/i, are
each based on only one single specimen; in other words, each specimen found has been made a
separate species, although the localities might have advised some caution; scarcely anybody will be
able to find reliable specific characters in the (otherwise very careful) descriptions of Danielssen, no
more than in his figures, and the specific diagnoses he himself gives, must be rejected by all, who
have had any experience with regard to the Pennatulids. The fact is that the differences really
found in these four specimens are individual ones, either such as are due to different development
and growth (as the number and particular arrangement of polyps and zooids), or such as owe their
origin to a casual degree of retraction (for instance, the features that have evidently given rise to
the specific name carinatum\ further, that the polyp-calyxes may be narrower, more or less projecting
and marked off from the rhachis etc.). That it is injudicious to seek characters for the division
of species in such features is immediately seen, when new specimens are found. In this respect I
may refer to the fact that the two specimens found by Dr. A pp ell of (Nr. 3 and 7 of the table), when
judged in the same way as has been done with those of the North-Atlantic Expedition, would of
necessity have to be established as two new species, neither of them agreeing exactly with any of the
species of Danielssen, but being more or less closely related to each of them and being, besides, provided
with something new. Grieg, to be sure, has seen this (at all events with regard to one specimen,
comp. Norges Penu. p. 21), but he has also seen that it would be dangerous to create new species; the
more so since the Swedish specimen (Nr. 11 of the table), which Grieg has also had the opportunity
of seeing, might as well lay claim to the same title, as it cannot be said absolutely that it seems to
be cpiite similar to the type-specimen of Pr. inohni "■■■ ; only with the specimens of Appellof in mind
could there be from Grieg's standpoint any question of referring it to Pr. mohnh. But when it is
understood that 'Pr. iiioluu may assume so many different shapes, the necessary consequence will be
the incorporation of the three other forms of Danielssen into the same species. I am therefore quite
persuaded that every one, who can now inspect and compare all the Scandinavian specimens I have had
at my disposal, will arrive at the same view with regard to their specific identity. Strange to say. no
comparison with the species Pr. thomsoni seems hitherto to have been attempted; at all events this
species is not mentioned at all by Danielssen or Grieg; nevertheless, the agreement between the
Sandinavian forms and this one is in all essential features so close, that I think myself fully justi-
fied in using the specific name of Kolliker. Also, the features of the inner structure mentioned by
Kolliker (Monogr. p. 371), have been found by me in the Scandinavian specimens I have examined
in this respect. Thus, I find the radiate canals, to which Kolliker especially calls our attention as

PENNATULIDA. 59

being found haufenweise an der Dorsalseite des dorsalen Kanales , developed very beautifully and in
considerable numbers. I shall add that it was easy for me to ascertain that these canals, the connec-
tion of which with the polyps K. could not explain, open into a network of other canals which are
again directly connected with the polyps and with the gastric sacs of the zooids placed between the
polyps; similarly, the dorsal zooids are connected with the dorsal main canal; on the other hand, the
narrow lateral canals do not here, any more than in most other Pennatulids that I have examined
in this respect, show any connection with polyps or zooids.

After to what has been said above, it will now be possible to give the following altered diag-
nosis of the species Pr. thomsoni K611. :

Rhachis long (longer than the peduncle), cylindrical or somewhat compressed; the dorsal
median streak distinct, the ventral one forming a fine line often indistinct or interrupted; the polyp-
calyxes with the whole axial side of the polyps embedded in the ccenenchyma, the abaxial side
provided with slightly marked teeth, most frequently three; all the tentacles with a layer of spicules
along the aboral side of the stem. The polyps of either side arranged in up to four longitudinal
rows; in developed colonies the polyps further form oblique rows rising towards the dorsal sur-
face, and containing up to six polyps. The colour of the calyxes of the polyps and zooids is red in
different shades, sometimes bright red; the retractile part of the polyps, the corresponding part of
the zooids, the dorsal median line on the rhachis, and the peduncle colourless or yellowish white.

Occurrence. This large species is only known from the warm area within the northern part
of the Atlantic, viz. from 36 37' N. Lat, 7 38' W. Long, 322 fathoms (the « Porcupine Expedition, the
four type-specimens of Kolliker); from the fjord-regions of Norway and the adjoining region: the
Vest Fjord {P. <dofotensc , P. mohni , P. carinatum) 160—341 fathoms; west of the mouth of the Sogne
Fjord {P. armahtm ■■), 206 fathoms; the Bergen Fjord (Nr. 3 and 7 of the table, taken by Dr. Appellof),
150— 200 fathoms; mouth of the Sulen Fjord (Nr. 8, 9, 10, taken by the « Michael Sars»), 215— 220 fathoms;
finally from the Skager Rak ( Pr. Gunkildcgi>, Nr. 11, taken by Bovallius and Theel), 440 fathoms.

This species therefore, does not seem to be a marked deep-sea form. It will probably - - like
other European Pennatulids from the same region — prove to be indigenous also on the American
side of the Atlantic, on the slopes of the banks at the Fseroe Isles and Iceland etc.

Protoptilum denticulatum n. sp.

PI. I, Figs. 9— 11.

(Rhachis short; the polyps arranged in one row on either side; dorsal zooids likewise, separated
by a narrow, naked dorsal streak; ventral zooids also in one row on either side, but a ventral median
streak is not distinct). Polyp-calyx with 8 (the terminal polyp) or 6 distinct and long teeth
supported by spicules; the polyps with a strong layer of spicules on the tentacle-stem; the calyx of
the zooids provided with two teeth (the terminal zooid with three teeth on the calyx); the spicule-
formation very abundant; the colony colourless.

In the Atlantic, to the south-east of the southern point of Greenland, the «Ingolf has taken
a small Protoptilum differing so much from the hitherto described species, that it must be established

6q PENNATULIDA.

as a new species. The specimen is quite complete, but it is a young stage and even very little
developed, so that it is impossible to give a complete diagnosis. The information given above
as to the arrangement of polyps and zooids is such as is known to change with the growth, and it
is therefore given in parentheses. The feature by which we are especially justified in supposing
this specimen to represent a separate species is the form of the polyp-calyx. This indeed, is
formed like a horn, as in the preceding Protoptilu w-species, but its upper end projects to a far higher
degree from the cceneuchyma of the rhachis, the axial edge being also distinctly marked and freely
projecting; this edge however has no teeth — at all events in most of the polyps — whereas the other
part of the edge runs out in six long, pointed and strongly spiculed teeth, of which the four middle ones
are as well-developed as the calyx-teeth for instance in Pennatula phosphorca. The terminal polyp
(and, I think, also the two upper lateral polyps nearest to it) is provided with a complete circle of
eight developed, long calyx-teeth. The calyx is abundantly furnished with long, strong spicules; some
of these form six slightly elevated lines continuing into the long calyx-teeth. The polyps are naked
with the exception of the tentacle-stem, the aboral side of which shows a very strong formation
of spicules. All the polyps, also the terminal one, contain large, developed sexual organs; even the
lowest, still cpiite undeveloped polyps contain large sexual organs, so that the whole rhachis (by
clearing in oil of cloves) conveys the impression of being quite filled with these products. The
polyps decrease in size gradually from above downwards; the lowermost ones are still quite undevel-
oped, and form a rather short row of 4 — 5 individuals on either side.

In the zooids, the abaxial side is richly provided with spicules which form a calyx, generally
with two lateral points; the terminal zooid referred to later has a more strongly marked calyx,
provided with three calyx teeth.

The arrangement of the polyps and zooids shows quite distinctly that the colony is in
an undeveloped state. At the top, the rhachis ends in a terminal polyp whose calyx, as mentioned
above, is complete all round and provided with a circle of eight teeth. The other polyps generally
form a single row on either side of the rhachis; in this row each polyp more or less distinctly forms
a pair with that of the opposite side, but in reality the rows are a little alternating. On each side
larger polyps (a) alternate with others a little smaller (b); further, the latter are placed a little more
ventrally than the former. Each pair of opposite polyps consists of a larger, placed more dorsally (a),
and a smaller, more ventral, e. g. of an <?-polyp to the right and a £-polyp to the left; then the pairs*
above and below have a £-polyp belonging to the right side, and an (/-polyp belonging to the left
side and so on. At the upper end of the rhachis an arrangement in oblique rows of two polyps is
indicated, the two uppermost ^-polyps being placed much nearer to the ventral median line than the
others of the same pairs farther down the rhachis. Otherwise, the polyps of the same side join each
other at their calyxes. The dorsal zooids are arranged in a row on either side of a slight
dorsal median streak; at the top the double row terminates in a somewhat larger, unpaired terminal
zooid placed at the base of the calyx of the terminal polyp; as stated above, this zooid is provided
with a more marked and three-pointed calyx. The ventral zooids are in a similar way arranged in two
rows, but a ventral median streak is not distinctly seen, excepting at the lower end of the rhachis.

The spieulation is highly developed; spicules are only wanting on the terminal bulb of the

PENNATULIDA. 6l

peduncle and the retractile part of the body of the polyps, as well as on the pinnulse. The longest
and strongest are found on the abaxial side of the calyxes of the polyps, the shortest and
thickest (of almost oval contour) on the peduncle; on the lower part of this these spicules are
arranged, partly somewhat irregularly, partly with the longer axis placed transversely: farther up
they form longitudinal layers with finer and longer forms interspersed; still farther up elongated
spicules longitudinally arranged predominate. The long spicules are of the common triangular form,
sometimes the edges curve irregularly in and out. As I was not willing to sacrifice any part of this
one specimen for a more thorough examination of the spicules, only such have been measured as
could be seen distinctly in situ, or dropped off during the manipulation of the specimen. The longest
and biggest may be given as ca. o.96o mm long, and a greatest breadth of 0.064™"'; others (measured
when isolated) are 0.464— o.8o4 m,n long and o.o32 ram broad; the smallest tentacle-spicules measured are
o.400' nm long and o.0240 mi " broad; the short and thick spicules of the peduncle are generally 0.096
— o.i44 ,nm long, o — 032 mm broad.

The calcareous axis is round, ending below in a bent hook far from the lower end of the
peduncle (at * in the figure); this feature, however, is due to the accidental, much stretched condition
of this part of the body of the colony, and so it cannot here, any more than elsewhere, be regarded
as being of any value as a specific character. I have not been able to find the upper end of the
calcareous axis, but I have followed it to about the height of the terminal zooid.

The whole colony is white, all the spicules colourless.

Occurrence. St. 20, 58 20' N. Lat, 40 48' W. Long., 1695 fathoms. Bottom temp. 1 '.5 C,
the bottom light gray mud, 1 specimen.

The specimen in hand shows the following measurements:

Total length 75 mm -

Length of peduncle 4°

Greatest thickness of peduncle (close below the rhachis) 1.8

Length of terminal bulb 3

Breadth *-5 8

Breadth of rhachis measured across the mouths of two opposite polyp-calyxes... 3

Length of a polyp calyx, from the base to the point of a tooth 4—4-5

» notch between two teeth 3

The calyx-teeth 1 ~ 1 -5 *

Diameter of the mouth of the calyx i-3 »

Since, according to the description by Kolliker of Protoptiluiu smittii, there might be reason
to suppose that the specimen of the Ingolf was perhaps the same species, I have, by the kindness
of Professor The el, examined the type specimen of Pr. smittii. This specimen is also at a little
developed stage, mainly with one row of polyps on either side; but the top is broken off (it is upon
the whole rather damaged); some agreement is present in the outer appearance, but the teeth of
the polyp-calyxes are broader and vary more irregularly in size and number (3—5), the zooids are

g 2 PENNATUUDA.

much more numerous, the retractile parts of the polyps very dark-coloured, the spicules more slender
and fewer, especially on the peduncle. Although several of these differences are of such a nature,
that in some other Penuatulid-genera they would not be regarded as specific differences, I dare not
overlook them with regard to the genus Protoptilitw, our knowledge of the variation and the change
in growth of this genus being still very incomplete. Accordingly I take - - for the present - Pr.
smittii to be a young stage of another species than my Pr. dcnticulatiim. They belong both of them
to the Atlantic Ocean, and the great distance between the localities, respectively ca. 36 N. Lat. 14° W.
Long, (the Josephine Bank), and ca. 58 N. Lat. 40 W. Long., need not in itself -- any more than the
difference in depth (228 — 1695 fathoms) -- prove any specific difference, as may be seen from other
cases (e. g. Anthoptilum gra/idi flora m a. o.).

Distichoptilum Verrill.

The genus has been established by Verrill on one specimen of the species mentioned below,
and characterized in the following way: «Slender pennatulids, with an axis through the whole length.
Polyps arranged alternately, in a simple row, on each side. Calicles bilobed, appressed. Zooids three
to each polyp, one in front and one on each side of each cell. Spicula abundant in the calicles,
rachis, and stalk; those in the stalk are small, oblong, triquetral, interwoven*. In this diagnosis,
however, the following alterations have to be made: the calyxes of the lateral potyps are provided
with several (6) teeth on the abaxial edge; the axial edge is toothless, and the axial side of the calyx
fuses with the rhachis. The number of zooids is two at each polyp; they are placed directly above
the mouth of the calyx of the polyp, one on the dorsal side of the rhachis, and one on its ventral
side. The peduncle increases evenly in thickness downwards. This genus is especially interesting
in that it retains as a constant feature, an arrangement of the polyps which in other Pennatulids is
only found as a transitory one in young stages, and which may really be more or less distinctly
traced in all the Pennatulids of which we know sufficiently young stages; it is seen most distinctly
in the &Protocatilony>-sta.ge of the Virgularise and in Protoptilum. This simple arrangement in Disti-
choptilum is not only a constant feature, but persists throughout with the utmost regularity, which
gives to the whole colony an exceedingly emphatic character of simplicity.

Distichoptilum gracile Verr.

(PI. I, Figs. 12, 13, 14).

Distichoptilum gracile Verrill. Am. Journ. of Sc. 24, 1882, p. 362, Note.

Bull. Mus. Comp. Zool. Harvard Coll. Vol. XI, Nr. 1, 1883, p. 8, PI. I, Fig. 1.
Verrillii Studer. Ibid. Vol. 25, Nr. 5, 1894, p. 59.

At two stations, very far from each other, one in the Davis Straits, the other south of
Iceland, the Ingolf has obtained several specimens of a Distichoptilum which, in spite of some

PENNATULIDA. 63

differences, will have to be referred, I think, to the species D. gracilc of Verrill. The features in which
all the specimens differ from this species are the following: the polyp-calyxes are only here and there
seen with a two-lobed edge; and when this is the case it has only arisen by a closing of the mouth
of the calyx round the completely retracted polyp 1 ); in other cases — where the contraction of the
retractile parts is not so complete — the abaxial edge of the mouth of the polyp-calyx shows six
pointed teeth supported by spicules. Next, the polyps are not placed so close to each other, as given
in the description of Verrill and in the figure I.e. 1 a; generally, the front edge of a polyp-calyx does
not reach farther upward than to the lower end of the calyx placed next above it, and consequently
it covers no part of this next calyx; often, especially in the upper part of the rhachis, one polyp is
far from reaching to the base of the next. With regard to this point, however, it is to be remem-
bered that the sarcosoma of the rhachis in all Pennatulids may be shortened by a contraction of its
longitudinal musculature, so that the distance between the polyps ma}- be altered; as in Virgularia
1 in ira bilis), Sly lain la, etc., the appearance of Distichoptiliini may also be influenced by this fact.
Further, there are found at each polyp, not three but only two zooids (each with a small calyx provided
with spicules); they are placed close to the (axial) upper edge of the calyx, one on the dorsal, one on
the ventral side of the rhachis, and (very nearly) of the same relative height. When Verrill states
that one more zooid is placed in front of» i. e. above the calyx, such a zooid is absolutely wanting
in all the specimens of the Ingolf >; in this respect as in all others these specimens agree with
Studer's D. vcrrillii; in V err ill's figure also, this third zooid is not seen, and I think its existence
quite problematical. Finally, a more or less distinct longitudinal stripe or furrow is found both on
the dorsal and on the ventral side of the rhachis; the appearance of this furrow may however, I think,
be quite dependent on the degree to which the longitudinal canals are filled, and to the casual contrac-
tion of the sarcosoma. The colour is stated by Verrill to be bright orange red, due to the spicula;
end of bulb yellowish ; in the specimens of the Ingolf the colour is only red in the upper, abaxial
part of the polyp-calyxes, otherwise it is yellowish to yellowish- white; the red colour of the calyxes
extends over two only of the lateral points, whilst the other teeth contain colourless spicules (a further
reason for overlooking these teeth). In all the features stated, the D. verrillii of Studer agrees with
the Ingolf-specimens, except that the calyx is said by Studer also to be provided with two teeth; he
declares himself that his species is very closely allied to that of Verrill - - the only differences
being the greater distance between the polyps, and the greater size of his specimens -- but these
features are of no value as specific characters. So far as I can see, both Studer's specimens and ours
will have to be referred to the species D. gracilc first described.

The calcareous axis is round, in some specimens a little angular or compressed in part of
the rhachis.

In the upper retractile part of the polyps, spicules seem to be wanting, but they are found in
the tentacles on the aboral side of the stem, not, however, to the very point; pinnulae are naked.
The specimens taken by the Ingolf show the following proportions:

' 1 Verrill's fig. 1 a, which, by the way, is not very detailed, shows only quite retracted polyps.

6 4

PKNNATULIDA.

Nr.

4-

6.

7-

S.

9-

10.

9 S

S3

75

70

47

47

40

33

25

22

I

1

0.8-1

1

1

i.S

1.8

i.S

i.S

2-3

2

2-5

2-5

2

o.S

1

0.8

Total length in mm.

Length of peduncle

Greatest breadth of peduncle

Breadth of rhachis measured across a polyp-
calyx in mm.

Length of polyp-calyx

Breadth across the mouth

289
74
1-3

i.5

3

1

227
70
i-3

i-7
3

1

106

32

1— 1-3
3

i

104

25
i-3

101

35

T

i-3

3

1

36
15

1

1-5

2-5
o.S

As a matter of comparison I ma) - note that Verrill's specimen is 456"™ long, the breadth in
the middle 2 mm , the length of the peduncle ioo ,nm , whilst Stnder says that his species probably
reaches to a length of upwards of a metre; the single entire specimen, however, measured only 47o m,n .

Of the specimens before me Nr. 6, 7, and 10 are damaged, more or less of the upper part of
the calcareous axis having been denuded; the others look as if they were entire, but a more thorough
examination of the upper part of the rhachis makes it doubtful whether the original upper end of
the colony is preserved. With exception of the specimen Nr. 9, which I take to be abnormal, and
which will be more particularly mentioned hereafter, the rhachis ends in a solitary polyp which,
moreover, on its axial (dorsal) side carries a solitary and rather conspicuous large zooid (see PI. I,
fig. 13 z). It would be natural therefore to suppose these to be the terminal (i. e. the original) polyp
and a terminal zooid, such as we know to occur in young Pcnriafti /a-co\onies. But so far as I
can see, only six calyx-teeth are found in this apparently terminal polyp, as in the other polyps,
the lateral ones; now we might, with some probability, expect eight calyx-teeth and a fully developed
calyx-edge in the real terminal polyp, but the calyx-edge does not appear to be developed on the
side towards the apparently terminal zooid which, however, is difficult to decide with perfect

certainty, as the polyp in question is always strongly contracted, so that the calyx-teeth join each
other. At all events, the terminal zooid is placed close to the calyx-teeth of the end-polyp, whilst
in Pennatula, as is well known, it is placed below at the base of the calyx of the terminal polyp.
In other words, it is placed exactly where the one zooid would be found, if the apparent end-polyp
is a lateral polyp. I am most inclined to suppose therefore, that the «terminal polyp» is the upper-
most lateral polyp present, and the terminal zooid one of the two zooids of this polyp together
with an adjoining small part of the rhachis, so that the real upper termination of the colony is close
to this zooid. But such a termination of the rhachis can scarcely be the original one; it is probable
that in all these specimens an upper part of the colony has dropped off either as a consequence
of earlier mutilation or perhaps through atrophy in a somewhat similar way as in the Yirgularise.
That at all events, the original features are not found at the upper end of the longest specimen but
one (Nr. 2), although this end looks as in the others, may be concluded with certainty from the fact
that the calcareous axis ends immediately at the terminal zooid , cut off quite abruptly, and is there
of the same thickness as farther down. With regard to the others I have not been able -- even by
clearing (in oil of cloves and the like) — to get a clear notion of the upper end of the calcareous axis,
but it seems to be tapering, and does not appear to be broken. On the other hand, the lower end may
easily be observed; it is found in the outermost point of the peduncle, and forms here a curved hook.

PKNNATULIDA.

The specimen Nr. 9 deviates from all the others in having the lateral polyps in the uppermost
part of the rhachis arranged in (almost) opposite pairs; three such pairs are present (PI. I, fig. 14);
between the two polyps of the uppermost pair the rhachis ends as a little knob (k) containing the upper,
fine end of the calcareous axis, and provided with one ventral zooid at all events for one of the
two polyps.

The spicules on the calyx have a length of 0.528 — o.8oo mm and a breadth of 0.032 — o.048 mm ; on
the tentacles, the spicules are 0.096 — o.i76 mm long and o.oo8' mn broad.

Occurrence. The specimens of the Ingolf have been taken at the following places: St. 24,
63 06' N. Lat, 56° W.Long., 1199 fathoms, bottom-temp, -j- 2° 4 C. (10 specimens, Nr. 1 — 7, 9 — n, and
some fragments of the rhachis of other specimens); St. 41, 61 r 39' N. Lat., 17' 10' W. Long., 1245 fathoms,
bottom-temp. + 2" C. (1 specimen, Nr. 8).

The specimen of Verrill was obtained south-west of Nantucket Island, 39 59' 45" N. Lat, 6S
54' W. Long., ca. 700 fathoms; those of Studer, on the other hand, are from the Pacific, o° 4' S. Lat.,
90° 24' 30" W. Long., 885 fathoms, 23 59' N. Lat., 108 40' W. Long., 995 fathoms, i° 7' N. Lat,, 80 21'
W. Long., 1573 fathoms. With regard to the Atlantic I imagine that the distribution will be found to
comprise the whole of the warm area at great depths and on soft bottom.

Fam. Anthoptilidce Koll.
Anthoptihun Koll.

To the characteristics of the genus pointed out by Kolliker may be added, that the stem of the
colony is very much bent so as to form a large curve almost in the middle, and further that it may show
smaller curves, so that its form may be somewhat like an S; the concave side of the chief curve is
formed by the naked dorsal surface of the colon}-. All the figures of Kolliker show this curve, but
as it is not mentioned, it might be supposed to be individual, or even to have appeared in the pre-
served state. The many specimens that I have had the opportunity of seeing, taken directly from
the bottom of the sea, show with certainty that the curved form is a natural feature of the colony,
being already distinctly marked in small specimens. The stem ends above in a naked, longer or shorter
cone without any terminal individual. The sexual organs are found in the polyps themselves.

The calcareous axis, as stated by Kolliker, is round, in so far as it has no edges; but it is
distinctly compressed from the sides; it is thickest at the beginning of the chief curve or a little below
this, but a more conspicuous swelling of the calcareous axis, corresponding to the elliptical thickening
of the upper part of the peduncle which is a peculiarity of the genus, is not found (as in Pavonnrin
jinmarcliica and Halipteris chrisfii).

The genus is hitherto known only from the Atlantic, where it occurs from the most northern
to the most southern regions, but only at great depths.

The Ingolf-Expedition. V. t. n

N,

PENNATULIDA.

Anthoptilum grandiflorum (Verr.).

Virgularia grandiflora Verrill. Am. J. Sc. (3) Vol. 17, 1879, p. 239.
Anthoptilum thomsoni KSlliker. Challenger Rep. Penn. 1880, p. 13, PI. V, Figs. 16— 18.

grandiflontm Verrill. Am. J. Sc. (3), Vol. 23, 1882, pp. 312, 315; Bull. Mus. Comp. Zool.
Vol. XI, p. 5, PL I, Fig. 6.

The Ingolf has obtained six specimens at one station in the Davis Straits; three of these
are well-preserved, two rather large, the third small; two are more or less damaged, and one specimen
is quite denuded (Nr. 6); two specimens (Nr. 7, 8) have been taken at another station, both of them
in bad condition.

Nr.

Total length in mm.

Length of peduncle »

Diameter of the upper enlargement of the peduncle . »
Breadth of the naked dorsal surface of the rhachis »

Number of polyps in the wings

Length of an extended polyp with the tentacles in mm.

415

52

8

c.4

5-7
16

300

41
5-5
4
5-7

205

32

3

3

3—5

172

122

23

2

1 — 2

2-3
5

1 So 1)

180

130

31

20

4

3

c. 2

2

3~5

2-3

5

5

As is generally the case in the species, spicules are only found in the end of the peduncle;
in one of the specimens the largest measure to 0.024 — o.030 mm , whilst others show a size of ca. 0.012
— 0.020 mm ; the smallest seem to be even smaller than the minimum size of 0.007™™ given by Kolliker;
a regular combination of four spicides is rather common, but more irregular combinations of three or
four spicules are also found.

The number of polyps increases during growth, not only in the lower, boundary part of
the rhachis, but also farther up in the older part, where the oblique rows here representing
the wings are often seen on the ventral side of the colony to be increased by a young polyp
or a bud.

The colour of the polyps when living is violet, that of the stem reddish; in spirit this sea-pen
loses its colour only the tentacles of the polyps remain coloured, and there it is turned brown. In several
of the specimens from both stations peculiar, worm-like parasites were found, partly projecting from
the mouth of the polyps; they were parasitic Copepoda of the genus Lamippe Bruz. ; evidently a
distinct new species, to which I give the name of L. anthoptili.

Anthoptilum grandiflorum has hitherto only been mentioned from the American side of the

Atlantic; from North America it has been known from off Guadaloupe to off Nova Scotia (42° 46' N. Lat.

63 45' W. Long.) (Verrill, Agassiz); from South America from 37 17' S. Lat, 53° 12' W. Long., south of

Buenos Ayres (the Challenger •). To these localities two new ones are to be added, showing that the

species ranges over a far wider geographical territory, as well to the north as to the east, in the

Atlantic. The specimens of the Ingolf ■> have been obtained in the Davis Straits, partly south-west

of Godthaab, at St. 25, 63 30' N. Lat., 54 25' W. Long., 582 fathoms (soft mud, somewhat like potter's

M The greatest breadth of the naked calcareous axis on the compressed lateral surface 2™'", on the narrow sur-
face- 1.5mm.

PENNATULIDA.

6 7

clay), partly west of Sukkertoppen, at St. 28, 65 14' N. Lat., 55 42' W. L., 420 fathoms (soft, brown-gray
mud with innumerable arenaceous shells of Rhizopoda); and from the Manchester Museum our Museum
has obtained a gigantic specimen 1 ) taken near the Cape of Good Hope (■ Lion's Head N. 67°, E. 25
miles , I think, ca. 34° S. Lat.) at 131— 136 fathoms, together with several other specimens of lengths
of about 3 feet, and with polyps 3o mm long (according to information from Prof. S. Hicks on) 2 ).
Otherwise the species seems to be a deep-sea form; on the American side it has not been taken at
depths under 220 fathoms, but on the other hand at depths up to 1106 fathoms (U. S. Comm. of Fish.
and Fisheries Rep. 1883 (1885) p. 510).

Anthoptilum murrayi Kolliker.

Anthoptilum murrayi Kolliker. Chall. Rep. Perm. 1880, p. 14, PI. VI, Figs. 19 — 21.

Of this species the <Ingolf has brought home 43 specimens from four different stations, and
about half of them in a well-preserved state. In all chief characters they agree with the description
of Kolliker; at the upper end of the peduncle, however, a distinct elliptical swelling is found in most
of the specimens. As will be seen from the table below of the proportions of all the specimens,
several small ones are included in the number; but they differ so little from the larger ones, that no
doubt whatever can arise with regard to the determination of the species 3). In a few specimens the
circle of tentacles of the polyps is retracted; as a general rule, however, this is not the case, and the
polyps and their tentacles are only contracted in different degrees; the proportions given below are
of the polyps including the tentacles. Spicules are only found in the end of the peduncle, of a similar
size and form as in the preceding species, but in somewhat greater numbers, at all events sometimes.
Sexual organs are present, even in small specimens. Large eggs (or larvse?) are often found in the
tentacles.

Nr.

Total length in mm.

Length of peduncle

Diameter of the upper enlargement of the peduncle. . . »

Length of a polyp »

Breadth of the naked dorsal streak »

St. 83.

I.

2.

3-

405

310

275

47

37

40

5

4

4

8

8-9

6

2

2

2

St. 40.

St. 65.

55

15

1-5

4-5

c. 1

230

3°
4
4
[-5—2

145

27

3-5
5
1-5—2

') It measures 1350mm, the peduncle 190mm, its enlargement about 30mm in diameter. The polyps are coalesced into
groups of 5—8 individuals, and measure 17— 25mm. American specimens grow to a height of 2 feet, and a diameter of ca.
25mm (Verrill, U. S. Fish. Comm. p. 510).

2) West of Tristan d'Acunha, at 35 ° 41' S. Lat, 20 55' W. Long., depth 1500 fathoms, the Challenger obtained an
Anthopt. simplex KolL (a damaged specimen of a length of 400mm), which resembles A. grandiflorum, but with only two
polyps in each row; this specimen is possibly only an A. grandiftoyum the development of which has been delayed, and in
which the number of polyps for some reason or other has remained small; in other regions, however, the number of polyps
may be greater at a far smaller size of the colony, which is proved by the specimens of the «Ingolf., especially the smallest
of a length of only I22 m m; therefore I think it rather hazardous to suppose A. simplex and A. grandiftorum to be identical.

3) I am therefore able to declare with complete certainty that Roule's iTrichoptilum* sp. (I.e. p. 307) has nothing
to do with the genus Anikopiilutn, but is really, as stated on p. 50, Funiculina quadrangtilaris.

9*

68

PENNATULIDA.

Xr.

Total length in mm.

Length of peduncle

I )iam. of the upper enlargement of the peduncle »

Length of a polyp

Breadth of the naked dorsal streak

St. 47.

2S5

43
5
6

r-5-2

2.

255

35

5

S

2

5-

250

34

5

8

2

245 245

37 37

5 5

8 S

2 2

245

32

5

11

2

240

240

33

33

5

4

10

S-12

2

2

235

31

4

8-9

2

in.

11.

230

31
5

7

230

32
5
S

230

28

4
7

Nr.

L3-

14.

15-

16.

17-

18.

r 9-

20.

21.

22.

23-

24.

Total length in

mm.

225

225

220

220

220

220

220

210

205

200

195

185

Length of peduncle

»

34

32

3i

32

32

32

31

3i

29

24

2S

Diam. of the upper enlargement of the peduncle

5

5

4—5

5

4

5

5

4

5

5-5

4

4

Length of a polyp

I

8

8

12

6

9

10

10 — 11

S

6

10

Breadth of the naked dorsal streak

»

2

2

1-2

Nr.

25-

26.

27.

28.

29.

30.

3i-

32-

33-

34-

35-

36.

37-

Total length in mm.

1*

175

175

175

175

175

'55

145

125

108

102

85

So

2S

3°

2S

26

27

21

25

26

21

20

20

17

18

Diam. of the upper enlargement of the peduncle »

4

4

3

4

4

4

4

4

3

2-5

2

Len gth of a polyp »

6

7

5

6

7

7-8

6

Breadth of the naked dorsal streak i>

Anthoptilum murrayi has originally been taken by the Challengers, south of Halifax (one
specimen, 510"™ long), 48 8' N. Lat., 63 39 W. Long., depth 1250 fathoms; later, it has proved to be of
frequent occurrence off the east coast of North America, partly at the same localities as the preceding
species, in depths from 640—1362 fathoms (Verrill, Am. J. Sc. (3), vol.28, p. 220; U. S. Fish. Comm. Rep.
1883, p. 511). On the European side of the Atlantic, it was found almost at the same time at two
rather widely separated localities, viz. by the «Caudan in the Bay of Gascony (depth 1410 metres,
45 57' N. Lat, 6° 21' W. Eon g. (Paris), one specimen, 400'""' long), and by the «Ingolf» south of Iceland,
at four stations situated on almost the same line west to east between Greenland and the Freroe Isles,
viz. St. 83, 62 25' N. Lat, 28 30' W. Long., 912 fathoms; St. 40, 62° N. Lat, 21 36' W. Long., 845 fathoms;
St. 65, 6i° 33' N. Lat, 19 W.Long, 1089 fathoms; and St. 47, 6i° 32' N. Lat, 13 40' W. Long., 950 fathoms.

Thus it is evident that the species belongs to the deep part of the whole northern Atlantic
inside the territory of the positive bottom temperatures.

Fam. KophobelemnonidcB K611.

Kophobelemnon Asbjorns.
Kophobelemnon stelliferum (O. F. Midler).

Pennatula stellifera O. F. Midler. Zool. Dan. Prodr. 1776, Nr. 3076; Zool. Dan. 1788, p. 44, PI. XXXVI.
Kophobelemnon Mullen Asbjornsen. Fauna litt Norvegite II, p. 81, PI. X, Figs. 1 — 8.

PENNATUUDA. 69

Kophobelemnon stelliferum Koll. Monogr. 1872, p. 304, PI. XXI, Figs. 179— 181.
» Leuckartii Koll. Ibid. p. 306, PI. XXI, Fig. 182.

Moebii Kor. & Dan. Berg. Mus. Nye Gorgon, etc. 1883, p. 25, PI. XII.

abyssorum Kor. & Dan. N. North-Atlantic-Exp. Penn. 1884, p. 10, PI. IV, Figs. 17 — 20.

scabrum Verrill. Bull. Mus. Conip. Zool. Vol. XI, 1883, p. 7, PI. I, Fig. 5.
? tenue Rep. Comm. Fish and Fisheries 1883, p. 510, PI. Ill, Fig. 5.

Gunneria borealis Dan. & Kor. N. North-Atlantic-Exp. 1884, p. 58, PI. IV, Figs. 8 — 16.

Of the genus Kophobelemnon several species have been established, some from the Atlantic
(and the different branches of that ocean), others from the Pacific. Of these species as of so many
other sea-pens, the same things holds good, that the material has been too scarce to allow their determ-
ination to be reliable. Here, however, only the Atlantic species are to be mentioned in detail. As will
appear from the above list of synonyms I am of the opinion that they must all be united into one
species, which then receives the old specific name stelliferum given it by O. F. M filler. To this
must further be added the genus and species Gunneria borealis of Danielssen and Koren (as to this
see p. 72). The specimen of O. F. Muller was taken at Drobak, the numerous specimens of Asbjornsen
at different places in the Christiania Fjord. When Kolliker worked up the genus in his monograph,
he justly referred all the Scandinavian specimens at his disposal to one species, not only those origi-
nating from about the same localities as the specimens of Muller and Asbjornsen, but also a few
from somewhat great depths in the Atlantic proper, north-west of Scotland (even if he thought them
to be a variety durum on account of the size of the spicules; later examination of another specimen
from another part of the Atlantic attached this variety still more closely to the type (1. c. p. 370)). At
the same time, however, he established a new species K. leuckartii on some specimens from the
Mediterranean (Nizza); to be sure he declares this form to be very closely allied to the former, immerhin
mahnt der Fundort zur Vorsicht , and he enumerates a number of differences from his K. stelliferum.
These differences are: the size of the whole colony and of its individual animals, the number of the
polyps, the fact of the calcareous axis reaching to the distal end of the peduncle, a richer provision
of spicules and their larger size, the lack of colour in the epithelium of the stomach, and the bound-
ary between the peduncle and the rhachis being less distinct. I can, however, declare all these differ-
ences to be individual peculiarities; one of them - - the extent downward of the calcareous axis —
is quite casual, owing to a contraction of the soft tissues in this region ; and a comparison of the two
specific diagnoses will show that no difference is mentioned which is not individually variable according
to the knowledge we now possess of other Peunatulids. Nevertheless, the species K leuckartii of Kolliker
has been preserved without any objection ; only Paul Fischer, relying presumably on the descriptions of
Kolliker, regards the Kophobelemnon of the Mediterranean as the same species as stelliferum dont le
K. Leuckartii u 'est qu'un race (Note sur le Pavonaria quadrangularis etc., Bull. Zool. de France, Vol. 14,
1889, p. T,y). That it does not, at all events, represent a race peculiar only to the Mediterranean -
in this case the question of species or race would be rather a difficult one to solve — is shown by the
fact that it is said to occur also in the north, inside the territory of the genuine. K. stelliferum.

In 1872 the German Pommerania Expedition found some specimens of Kophobelemnon in the
Skager Rak which were determined as K leuckartii Koll. by F. E. Schultze (Jahresber. der Komm.

y PENNATUUDA.

zur wissensch. Unters. deutsch. Meere f. d. J. 1872—73 (1875) p. 142). Koren & Danielssen, however,
found that these specimens completely agreed with others, partly taken in the Skager Rak also (by
the Swedish gunboat Gunhild»), partly in fjords of the Norwegian west coast (Kors Fjord, Vest Fjord),
which specimens they now made to represent a new species K. moebii (Nye Gorg. etc. p. 25); but at
the same time they thought they had found the genuine K. leuckartii in a specimen from Trond-
hjem Fjord (1. c. p. 28), where this Peunatulid is said to be very common. Later, the same authors
described still another form as K. abyssorum from various Norwegian fjords, in some of which K. moebii
had been found. Thus we had no less than four Scandinavian species, of which only one seemed
the same as that from the Mediterranean '). If we now examine the descriptions of these supposed
species we shall search in vain for characters that might be taken as specific; only such differences
are mentioned as change with age and size; this holds good of such features as the length of the
colony, the number of polyps, the proportion between the length of the rhachis and of the peduncle,
the number of rows of polyps, the size of the polyps; also, the size and numbers of the spicules are
very varying; other specific characters arise from mere chance (e. g. a longitudinal furrow on the
rhachis or the like). As very characteristic examples of the value of the supposed specific characters,
the following may serve: in KopJi. abyssorum the stalk is almost twice the length of the rhachis; in
A', leuckartii it is only a little longer than the rhachis; in A', moebii the stalk is generally half the
length of the rhachis -- but in two very long specimens the stalk was relatively much longer, and
in young specimens it was as long as or even longer than the rhachis! I have carefully compared all
the type-specimens of these Scandinavian species, and I have not been able to keep them distinct;
further, I have examined the specimens of the Christiania Museum and a very large material of
Kophobelemnon in our own, and finally I have had the opportunity of examining a very large number
of K., partly collected in the deep channel of the Skager Rak by Dr. Joh. Petersen and Mag. A. C. Jo-
hansen, partly from different regions off Norway, taken by Dr. Hjort on the Michael Sars . In all
this large material I have only been able to see one single species; but of course it appears rather
different, if we compare extreme forms without an)- intermediate links: young colonies with giants
up to 75o mm long; specimens with quite few and relatively small polyps with others provided with
ca. 150 large polyps with bodies of a length of ca. 20 mm ; specimens with fully stretched polyps with such
where the polyps are quite retracted (for even if the latter case is rather rare it does occur, and shows
accordingly that the polyps are quite retractile); big and clumsy specimens with slender and thin
ones etc. All the K. hitherto found at Norway and in the Skager Rak are, beyond a doubt, the
same species as the K. stelliferum of O. F. Midler, Asbjornsen, and Kolliker; and the different authors
have also referred to this species all the specimens of K. taken at different localities in the Atlantic
by the English expeditious of the « Lightning*, the « Porcupine , the « Triton , and the .Knight Errant .
From the American side of the Atlantic, Verrill has established two species: K. scabrum
(Bull. Mus. Comp. Zool. XI, p. 7, PI. I, fig. 5), and A', tenue (Report Comm. Fish and Fisheres 1883,
p. 510, PI. Ill, fig. 5 (1885)). The former is short and clumsy, a young stage (56 mm long) with few (8)
polyps, rough on account of its rich provision of spicules; from the description I must regard it as

i) Panceri, however, had already in 1S71 found a A', at Naples which he had determined as K. stelliferum, but
which Kolliker supposed to be identical more probably with A', leuckartii (Monogr. p. 370).

PENNATULIDA. 71

identical with K. stelliferum 1 ); of the second it is only said that it is long, slender, and smooth,
with a number of large polyps ; neither this description nor the figure, which is very meagre in
details, exclude the possibility that it may be the same species. The figure gives the natural size
and shows by this and by the proportion as to length between the peduncle and the rhachis (they
are of about equal length) that the question is only of a young form. It is my conviction that hardly
more than the one species of K. stelliferum (Mull.) is known from the Atlantic Ocean. But on the
other hand, it is known in a more complete series of stages than most other Pennatulids; for my
own part I have had the opportunity of examining and comparing an almost continuous series
from 7.5 mra with one polyp up to 75o mm with more than 150 polyps; such a size as the last is
scarcely to be met with in previous literature; neither the first author of the species, O. F. Midler,
nor Asbjornsen, who established the genus Kophobelemnoit, nor Kolliker, had any idea of how far
their specimens — in spite of the full development of the sexual products — were from having
reached the maximum development. The very circumstance that specimens of 130 — 135""" become
the types of the species K. stelliferum might be the reason why ether stages, and especially the much
larger ones, were interpreted as belonging to other species; the most unwarrantable to my mind was
the K. moebii established by Koren and Danielssen, because these authors had so many different
specimens, that its relationship to the species of Asbjornsen should have been sufficiently plain.
Asbjornsen knew already quite small specimens; he has (1. c. p. 82, PI. X, figs. 3 and 4) described and
figured a specimen with only one polyp and another with two polyps, both about 20 — 3o mm long;
later, Grieg (Bergens Mus. Aarb. 1893, Nr. 2, ibid., 1896, no. 3) has described quite a series of develop-
mental stages from a length of 6 mm upwards; young stages are mentioned also in several other authors
and sometimes figured (for instance F. E. Schultze, 1. c. p. 142, PL II, fig. 2, a specimen 48""" long
with six polyps; M. Marshall Triton -Penn. p. 138, Marshall and Fowler Porcupine -Penn.
p. 460). The very young colonies show already that no certain proportion exists between the number
of the polyps and the length of the whole colon}-, as will appear from the following table:

Number of polyps

1.

2.

3-

4-

5-

6.

7-

8.

9-

.

6—27

26—30

3°-4i

3 2 —47

?

45-62

70

56-69

71-88

Already at a length of 8o mm we have specimens with 16 polyps, and at a length of 88 — 90 11 ""
even 18 — 20 polyps. In the specimens with only one polyp, the primary polyp, this is never placed
terminally, but a zooid is found near the apex of the rhachis. In the least developed specimens the
whole colony consists of these two individuals only, but by and by the number of zooids is increased
along the dorsal side of the stem below the terminal zooid. This zooid, however, is not so markedly
different from the other zooids as it is, for example, in Prima f ///a or Renilla (comp. also Grieg. 1893).

The Ingolf has only obtained young stages from four different stations: south of Iceland
(St. 40), in Denmark Straits (St. 10), and in Davis Straits (Sts. 24 and 28); altogether 22 specimens. Of

*) In a remark in Rep. Coram. Fish. etc. p. 510, several specimens are said to have been taken later, but further informa-
tion of these is wanting. I may add that a specimen from the Skager Rak of quite the same length and also with 8 polyps,
agrees remarkably well with the type-specimen of Verrill.

72

I'KNNATUUDA.

these specimens 21 are quite small, provided only with the primary polyp and a few zooids; the number
of these zooids is given in the table below. None of the zooids in these specimens appears as a real
terminal zooid; I doubt therefore --as already hinted above -- whether a real terminal zooid corre-
sponding to that of Pennatula and Renilla, is on the whole to be found in the genus Kophobelemnon.
The zooids do not occur quite symmetrically, nor always in two regular rows. In one of the speci-
mens (Xr. 21 from St. 40) sexual organs seem to occur in the primary polyp, which, however, is at
variance with the observations of Grieg (1893, P- I ^)-

St. 24.

St. 28.

St.

40.

Nr.

1.

2.

3-

4-

5-

6.

7-

8.

9-

10.

11.

12.

13-

14- | 15-

16.

17-

1 8.

19.

20.

21.

11
1-1

11
2-1

13
2-1

13
2-2

14 15 , 15
1-0 3-3 ' 3-2

16
2-2

16
4-4

17
5-5

18

4-4

19

4-4

20
2-3

20
4-4

21
7-6

22

4-3

27
6-6

121)
5-5

21
6-6

23
5-5

27

7 -6

Number of zooids on

each side

The specimen from St. 10 is 4o mm long, the rhachis in its broadest part ca. 4 mm broad; the
peduncle is remarkably short, only i5" ,m long, ca. 2 mm broad, so that the whole form is unusually
clumsy. All the polyps are retracted; only five of them, among which is the primary one, are fully
developed, but more are beginning as buds; the zooids are numerous, and are not restricted to the
dorsal side of the colony.

It was observed above that Gunneria borealis Kor. & Dan. must also be referred to Kopliobcl.
stelliferum. These authors have given this name to a sea-pen in which they thought they had a
very peculiar form approaching in some features to the Gorgonids; they say of it that we are able
with considerable certainty to satisfy ourselves that this sea-pen can not be assigned to any of the
known genera , but that it is, however, to be referred to the family of the Protoptiles. The name,
however, ought to disappear, and the comments on the supposed peculiarities, to be consigned to
oblivion ; the question is only of a damaged fragment of Koph. stelliferum. Gunneria borealis was
established on one single specimen figured in Norw. North- Atlantic Exp. Penn. PL IV. fig. 8; I have
had this specimen for examination, and it was easy to determine that it is the lower end of a Kopho-
belemnon the greater part of whose rhachis has been scraped away from the calcareous axis, whilst
the rest has been much displaced downward towards the peduncle, the soft tissues of which have also
been compressed. The effect of this compression, however, is not seen in the figure, although it
is very conspicuous, and especially so on the opposite side of the one given in the figure; the strong
incrustation with spicules pointed out by the authors, is partly owing to this compression. As correctly
stated in the description, little more than young polyps and buds of these are preserved; at the top,
however, a few developed polyps are present, more or less retracted; two are seen in the figure (a, b).

The young polyps look the same as elsewhere in Kophobelemnon] they only differ from the
zooids accordingly in being larger and in having rudiments of tentacles (the authors have correctly
seen « rudimentary » tentacles); as always in this region, real zooids are found between them and are
seen quite distinctly in the figure (in the description, on the other hand, the authors pronounce the
specimen destitute of zooids, although they think that the part of the rhachis lacking has been

] 1 The lower end broken.

1'KXXATULIDA.

73

provided with such, as they have seen a zooid under one of the preserved developed polyps). When
the authors attribute <cells in the developed polyps to their < Gunneria , and figure such a polyp
with a v cell - in fig. 9, it is to be observed that this figure cannot exactly be called a true representa-
tion of fact, nor does the description correspond with it. Kophobelemnon has not a real calyx , nor has
this specimen; only a superficial examination could so misinterpret the features arising from the different
degree of retraction of the polyp-body and the tentacles. Otherwise, it will be found that the special
statements of the authors with regard to the form of the spicules, the strong equipment with spicules
of the tentacles and polyp-body, the pigmentation of the pharyngeal sac and gastric filaments
etc., correspond exactly with the features in Kophobelemnon. It might be supposed that Danielssen
and Koren have never made a comparison with this form, as any one who compares the Gunneria -
specimen with a Kophobelemnon ( morbii ■ ), must be struck immediately by the similarity; even by a
comparison of the not quite correct figure 8 with the corresponding part of a Kophobelemnon of a
suitable size, the agreement will easily be seen; it seems to me superfluous, therefore, to enter into
any further details.

Distribution and Occurrence. -- The stations of the Ingolf extend the known territory
of the species considerably:

St. 40: 62 00' N. Lat., 21 : 36' W. Long., depth 845 fathoms, bottom temp. 3 Q -3 C.
» 10: 64 : 24' 28 c 50' 788 3°-5

24: 63° 06' 56' 00' » 1199 2°4 »

28: 65 14' 55 42' 420 3°-5

The steamer Thor has further added as new localities: 63 05' N. Lat, 20 7' W. Long., 557
metres (St. 167 of the Thor , south of Iceland, H/ 7 1903). Here 14 specimens were obtained, of which
the largest is io5 mm long with 12 polyps, the smallest 50™™ with 7 polyps. Further: 62 57' N. Lat.,
19° 58' W.Long., 957 metres; here three specimens were taken: two small young stages with solitary
primary polyp, the one only 8 mm long and very small, with one large and two quite small zooids;
the other 24""" long, rather big (ca. 2 mm broad), with a polyp-bud just appearing on the right side
and with many and large zooids; the third specimen 55 mm long, damaged, with several polyps and
large zooids.

Apart from the coast-regions of Norway where the Vest Fjord (ca. 67 — 68° N. Lat.) is given as
its northernmost occurrence, the species has not hitherto been known farther north than 6o° 18'N. Lat.,
in the Fseroe Channel (the Triton- Exp. St. 8), whilst its most westerly occurrence on the European side
of the Atlantic did not extend much farther than io° W. Long, west of Brittany (the Porcupine Exp.
1870; see Kolliker, Monogr. p. 370). Now its territory is seen to extend, not only to Iceland but across
the Denmark Straits and quite into the Davis Straits, west of Greenland; its occurrence there further
corroborates my supposition that the Kophobelemnon occurring farther south on the east coast of
America belong to the same species. Formerly, it was known from many of the Norwegian fjords and the
adjoining coast-regions, viz. from the Vest Fjord and farther south through the channel of the Skager
Rak to the Christiania Fjord and to Bohuslan; also from many places in the Fa^roe Channel; off
Brittany; from the coast of Les Landes: from different places in the Mediterranean (Cape Beam, Nice

The Ingolf-Expedition. V. I. IO

- , PENNATUUDA.

74

Naples: K. leuckartii ); finally on the American side of the Atlantic near George's Bank (41 29' N. Lat,
65° 47' W. Long., <tK. scabrum ), and at several other places, not more exactly designated. Its distribu-
tion, then, probably comprises the whole northern part of the Atlantic (at all events north of ca. 40 )
within the territory of the positive bottom-temperatures. It must be observed, however, that a few
specimens have been taken by the Porcupine and the Triton outside the warm area, but at its
very border, in the peculiar narrow strip of water which stretches from the cold depths of the North At-
lantic into the Faeroe Channel towards the Wyville Thomson ridge. The question is of two localities, very-
close to each other: 6o° 14' N. Lat., 6° 17' W. Long., and 6o° 18' N. Lat, 6° 15' W. Long., where the respec-
tive bottom-temperatures are given as -=- 0.8° C. (the Triton , St. 8, Marsh, p. 120) and 4- 1.1 C. (the
Porcupine; 1869, St. 57; see < Depths of the Sea PI. IV and p. 143); only two specimens were taken
at either place. That the occurrence within the cold area must be regarded as a mere exception
and is only possible so close to its border, will appear from the fact that neither the North-Atlantic
Expedition nor the - Ingolf , any more than the < Michael Sars» or the Thor ■:■, have obtained any
Kophobelemnon in the really great depths of the cold area of the North Atlantic, and also from the fact
that K. stelliferum evidently reaches its highest development in such regions as the Norwegian fjords
and especially the Skager Rak where the largest specimens hitherto found have been obtained.

The ba thy metrical range of the species is evidently very wide: in the Skager Rak it
has been taken at depths from 26—400 fathoms; at Norway from 20—400 fathoms; in the Mediterra-
nean at ca. 30 fathoms, as also at greater depths; in the Atlantic itself it has upon the whole been
taken at considerable depths, between 420 and 1 199 fathoms, but most frequently, it seems, as small or
even very small specimens; at America, finally, in depths from 980 to upwards of 2000 fathoms (the
latter depth is stated by A. Agassi z in: Three Cruises of the Blake, », Vol. 2, 1888, p. 142, and
V err ill for his two species » gives depths from 499 to 2369 fathoms; Rep. U. S. Fish Coram.,
1883, p. 510).

I shall only add further that the genus Kophobelemnon is also distributed over the Pacific, and,
according to the statements in the literature, is represented there by several species; but the number
of these will probably also be reduced in future; of one of these species, K. affine Stud, its author
already states that it is very closely allied to K. stelliferum ( Albatross , Rep. Bull. Mus. Comp. Zool.
Vol.25, Nr. 5, p. 57), and the same is said by Kolliker (Chall. Rep., p. 16) of a young stage from the
seas north-east of New Zealand.

Fam. Umbellalidce Roll.
Umbellula Cuv.

This genus, which was down to the year 187 1 only known from Ellis's and Mylius's im-
perfect descriptions of the dried specimens of U. encrinus brought home from the Polar Sea in 1753
by the Jutland skipper Adrians, has been proved to have a worldwide distribution in the great depths
of the sea by the expeditions of recent times, and several species have now been established. The
knowledge of these species, however, leaves much to be desired with regard to accuracy; of several

PEXXATVLIDA. 75

— I think most — of them only young stages are known, and we know from U. encrinus that the
appearance of the species changes much during growth; no single investigator has been able to com-
pare to any considerable extent the many species established on more or less scarce material etc.
So much seems to be certain, however, that there is a number of species with well-developed spicula-
tion, and others where the spiculation is quite reduced, only the peduncle containing quite micro-
scopic spicules; others again are stated to be quite devoid of spicules, but, as will be shown with
regard to the two species more particularly mentioned below, not all the statements in this respect
are correct.

The following species have been described as devoid of spicules: U. encrinus (L.), magniflora
K611., lindahlii K61L, gracilis Marsh., gcnicitlata Stud., and (I suppose) bairdii Yerr. Apart from U. geni-
culate with regard to which I can say nothing, I believe all these species to be closely allied; but as
long as the knowledge of the characteristics of the species, and their variability and changes during
growth is so uncertain, I think that some stress is to be laid on the geographical occurrence; thus, I
believe we may take it for granted that those of the mentioned species which have been found in the
Pacific, viz. U. magniflora and geniculata (and Studer's U. encrinus ) are specifically different from the
others which belong to the Atlantic and the Arctic Oceans; and further, as the whole group of Penna-
tulids in its distribution within the northern parts of the ocean has proved to be remarkably depen-
dent on the bottom-temperature, there is every reason to suppose that the form which has been found
well- and even enormously developed within the cold area , is specifically different from those found
in the warm area of the Atlantic and the adjoining seas. On the other hand, I am inclined to regard
these latter U. lindahlii, gracilis, and bairdii as one and the same species.

Umbellula lindahlii K6U.

Tab. Ill, Figs. 37—46.

Umbellula miniacea Lindahl. Kgl. Sv. Vet.-Akad. Handl. Bd. 13, Nr. 3, 1874, p. 12, PI. I, II, Figs. 12— 19.
» pallida ibid. p. 13, PI. III.

Lindahlii Kolliker. Festschr. Phys.-Med. Gesellsch. in Wurzburg 1875, p. 10.
gracilis Marshall. Rep. « Triton -Penn. 1883, p. 143, PI. XXV.

Bairdii Verrill. Am. Jour. Sc. (3) Vol. 28, 1884, p. 219 (Note) and U. S. Fish. Comm. Rep. 1883,
(1885), p. 509, PI. I, Figs. 1— 2.

Two specimens of the genus Umbellula, taken by the Ingolf respectively east and west of
Greenland in the warm area, I consider to belong to the same species as the two specimens of
Lindahl from West Greenland.

One of the specimens, from the deep part of the Davis Straits, is, I think, the smallest and least
developed stage hitherto known of the genus Umbellula. It has a total length of 48""", and appar-
ently consists only of the primary polyp (PI. Ill, fig. 37); a closer inspection shows, however, that it
is provided with a somewhat great number of zooids (z| and a small bud of another polyp (p). The
developed polyp is supported by a fine narrow stalk, of which the lowermost 2 mra form an egg-shaped
swelling, of a diameter hardly r mm ; this swelling is really the whole peduncle, as zooids appear

7 6

PENNATULIDA.

immediately above it (comp. fig- 39); accordingly, the remaining part of the stalk must be regarded as
corresponding to the rhachis of other Pennatulids. This part decreases regularly in thickness upwards
- the middle thickness is ca. 0.448"™ — until ca. 8.5' nm from the mouth of the end-polyp it rather
suddenly tapers to a thin part - only o.24 mm broad, — which again widens quite evenly up-
wards and extends to the body of the primary polyp. The calcareous axis shines distinctly through
the thin sarcosoma, and shows the characteristic quadrangular form, with rather deeply concave surfaces
and ridge-like projecting edges; at the transition to the thinner part of the stalk it narrows abruptly;
from this point, where the diameter is o.io,2 mn \ it continues quite thin up through the dorsal part
of the polyp-body, and ends in a rather curiously shaped double curve. The polyp, unfortunately, is
somewhat damaged, some of the arms being broken off, and the preserved arms are twisted together
and even rather damaged; they are longer than the polyp-body; pinnules of different sizes alternate
without any order. The polyp is somewhat curved in a halfmoon-shaped way and compressed late-
rally; reckoned from the mouth to the polyp-bud on the left side it is ca. 4 mm long; the dorso-ventral
diameter of the part in front of the calcareous axis is somewhat more than i mm . The mouth is di-
stinctly oval with long axis dorso-ventral, and placed on a low oral cone very sharply divided into
eight lips, each corresponding to an arm and gastric cavity; eight distinct lines corresponding to the
edges of the eight septa, run down the polyp-body, and disappear at its hinder end; the much folded,
dark-pigmented pharyngeal sac is seen faintly through the wall of the body, and below this the six
much twisted gastric filaments on the ventral and lateral septa. By a suitable clearing (in glycerine
or bergamot) these filaments are seen suddenly to cease in the lower part of the gastric cavity, where
their septa may still be traced for a short distance corresponding to the part which carries the sexual
organs in developed colonies; of such organs no trace is seen; the lower end of the two dorsal septa
which are turned towards the calcareous axis may also be seen; farther upwards they are hidden by
the twisted filaments of the other septa.

On the left side of this primary polyp, immediately below the place where the septa cease, is
a rudiment of a polyp Nr. 2 (p, fig. 38). This rudiment projects only quite slightly as a low truncate
cone with a cleft-like oral aperture; the eight septa with their filaments are distinctly begun, but no
trace of arms is seen. The length of this rudiment is 0.32™™, the breadth o.i54 ram .

Along the primary polyp's dorsal side, which is distended by the calcareous axis, is a row of
zooids (z) on each side, all placed in the usual way, i. e. with the dorsal side toward the apex of the
axis. No terminal zooid is seen above the end of the calcareous axis. Zooids of quite the same
appearance are further found down the stalk with intervals of 0.5— i mm , down to ca. 2 mm from the
lower end, as before mentioned (fig. 39). The zooids vary in size between 0.096 and o.240 mm ; the
lowermost ones are generally the smallest; on the stalk they are in the main placed on the lateral
surfaces, the ventral side, at all events, quite wanting zooids. None of the zooids show any trace of
the tentacle which the larger zooids always possess in developed Umbcllula elsewhere.

On the lower egg-shaped end of the stalk, which represents the peduncle proper, quite small
oval calcareous spicules are seen under high magnifying powers and by suitable clearing (as I did
not wish to destroy the specimen, I had to be content with only convincing myself of the existence
of these spicules); the other part is quite devoid of spicules.

PENNATTJLIDA. 77

The other specimen, from the Denmark Straits, was unfortunately not preserved until it was
quite shrunk; it had long escaped attention, because it was jammed in at the iron frame of the trawl,
and was quite wrapped up in threads of swabs; the specimen was otherwise quite complete, the
slender calcareous axis not even broken. The total length, the stretched polyps with their tentacles
included, is (fully) 530™'"; ca. 500 mm from the lower end of the peduncle to the base of the polyp-
bodies. The length of the peduncle is only 40 mm , its largest diameter somewhat more than 3 mm ; from
the peduncle the stalk decreases in thickness upward from ca. 2 mm to 0.5" 1 ™ and then it again in-
creases evenly in thickness towards the cluster of the polyps; it is somewhat spiral-shaped longitudi-
nally, and bent in a large flat curve ca. 20 nim below the polyp-cluster whose longitudinal axis is a
direct prolongation of the upper part of the stalk. Even if respect is paid to the completely shrunk
condition of the specimen, which disparages, of course, the value of all the measurements to some
degree, it will be seen that the form is remarkably long and thin. The polyp-cluster is relatively
small and slender, ca. 37 mm long, measured from the point where the stalk begins to spread into the
polyp-bearing part, to the point of the stretched tentacles; it consists of six developed polyps, and no
rudiments are found; the polyps are of somewhat different sizes, 6 — n mra long, with tentacles longer
than the polyp-body, the longest ones up to ca. 20 mra , with irregularly alternating larger and smaller
pinnulse. The primary polyp is easily recognisable from the fact that the calcareous axis may be traced
distinctly into the dorsal side of its base: it is smaller than the polyps placed nearest to it laterally.
The latter are arranged in such a way, that two are placed to the right of the primary polyp and
three to the left; of the latter, the one placed nearest to the ventral median line is the smallest, it
is even considerably smaller than the primary polyp; the others are of about equal size. The polyp-
bearing part of the stalk (the rhaehis: of the authors) is about 13"™ long, 5 mm from side to side,
and 3 mm from the dorsal to the ventral side in the broadest place.

The zooids form tongue-shaped areas between the polyps, two on the left side, one on the
right; between the polyps of the two sides on the ventral aspect there is a narrow naked streak,
whilst the zooids of the dorsal aspect continue up on the base of the primary polyp. On the upper
part of the stalk, below the cluster, the zooids are placed all round, but they soon pass into a single
row on either side, and they continue to be arranged chiefly on the two lateral surfaces quite down
to the peduncle; a few may, however, be found here and there on the other surfaces. In spite of the
shrunk condition the two dorsal mesenteric filaments (the only ones) of these zooids are seen very distinctly;
on the other hand, no tentacle is seen on the zooids of the stalk. The calcareous axis throughout
its whole extent is quadrangular with deeply concave surfaces. Spicules are wanting, except in the
lower end of the peduncle, where there are numerous, very small -- o.oi6 mm long, o.oo64 mm broad -
oval calcareous bodies, often slightly constricted in the middle, so as to get somewhat the shape of an
8; still smaller, quadruple calcareous bodies are rather numerous. The colour of the polyps was a
dark chocolate-colour; in water a red-brown dye may be pressed out of crevices in the polyps.

This specimen agrees very well in appearance with Liudahl's two specimens from Baffin's
Bay and the Umanak Fjord, and especially with the one he has called U. pallida; the whole slender
form of the polyp-cluster the fact that the tentacles of the polyps are much longer than their body
— about twice as long — and the exceedingly long, slender, and graceful stalk, render it certain that

PENNATULIDA.

our specimen belongs to exactly the same species at that of Lindahl, and on the other hand also
very probable that it is a species different from U. encrinus. The young specimens of U. encrinus,
to be more particularly mentioned below, as well as the more or less young specimens with few
polyps in the cluster obtained by the Norwegian North-Atlantic Expedition, are far more robust and
short-stalked than this specimen or those of Lindahl; 1 think therefore that Danielssen's incorporation
of these latter into the species U. encrinus may, at least for the present, be regarded as unjustified,
the more so indeed as I cannot find the transference well-grounded by this author.

During the cruise of the Michael Sars in 1902, a specimen of an Umbellula has been taken
which is very short-stalked when compared with the specimen just mentioned (PL III, figs. 40 — 45); it
thus has a robust appearance similar to the young U. encrinus; therefore, if the locality had not told
against it, I would have been inclined to refer this specimen to this species. The locality is St. 76
of the Michael Sars , 59 29' N. Lat, 7° 51' W. Long, south of the Wyville Thomson ridge, accordingly
within the warm area , the depth 580—689 fathoms. The tentacles, however, are much longer in
relation to the polyp-body, than is the case in the specimens of U. encrinus I have seen, being about
twice as long as the body. The length of the stalk is io7 mm , the length of its lower swollen part
23 mm , the diameter of the peduncle and the swollen lower part of the rhachis ca. 2 mm , the thinnest
part of the stalk ca. o.5 m,n , the length of its upper widened part ca. g mm , the length of the cluster
ca. 25 mm , that of the club of the rhachis ca. 6 mm , its breadth 3.5 mm . The cluster has five developed
polyps and one rudiment; the length of the polyp-bodies is 5 — 6 mm , that of the longest tentacles i3 mm .
The terminal polyp (P, figs. 40—41) is easily recognisable as such; on its right side are two fully
developed polyps and one small one, on its left side two developed polyps; the largest of the devel-
oped polyps is the one nearest to the primary polyp on the right, the smallest the second on the
left counting from the primary polyp. The small polyp is remarkable from the fact that its dorsal
arms are much smaller than the ventral (comp. below under U. encrinus). Above the circle of the
lateral polyps, ventral to the primary polyp, are two large zooids, each with a well-developed tentacle
with small side-branches; a similar large zooid is found to the left of the small polyp. Not only the
zooids of the rhachis-club, but also many of the zooids of the other parts of the stalk quite to the
lower swelling, show a tentacle ; on the small zooids of the stalk it is, however, very fine and delicate.
(This explains the want of a tentacle in the zooids of this region in the other specimens as probably
due to some accident.)

This specimen has also the same numerous, small spicules in the lower end of the peduncle
(PI. Ill, figs. 44 and 45). In the living state the colour was reddish-brown passing somewhat
into violet.

Being of opinion that the specimens of Lindahl and those described here, ought to be kept as
a separate species I have designated it with the name U. lindahlii given it by Kolliker in 1875 (1. c.
p. 10). The reason for referring the above described, quite small stage from the Davis Straits to this
species, is partly that the tentacles of the primary polyp are so long, partly and more especially the
locality where this specimen has been taken. When further, I have given U. gracilis Marshall and
U. bairdii Verr. as synonyms, I have done so as a matter of judgement, as the description of these shows

PKNNATUUDA.

79

nothing opposed to it 1 ), whilst on the other hand, the long and slender form of both the stalk and
the cluster, and also the geographical occurrence speak in favour of this opinion. As to the latter,
U. gracilis has been taken (by the Triton ) at 59 29' 30" N. Lat., 7 13' W. Long, 555 fathoms (the
Atlantic, west of Rona), U. Bairdii at 38° 30' 30" — 38 53' N. Lat., 69 8' 25" — 69 24' 40" W. Long, 1731
—2033 fathoms, east of North America (south of Martha's Vineyard), both accordingly within a terri-
tory that has proved upon the whole to possess the very same species of Pennatulids as the norther-
most part of the Atlantic, so far as the warm area extends.

The species U. lindahlii is now known from the mouth of the Umanak Fjord, 71 27' N. Lat,
53 58' W. Long., 122 fathoms, from Baffin's Bay, 70 43' N. Lat., 52 3' W. L, 410 fathoms (Lindahl). from
the Davis Straits, the «Ingolf 's St. 36, 6i° 50' N. Lat., 56° 21' W. Long., 1435 fathoms, from the Denmark
Straits, St. 90 of the Ingolf », 64 45' N. Lat., 29 06' W. Long., 568 fathoms, and from the Fseroe Channel,
St. 76 of the Mich. Sars», 59 29' N. Lat, 7 51' W. Long., 580—689 fathoms; probably it is distributed
over the whole northern part of the Atlantic. Farther south it is perhaps replaced by other species
that cannot be confused with it, for one thing on account of their abundance of developed spicules,
as U. giintheri K611. (found in the Atlantic almost under the equator, at the West Indies, and off the
east coast of North America, sometimes together with U. bairdii), and U. thomsoni K61L, found
between Portugal and Madeira (the Challenger I and in the Bay of Gascony (the Caudan.) 2 ).

Umbellula encrinus (L.).

PI. Ill, Figs. 46—51.

his (lie riu us Linue. Syst Nat. Ed. X. 1758, p. 800.

Umbellula encrinus Danielssen & Koren, Norw. North-Atlantic-Exped. Penn. 1884, p. 13 — 56, Pis. V — XII.

The polyps (in large specimens) form a dense, hanging cluster; this cluster is apparently radiate,
the largest polyps forming an (apparently) closed, lower circle of 9 — 12 (13) individuals which may hide
the other polyps more or less completely; the tentacles are of about the same length as the polyp-
body often somewhat shorter. The part of the rhachis-club seen below the polyps is formed like a short
calyx covered with zooids; each of these is provided with a ventral tentacle; between the bases of the
polyps in the lower circle the zooids form regular, tongue-shaped areas; among the zooids in the cluster
are always found a larger or smaller number of very considerable size. The lower part of the stalk
passes rather abruptly into a swelling that is continued into the peduncle without any sharply marked
boundary; the upper part of the stalk towards the cluster, passes evenly into a somewhat compressed
and widened part (the sheath-formed dilation Dan.); this part again widens rather abruptly into the

I ) With the exception, however, that the calcareous axis in U. gracilis is said to be cylindrical for a great part of
its length. This statement I think to be incorrect, but the exact shape has probably been indistinguishable through the soft
tissues. The statement of a complete want of spicules I take to be owing to the fact that the skin of the peduncle has not
been examined under sufficiently high magnifying power.

-') It is perhaps this Umbellula which is mentioned by P. Fischer (1889, 1. c. p. 37) as U. ambigua Marion taken off
Bidassoa. The Umbellula found by the «Gazelle>, which Studer (Anthoz. Gazelle, p. 6741 designates as U. thomsoni K6U.?,
is perhaps correctly determined, but no description is given of it; it is only said that the two specimens obtained were in a
bad state, so that the specific identity could not be made out; the colour was brown in the living state. The locality: west
of Africa, io° 12.9' N. Lat, 17° 25.5' W. Long., depth 360 fathoms.

8o

PENNATUUDA.

polyp-bearing rhachis-club. The whole stalk is covered with zooids, downward to the upper part of
the lower swelling, where the cessation of the zooids shows the boundary to the peduncle. Of the
zooids of the stalk, those of the upper swelling are almost always provided with a tentacle like those
of the rhachis-club; but also farther down the stalk, for instance in the thinnest part of it, the zooids
may be provided with a tentacle, but it is then a very thin one; most likely all zooids have this
equipment, but the thin tentacles are easily torn off, and this I take to be the reason why it seems
to be wanting in most of the zooids of the stalk. The upper part of the stalk, below the rhachis-club,
is twisted and bent in various ways, but always in such a manner that the cluster remains pendent;
further the whole stalk, above the lower swelling, is turned like a spiral round its longitudinal axis.
The calcareous axis is quadrangular with deeply concave surfaces and rounded edges. Very small
spicules are found, but only in the lower end of the peduncle. The colony grows to a
height of more than two metres.

Of this species the <Ingolf > brought home five specimens (the Nrs. 2, 3, 5, 6, and 7 of the table
below), and later the Amdrup — Hartz Expedition has brought three more specimens (Nrs. 1, 4, and 8),
Caud. Jensen from the cruise of the «Mich. Sars>, two specimens (Nrs. 9 and n), and the steamer .Thor ,
one (Nr. 10). The particulars with regard to these specimens are given in the table below, which shows
that only six specimens have kept the stalk entire, or kept it so far, that it has been possible to
measure its length (Nrs. 4 and 10 are broken and parts of the calcareous axis wanting), and that two
of these six are young stages; the specimens Nrs. 8 and 11 are torn off clusters and in a very bad
condition, and therefore some of the measurements given are rather uncertain. I think that the spec-
imen Nr. 3 is the longest one hitherto measured.

Nr.
Locality

Cape
Brewster

St. 105 ; St. 116

4-

Canning
Land

St. 116

St. 1 16 St. 116

Canning
Land

9-
M. Sars
St. 34 a

< Thor>
St. 51

11.

«M. Sars
St. 34

Length of the stalk . in nun.

Length of the bulbous part
in 111111.

Diameter of the lower end of

the peduncle in mm.

Diameter of the swelling above

the peduncle in mm.

Diameter of the thinnest part

of the stalk in mm.

Length of the sheath-formed

dilation in mm.

Length of the cluster . in mm.

Length of the rhachis-club

in mm.

Breadth of the rhachis-club

in mm.

Number of polyps

in the lower

circle

148

215
40

2 2.5

2 3

0.5 0.5

10 9

17 23

2-5—3 3-5

4(+i) 7(+4)

2350
312

17
23

1090

24
5-6

c. 100

170

40
40

no 125

38 43

1090
230

17
iS

4.5
c.6o(?) c. 84

105 c. 85

150
33

35

4o 39(37 + 2)] 38 3S1) 29 37(31+6)

128

c. 40

37
25

3(+i)

6(+4)

1410
290

17
c. 25

c-5

c. 100
c. 130

40

42
41

c. 90 — 92

30

21
25

T ) Several small polyps, but none still in the rudimentary stage.

PENNATULIDA.

81

Nr.
Locality

I.

Cape
Brewster

2.

St. 105

3-
St. 116

4-

Canning

Land

5-
St. 116

6.
St 116

7-
St. 116

8.

Canning
Land

9-

«M. Sars»

St. 34 a

10.

«<Thor.>
-St. 51

11.

«M.Sars ■
St. 34

Length of the body of the lar-

8-9
9-10

5

+

'12—13

IO — 12

7
1

_L

77
c. 70

13
4

100
So— 90
20—27

5

5°
20

+

67
60—65

7

65
70
20

c 42—55

c 40-55

17

_L_

58
49
36

2

60-63

60 — 62

26

c 1.5

+

41

6

c. 1

+

Length of the tentacles of the
largest polyps in mm.

Length of the body of the least
developed polyps ... in mm.

Length of the body of the lar-
gest zooids in the cluster
in mm.

Terminal polyp certainly recog-
nisable as such

After the comprehensive description which Danielssen has given of this species, on the basis
of the excellent material of the North-Atlantic Expedition, its identification will generally be an easy
task, and the principal features of its structure be ascertained. There are, however, several places
where additions or alterations have had to be made; in the above short description some of these
have already been mentioned, and in the following they will be more particularly discussed. Thns, it
may be pointed out that Umbcllula encrimts like the preceding species (and probably most Umbellula-
species), is provided with zooids on the whole stalk from its very earliest stage, at all events in all the
stages hitherto known. Danielssen says indeed, 1. c. p. 52, that zooids occur < everywhere on the stalk
with exception of the inferior part of the bulbous portion , but by this he has evidently been thinking
of the large full-grown specimens, as none of his figures of young stages show zooids on the stalk
much below the cluster; only in fig. 20, PL VII zooids seem to be indicated down the thin part of the
stalk, and in the text it is for the first time mentioned in the detailed description of this specimen
(Nr. 5, 380™'", with thirteen developed polyps in the cluster) that zooids are generally found «further
down the stem, they become stdl more dispersed, and are situated, partly, in a series on each side
(I.e. p. 23); in four of the large specimens later described, zooids on the stalk are especially mentioned,
most thoroughly for Nr. 12. A very minute examination shows, however, as before mentioned, that
they are found in the very youngest stages, which I have been convinced of by inspecting Danielssen's
material. My two young stages mentioned above (Nr. 1 and 2 of the table) with respectively four and
seven polyps in the cluster carry zooids all along the stalk to the bulbous part; these zooids are
very small, much smaller than those placed near the cluster which feature is preserved also in
the older colonies, where the number becomes very large; they certainly do not seem to have any
tentacle; I have succeeded, however, in seeing such a tentacle in a single specimen (Nr. 10), in zooids
placed below the sheath-formed upper dilation . In the youngest stages these zooids of the stalk are
placed regularly in two longitudinal stripes, so that the dorsal and ventral sides of the stalk are naked;
this latter fact, however, is only to be decided with difficulty on account of the spiral-like twisting
of the stalk; frequently, individuals a little larger alternate with quite small ones; the latter have
evidently just appeared. By and by, as the colony grows, the number of zooids in the stalk increases
greatly; in large specimens, as Nrs. 3 and 4 of the table, they are arranged in short longitudinal rows,

The Ingolf-Expedition. V. i. II

3 2 PENNATULIDA.

right down on the upper portion of the bulbous part; in the narrow part of the stalk the small rows
unite and form two broader bands; but outside of these are also found more scattered rows of zooids,
so that there is hardly any quite naked dorsal and ventral surface of the stalk. From this fact it
follows, that only the zooid-less part of the bulb corresponds to the peduncle of other Penuatulids, and
that the greater part of the stalk from the beginning is to be regarded as the rhachis; to restrict this
name to the widened upper end bearing the polyp-cluster, as has been done hitherto by all authors, is
in reality misleading; I therefore designate this part as the rhachis-club.

The polyp-cluster in the younger stages is always quite distinctly bilateral, often (oftenest?) also
asymmetric, and the terminal (primary) polyp is always evident as such; in the large colonies the
cluster is apparently radiate, and the terminal polyp is often not to be determined with certainty. A
closer inspection shows, however, that the polyp-cluster is still bilateral, and this fact may be discerned
whether the terminal polyp may be made out with certainty or not.

In the younger colonies the upper end of the rhachis-club is seen to continue as a broad belt
of zooids directly to the base of the polyp, that is the primary individual. The upper end of the cal-
careous axis runs originally under this belt of zooids; from the structure of the primary individual
this belt must be determined as dorsal (Danielssen, like all other recent authors, follows Kolliker here
also, and uses the expression ventral); on the opposite -- accordingly ventral -- side of the rhachis-
club new polyps arise, often not in the ventral median line, but beside it. In my youngest specimen,
Nr. i, three developed polyps and a small rudiment (i-5 mm high) are placed below the terminal polyp
(fig. 46 P.), but otherwise of the same mutual height; of the developed polyps the terminal polyp is
the smallest. To the left of the terminal polyp are two developed polyps, to the right, one polyp and
the small rudiment; these (3^1) polyps represent evidently only the lower circle of polyps in a devel-
oped cluster; the tongue-shaped groups of zooids on the rhachis-club between the bases of the polyps
of the lower circle of the cluster are indicated in the following way: besides the dorsal belt of zooids,
which has not yet assumed the tongue-shape, there is only a distinct tongue-area between the two
polyps of the left side (fig. 46 t), and a quite slight indication on either side of the rudiment.

The other young stage, Nr. 2 (fig. 47), has six polyps below the terminal polyp somewhat dif-
ferently developed, and in addition four small rudiments; all these six polyps and the rudiments still
only represent the lower circle of polyps of the large clusters; nor are the rudiments found here in
the ventral median line of the rhachis-club; the fact is that four developed polyps are found to the
left of the terminal polyp, to the right only two (figs. 47—49: 1, 2); the two polyps between which
the rudiments are found are evidently last developed, being both shorter and more slender than the
others, the terminal polyp included; this latter is likewise also smaller than those nearest to it. The
rudiments are of different sizes, so that they are seen to have arisen alternately to the right and
to the left. Corresponding to the arrangement of the polyps, the rhachis-club — besides the dorsal
belt (dt in figs. 47, 48, 49) — shows three distinct tongues of zooids (t, fig. 48) on the left side decreas-
ing in size towards the rudiments, one on the right side; in the rudiment-zone (fig. 49) there are slight
indications of as many tongues as there are buds. In the cluster, at the ventral base of the terminal
polyp are two zooids provided with a tentacle; one of them is very large (fig. 47, z), and its tentacle
has one lateral branch.

PENNATULIDA. 83

Unfortunately intermediate stages are wanting in my material between these young stages and
the large ones; but the break is filled, at all events partly, by the material of the Norwegian North-
Atlantic Expedition, which I have had the opportunity of seeing in the Bergen Museum. If we examine
the specimens described by Danielssen as Nrs. 2, 3, and 4, we shall find that besides the terminal polyp
and those representing the lower circle, 2 — 3 polyps are further found above the latter, which show
by their smaller size that they have only arisen recently; and in the specimens described by Danielssen
as Nr. 5 and 6, the polyps placed above the lower circle have increased in number and size, whilst at
the same time the number in the lower circle has increased ; some of the specimens mentioned show
also that this latter increase is due to the fact that new rudiments of polyps arise in one certain
place in the lower circle, on the ventral side of the rhachis-club 1 )- The lower circle in all young
stages is really interrupted in two places, viz. at the dorsal side of the colony where the rhachis-club
with its dorsal belt of zooids reaches to the base of the terminal polyp, and on the opposite side
where the rudiments arise; in other words: the lower «circle> is formed of two lateral rows, a right
and a left, corresponding to the two lateral rows of polyps which are placed along the dorsal side of
the rhachis in sea-pens of the common long form; as in such forms the polyp in each of the two
rows which is placed nearest to the terminal polyp (uppermost) is the oldest one, the polyp placed
farthest therefrom (lowermost) the youngest — such is also the case here in Umbellula.

My two young Umb. encrinus (Nr. 1 and 2) — in spite of their considerable size — thus represent
just the stage which in Virgularia I have called the Protocaulon-stage, and the stage which I have
formerly described and figured for Pennatula phosphorea (Vid. Medd. Nath. For. Kbhvn. 1888). This is
also seen quite easily and convincingly, if we take into the comparison such Umbell/tla-species where
the rhachis-club is more elongated, e. g. U giintheri (partly U. leptocaulis, simplex, and carpenteri).
The other polyps, which arise inside (above) the lower circle , must then of necessity correspond to
those placed on the ventral surface of the rhachis in other sea-pens, which is bordered by the two
dorso-lateral rows of polyps; accordingly, they correspond to the polyps which in most sea-pens are
arranged in oblique rows. Now the appearance and arrangement of these polyps is irregular in
several sea-pens (as in Anthoptilum murrayi, the species of the genus Kophobelemnon a. o.); sometimes
not even a grouping on either side of the ventral median line is preserved. In Umbellula encrinus
the case is as in these latter; at all events, I have not been able to find any invariable regularity in
the appearance and arrangement of the upper or inner polyps. So much is evident, however, that
Umbellula like the typical Pennatulids has two regions of growth where new individuals arise: one
corresponding to that where, in forms with wings (as a Pennatula), new wings arise, and one corre-
sponding to that where the number of individuals in the wings increases. And as in all other bilateral
sea-pens those polvps are largest which form the two most dorsal longitudinal rows, so in Umbellula
the corresponding polyps, i. e. those of the lower circle, are the largest.

In large specimens of Umb. encrinus the polyp-cluster appears to be radiate; but a closer inspec-
tion of the tongues of zooids will show that the bilateral structure is preserved in reality. The fact
is that one of these zooid-areas is always a little larger than all the others, especially somewhat

*) This cannot be seen from the figures of Danielssen or be gathered from his text; in some of these specimens he
has overlooked the rudiments or not mentioned them.

8 4

PENNATULIDA.

broader; it is the dorsal belt of zooids, or, more correctly, the lower part of this belt; in several large
specimens it may easily and unmistakably, though as a narrow stripe, be traced between the npper
polyps to the base of a certain one, the original terminal polyp; in other specimens this tongue may
be traced to the end of the rhachis among the polyps, but which of these is the terminal polyp can-
not be determined with absolute certainty; not so much because this polyp has been atrophied, but
that it simply has been somewhat displaced during growth. The other zooid-areas are, as mentioned,
all somewhat smaller (narrower) than the dorsal one, but most conspicuously smaller are one, two,
or three areas on the opposite — ventral — side of the rhachis. In my specimen Nr. 5 the fifth zooid-
area to the left of the dorsal one is the smallest; in Nr. 6 the fourth and fifth are smallest, in Nr. 3
the fourth, fifth, and sixth; in Nr. 4 the sixth and seventh areas are quite narrow, and between their
upper end is a quite young rudiment (in form like a zooid, see below); in Nr. 8 the fifth and sixth
areas are also quite narrow, and between them is seen a quite young polyp; in Nr. 10 the sixth area
is divided above into two narrow points by a naked streak; at the upper end of this streak is found
an isolated large zooid (or a rudiment of a polyp).

These smallest zooid-areas must show the region where the latest appearing polyps of the
lower circle are found, without regard to whether these polyps may have reached the same size as
those placed nearest to the dorsal tongue or not. The place corresponds also to the zone in the
younger stages where new individuals of the lower circle arise; and in the specimens Nr. 4 and 8
new individuals are seen to arise here, as well as new, quite narrow zooid-areas; but in Nr. 4 the
circumstance is to be noted that the new individual is formed like a zooid, whilst rudiments of polyps
of the same size elsewhere — the zooid in question is ca. 7 mm high on the side turned outward, ca.
2 mm on t ] ie opposite side — already prove themselves to be polyps by being provided with eight arms.
The question then arises here; do not in Umb. encrinus, some of the later polyps begin as zooids, or
- what is essentially the same thing — cannot some zooids by degrees be turned into polyps? It has
been indicated above that, among the numerous tentacle-bearing zooids which look like those of the
tongue-areas, are found a larger or smaller number of much larger zooids, several more than i mra high, a
few still much larger up to 5 mm , on that part of the rhachis-club which is more or less hidden by the
polyps (they begin to appear already in young clusters, see above Nr. 2); at the first glance, one cannot
help regarding them as young rudiments of polyps; but at the same time, individuals of a similar size
or not much larger may be found, which are decidedly rudiments of polyps, as they have eight
distinct arms, and corresponding to this another form of the mouth-lips. I have only in one case seen
a zooid which seemed to me to be in the act of turning into a polyp; in the large specimen Nr. 3,
directly at the upper end of the rhachis-club, is an individual, 4 ,nm high, provided with the two
mouth-lips of the zooids (but unusually large and conspicuous), and carrying four tentacles, two very
long and with pinnulse, two quite short and without lateral branches; the ventral tentacle is the
longest and most developed; next to it comes the right ventro-lateral one, whilst the left ventro-
lateral and the left lateral are quite small. This case, together with the one mentioned above in Nr. 4,
where a gigantic zooid is found in the very place of a polyp, might tempt one to answer the question
put above in the affirmative; it ought not to be forgotten, however, that these two cases may be

PENNATULIDA.

85

exceptions, i. e. abnormalities 1 ), as it seems everywhere else in the octaetinia to be a law that the
eight arms of the polyps are formed at the same time.

It has already been stated above that Un/b. encrinus is not quite devoid of spicules, as has
always been maintained hitherto. Just as in the preceding species and in some others, the lower
end of the peduncle has indeed numerous, but very small spicules which cannot be seen with the
lens. These spicules (PI. Ill, fig. 51) are sometimes oval, 0.012— o.oi6 mm long, and o.oo8 mm broad,
sometimes somewhat dumb-bell-shaped, sometimes finally, small, and combined into numerous groups
of four measuring o.on mra . Some of the oval spicules show an indication of the common triangular
form of the Pennatulids. I have seen no spicules in other parts of the colony

With regard to the construction of the polyps I may mention that the tentacles are not so
long in proportion to the body of the polyp as in the preceding species. These organs, to be sure,
are exceedingly variable as to length, and their chance degree of contraction plays, of course, a very
great part; but in the preserved specimens they are upon the whole of about the same length as the
bodies of the polyps — sometimes somewhat longer, sometimes somewhat shorter, but never, for
instance, more than twice as long.

Of the sexual organs Danielssen 1. c. p. 51, states the following: The generative organs
develop themselves on the gastral filaments, a circumstance, that in a material degree differs from what
has previously been known relative to Pennatulidcr. where the sexual organs are developed on the septula*.
Later, on p. 52, it is said : All of the eight gastral filaments are occupied by sexual organs ; but, upon the
two dorsal ones, they begin a little lower down than on the other six». I cannot corroborate these state-
ments. The two dorsal septa do not bear sexual organs here, any more than in other octocoralla. These two
septa are as always narrower than the others, and their mesenteric filaments arising from the ectoderm are
of a simpler form than in the other septa: they form a finely sinuous line running from the base of the
stomodseum along the edge of the septum. They retain the same appearance through the whole gastric
cavity of the polyp-body, and into the continuation of this cavity in the rhachis; here they cease, but
their septa still continue some way without filaments. All the other 6 septa, on the other hand, carry
sexual organs; these septa have thick, much folded endodermal mesenteric filaments which begin at
the stomodseum, but reach only a little way down on their respective septum, when the}' are replaced by
the sexual organs, which accordingly take their place on the edge of the septum; the filaments as such
have dissappeared in this region. In a polyp whose body is 65™™ long (from Nr. 7, a g), the much
folded mesenteric filament occupies a space of 8 mm , then it is finely sinuous for quite a short space,
almost as in the two dorsal septa, and the sexual organs then succeed. The sexual organs remind
one of grapes, composed of small clusters, often of groups of three sexual organs (testicles or eggs),
and the middle one is the oldest and most developed. Accordingly, the sexual organs are here in
all essential features as in other Pennatulids. Marshall has in Umb. gracilis (= lindahlii) also found
the six septa carrying genital organs below the filaments (1. c. p. 144); when, however, in his figure
(PI. XXV, fig. 34) he makes the sexual organs occupy a relatively considerable part of the lateral sur-
face of the septum, this is hardly correct, and does not, at all events, agree with U. encrinus. In
U. thomsoni, Kolliker (Festschr. Phys. med. Ges. Wiirzb. 1875, p. 9) states that two pairs of filaments:),
1) We know from Penn. pliosphorea (and Virg. miraiilis) that zooids may turn into polyps, but only as an exception.

36 PENNATUUDA.

i. e. the lateral ones, carry the sexual organs, but he says also that he has not been able to make a
thorough examination on this point.

As to the construction of the zoo ids, most frequently I have been unable to see any ten-
tacle in any of the zooids of the stalk, except in those placed on the «sheath-formed dilations and on
the rhachis-club. When also many of the zooids in these latter regions want a tentacle, I take it to
have been lost; the other zooids of the stalk have, I think, also had such a tentacle. I cannot acknow-
ledge Danielssen's figure 56, I.e. PL X, to be correct; more especially, I must remark that the feature
given at e of <au oblong aperture in the slightly retracted tentacle (comp. the explanation of the
figures, 1. c. p. 82), is surely a misinterpretation; but several of the figured stages of retraction are
also hardly correct, at all events I have never been able to find them. The tentacles may often be
several times longer than the body of the zooid, and they may --as correctly stated by D. — have
two rows of piunulse, or one, or none at all.

With regard to its distribution, Umbellula encrinus is limited to the Arctic Ocean and to the
deeper part of the Atlantic which has the character of the Arctic sea by being situated west of the
coast-banks of the Scandinavian peninsula and north of the submarine ridges connecting Greenland
with Iceland, Iceland with the Faeroes, and these latter with the Shetland Islands. Originally (1753) it
was taken in the Arctic Ocean at 79 N. Lat., 80 miles from the coast of «Greenland> 1 ) (Ellis's and
Mylius's specimens); then it was found again in 1873 by the Austro-Hungarian Expedition (Maren-
zeller: Denkschr. Ak. Wien 1878, p. 377) near 79 N. Eat, 62 29' E.Long, at a depth of 210 metres, east
of Franz-Joseph Land (the specimen, 630 mm long, was lost with the ship); later by the Vega Expedi-
tion in the Kara Sea (the «Vega»'s Station 54) at 130 fathoms (Stuxberg: Vega-Exp. vetensk. Arb. I,
1882, p. 692, and V, 1887, p. 163) ; by the Norwegian North-Atlantic Expedition at four different places
between 79° 59' N. Lat, 5 40' E. Long., west of Spitzbergen, and 62 44' N. Lat, i c 48' E. Long., between
Norway and the Faeroes, in depths from 412 — 498 fathoms (a dead axis was further found at a fifth
place at a depth of 536 fathoms) all within the territory of the tcold area . To these localities are to
be added those of the dngolf : St. 105, (1 specimen, Nr. 2) 65 34' N. Lat, 7 31' W. Long., 762 fathoms,
about midway between the Faeroe Islands and Jan Mayen, and St. 116, 70 5' N. Lat, 8° 26' W.Long.,
371 fathoms, south of Jan Mayen. At the latter place the trawl of the < Ingolf « evidently passed over
a plantation of large specimens; one was caught by the crow-foot in front of the trawl, and was saved
entirely (Nr. 3) 2 ), of three the tops were cut off (Nrs. 5, 6, 7), and one was hanging on the outside of
the trawl-bag, but slipped into the sea again; at St. 126, 67 19' N. Lat, 15 52' W.Long., 293 fathoms,
north of Iceland, was further taken a dead piece of the calcareous axis, 26o mm long, of a large
specimen. Finally, Nathorst (Nat. Science, Nov. 1899, p. 319, and «Twa somrar etc*, 2, p. 93) has, in
1899, taken a large specimen, 2 metres, i2 ct,n long, close to Jan Mayen, and from the Danish East-
Greenland Expedition in the summer of 1900, Mag. Sc. Soren Jensen brought home three specimens
(Nrs. 1, 4, 8), a young stage from Cape Brewster, depth 250 fathoms, a large specimen and a torn off

1 ) i. e. either East-Greenland or Spitzbergen (corup. Lindahl 1. c. p. 3).

2 ) As there was, of course, no glass tube of such dimensions on board, one of our spare trawl-beams — an iron tube
15 feet long — was, b}- the excellent proposal of Captain Evers, used as a reservoir, filled with spirit and properly closed
at both ends.

PENNATULIDA. 87

cluster from Canning Land, 202 fathoms; the contemporary Swedish expedition got a mycket vackert»
specimen on the 14 th of Aug., in the mouth of the Franz-Joseph Fjord, at a depth of 200 — 300 metres
(Kolthoff, p. 176). On the cruise of the Michael Sars> in 1902, one entire specimen (Nr. 9) and a torn-
off cluster were taken at St. 34, north-east of the Fseroes, 62 53' N. Lat., 4 14' E. Long., 384 fathoms,
and in 1903 one entire, but broken specimen was taken by the Thor at St. 51, east of Iceland,
66° 2' N. Lat, n° 5' W. Long., at a depth of 900 — 1040 metres.

The localities enumerated show that this species, in contrast to most of the species of the
genus, does not seem to thrive in very greath depths, as the known depths reach from ca. 105
fathoms to 762 fathoms. It seems to be most luxuriantly developed and most numerous on the slopes
towards the cold depths, but in places and depths where the temperature is already below zero; whether
it is found, otherwise than exceptionally, in the really great depths must for the present be left
undecided; hitherto, we have only one instance of its being found there, viz. the specimen from
St. '105 of the Ingolf .

I doubt very much whether the species Uvib. encrinus would be found outside the territory
so well bounded geographically where it is really known; I am quite persuaded that the determination
is erroneous, when Studer mentions the species U. encrinus from the Pacific, o° 19' N. Lat, 90° 34'
W.Long., at a depth of 331 fathoms. («Albatross», Bull. Mus. Comp. Zool. vol. XXV, no. 5); Studer's
species is perhaps identical with Kolliker's U. magniflora. from the Southern Ocean, east of Kerguelen
Island which species I cannot, however (as Danielssen is inclined to do), regard as identical with
U. encrinus, even if it is closely allied to it. This species of Kolliker (Chall. Rep. p. 24) is said, among
other things, to want spicules entirely. That there can be no doubt of the correctness of this observa-
tion arises from the fact that Kolliker was the first who discovered that microscopically small spicules
may be found only in the lower end of the peduncle in several other U tube I hi la-species (and in Anf/iop-
tiluiu). Kiikenthal (Zool. Anz. 1902, p. 596) has described a new variety of U»ib. encrintis, a
var. antarctica, taken east of Bouvet-Island, at a depth of 450 metres. As it differs in several respects
from the northern genuine Umb. rue 7 -in us, I take it to be another species; perhaps also identical with
U. magniflora Koll. (which Kiikenthal however regards as another variety of U. encrinus).

General Review of the Occurrence and Distribution of the

Northern Pennatulids.

Of the 21 species which, according to the preceding account, constitute the whole Pennatulid-
fauna within the territory treated of here, the following six have not hitherto been taken in any
locality north of 49 N. Lat: Protoptiluni carpenteri, Anthoptilum grandiflorum y A nth. murrayt, Disti-
choptilum gracile, and the two new species Protoptiluni denticulatum and Pennatula prolifera.

To the fauna of Norway 13 species are to be referred, as the large number of 29 species

88 PENNATULIDA.

given') in Grieg's review (Berg. Mus. Aarb. 1891) must be reduced to the following: Pennatula
phosphorea, Penn. aculeata, Penn. grand is, Virgularia affinis, Virg. mirabilis, Virg. cladiscus, Stylatula
(Dubenia) elegans, Pavonaria finmarchica, Halipteris christii, Protoptilum thomsoni, Funiculina qua-
drangularis, Kophobelemnon stelliferum, Umbellula encrinus.

From the Fseroe Islands, Iceland, and Greenland no Pennatulid was hitherto known
(apart from Umbellula lindahli and the statement concerning Pavonaria finmarchica, see p. 39). It has
now been proved that several species are found in the depths of the sea near these countries, also a
few in more shallow water. From the banks at the Fseroes are now known Pennatula grandis
(numerous) and Halipteris christii. From more shallow water (less than 100 fathoms) at Iceland (the
Vestman Islands) are now known: Pennatula pliosphorca (var. Candida), Virgularia mirabilis, Virg.
cladiscus, and Stylatula (Dubenia) elegans. From the fjords of the Fseroes and Iceland — in contrast
to those of Norway - - sea-pens are now as little known as hitherto. This holds good also with
regard to the western fjords of Greenland. From the sea between Greenland and the American
polar islands only Umbellula lindahlii was previously known; this species has now been found again
farther south , in the Davis Straits, and to it are further to be added Pennatula aculeata, Penn.
prolifera, Authoptilum grandipZorum, Distichoptihim gracile, and Kophobelemnon stelliferum, so that for
the present six species of sea-pens are known as « West-Greenland: species. From East-Greenland
only one species, Umbellula encrinus, is known, and only from its northern part.

The fact that the species upon the whole divide into an Atlantic and an Arctic territory seems
to me, however, of greater interest than the reference of the species found to the fauna of the nearest
land. The boundary between these regions is very far from coinciding with the purely geographical
boundary of the Polar Circle between the Atlantic and the Arctic Oceans; it is contingent upon the
configuration of the bottom and the influence of this upon the temperature and currents of the sea.

I have previously (Geograf. Tidsskrift, vol. 14, 1896 — 98, p. 38, and Forhdl. 15 Skand. Natur-

forskaremote i Stockholm 1898, pp. 271 — 74) tried to show that the submarine ridges connecting

Greenland with Iceland, Iceland with the Fseroe Isles, and these with the Scottish Isles, form a

boundary between an Arctic and an i\tlantic deep-sea fauna; on the one hand, they prevent directly,

by their height, the wanderings of many forms, that is to say forms that may be supposed always

and in all ages to be confined to depths greater than 300 fathoms; on the other, they influence the

distribution of animals by producing a great climatic contrast in the sea-water and by deflecting the

currents. North of the ridges the temperature of the water from the bottom to about 300 fathoms

below the surface is constantly below zero, whilst south of the ridges it is always positive. The cold

northern basin forms part of a large and deep polar water which spreads farther north beyond the

large land-masses of the northern hemisphere, but in other places is also shut out from other ocean

depths by ridges or regions of more shallow water (Smith Sound etc., and the Behring Straits); the

deep, warm waters occurring south of the ridges pass evenly into the great depths of the true

Atlantic, only forming the northern part of this Ocean; west of Greenland it passes far north right

through the Davis Straits and even some way into the Baffin Sea, constantly with positive bottom

') This account contains 30 species; but Leplopiilum gracile K611. var. norvegica Dan. Kor. has already been abandoned
by <",rieg himself as a young stage of Funiculina quadrangularis. Comp. p. 50. The account also, at the time it appeared,
should have contained 31 «species», as Koren's & Danielssen's Cladisczis Kollikeri has been forgotten.

PENNATULIDA.

8 9

temperatures. The climatic influence of the submarine barriers reaches, however, to the upper layers
of water also; these are considerably colder on the north than on the south side. The influence on
the currents is briefly thus: the warm body of water of the Atlantic streams north-east towards
Iceland and the ridges; here it is turned off in such a way, that it bends towards the west at the
ridge of the Denmark Straits, and then streams south alongside the south-going East-Greenland
Polar Current, forming the principal part of the Irminger Current; here accordingly, it returns to
the Atlantic; at the ridge between Iceland and the Faeroes the warm bod}- of water is turned east-
ward and led towards Norway, where it streams northward. This water is only cooled far to the
north, then sinks to the bottom, and is led southward in the Arctic basin as cold water towards the
north side of the ridges without being able to pass these to any considerable degree.

From this it follows that the deep-sea animals which rise so much above the bottom as to live
in this Atlantic water (which ma}- be thought to apply among others to the larvae of bottom animals),
are partly retained in the Atlantic, and may partly wander towards Norway; here some of them may
take root, as the conditions to a depth of ca. 300 fathoms agree with those of corresponding depths
south of the ridges; and this holds good far north of the Polar Circle. Thus, the Atlantic fauna
reigns not only south of the mentioned boundary of the cold region, but also east of this region along
the whole Scandinavian peninsula. It is quite wrong to regard some deep-sea animals as arctic
forms, because they were first known from high northern latitudes at Norway; like several «southern .
animals from the same regions they belong really to a widely spread Atlantic fauna. A great number,
at least, of the animals found at the Scandinavian peninsula even to the North Cape, are also found
along the Atlantic side of southern Europe; some of them even pass into the Mediterranean; several
occur at the Azores, and still more on the east side of North America (at latitudes corresponding to
southern, and partly to Central Europe); and now the Ingolf Expedition has been able to show further
that some follow the warm layer of water far up on the west of Greenland.

When we now turn to the Peunatulids, we find that of the 21 species enumerated here, no
less than 20 have been found in the Atlantic territory; only one, Umbellula encrinus, is an exclusively
Arctic species. But among the species from the Atlantic region there are two which may occur
beyond the boundary towards the cold region, viz. Virgularia cladisciis and Kophobelemnon stettiferum.
With regard to the latter, however, it has only been found in single instances and only in the narrow
branch sent off from the cold deep waters towards the Wyville Thomson ridge in the Faeroe Channel;
this species has otherwise the widest possible distribution inside the «warm area-. Virgularia cladiscus,
certainly, has been found more frequently and in several places inside the cold area, but always near
to the boundary towards the warm area; both these species must assuredly be regarded as Atlantic
species endowed with more than common hardiness with regard to temperature.

The preponderance in richness of forms shown by the Atlantic territory in respect to Penua-
tulids, is a general feature; the cold territory of the depths of the North Atlantic is upon the whole
poor; but to be sure, the contrast is not so large in all groups as in this.

The common characteristic of the fauna of the Atlantic territory with regard to the northern
Pennatulids will be seen plainly from the table below.

The Ingolf-Expcditioii. V. I. 12

go

PENNATULIDA.

Pcnnatula phosphorea ... .

» aculeata

» prolifera

grandis

I Irgularia ajfinis

i> mirabilis

cladiscus

Stylatula (Diibenia) elegans
Pavonaria finmarchica . . .

Haliplcris christii

Protoptilum thomsoni

Protoptilum cavpeuteri ....

» denticulatunt .

Funiculina quadrangularis

Distichoptiltim gracile

Kophobelemnon stelliferum
Bathyptilum carpenteri ....
Anlhoptilum grandiflornm .

> murrayi

Umbellula lindahli

Davis Str.

East side of
N. America

+

+

+

+

+
+
+

+

South of
the ridges

Norway-

West of
Europe

The Medi-
terranean

The
Azores

+
+

+

+
+
+

+
+

+

+
+
+

+
+

+

+

+
+
+
+

+

+
+

-

+

+

+

+

+

+

A ? in a column means that there are indications which make it probable that the species in
question might be entered as found.

The table shows how great the agreement is between the most widely separated parts of the
North Atlantic territory, the west side of Norway and the east side of North America. Six of the
twelve species of Norway are known from America, and there is some probability that two more
may now be added. Species which previously had to be regarded as « American > (Anthoptilum murrayi,
Distichoptilum gracile) have proved to be distributed quite to Europe ; species which were hitherto only
known as far more southern ones (even from south of the equator), as Anthoptilum grandiflorum, are
found in the Davis Straits; of the other five species found there, four occur both at America and
Europe; but one (Penn. prolifera) is as yet only known from these Straits. Apart from the quite imperfectly
known Bathyptilum carpenteri (only taken in one mutilated fragment) and Protoptilum denticulatuiii,
no form peculiar to the northern part of the Atlantic south of the ridges has appeared; of the other
14 species from this region six are found both at Norway and America, four (for the present) at Nor-
way alone, and three at America alone; but of the last-mentioned species two also occur off the west
coast of southern Europe.

It seems likely that continued researches inside the Atlantic area will show that the uniformity
is still greater than I have been able to show here; so wide a distribution as is now known for
. Inthoptilum grandifloruiii, which has been found from off Buenos Ayres to far up in the Davis Straits,
further at the African coast off the Cape (and possibly near Tristan d'Acunha) will scarcely prove an
isolated phenomenon; several other Pennatulids of which we know that they may thrive in great

PENNATULIDA. 91

depths, will certainly be found distributed over the greater part of the widely-extended depths of the
Atlantic, as far as the positive bottom-temperatures are found.

With regard to the bathymetrical range, the species treated off here show the peculiar
feature that most of them may occur at very different depths; many of them have a surprisingly great
vertical range. In the following list the least and the greatest depth at which each species has
hitherto been found, is given as far as possible.

Fathoms

Pennatula phospkorea 5-10 — 114

» aculeala 30—1255

prolifera I:t 99 — x 435

grandis 50 — 1255

Virgularia affinis 60 — 100

mirabilis 5 — 200

» cladiscus 40 — 555

Stylatula (Diibenia) elegans 30 — 550

Pavonaria finmarchica 60—980

Halipteris christii c. 200

Protoptilum thomsoni 150 — 440

P'athoms

Protoptilum car pe uteri 690 — 1700 ')

denticulatum ID 95

Funiculina quadrangularis 10 — 1135

Distichoptilum gracile 700 — 1245 2 )

Kophobelemnon stelliferum. . . . 20 — over 2000 (2369)

Bathyptilum carpenteri 650

Antlioptiliiin grandiflorum 131 — 1106

uiurrayi 640 — 1362

Umbellula lindahli 122 — 1435 3 )

eucrinus c. 105 — 762

According to this list only four specimens would seem to be exclusively deep-sea forms: Pen-
natula prolifera, Anthoptilum uiurrayi, Distichoplilum gracile, and Protoptilum denticulatum; as only
one specimen of the latter has been taken, however, I think it better to make a reservation with
regard to it. Several of the species which are known to reach to very great depths, seem, however,
to have their natural home in somewhat higher regions: Pennatula aculeata occurs most frequently
between 150 and 300 fathoms; Penn. grandis, Funiculina quadrangularis, Halipteris christii, Pavonaria
finmarchica, Anthoptilum grandiflorum, Kophobelemnon stelliferum. Protoptilum thomsoni, Umbellula
encrinus, perhaps still more, seem to occur in greatest numbers and to be largest and most developed
in similar small and moderately great depths. With regard to these and several other sea-pens the spe-
cimens taken at very great depths are only young stages; even if they are provided with sexual
organs, sometimes even with ripe sexual products, their outer form has either not reached full
development, or may even be so little developed, that in several instances these specimens have been
interpreted as independent species or genera of simple structure. From this fact originates the view
advanced by Kolliker (Monogr. p. 449 and Chall. Report p. 39), that it is upon the whole the simple
forms of Pennatulids which live in the great depths, whilst the more ■; complicated or differentiated
forms belong to less deep waters ; and since the forms interpreted as primitive ones are as usual regarded
as the oldest, he has also thought he found the supposition corroborated in the distribution of the
Pennatulids, that the remainder of an otherwise extinct, ancient fauna was living in the great depths.
This latter supposition is upon the whole not very probable ; I shall not, however, treat of this question

1) Verrill states that he has found it at less depths, comp p. 54; but he gives no depth.

2) In the Pacific 885—1573 fathoms.

3) U. bairdi, which is presumably the same species, reaches to 2033 fathoms.

„ 2 PENNATULIDA.

here, but only point out that the Pennatulids cannot be used to support it. But how is the fact to
be explained that the specimens of many species taken at great depths are exclusively or most
frequently young stages?

The reason may possibly be that the severe conditions of life which are undoubtedly
found at great depths, check the growth, so that the few individuals which have emigrated from
better regions, become more or less dwarfed. But the reason may possibly be the more casual one
that hitherto the great depths have been fished so little and over so small an extent, and that the
apparatus used have been very defective. The Pennatulids found at the coast and in shallow water
and there regarded as « common , often occur much scattered, sometimes in strictly local and limited
plantations-. Similar features are undoubtedly also found in the great depths; it has been pointed out
by Kolliker (Chall. Rep. p. 36) that the Challenger » obtained not a single Pennatulid in wide regions
of the oceans; of the 118 stations, at which the Ingolf» used fishing gear on the bottom, only 17
have yielded Pennatulids; and large specimens of forms such as Penn. grandis, Funiciilina, Halipteris,
Protoptilum etc. are seldom if ever got in the dredge, rarely enough in a trawl of the types generally
used in deep-sea researches; generally, we have to thank the lines of fishermen or their large, wide-
spreading nets for the large and entire specimens which give us correct ideas of these forms; and it
may upon the whole be said that we have only a full representation of these forms from regions,
where fishing has been far more intensely carried on than in the great depths.

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Mediterraueo. Rendiconto dell' Acad. R. delle sc. fis.
e matemat. di Napoli. 1S71.

68. Prouho, H.: Observations sur la Goniactinia prolifera (Sars)

draguee dans la Mediterranee. Arch. Zool. exp. et gen.
(2) T. IX. p. 247. 1 89 1.

69. Richiardi, S.: Monografia della Famiglia dei Pennatularii.

Archivio per la Zoologia, l'Anatomia e la Fisiologia
Ser. II. Vol. 1. 1869.

70. Roule, L. : Coelenteres i : Kohler, R. : Resultats scientifiques

de la Carnpagne du «Caudan - dans le golfe de Gas-
cogne. 1896. Annales del' Universitede Lyon. T. XXVI.
1896.

71. Sars, M. : Beschreibung der Pennatula borealis. Fauna

littoralis Norvegiae. 1st part, p. 17. 1846.

72. — Beretning om en i Sommeren 1849 foretagen zoologisk

Rejse i Lofoten og Finmarken. Separate copy. p. 19. ( Virg.
finm.). NytMagazin for Naturvidenskabeu 1851. p. 139.

73. — Nye Polyper (Virgul. finmarchica). Fauna litt. Norvegise.

2nd part. p. 68. 1S56.

74. — OmSofjasrenesUdvikliug. Forhdl. 7. skand. Naturforsker-

mode i Kristiania 1856.

75. Schultze, F. E. : Coelenteraten in : Jahresber. der Kommission

zur wissensch. Uutersuchung deutscher Meere f. d. J.
1872-73. 1875.

76. Semper, C.: tiber Generations wechsel bei Steinkorallen.

Zeitschrift fur wissensch. Zool. 17 Bd. 1867.

77. Skyrsla um hi& islenzka naturufrsebisfelag arid 1897 — 98.

Reykjavik 1S9S. p. 12.

78. Stearns, R. E. C. : Description of a new Species of Alcyonoid

Polyp (Pavonaria Blakei). San Francisco «Mining
and Scientific Press*, Aug. 9. 1873.
- Remarks on a New Alcyonoid Polyp, from Burrard's
Inlet. Proc. Calif. Acad. Sc. Vol. 5, 1. pp. 7 — 12.

— Description of a new Genus and Species of Alcyonid
Polyp. (Verrillia Blakei). Proc. Calif. Acad. Sc. 1873.

— Verrillia Blakei or Halipteris Blakei. Amer. Naturalist.
Vol. 16. 1882. p. 55. (1SS1).

— Description of a new Genus and Species of Alcyonid

Polyp (Radicipes pleurocristatus n. g. n. sp.) etc. Pro-
ceedings U. S. Nat. Museum. Vol. 6. p. 96. 1883.

83. Studer, Th. : Ubersicht der Anthozoa Alcyonaria, welche
wahrend der Reise S. M. S. Gazelle um die Erde
gesammelt wurden. Monatsber. der Akad. Wissensch.
zu Berlin. 1878. pp. 632 — 75.

S4. — Versuch eines Systemes der Alcyonaria. Archiv f. Natur-
gesch. 1. 1SS7. pp. 1 — 74.

85. — and Wright, P.: Report on the Alcyonaria. Introduc-

tion, p. XXVII. Report Scient. Res. of the Voyage of
H. M. S. Challenger. Zoology Vol. 31. 18S9.

86. — Note prelim, sur les Alcyonaires provenant des cam-

pagnes du yacht l'Hirondelle (18S6— 88). Mem. Soc.
Zool. de France. Vol. 3. 1890.

87. — Note prelimin. sur les Alcyonaires. Report on the dred-

ging operations by the steamer Albatross. X. Bull.
Mus. Comp. Zool. Vol. 25. 1894.

79-

So.

81.

82.

PENNATULIDA.

95

88. Studer, Th.: Alcyonaires provenant des campagnes de

l'Hirondelle ( 1886— 88). Resultats des camp, scienti-
fiques ace. sur son yacht par Albert i Kr , Prince de
Monaco. Fasc. XX. 1901.

89. Stuxberg, A.: Evertebratfaunan i Sibiriens Ishaf, and:

FaunanpaochkringNovajaSemblja. In: Nordenskiold,
A. E.: Vega-Expeditionens vetensk. Iakttagelser. Bd. 1,
1882, p. 692, and Bd. 5, 1S87, p. 163.

90. Thomson, C. Wyville: The Depths of the Sea. 1873.

91. Verrill, A. E. : Notice of recent Additions to the Marine

Fauna of the eastern coast of North America. No. 2.
The American Journal of Sc. & Arts. (Silliman &
Dana). 3. Ser. Vol.16. 1S78. pp. 371 — 7S.

92. — Notice etc. No. 3. Ibid. Vol.17. 1S79. pp. 239— 43.

93. — Notice of recent Additions to the marine lnvertebrata,

of the northeastern Coast of America etc. Part. 1.
Proc. U. S. Nat. Mus. 1879. p. 165. (1880).
94- — Notice of the remarkable Marine Fauna occupying the
outer banks off the Southern Coast of New England.
No. 5. Am. Journ. Sc. VoL 23. 1S82. pp.309 — 16.

95. Verrill, A. E. : Reports on the Results of Dredging,

under the Supervision of Al. Agassiz, on the East
Coast of the Un. States, during the Summer of 1880,
by the U. S. Coast Survey Steamer Blake . Report
on the Anth'ozoa, and some additional Species
dredged by the « Blake ■> in 1877 — 79, and by the I*.
S. Fish Comm. Steam. Fish Hawk in 1880—82.
Bull. Mus. Comp. Zool. Vol.11. No. 1. 1883.

96. — Noticeetc. N0.9. Am. Journ. Sc. V0I.2S. 18S4. pp. 213 — 220.

97. — Results of the Explorations made by the Steamer

Albatross-* off the Northern Coast of the United
States, in 18S3. U. S. Commission of Fish and
Fisheries. Comm. Rep. 1883. 1885. pp. 504-601.

98. — Notice of the remarkable Marine Fauna etc. No. 11.

Am. Journ. Sc. Vol.29. T 8S5. pp. 149— 157.

99. Whiteaves, I. F. : Catalogue of the Marine lnvertebrata

of Eastern Canada. Geological Survey of Canada.
1901. pp. 34—35.

100. Wilson, E. B. : The development of Renilla. Phil. Trans.

Roy. Soc. 1883.

101. Worm, O. : Museum Wormianum s. hist. rer. rar. etc. 1655.

Plate I.

Fig.

3-

Fig.

4-

Fig.

5-

Plate I.

Fig. i. Pennatula aculeata Kor. Dan. from Davis Straits. The specimen, which is seen from the
ventral side, is of an unusual violet colour. The pen is highly contracted and somewhat
damaged. The specimen when just taken, was painted on board the Tngolf by the artist-
painter Sigurd Wandel; somewhat enlarged (See p. 12 No. 1).

Fig. 2. Protoptihim carpenteri K611. (the «Ingolf s specimen No. 3, p. 52); seen from the ventral side;
x 3. b younger polyps which on developing will make the pen two-rowed.
Id.: seen from the dorsal side; the same enlargement, n the naked streak.
Protoptilion thomsoni K6U. The top of a specimen taken by the « Michael Sars» (No. 10, p. 57),
seen from the left side; x 3; n the naked dorsal streak, t the top of the rhachis.
Id.: a piece from the middle of the rhachis seen from the left side; X3; from a specimen
taken in the Skager Rak (No. 11, p. 57), belonging to the Riksmuseum in Stockholm.

Fig. 6. Id.: the lower part of the rhachis, seen from the left side; x 3; from the same specimen
(No. n); // the dorsal, //' the ventral naked streak.

Fig. 7. Id.: a piece of the middle part of the rhachis, also seen from the left side, but turned in
such a way, that the ventral streak n' is seen, whilst the dorsal streak is hidden; V ventral
side, D dorsal side; x 3; of the polyps only the calyxes have been drawn so that the arrange-
ment of the polyps may be distinctly seen; the same specimen as in figs 5 and 6.

Fig. 8. Id.: the lower part of the rhachis, seen from the right side; x 3; 1, 2, 3, young polyps form-
ing together an oblique row rising towards the dorsal side Z>; 1, 1, 1 the oldest polyps
in each oblique row (comp. fig. 7). The lower part of the rhachis (together with the peduncle)
is twisted round the calcareous axis in a somewhat spiral-shaped manner. The same specimen
as in figs 5—7.

Fig. 9. Protoptihim denticulatum n. sp., seen from the dorsal side; x 2. P terminal polyp, Z terminal
zooid; a somewhat larger, b somewhat smaller and more ventrally placed polyps. * the place
where the lower bend of the inner calcareous axis is found.

Fig. 10. Id.: seen from the ventral side; x 2. The signification of the letters as in the preceding figure.

Fig. 11. Id.: the top seen from the left side; x 3. P terminal polyp, Z. terminal zooid, z dorsal zooid,
/ lateral polyp.

Fig. 12. Distichoptilujii gracile Verr., part of the rhachis seen from the dorsal side. The colour taken
from a coloured sketch made on board the Ingolf » by the artist-painter Sigurd Wandel. The
polyps retracted; 2 dorsal zooids; x 3.

Fig. 13. Id. : the upper end of the rhachis of another specimen (No. 6, p. 64) x 5 , to show the teeth
of the calyx-mouth, the (apparent) terminal zooid z, and the (likewise apparent) terminal polyp;
comp. p. 64.

Fig. 14. Id.: the upper end of a third, somewhat deviating specimen (No. 9, comp. p. 65); x 2; k the
upper, knob-shaped end of the rhachis.

Figs. 2— 14 drawn by Th. Bloch.

Ingolf Exp.editioiven \. I

Jungerseri: Peruiatnlida Tab. I.

k

13

■

■

*••

12

m

*

10

b - aP

mst

J

Proiojxlilum Fig. 2-11 Distichojitiltin i

Plate II.

Plate II.

Fig. 15. Pennatula prolifera n. sp., young specimen (comp. p. 18), seen from the ventral side; x 5. Z the

top-zooid, only partly visible, part of it being hidden between the two uppermost lateral

polyps. I — VIII the more developed pairs of wings; 1, 2, 3 the single polyps of these,

according to age; IX — XII the four rudimentary wings.
Fig. 16. Id.: from the dorsal side, x 5. ;/ zooid-less (« naked ••>) dorsal streak.
Fig. 17. Id.: a very young specimen, x 5, from the ventral side; there are two pairs of wings (I, II)

consisting of two individuals, 1 and 2; below these, two rudiments on each side.
Fig. 18. Id.: the same specimen as the preceding one, seen from the dorsal side. Z top zooid, z

dorsal zooid.
Fig. 19. Id.: a still younger specimen from the ventral side, x 5. 1, 2 the two individuals of wing I,

the only wing as yet present.
Fig. 20. Id.: the same specimen as the preceding one, seen from the dorsal side; Z and 2 as above.
Fig. 21. Id.: the pen of a young specimen with five pairs of wings, seen from the dorsal side, x 5.

The top is in process of being separated-off.
Fig. 22. Id.: the upper part of the pen of another specimen with the same number of wings, from

the dorsal side; also x 5; the upper end of the calcareous axis is seen in the stem (seep. 22);

the top is likewise in process of being separated-off.
Fig. 23. Id.: the top of the pen of a third specimen, from the dorsal side, x 6 — 7; only the upper-
most polyps (I) are drawn; the constriction is present, but the top part of the stem is not yet

prolonged to any considerable extent.
Fig. 24. Id.: the upper end of the pen of a specimen without an}- constriction at the top; from the

dorsal side, x6; in the place where other specimens are constricted, there is a large zooid Z

(comp. p. 23).
Fig. 25. Virgularia mirabilis (O. F. M.), a young ( Protocaulo)n>-) stage, 10,5™"' long (belonging to the

Riksmuseum in Stockholm); from the dorsal side, x ca. 8; all polyps, with the exception of

one, completely retracted; p rudimentary (i. e. atrophied) polyps; Iz lateral zooid, sz stalk-zooids.
Fig. 26. Id.: a somewhat older young stage, 22 mm long, from the ventral side, x ca. 6. At p 2 the

polyp in question has been cut away (comp. p. 27).
Fig. 27. Id.: a still older stage, 45 mm long, from the ventral side, x 4. / the uppermost rudimentary

wings; 5 wings on one side, 6 on the other, have two developed polyps; the succeeding wings

below these are three-polyped. a the region of the developed wings, b that of the rudimentary

wings, c that of the stalk-zooids.
Pig. 27 a shows a part of the specimen figured in 27, under higher magnifying power, so that the

three wing-polyps may be seen more distinctly; in the lowermost wing figured here, the

innermost (most ventral) polyp * is very small.
Fig. 28. Pavonaria finmarchica (M. Sars), young (iMicroptilum,-) stage; a small portion of the middle

part of the rhachis of the specimen, figured on PI. Ill, fig. 33, from the ventral side, x 5;

2 zooids.
Fig. 29. Id.: a small part of the rhachis, seen in profile from the right; of another specimen

(No. 2, p. 40).
Fig. 30. Halipteris christii (Kor. & Dan.) , young ( Lygomorpha*-) stage, ca. 73""" long, x 3; from the

ventral side; a young polyps recently developed. At * the soft parts have been rubbed away.
Fig. 31. Id.: a small part of the rhachis of another specimen, seen in profile from the right side, so

that it may be compared with fig. 29 of Pavonaria. The specimen is the smallest one of

Koren and Danielssen's two type-specimens of i.Lygomorplia Sarsii>, io6 nun long (see p. 46).
Fig. 32. Id.: the top of a full-grown specimen, 400 mm long, from the Faeroes, from the ventral side;

shows the different appearance of the calyx-points in different individuals.

Figs. 15 and 16 are drawn by C. Cordts, figs. 23, 24, 25 and 26 by the author, the others by Th. Bloch.

Ingolf Exfu ditionen I ' /

Jungcrscn Peniiatulida Tab. II.

Pennatula p.rolifera Virgularia mirabilis Pavonaria finmarchica

llillljlh I IS ill. U t t

Plate III.

Plate HI.

Fig. 33. Pavonaria fiumarcliica (M. Sars), young (i-Microptilum,-) stage, from the ventral side, x 2.

(The specimen No. 3, p. 40). The zooids are not represented in the figure; they can scarcely

be seen with this enlargement; the bodies looking like zooids are young polyps.
Fig. 34. Id.: the upper part of the pen of another specimen (No. 2, p. 40), from the ventral side, x 3.

a are quite young polyps, / the conical end of the stem.
Fig. 35. Id.: the lower part of the rhachis of the same specimen as in the preceding figure, a young

polyps; x 3.
Fig. 36. Id.: the lower part of the pen of the same specimen as figs. 34 and 35, seen in profile from

the left to show the Micr opt Hum-character; comp. the lower part of the pen of Kollikers figure,

Chall. Rep. PI. VII, Fig. 27 b, of Microptilum •willemoesii.
Fig. 37. Umbellula lindahlii K611., young specimen, seen from the left, natural size; from Davis Straits

(see p. 75).
Fig. 38. The same specimen; the primary polyp, x 8; most of the tentacles torn off; at k part of the

upper end of the calcareous axis shines through; p bud of a new polyp; z zooids.
Fig. 39. The same specimen; the lower part of the rhachis, with zooids z, and peduncle; x 8.
Fig. 40. Umbellula lindahlii, young specimen, natural size; the cluster seen so much from the dorsal

side, that P, the terminal or primary polyp, is distinctly seen.
Fig. 41. The same specimen, natural size; the cluster is seen from the ventral side.
Fig. 42. The same specimen, part of the cluster, x 3, seen from the ventral side to show the tentacle

of the zooids, and that the ventral tentacle v of the small polyp is larger than the others.
Fig. 43. The same specimen; the lowermost part of the stalk, x 6; shows zooids with fine tentacles;

some zooids have a tentacle with lateral branches (t).
Fig. 44. The same specimen; a group of spicules from the lower end of the peduncle; enlargement:

Zeiss, Obj. apochr. 8.o mm , Comp. ocular 4, length of tube i6o mm .
Fig. 45. The same specimen; spicules, under higher magnifying powers. Zeiss, Obj. apochr. 8.o mtn , Comp.

ocular 12.
Fig. 46. Umbellula encrinus (L.). The cluster of a young specimen (No. 1, p. 80), seen from the rudiment-
side (the ventral side), x 3; P the primary polyp; t tongue-shaped zooid-area.
Fig. 47. The same species; another specimen, young stage; natural size (No. 2, p. 80), from the right

side of the terminal polyp. P the terminal polyp; z large zooid; a rudiment of a polyp; 1, 2

the two polyps on the right side of the terminal polyp; dt the dorsal zooid-belt reaching up

to the base of the body of the primary polyp.
Fig. 48. The same specimen as in the preceding figure; part of the cluster, x 3; dt, P. 1, 2 as in the

preceding figure; t the three tongues of zooids on the left side.
Fig. 49. The same specimen as in figs. 47 — 48; part of the cluster, seen from the rudiment-region (the

ventral side), x 3. P, 1, 2, dt as above; a rudiments of four polyps; between these rudiments

stripes of zooids shoot up, the first rudiments of zooid-tongues.
Fig. 50. Umbellula encrinus; a large zooid from the inner part of the cluster of a fully formed, large

specimen, enlarged (of No. 7, p. 80); t tentacle with lateral branches; m mouth.
Fig. 51. Id.: spicules from the lower end of the peduncle (of the specimen No. 10, p. 80). Zeiss: Obj.

apochr. 8.o mm , Comp. ocular 12, length of tube i6o mm .

Figs. 37, 38, 44, 45, 50 and 51 drawn by the author, the others by Th. Bloch.

Ingoll /' i/ii ditionen . \. I

Jungersen : PenncUulida Tab. III.

■

>/

V

/'

/

w

\

n

- - «^?v

,56

• !

w

w

"

/

'

50

/

?

.Tv

-"'.' f'inniarchica ■ i Umbelliila lindahl Umbcllula encrinus

THE INGOLF-EXPEDITION

1895 — 1896.

THE LOCALITIES, DEPTHS, AND BOTTOMTEMPERATURES OF THE STATIONS.

Station
Nr.

Lat. N.

Long. W.

Depth

in
Danish
fathoms

Bottom-
temp.

Station
Xr.

Lat. N.

Long. W.

Depth

in
Danish
fathoms

Bottom-
temp.

Station
Nr.

Lat. X.

Long. W.

Depth

in
Danish
fathoms

" Bottom-
temp.

I

62 ° 30'

8° 21'

132

7°2

24

63° 06'

56° 00'

1199

2°4

45

61° 32-

9° 43'

643

4°i7

2

63° 04'

9 22'

262

5°3

25

63- 30'

54° 25'

5S2

3°3

46

61° 32'

11° 36'

720

2°40

3

63° 35'

10° 24'

272

o°5

63° 5i'

53° 03'

136

47

6i°32'

13° 4o'

95°

3°23

4

64° 07'

11° 12'

237

2°5

26

63° 57'

52° 41'

34

o°6

48

61° 32'

15°"'

1 150

3°'7

5

64 40'

12° 09'

155

64° 37'

54° 24'

109

49

62° 07'

15° "7'

1 1 20

2°9I

6

63° 43'

14° 34

90

7°o

27

64° 54'

55° lo-

393

3°S

SO

62° 43-

15° 07'

1 020

3°io

7

63° 13'

15° 41'

600

4°5

28

65° 14'

SS 42'

420

305

5i

64° 15'

14° 22'

68

7°32

S

63 56'

24 40'

! 3 6

6°o

29

65° 34'

54° 31'

6S

O 2

52

63° 57'

13° 32'

420

7 °8 7

9

64 iS'

27 00'

295

5°S

30

66° 50'

54° 28-

22

i°o5

53

63° 15'

15° 07'

795

3°o8

io

64 24'

2S 50'

788

3°5

31

66° 35'

55° 54'

ss

i°6

54

63° 08'

15° 4"'

69:

3°9

i 1

64° 34'

3i° 12'

1300

i°6

32

66° 35'

56° 38'

318

3°9

55

63° 33'

15° 02'

316

5°9

12

64° 38'

32° 37'

1040

o°3

33

67° 57'

55° 3°'

35

o°S

56

64° 1 N .'

15° 09'

68

7°57

13

&4° 47'

34° 33'

622

3°o

34

65° 17'

54° 17'

55

57

63° 37'

13° 02'

35"

3°4

14

64° 45'

35° 05'

176

4°4

35

65° 16'

55° 05'

362

3°6

5S

64° 25'

12° 09'

211

o°S

'5

66° 18'

25° 59'

33°

-o°75

36

6i°50'

56° 21'

1435

i°5

59

65°oo'

Il c I 6'

3'0

— o°i

16

65° 43'

26 58'

250

6°i

37

6o° 17'

54° 05'

1715

i°4

60

65° 09'

12° 27'

124

9

17

62 49'

26° 55'

745

3°4

3S

59° '2'

5i° 05'

1S70

i°3

61

65° 03'

I 3 ° 06-

55

o°4

iS

61° 44'

3°° 29'

"35

3°o

39

62° 00'

22° 3S'

S65

2=9

62

63° 18'

19° 12'

72

7°92

19

60° 29'

34° 14'

1566

2°4

40

62° 00'

21° 36'

S45

-,o„

63

62° 40'

19° OS'

Soo

4 °o

20

58° 20'

40 48'

i695

i°5

4i

61° 39'

I7 ; to'

1245

2°0

64

62° 06'

I9 OO'

1041

3°'

21

5 8°or'

44° 45'

1330

2°4

42

6i°4i'

io° 17'

625

o°4

65

6i°33'

19° OO'

1 089

3 °o

22

58 10'

48° 25'

1845

i°4

43

6i°42

IO° 11'

645

o°o5

66

61° 33'

20° 43'

1 128

3°3

23

60° 43'

56° 00

Only the

Pltinkton-N'et

used

44

61° 42'

545

4 : ^

67

61 ° 30'

22° 30'

475

3°o

Station
Xr.

Long. W.

Lat. N.

Depth

in
Danish
fathoms

Bottom-
temp.

.Station
Nr.

Lat. N.

Long. W.

Depth

in
Danish
fathoms

Bottom-
temp.

Station

Nr.

Lat. N.

Long. W.

Depth

in
Danish
fathoms

Bottom-
temp.

68

62 06'

22° 30'

S43

3°4

92

64° 44'

32° 52'

976

i°4

11S

68° 27

S°20 -

1060

— i°o

69

62° 40'

22° 17'

589

3°9

93

64° 24'

35° 14'

767

i°46

119

67° 53'

10° 19'

1010

— 1°0

7"

63° 09'

22° 05'

134

7°o

94

64° 56'

36° 19'

204

4° 1

120

67° 29'

n° 32'

ss 5

— 1°0

71

63 46'

22° 03'

46

65° 31'

30° 45'

213

121

66° 59

13° n'

529

— o°7

72

63 12'

23° 04'

! 97

6°7

95

65° 14'

30° 39

752

2°I

122

66° 42'

14° 44'

US

i°8

73

62° 58'

23° 28'

486

5°5

96

65° 24'

29° 00'

735

I°2

123

66° 52'

15° 40'

145

2°o

74

62 17'

24° 36'

695

4°2

97

65° 28'

27° 39'

45o

5°5

124

67° 40'

15° 4°'

495

— o°6

6i°57'

25° 35'

761

9S

65° 38'

26° 27'

138

5°9

125

68° oS'

16° 02'

729

— o°8

61° 2S'

25° 06'

829

99

66° 13

25° 53'

1S7

6°i

126

67° 19

15° 52'

293

-o°5

75

•6i°2S'

26° 25'

780

4°3

100

66° 23'

14° 02'

59

o° 4

127

66° 33'

20° Oj'

44

5°6

76

60° 50'

26° 50'

806

4°i

101

66° 23'

12° 05'

537

— o°7

12S

66° 50-

20° 02'

194

o°6

77

6o° 10'

26° 59'

95i

3°6

102

66° 23'

10° 26'

75o

— o°9

129

66° 35'

23° 47'

"7

6°5

78

60° 37'

27° 52'

799

4°5

103

66° 23'

8° 52'

579

— o°6

130

63° 00'

20° 40'

338

6°55

79

60° 52'

2S° 58'

653

4°4

104

66° 23'

7° 25'

957

— i°i

I3i

63° 00'

19° 09'

69S

4°7

So

6l°02'

29° 32'

935

4°o

i°5

65° 34'

7° 31'

762

— o°S

132

63° 00'

17° 04'

747

4°6

81

6i°44'

27° 00'

485

6°i

106

65° 34'

S°54'

447

— o°6

133

63° M

11° 24'

230

2°2

82

6i°55'

27 28'

S24

4° 1

65° 29'

8°4o'

466

134

62° 34'

io° 26'

299

4° i

S3

62 25'

2S 30'

912

3°5

107

65° 33'

10° 28'

492

-o°3

135

62° 4S'

9 ° 4 S'

270

o° 4

62° 36-

26°0l'

472

108

65° 30'

12° 00'

97

i°i

r36

63° 01'

9° 11'

256

4°8

62° 36'

25° 3°'

401

109

65° 29'

13° 25'

38

i°5

137

63° 14'

8° 31'

297

— o°6

84

62 58'

25° 24'

633

4°8

no

66° 44'

"° 33'

78i

— o°S

1 38

63° 26'

7°56'

471

— o°6

85

63° 21'

25° 21'

170

III

67° u

8° 48'

S60

— o°9

139

63° 36'

7° 30'

702

— o°6

S6

65° 03' 6

23° 47' 6

76

112

67° 57'

6° 44'

1267

— i°i

140

63° 29

6° 57'

780

-o° 9

87

6 5 02' 3

23 56' 2

no

"3

69° 3I '

7° 06'

1309

— i°o

141

63° 22'

6° 5 S'

679

-o°6

ss

64° 58'

24 25'

76

6° 9

114

70° 36'

7° 29'

773

— i°o

142

63° 07'

7° 05'

5S7

— o°6

89

64° 45'

27° 20'

310

S°4

"5

70° 50'

8° 29'

86

o°i

U3

62° 58'

7° 09'

38S

-o° 4

90

64° 45'

29° 06'

568

4°4

116

70° 05'

S°26'

37i

-o°4

144

62° 49

7° 12'

276

i°6

91

64 44'

31° 00'

1236

3°i

117

69° 13'

S°2 3 '

1003

— 1°0

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