Harriman Alaska Expedition (1899).

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been made. EDITOR.



d74)




HYDROIDS OF THE EXPEDITION



BY C. C. NUTTING

CONTENTS

Introduction 175

Geographic distribution 176

Systematic discussion 181

INTRODUCTION.

The collection of Hydroida secured by the Harriman Ex-
pedition is of exceptional interest, and proves to be one of the
most important and most extensive collections of these beautiful
forms of marine life thus far made in Alaskan waters. Our pre-
vious knowledge of the Hydroid fauna of this region rested almost
exclusively on the collection made by Dr. W. H. Dall and his
associates during the years 1871-1874 and reported on by Dr. S.
F. Clark. 1 The number of species listed in Clark's report is 41,
in which was included Coppinia arcta^ now known to be merely
the gonosome of Lafosa. Of these 40 species, 15 are well known
British forms, and only one was then known from the Atlantic
coast of the United States. The remaining 24 species were new.

1 Report on the Hydroids collected on the coast of Alaska and the Aleutian
Islands by W. H. Dall, U. S. Coast Survey, and party, from 1871 to 1874,
inclusive. Proc. Acad. Nat. Sciences, Philadelphia, 1876.

[157] (175)



176 NUTTING [ I 58]

In 1878 C. Mereschkowsky l added a single species to the
Alaskan fauna, bringing the total up to 41.

No other additions were made until 1899, when I added eight,
of which three were well known British species and five were
new. 2 That made a total of 49 species reported prior to the
Harriman Expedition.

The collection here treated of comprises 53 species, 24 of
which had been previously reported. Of the remainder, 9 were
previously recorded from other localities, and 20 are new.
Thus the Harriman Expedition has added about 60 percent to
the number of species hitherto known from Alaskan waters.
More than half of the species secured are new to Alaska and
nearly 40 percent are new to science.

The whole number of species of hydroids now known from
Alaska is 78. Considering the small amount of collecting that
has been done in that region, compared with the extensive ex-
plorations of the coasts of Europe and the Atlantic coast of the
United States, one may confidently expect that the waters of
the far Northwest will prove to be very rich in hydroid life.

GEOGRAPHIC DISTRIBUTION.

A table is here given to indicate, first, the localities at which
each species was collected by the Harriman Expedition, and
second, the extent to which Hydroids have been distributed south-
ward along meridional lines from what appears to have been
a polar center of distribution. No attempt has been made to
represent the complete distribution of the species.

A glance at the part of the table showing the distribution as
represented in the collection secured by the expedition, shows
an apparent poverty of the Hydroid fauna of the western, as
compared with the eastern, portion of the territory explored.
For convenience in such comparison the stations are arranged
consecutively from east to west. The largest series were ob-
tained at Berg Inlet in Glacier Bay ; Yakutat Bay ; and at Orca
in Prince William Sound. These localities are all in deep bays,
sheltered from storms and surrounded by rocky shores. On

1 New Hydroida from Ochotsk, Kamtschatka, and other parts of the North
Pacific Ocean. Annals and Mag. Nat. Hist., Dec., 1878.

2 Hydroida from Alaska and Puget Sound. Proc. U. S. Nat. Museum, Vol.
xxi. (No. 1171.)



GEOGRAPHIC DISTRIBUTION OF THE HYDROIDS COLLECTED BY
THE HARRIMAN EXPEDITION.



Name. 1


Distribution of Specimens in the Harri-
man Collection.


General Distribution.




1

a

3


3

in


1

bo

E


Yakutat.


CO

a


Kadiak.


Popof Island.


a

I


Dutch Harbor.


I

(2


Arctic Regions.


East Coast U.S.


Pacific Coast
South of Alaska.


Alaska only.


Syncoryne eximia.
*Coryne brachiata.
*Garveia annulata.
Garveia nutans.
Eudendrium vaginatum.
*Tubularia harrimani.
^Campanularia ritteri.
Campanularia denticulata.
Campanularia verticillata.
Campanularia lineata.
Campanularia speciosa.
Campanularia urceolata.
*Campanulariareduplicata.
*Campanularia regia.
Clytia caliculata.
Clytia compressa.
Obelia plicata.
Obelia dichotoma.
*Obelia borealis.
*Obelia dubia.
Hebella pocillum.
*Gonothyra:a inornata.
*Campanulina rugosa.
Calycella syringa.
Lafoea dumosa.
Lafoea gracillima.
Lafoea fruticosa.
*Lafcea adhaerens.
*Grammaria immersa.
Filellum serpens.
Halecium halecinum.
Halecium muricatum.
Halecium scutum.
*Halecium reversum.
*Halecium robustum.
*Halecium ornatum.
*Halecium speciosum.
Sertularella tricuspidata.
Sertularella polyzonias.
*Sertularella saccata.
Thuiaria argentea.
Thuiaria similis.
Thuiaria variabilis.
Thuiaria cupressoides.
*Thuiaria coei.
Thuiaria fabricii.
Thuiaria turgida.
Thuiaria gigantea.
Thuiaria thuiarioides.
*Thuiaria elegans.
*Thuiaria costata.
Plumularia lagenifera.
Plumularia palmeri.
Totals.


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1 Species marked by a * are new.



178 NUTTING

account of the presence of perpetual ice in the form of glacier
fronts and bergs, the water must be very cold the year around.
Such a combination of conditions is particularly favorable to
Hydroid life and accounts for the remarkably rich collections
made at these places and also for the presence of so many arc-
tic species. In this connection it is interesting to note the fol-
lowing paragraph written about twenty five years ago by Dr.
Dall : " The material derived from the northwestern coasts of
America, from Cook's Inlet south and east, indicates a series of
Arctic colonies in favored localities, the future exploration of
which offers a labor of the highest interest. These colonies are
situated where the depth of water, the drippings of glaciers, and
the high and adjacent shores of the Great Archipelago combine
to reduce the temperature of the water below its apparently
normal isotherm. Cook's Inlet affords one of them, one exists
in the Gulf of Georgia, and others only await further explora-
tion." 1 It should be noted, however, that nearly all of the
Arctic species are well known forms belonging to the ' Holarctic
Province ' of authors, and that these species are of practically
continuous distribution on all coasts in northern regions so far
as explored.

In the same paper, Dr. Dall divides the coasts of America
from Monterey, California, north and west, into three faunal
areas, as follows : (a) the Oregonian, extending from Monterey
to the Shumagin Islands ; () the Aleutian, extending from the
Shumagin Islands to the end of the Aleutian chain, and north-
ward to the winter line of floating ice in Bering Sea ; (c) the
Arctic, limited on the shore line to the winter line of floating
ice and passing southward indefinitely in deep water.

This paper deals chiefly with what Dr. Dall would call the
Oregonian Fauna, only seven species having been secured to
the westward of the Shumagin Islands. Of these seven species
five are also found in his Oregonian Fauna, and the other two
are new and known, thus far, from only one locality.

Dr. Clark, in reporting on the collection made by Dr. Dall,
enumerates 25 species that occur west and north of the Shu-
magin Islands. Of these 25 species we now know that sixteen

Proc. Acad. Nat. Sciences, Philadelphia, p. 206, 1876.



[l6l] THE HYDROIDS 179

also occur to the eastward of the Islands, while five have not
been reported from any locality other than the ones where they
were originally discovered. Our present knowledge therefore
does not support the validity of Dr. Ball's division of faunae at
the Shumagin Islands. It rather indicates a continuity of fauna
from southern Alaska to the end of the Aleutian chain. Hydroid
life appears to decrease as we go westward, but this may be only
apparent and due to the more extensive exploration of the shores
east of the Aleutian Islands.

Dr. Dall extends his Oregonian fauna down to Monterey,
California. Reasoning again merely from the known distribu-
tion of hydroids, it would seem that Puget Sound is a natural
region of demarcation between faunas, although the region from
Puget Sound to San Francisco has been very little explored.
In 1876 Dr. Clark published a paper on * The Hydroids of the
Pacific Coast of the United States south of Vancouver Island,' l
in which he gives a list of twenty-four species ; of these only
two, Lafcea dumosa and Sertularia argentea, have as yet been
reported north of Puget Sound. The same author, in reporting
on Dr. Dall's collections from Alaska, notes as one of the main
points of interest, the " small number of species that are com-
mon to the Alaskan coast and the western shores of the United
States from Vancouver Island southward. 2 In 1899 t " ie P resen *
writer published a paper on ' Hydroida from Alaska and Puget
Sound ' 3 in which it appears that out of twenty-two species from
Puget Sound, only four have been reported farther south, while
fifteen are now known to occur in Alaska. In the same year
Mr. G. N. Calkins published a paper entitled ' Some Hydroids
from Puget Sound,' 2 in which some thirty species are noted,
only two of which are known to occur south of Puget Sound.

From this study of the distribution of the Hydroids of the
northwest coast of America, therefore, I am strongly persuaded
that the region south of Puget Sound constitutes one distinct
faunal area, and that the region from Puget Sound north and
west to the end of the Aleutian chain constitutes another un-

ir Trans. Conn. Acad. Sci., Vol. ni, pp. 250-251, 1876.
*Proc. Acad. Nat. Sciences, Philadelphia, p. 212, 1876.
3 Proc. U. S. Nat. Museum, Vol. xxi, No. 1171, 1899.
*Proc. Bost. Soc. Nat. Hist., Vol. xxvui, No. 13, 1899.



l8o NUTTING

broken faunal area that might properly be designated as Alaskan.
From the number of arctic species included in this area it is not
improbable that it extends northward along the shores of Bering
Sea.

Dr. Clark agrees with Dr. Dall that there is a distinct faunal
difference between the region east of the Shumagin Islands and
that west of them. The material added since the publication of
his paper, however, seems to prove that this difference is only
apparent and due solely to lack of exploration.

The most important thing to be noted in that part of the table
devoted to general distribution is the Holarctic distribution of a
number of species. Of the eighteen species known to occur in
the Arctic region, no less than fifteen also occur on the European
coast, fourteen on the Atlantic coast of the United States, and
thirteen on the Pacific coast as far south as Puget Sound. An
examination of the table shows further, that the Hydroid fauna
of Alaska, as represented by the Harriman collection, includes
fifty-three species in all, of which eighteen are Arctic in fact,
having been secured in Arctic waters ; four others are in all
probability Arctic, being found both in European and American
waters ; four are, so far as is known, confined to the Alaskan
and Pacific coast south to Puget Sound ; twenty-five are thus far
known from Alaska only, and two are Californian. If we rec-
ognize the Alaskan faunal region as extending to Puget Sound,
and include those species actually known to be Arctic, together
with those in all probability Arctic, in a group which may justly be
called Arctic, the following significant analysis of the faunal rela-
tions of the collection may be made : Alaskan species, twenty-
nine ; Arctic species, twenty-two ; Californian species, two. This
shows that fifty-five percent of the hydroid fauna as a whole is
peculiar to Alaska, but that there has been a strong invasion from
the Arctic regions of the Holarctic species constituting about
forty-one percent of the collection, and that only two species,
or less than four percent, are Californian. If all the species
known to occur in Alaska were included in the computation the
result would be a larger percentage of Alaskan species, a cor-
responding decrease of the Arctic species, and the addition of
one or two Californian species.



[163] THE HYDROIDS l8l

SYSTEMATIC DISCUSSION.

The writer deems it unnecessary to attempt a complete syn-
onymy of the well known European species contained in the
Harriman collection and considers it sufficient to give,Jirst, and
in all cases, the original reference to the species ; second, all
obtainable references to the occurrence of the species on the
Pacific Coast of America, and, third, a reference to verify the
' General Distribution ' as given in the table just discussed. In
this latter case only one reference will be given to verify the
occurrence of a given species in each of the regions included
in the right hand portion of the table.

As to the classification employed in this report, it seems best,
on the whole, to pursue a conservative course, following pretty
closely the lines laid down by the able British naturalists,
Hincks and Allman. While it is true that the classification is
in an unsatisfactory state, the writer does not feel that a thor-
ough revision of the entire group of Hydroida should be at-
tempted here, and frankly confesses his conviction that recent
attempts in that direction have not been successful, though each
contains valuable suggestions. Levinsen, for example, has
made a notable contribution to our knowledge of the Campanu-
linidae in his able and careful exposition of the differences in
the opercula of various species, but his genera founded solely on
these structures appear to be artificial, as usually happens when
a single charactef is made the basis of classification. 1 In his
terse characterization of the genera of Sertularidae this author
has been most fortunate, as well as in his masterly clearing up
of the mystery concerning the gonosome of Lafcea.

Schneider, 2 also, has attempted to rearrange the Hydroida on
a logical basis. Instead of multiplying groups, as has been the
tendency of late, he has, in my opinion, gone far to the other
extreme, uniting families that almost any other student acquainted
with the group would regard as surely distinct. It seems un-
likely that he will be followed in uniting such groups as the
Tubularidse and Pennaridae in a single family, although one

1 Meduser, Ctenopherer og Hydroider fra Gronlands Vestkyst, Copenhagen,
1893.

2 Hydropolypen von Rovigno, nebst iibersicht iiber das system der hydro-
polypen in allgemeinen. Zool. Jahrb., Syst. Abth., Vol. x, 1897.



l82 NUTTING

writer, Calkins, 1 has followed his classification quite closely, and
includes representatives of what would ordinarily be regarded as
at least eleven families in five families as defined by Schneider.
It is not likely that classifications will ever represent anything
but individual opinion, and it is probable that there will always
be two sets of extremists who on the one hand will be too ready
to multiply groups, and on the other will be too conservative to
recognize real progress. The Hydroida offer unusual difficul-
ties and consequently students of that group find agreement,
even along the most general lines, practically impossible. The
objective point of systematic discussion has been the attainment
of a system of classification by which genera could be distin-
guished by means of the trophosome alone. This end, although
in theory greatly to be desired, appears to be unattainable. The
writer has chased this will-o-the-wisp for years, and is ready to
abandon its pursuit as unprofitable. Abler men have had the
same experience, and it appears to be pretty well established
that in practice we must base generic distinctions on the gono-
some alone, although the judgment of students will inevitably
differ as to the extent to which this can be profitably done.
Botanists have encountered the same difficulty in their study of
the lower plants, such as the fungi, and have come to the same
conclusion. In neither case has nature been working for the
convenience of naturalists, and the fact should be accepted
without a bootless chase after the unattainable.

G TMNOBLASTEA.

Hydroida in which well differentiated hydrothecae and gonangia are
not present. What might be called ' pseudo-hydrothecae ' are found in
some species as in Eudendrium vaginatum (see description of that
species on pages 167-168).

Family CORTNID^E.

Trophosome. Hydranth with a terete body and proboscis and scat-
tered capitate tentacles only.

Gonosome. Fixed sporosacs, or free medusae with a very long manu-
brium, four marginal tentacles and four sense-bulbs with eye-spots.



Hydroids from Puget Sound, Proc. Bost. Soc. Nat. Hist., Vol.
xxvin, No. 13, 1899.



[165] THE HYDROIDS 183

CORYNE.

Trophosome. Characters of the family.

Gonosome. Reproductive elements produced in fixed sporosacs
growing on the hydranth body.

CORYNE BRACHIATA sp. nov.
(Plate xiv, figs, i, 2.)

Trophosome. Colony forming a dense tuft of irregularly branching
stems, sometimes attaining a height of about ^ inch. Stems and
branches profusely and regularly annulated throughout, fairly stout ex-
cept at the proximal ends where they taper gradually to their point of
origin ; distal ends of many of the branches bear a more or less regular
whorl, or radiating cluster, of annulated branchlets just below the
hydranth body, reminding one of the whorls of cirri around the stems
of the stalked crinoids. Hydranths large, with long, slender body and
proboscis and numerous (20-35) capitate tentacles arranged in a
scattered or sub-verticillate manner over nearly the whole surface.

Gonosome. Gonophores very numerous, borne among the tentacles
on the hydranth bodies, globular in outline and showing no traces
of radial canals or other medusoid structures. The specimens
secured were females and the gonophores were packed full of develop-
ing ova.

Distribution. All the specimens were secured in Yakutat Bay,
Alaska, by Dr. W. R. Coe of the Harriman Expedition.

This interesting species seems to be nearest to C. pusilla Gaertner, if
Allman has properly identified that species. It differs from other
members of the genus in the curious whorl of short branchlets which
bear neither hydranths nor gonophores and are situated a short distance
below the terminal hydranth of the stem or branch to which they are
attached. Another character not shown in the figures of this genus
published by Hincks and Allman, is the narrowing at the proximal
ends of the stems and branches. The specimens were found immersed
in sponge so far that only the hydranths extended above the surface of
the sponge.

SYNCORYNE.

Trophosome. Characters the same as those given for the family.

Gonosome. Reproductive elements produced in free medusae with
a long manubrium and four marginal tentacles, each having a sense
bulb with an eye-spot at its base.



184 NUTTING



SYNCORYNE EXIMIA Allman.
(Plate xiv, figs. 3, 4.)

Coryne eximia ALLMAN, Annals and Mag. Nat. Hist., 3d Series, Vol. iv, p.

141. Aug., 1859.
Syncoryne eximia ALLMAN, Annals and Mag. Nat. Hist., 3d Series, Vol. xm,

p. 357. May, 1864.

This appears to be the first record of the occurrence of this species
in American waters. There are many specimens in the collection, but
all are from the same locality.

Distribution. Juneau, Alaska (Harriman Expedition) ; Great
Britain (Allman and Hincks) ; Lofoten Islands, Norway (Sars).

Family BIMERID^E.

Trophosome. Hydranths with a conical or dome-shaped proboscis,
around the base of which is a whorl of filiform tentacles.
Gonosome. Sexual products developed in fixed sporosacs.

GARVEIA.

Trophosome. Colony branched ; perisarc conspicuous.

Gonosome. Gonophores borne on distinct branchlets which have a
chitinous investment ending in a cup-like expansion just below the
gonophore.

GARVEIA NUTANS Wright.

Garveia nutans WRIGHT, Edinburgh New Phil. Jour., p. 109. July, 1859.
Eudendrium bacciferum ALLMAN, Annals and Mag. Nat. Hist., 3d Series,
Vol. iv, p. 52, July, 1859.

This is another species that has not before been reported from
American waters. The specimens were fragmentary, making the
identification somewhat uncertain, although I have little doubt of its
correctness.

Distribution. Berg Inlet, Glacier Bay, Alaska. (Harriman Ex-
pedition.) Originally described from the British Coast.

GARVEIA ANNULATA sp. nov.

(Plate xv, figs, i, 2.)

Trophosome. Colony attaining a height of ij^ inches, consisting of
a number of closely aggregated and sparingly and irregularly branched
stems. Stems strongly and evenly annulated throughout, the perisarc
expanding distally into thin chitinous pseudo-hydrothecae which cover
the hydranth body nearly to the level of the tentacles. Hydranths with



[167] THE HYDROIDS 185

a conical, or rather conoid, proboscis and about sixteen tentacles all of
which appear to be held more or less erect.

Gonosome. Gonophores borne either on the stem or hydrorhiza,
more frequently the latter, oval in shape, borne on pedicels enveloped
in a chitinous perisarc which ends in a slightly expanded collar a little
below the gonophore. The specimens collected were female and the
gonophores were packed with apparently mature ova.

Color. The label accompanying the specimens bore the following
statement : " Bright orange throughout, heads, stems and all."

Distribution. Yakutat and Sitka, Alaska. Collected by the Har-
riman Expedition in considerable quantities.

This species can be sharply distinguished from its British relative
by the very distinct and beautiful annulation which covers the entire
stem and branches. It is much less extensively branched than the British
species, and the gonophores are more generally borne on the roots.

The structure that I have designated above as a ' pseudo-hydrotheca '
is of considerable morphological interest, for it may throw light on the
origin of the hydrotheca. The extension of the chitinous perisarc of
the stem over the body of the hydranth appears to be attached to the
latter. A true hydrotheca would be formed if the perisarc around the
hydranth body should become thicker and detached.

Family EUDENDRIDsft.

Trophosome. Colony branching. Hydranths with a single whorl
of filiform tentacles and a trumpet-shaped or hemispherical proboscis
which is expanded distally and contracted proximally, thus being
sharply distinguished from the hydranth body.

Gonosome. Reproductive elements developed in fixed sporosacs at-
tached to a usually more or less degenerated hydranth body below the
tentacles.

This family contains but one genus, Eudendrium, which needs no
further definition.

EUDENDRIUM VAGINATUM Allman.
(Plate xv, figs. 3-6.)

Eudendrium vaginatum ALLMAN, Annals and Mag. Nat. Hist., Third Se-
ries, Vol. xi, p. 10, Jan., 1863.

As the gonosome of this species has not heretofore been described,
the following is inserted here :

Gonosome. Gonophores (female) in dense clusters around the
bodies of hydranths that are usually devoid of tentacles. Each gono-


1 2 3 4 5 6 7 8 9 10 11 12 13 14 16 18 19 20

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