Harriman Alaska Expedition (1899).

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phore is borne on a pedicel which resembles those of Garveta, having
a distinct expanded collar a short distance below the hydranth.

Distribution. Sitka Harbor and Yakutat, Alaska, abundant (Harri-
man Expedition) ; Shetland Island, Scotland (Allman).

This beautiful species bears considerable resemblance to Euden-
drium annulatum Allman, especially in its gonosome which Allman
describes as follows : " The gonophores are grouped in clusters, con-
sisting of from eight to twenty egg-shaped bodies attached around the
axis of gonoblastidea, which are of moderate length." 1

Were it not for a peculiar character of the trophosome, i. ., the ex-
panded pseudo-hydrotheca investing the body below the tentacles much
as in Garveia, there might be some suspicion that E. vaginatum and
E. annulatum are synonyms, particularly in view of the fact that both
were described from the Shetland Islands.

It also seems not improbable that Eudendrium pygmceum Clark *
may be another synonym of E. vaginatum, as Clark's description of
the gonosome agrees well with the gonosome described above. If
this be true, it is also likely that the dried stems described by Clark
from Santa Cruz, California, will be found to belong to this same


Trophosome. Hydranths large, with a basal whorl of filiform ten-
tacles and a distal set of closely crowded shorter filiform tentacles.

Gonosome. Reproductive elements developed in sessile medusae
borne in clusters just above the basal tentacles and producing actinules
instead of planulae.


The only genus included in the family in the sense here used.

(Plate xvi.)

Trophosome. Stem usually unbranched, attaining a height of i ^
inches, irregularly and sparingly annulated and increasing in size from
the proximal to the distal end, but more rapidly on the basal portion ;
a marked constriction some distance below the hydranth body ; stem
canaliculated between the constriction and the hydranth. Hydranth
with forty to fifty basal tentacles and about twenty in the distal set.

Gonosome. Gonophores borne in about twelve very long and
densely crowded racemes, which are supported by long, tentacle-like

'Ann. and Mag. Nat. Hist., 3d Series, Vol. xni, p. 83, Jan.. 1864.
2 Proc. Acad. Nat. Sciences, Philadelphia, p. 232, 1876.

[169] THE HYDROIDS 187

pedicels arising above the proximal row of tentacles. Gonophores (fe-
male) with three to five long tentacular processes which are sometimes
half the length of the gonophore. The actinule at birth is without a
distal row of tentacles, and the gonophore has no indication of radial

Distribution. Orca, Prince William Sound, Alaska (W. E.Ritter).

Exclusive of this species there have now been described four species
of Tubularia from the Pacific Coast of North America ; Parypha
microcephala (A. Agassiz), 1 which differs from the present species in
having flattened, instead of round, tentacular processes to the gono-
phores ; Tubularia elegans Clark, 2 which has mere tubercles instead
of the filiform processes to the gonophores; Tubularia borealis
Clark, 3 which differs in having laterally compressed processes to the
gonophores; and Tubularia larynx Ellis and Solander* (reported by
Gary S. Calkins), which differs from T. harrimani in the number of
both proximal and distal sets of tentacles, as well as in several other

The species seems to be abundant at Orca, as numerous specimens
were found.


Hydroida in which hydrothecae are developed for the protection of
the hydranths and gonangia for the protection of the gonophores.


Trophosome. Hydrothecae well developed, non-operculate, never
adnate nor immersed in the stem and always with a septum partially
dividing the hydrothecal cavity from the stem cavity. Hydranths
usually with conical proboscis and a single whorl of filiform tentacles.

Gonosome. Gonophores producing planulae or free medusae.

It would be hard to find two authorities who would agree as to the
genera of this exceedingly perplexing family. The arrangement here
adopted is substantially the same as that used by the writer in another
work now in press. 5 It is not offered as a final solution of the diffi-
culty, but as a convenience in discussing the group in the present

1 North American Acalephse, p. 195.
transactions Conn. Acad. Sci., Vol. in, p. 253, 1876.
3 Proc. Acad. Nat. Sci. Philadelphia, p. 231, 1876.
4 Proc. Boston Soc. Nat. Hist., Vol. xxvin, No. 13, p. 335.
6 Handbook of the Hydroids of the Woods Hole Region. To be published by
the U. S. Fish Commission.



Trophosome. Stem not regularly branched. Hydrothecae with
toothed margins, or with excessively thick walls and with long

Gonosome. Reproduction by means of free medusae.

(Plate xvn, figs, i, 2.)

Campanularia caliculata HINCKS, Annals and Mag. Nat. Hist., 2nd ser. , Vol.
xi, p. 178, March, 1853. VERRILL, Preliminary check-list of Marine
Invertebrates of Atlantic Coast, etc., p. 16, 1879. MARKTANNER-TURN-
ERETSCHER, Hydroiden von Ost-Spitzbergen, Zool. Jahrb., Vol. vm, p.
406. 1895. CALKINS, Some Hydroids from Puget Sound, Proc. Boston
Soc. Nat. Hist., Vol. xxvm, No. 13, p. 351, 1899.

Some authors, as Levinsen, regard this species as identical with
C. integra Macgillivray. The mode of reproduction is so different,
however, that the two would go into different genera in the classifica-
tion here adopted.

Distribution. Yakutat, Alaska (HarrimanExped.) ; British Coast
(Hincks) ; Spitsbergen (Marktanner-Turneretscher) ; New England
Coast (Verrill) ; Puget Sound (Calkins) .

(Plate xvn, figs. 3, 4.)

Campanularia compressa CLARK, Proc. Acad. Nat. Sciences, Philadelphia,

p. 214, 1876.
Eucopella Campanularia (VON LENDENFELD)? Uber Coelenteraten der Sudsee,

iv, Mitth. Zeitsch. Wiss. Zool., xxxvm, p. 497, 1883.

Distribution. Orca, Alaska (Harriman Exped.) ; Shumagin
Islands (Clark) . The figures given of this species well illustrate the
great variation in thickness of the hydrothecal walls. All the speci-
mens thus far discovered were found attached to Laminaria, over
which they creep in great profusion.

Von Lendenfeld makes his Eucopella campamilaria the subject of
one of his masterly monographic papers and it appears to agree in
every particular with the species under discussion. If I am correct in
supposing the two species identical, the name Eucopella companularia
will become a synonym and a very exceptional distribution will be re-
corded for Clytia compressa. It is interesting to note that von
Lendenf eld's species was also found growing on Laminaria.

The present writer does not agree with Calkins in his suggestion
that C. compressa is a synonym of C. caliculata.



Trophosome. Colony unbranched or regularly branched ; stem
simple or fascicled ; hydrothecae campanulate, never completely ses-
sile nor with operculum.

Gonosome. Gonophores producing planulaa without the interven-
tion of medusae.


Sertularia verticillata LINN., Syst. Nat., loth ed., p. 811, 1758.
Campamilaria verticillata SARS, Bidrag til Kundskaben om Norges Hydroider,

p. 46, 1873. VERRILL, Preliminary check-list of Marine Invertebrates

of Atlantic Coast, p. 16, 1879.

Distribution. Kadiak, Alaska (Harriman Exped.) ; North Cape,
Norway (Sars) ; New England Coast (Verrill).


Campanularia denticulata CLARK, Proc. Acad. Nat. Sciences, Philadelphia,
p. 213, 1876.

Distribution. Orca, Alaska (Harriman Exped.); Port Etches,
Alaska (Clark).


Campanularia lineata NUTTING, Hydroida from Alaska and Puget Sound,
Proc. U. S. Nat. Mus., Vol. xxi, p. 744, 1899.

Distribution. Berg Inlet, Glacier Bay, Alaska (Harriman Ex-
ped.) ; Puget Sound (Nutting).

(Plate xvn, fig. 5.)

Trophosome. Colony usually consisting of unbranched pedicels
growing directly from a creeping rootstock. Pedicels long and
slender, usually with a single distinct annulation at the distal end just
below the hydrotheca and about three less distinct ones at the proximal
end, the middle portion not being annulated. Hydrothecae cylindrical,
large, delicate in structure and with a perfectly even rim.

Gonosome. Unknown.

Distribution. Juneau, Alaska, 20 fathoms. Collected by Prof.
Wm. E. Ritter to whose efforts the fine series of Hydroids here dis-
cussed is so largely due, and for whom this species is named.


(Plate xvni, fig. 3, Plate xix, fig. 3.)

Campanularia speciosa CLARK, Proc. Acad. Nat. Sciences, Philadelphia, p.
214, 1876. LEVINSEN, Meduser, Ctenophorer og Hydroider fra Gron-
lands Vestkyst, p. 25, 1893.


Campanularia crenata ALLMAN, Diagnoses of new Genera and Species of
Hydroida, Linnaean Society Journal, Zoology, Vol. xi, p. 258, 1876.

Distribution. Orca, Alaska (Harriman Exped.) ; Yukon Harbor,
Big Koniuji, Shumagin Islands, Alaska (Clark) ; Japan Coast (All-
man) ; Greenland (Levinsen) .


(Plate xvin, fig. 2.)

Campanularia urceolata CLARK, Proc. Acad. Nat. Sciences, Philadelphia,
p. 215, 1876.

Distribution. Yakutat Bay, Alaska (Harriman Exped.) ; Lituya
Bay, Alaska (Clark) .

The specimens collected by the Harriman Expedition were growing
profusely over the stems and branches of Thuiaria costata in com-
pany with another parasitic species.

(Plate xvin, fig. i.)

Trophosome. Colony consisting of unbranched stems or pedicels
springing from a creeping rootstock. Pedicels one to three times as
long as the hydrothecae, and strongly annulated throughout. Hydro-
thecae deeply campanulate, thick-walled ; margins armed with twelve
to fourteen rather pointed teeth, and reduplicated once or twice, giving
a striking and unusually ornate appearance.

Gonosome. Gonangia roughly ovate, irregular in outline, with a
short neck, small terminal aperture and a very short pedicel. They
were empty in the specimens examined, so that it was impossible to
ascertain whether they produced planulas or medusas.

Distribution. Yakutat, Alaska (Harriman Exped.). All the
specimens were found growing in a parasitic manner over colonies of
Thuiaria costata^ in company with Campanularia urceolata. The
two species were often so intimately interwoven as they crept over the
stems and branches of the sertularian that I, at first, thought them
dimorphic forms of one species. However, in all cases careful dissection
showed that they were entirely separate colonies. The reduplication
of the hydrothecal margins seems to be a constant feature and adds
peculiar beauty to this striking little campanularian.


(Plate xix, figs, i, 2.)

Trophosome. Colony consisting of a creeping rootstock without
annulations, giving forth strong pedicels that are sometimes longer

[173] THE HYDROIDS 191

than the hydrothecae and sometimes considerably shorter, without a
definite swelling below the hydrothecae. Hydrothecae immense, in
one case nearly -fa of an inch in height, long, tubular-urceolate, ex-
panded distally, with slightly everted and broadly sinuous margin.
The margin is reduplicated in one specimen. Hydranth with about
twenty tentacles.

Gonosome. Not known.

Distribution. Orca, Prince William Sound, Alaska (W. R. Coe,
Harriman Exped.).

This species is closely allied to C. grandts Allman, 1 and may be
identical with it although it differs from Allman's description in the
character of the pedicels which he describes as having distinct node-
like enlargements immediately below the hydrothecae. The hydro-
thecae are larger than those of any other campanularian known to me.
But one small colony was found and it was creeping over another


Trophosome. Colony regularly branching; stem simple or com-
pound. Hydrothecae campanulate, thin, never with greatly thickened

Gonosome. Reproduction by means of medusae which are disk-
shaped, with four radial canals, more than eight marginal tentacles,
eight interradial lithocysts and a short manubrium without mouth ten-


Sertularia dichotoma LINN., Systema Naturae, Ed. x, p. 812. 1758.

Obelia dichotoma SCHULZE, Nordsee Expedition, Hydroida, p. 129, 1872.
VERRILL, Preliminary check-list Marine Invertebrates of Atlantic coast,
p. 16, 1879. CALKINS, Some Hydroids from Puget Sound, Proc. Boston
Soc. Nat. Hist., Vol. xxvm, No. 13, p. 256, 1899.

Distribution. Sitka, Berg Inlet, and Orca, Alaska (Harriman
Exped.) ; British Coast (Hincks) ; Helgoland (Schulze) ; Puget
Sound (Calkins).


Obelia plicata HINCKS, British Hydroid Zoophytes, p. 159, 1868. NUTTING,
Hydroida from Alaska and Puget Sound, Proc. U. S. Nat. Mus., Vol.
xxi, No. 13, p. 741, 1899.

Distribution. Orca, Alaska (Harriman Exped.) ; Puget Sound
(Nutting) ; British Coasts (Hincks) .

1 Diagnoses of new Genera and Species of Hydroida ; Linnaean Soc. Jour.
Zoology, Vol. xi, p. 259.

192 NUTTING [ J 74]

(Plate xix, figs. 4, 6.)

Trophosome. Colony sometimes attaining a height of eighteen
inches, but usually much shorter; stem not truly fascicled, although
several stems may be interwoven, exceedingly long and slender, sinu-
ous, giving off lateral branches in pairs on proximal portion and more
often singly on distal portion ; branches with a strong tendency to ver-
ticillate arrangement, forming oblique angles with the stem and
divided into numerous branchlets in a flabellate manner. Pedicels
short and completely annulated, or long and annulated only at ends,
set on broad shoulders of the stem. Hydrothecae funnel-shaped, the
sides usually straight, aperture with an even rim. Hydranths not well
preserved in specimens examined.

Gonosome. Gonangia borne in axils of branches and branchlets,
oblong-ovate, truncated above, having a collar in mature specimens ;
aperture apparently very large, pedicels strongly annulated. The
gonangia of the specimens examined were filled with developing
medusae of the regular Obelia type.

Distribution. Yakutat, Alaska (Harriman Exped.).

This fine species is related to O. Jlabellata, but the hydrothecae are
much deeper than in O. jlabellata, in which they are sub-triangular in
outline. It also bears some resemblance to O. commissuralis, which,
however, is a much more delicate species, with smaller and more
campanulate hydrothecae.

(Plate xx, fig. i.)

Trophosome. Colony attaining a height of about ^ inch; stem
sparingly branched, the main stem and larger branches sinuous or
slightly geniculate, giving forth pedicels singly or in opposite pairs at
the bends. Pedicels rather long and annulated throughout, the stem
also more extensively annulated than in most species of the genus.
Hydrothecae very large, deep, tubular, with very shallow undulations
around the margin, from between which lines run down for a short
distance on the surface of the hydrothecae.

Gonosome. Unknown .

Distribution. Orca, Alaska (Harriman Exped.).

This species bears some resemblance to O. bidentata Clark, found
on the New England Coast, but differs in the nature of the hydrothe-
cal teeth which are mere sinuosities, instead of being mucronate with
two denticles each as in the latter species.

[175] THE HYDROIDS 193


Trophosome. Pedicels arising from a creeping rootstock, very
short. Hydrothecae tubular, with entire margins, without opercula,
and having their cavities separated from those of the stems by a partial
septum. Hydranths with a conical proboscis.

This genus was originally instituted by Allman. 1 As here defined
it includes several species heretofore included in the genus Lafcea.


Lafcea pocillum HINCKS, British Hydroid Zoophytes, p. 204, 1868. CLARK,
Proc. Acad. Nat. Sciences, Philadelphia, p. 215, 1876. VERRILL, Pre-
liminary check-list Marine Vertebrates of Atlantic coast, p. 17, 1879.
BERGH, Goplepolyper (Hydroider) fra Kara-Havet, p. 333, 1887.

Distribution. Kadiak, Alaska (Harriman Exped.) ; Nunivak
Island, Alaska (Clark) ; Kara Sea (Bergh) ; British Coasts (Hincks) ;
New England Coast (Verrill) .


Trophosome. Much as in Obelia.

Gonosome. Planulae produced in sessile medusaform gonophores
which remain attached to the top of the gonangia until the spermatozoa
or planula3 are discharged.

(Plate xx, figs. 2-4.)

Trophosome. Colony attaining a height of about two inches and
consisting of a main stem which almost immediately breaks up into a
number of very slender, erect, almost straight branches which are
ornamented with about three annulations immediately above the origins
of the pedicels. Pedicels alternate, erect, much broader below than
above and with seven to ten annulations. Hydrothecae funnel-shaped,
with entire margins.

Gonosome. Gonangia borne in the axils of the pedicels, slender,
obconic, with a tendency to annulation. Each gonangium contains a
single sporosac which when mature rests upon the summit of the go-
nangium and has little indication of radial canals or tentacles.

Distribution. Yakutat Bay, Alaska (Harriman Exped.).

This species differs from others of the genus in having but one
sporosac to each gonangium and in the obliteration of most of the
medusoid characters of the sporosac. The entire margin of the hydro-
theca is also an exceptional character.

1 Allman, Challenger Report, Hydroida, Second Part, p. 29, 1888.



Trophosome. Colonies branched or unbranched. Hydrothecae
borne on pedicels, tubular, ending in an operculum composed of
several converging segments or triangular flaps. Hydranth with a
conical proboscis.

Gonosome. Gonangia producing free medusae or planulae.


Trophosome. Hydrothecae thin-walled, the upper portion cleft so
as to produce very long and slender teeth which form an operculum by
the convergence of their free ends.

Gonosome. Gonangia producing bell-shaped medusae, with four
radial canals, two or four marginal tentacles, and eight lithocysts.

(Plate xxn, figs, i, 2.)

Trophosome. Colony attaining a height of about YI inch. Stem
irregularly branched ; branches tending to an alternate arrangement,
straggling, geniculate ; stem and branches strongly and regularly annu-
lated throughout. Pedicels very short, with three to six annulations.
Hydrothecae ovoid-oblong, the distal third being composed of the oper-
culum consisting of ten or twelve segments. The hydranths have about
sixteen tentacles.

Gonosome. Gonangia in axils of the pedicels and branches, some-
times aggregated on certain branches to the exclusion of hydrothecae.
They are oblong-ovoid in shape and somewhat flattened on their distal
ends. Each gonangium contains a single medusa when mature.

Distribution. Juneau, Alaska (Harriman Exped.). The speci-
mens were found growing on Obelia.


Trophosome. Stem a creeping rootstock sending forth short an-
nulated pedicels. Hydrothecae tubular, thick-walled, with opercula
that are distinct from the hydrothecal teeth, and composed of several tri-
angular segments.

Gonosome. Gonangia borne on the rootstock, and, when mature,
bearing acrocysts.


Sertularia syringa LINN., Systema Naturae, Ed. xn, Tom. I, Pars n, p,
1311, 1767.

[177] THE HYDROIDS 195

Caly cella syringa CLARK, Proc. Acad. Nat. Sciences, Philadelphia, p. 210,
1876. VERRILL, Preliminary Check-list Marine Invertebrata Atlantic
Coast, p. 17, 1879. BERGH, Goplepolyper (Hydroider) fra Kara-Havet,
p. 335, 1887. NUTTING, Hydroida from Alaska and Puget Sound, Proc.
U. S. Nat. Mus., Vol. xxi, p. 741, 1899.

Distribution. Berg Inlet and Kadiak, Alaska (Harriman Exped.) ;
Coal Harbor, Shumagin Islands, Alaska (Clark) ; Kara Sea (Bergh) ;
British Coasts (Hincks) ; New England Coast (Verrill) ; Puget
Sound (Nutting).


Trophosome. Hydrothecae tubular, margins without teeth or oper-
cula, the hydrothecal cavity not divided from the stem cavity by a par-
tial septum.

Gonosome. Gonangia forming a ' Coppinia ' mass.


Trophosome. Colony with a fascicled stem, and with hydrothecae
either free or partially immersed in the stem, the distal portion not be-
ing abruptly turned upward.

Gonosome. A ' Coppinia ' mass.


Sertularia dumosa FLEMING, Edinburgh Phil. Jour., n, p. 83, 1828.

Lafoea dumosa SARS, Bidrag til Kundskaben om Norges Hydroider, p. 45,
1873. CLARK, Proc. Acad. Nat. Sciences, Philadelphia, p. 210, 1876.
VERRILL, Preliminary Check-list Marine Invertebrates of Atlantic Coast,
p. 17, 1879. NUTTING, Hydroida from Alaska and Puget Sound, Proc.
U. S. Nat. Mus., Vol. xxi, p. 741, 1899.

Distribution. Dutch Harbor, Unalaska (Harriman Exped.) ; Port
Etches, Alaska (Clark) ; North Cape, Norway (Sars) ; British Coast
(Hincks) ; New England Coast (Verrill) ; Puget Sound (Nutting) ;
California Coast (Clark).


Campanularia gracillima ALDER, Catalogue Zoophytes of Northumberland
and Durham, Trans. Tyneside Naturalists' Field Club, p. 39, 1857.

Lafoea gracillima CLARK, Proc. Acad. Nat. Sciences, Philadelphia, p. 216,
1876. VERRILL, Preliminary Check-list Marine Invertebrates of Atlantic
Coast, p. 17, 1879. MARKTANNER-TURNERETSCHER, Hydroiden von
Ost-Spitzbergen, Zoolog. Jahrbuch., Vol. vin, p. 410, 1895. NUTTING,
Hydroida from Alaska and Puget Sound, Proc. U. S. Nat. Mus., Vol. xxi,
p. 741, 1899.

Distribution. Juneau, Berg Inlet and Orca, Alaska (Harriman
Exped.) ; Shumagin Islands, Alaska (Clark) ; British Coast (Alder) ;


Spitzbergen (Marktanner-Turneretscher) ; New England Coast (Ver-
rill) ; Puget Sound (Nutting).


Lafcea fruticosa M. SARS, Bemaerkninger over 4 norske Hydroider Vid. Selsk.
Forh., 1862. G. O. SARS, Bidragtil Kundskaben om Norges Hydroider,
p. 26, 1873. CLARK, Proc. Acad. Nat. Sciences, Phila., p. 216, 1876.

Distribution. Juneau, Berg Inlet and Kadiak, Alaska (Harriman
Exped.) ; Kiska Harbor, Shumagin Islands, Alaska (Clark) ; British
Coasts (Hincks) ; Lofoten, Norway (G. O. Sars) ; New England
Coast (Verrill) ; Puget Sound (Nutting, MSS.).

(Plate xxi, figs. 3, 4.)

Trophosome. Colony forming an encrusting mass of adherent root-
stocks disposed both longitudinally and transversely over colonies of
other Hydroids, the tubes of the rootstock interwoven much like the
threads of a fabric. Hydrothecae sessile, tubular, often more or less
curved, aperture facing upward, entire; margin slightly expanded.
The hydrothecae are very irregularly disposed, being much more
crowded in some places than in others.

Gonosome. The ' Coppinia ' mass is much like that of Lafcea du-
mosa, being composed of closely packed gonangia interspersed with
long, tubular, variously curved modified hydrothecae. The gonangia
are flask-shaped, with a tubular neck and small aperture. Each
gonangium apparently contains a single ovum.

Distribution. Kadiak Harbor, Alaska. Growing over stems of
Thuiaria turgida (Harriman Exped.).

This interesting species is so different in appearance from the others of
the genus that I was at first inclined to make it the type of a new genus.


Trophosome. Stem fascicled, composed of an axial tube from
which the hydrothecae spring and to which they are partly adnate,
completely enclosed by a definite number of peripheral nonhydro-
thecate tubes.

Gonosome. A * Coppinia ' mass.


(Plate xxi, figs. 5, 6.)

Trophosome. Stem rigid, erect, giving forth scattered, stiff and
straight, alternate branches forming nearly a right angle with the stem.


Height of a fragmentary specimen about ^ inch. Stem and branches
sharply constricted proximally, composed of an axial tube which gives
off the eight or nine series of hydrothecae, and a number of peripheral
tubes enclosing the axial tube completely, thus burying all the hydro-
thecae nearly to their distal ends. Hydrothecae arranged in about eight
or nine longitudinal series, forming spirals. The distal ends of the
hydrothecae are abruptly bent outward, so that the round, even aper-
ture is vertical. When the peripheral tubes are removed the hydro-
thecae are seen to be long, tubular, doubly curved, narrowing prox-
imally, but without true pedicels, and all springing from the axial tube.

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