rete; REPORT

OF THE

CANADIAN ARCTIC EXPEDITION
1913-18

VOLUME IX: ANNELIDS, PARASITIC WORMS,
PROTOZOANS, ETC.

PART E: ACANTHOCEPHALA

By H. J. VAN CLEAVE

SOUTHERN PARTY—i913-16

OTTAWA

J. de LABROQUERIE TACHE
PRINTER TO THE KING’S MOST EXCELLENT MAJESTY

1920
Issued April 7 1920

wi

Report of the Canadian Arctic Expedition 1913-18.

VOLUME Ix: ANNELIDS, PARASITIC WORMS, PROTOZOANS, ETC.

Part A: OLIGOCHAETA..
Lumbriculidae. By Frank Smith.

Enchytraeidae. By Paul S, Welch.................. (Issued September 29, 1919).
Part B: POLYCHAETA. . By Ralph V. Chamberlain............ re csnvestaaonllne pread
Part C: HIRUDINEA. ‘By J. P. Moore................. 00 cece ee ees (In preparation).
Part D: GEPHYREA. By Ralph V. Chamberlin.. ane nauds aqusme reel a analebee tL (In press).
Part E: ACANTHOCEPHALA. By H. J. Van Cleave................04. “ese. (In press).
Part F: NEMATODA. By N. A. Cobb......... Shep aise plies ....(In preparation).
Part G: TREMATODA. By A. R, Cooper.........-.2...-.5-5 we naw B= (In preparation).
Part H: CESTODA. By A.R. Cooper................00 000 ee (In preparation).
Part I: TURBELLARIA. By A. Hassell...... SER er tail < hamid (In preparation).
Part J: GORDIACEA.
Part K: SPOROZOA. ByJ.W.Mavor............0.0. 0.0: e cece cece eens (In preparation).

Part L:
Part M: FORAMINIFERA. By J.A.Cushman.......... ....... (Issued February 6, 1920) ..

REPORT

OF THE

CANADIAN ARCTIC EXPEDITION
1913-18

VOLUME IX: ANNELIDS, PARASITIC WORMS,
PROTOZOANS, ETC.

PART E: ACANTHOCEPHALA

By H. J. VAN CLEAVE

SOUTHERN PARTY—1913-16

OTTAWA

J. de LABROQUERIE TACHE
PRINTER TO THE KING’S MOST EXCELLENT MAJESTY

1920
71929—1 Issued April 7, 1920

Cornell University

The original of this book is in
the Cornell University Library.

There are no known copyright restrictions in
the United States on the use of the text.

http:/Awww.archive.org/details/cu31924074096532

Acanthocephala of the Canadian Arctic Expedition, 1913-1918*

H. J. Van CuEave.
(With 2 Plates).

INTRODUCTION.

The acanthocephalan fauna of the Arctic regions has received little attention
from investigators, as evidenced by the meagre references to it in literature.
Records of some of the early expeditions contain references to the occurrence
of Acanthocephala in the Arctic but most of the descriptions are too vague
to be of value in any consideration of the taxonomy of the group. Species
described by some of the early explorers have become the objects of much
conjecture on the part of present-day investigators: Under the name Sipunculus
lendiz, Phipps (1774) described from an eider duck what is obviously a species
of Acanthocephala. Soon afterward, Goeza (1782:141) called attention to the
fact that this species of Phipps’ is in reality an acanthocephalan. Since that
time various investigators have endeavoured to determine the correct disposition
of this species within the group, but all of their attempts appear to be mere
guesses ostensibly fostered by the desire to distribute all of the species names
into groups which would at least give the appearance of a completely worked
out synonymy.

A considerable number of species of Acanthocephala have been described
by European parasitologists from hosts of arctic or sub-arctic range. Most
of the species have been found in whales and other marine mammals. Very
little evidence is available regarding the Acanthocephala of arctic birds and
fishes. Von Linstow (1905) published a report on the parasitic worms obtained
by the Russian Polar Expedition of 1900-1903 in which he gave descriptions
of three species of Acanthocephala representing the fauna of West Tajmirland.
Of these, a new species, Echinorhynchus pupa, was described from the King
Eider, Somateria (Erionetta) spectabilis; while the two remaining species Cory-
nosoma strumosum (=F. strumosus) and E. reductus, n. sp., were from the seal,
Phoca foetida.

In so far as the present writer is aware there are no published records of
the occurrence of Acanthocephala in the arctic fauna of North America. The
collections obtained by the Canadian Arctic Expedition thus offer an opportunity
of making a start toward the study of this interesting faunal region.

MATERIALS AND ACKNOWLEDGMENTS.

Through the kindness of Dr. A. R. Cooper the writer has received for
study the collection of Acanthocephala taken by Mr. F. Johansen, naturalist
on the Canadian Arctic Expedition. This collection is not extensive, either
with regard to the number of host species represented or in the regions repre-
sented. Nevertheless, since it represents the first arctic collection of these
parasites from the American continent it seems worth while to publish the
results of this investigation. Most of the specimens were collected in the vicinity
of Bernard harbour (Dolphin and Union strait), Northwest Territories, where

*Contributions from the Zoological Laboratory of the University of Illinois, No. 147.
71929—2

Canadian Arctic Expedition, 1913-18

the members of the expedition made their headquarters for a considerable period.
One lot of specimens was obtained from fish specimens taken by Dr. R. M.
Anderson on the east shore of Banks peninsula, Bathurst inlet, Northwest Terri-
tories. The collection comprises nine lots of specimens from six host species,
each lot including from a few to numerous individuals and representing, in some
instances, several host individuals. Three species of fresh-water fishes, two
marine fishes, and one bird constitute the entire list of acanthocephalan hosts
recorded by the expedition. In all, these harboured four species of Acanthoce-
phala. Echinorhynchus salvelini and E. coregoni are characteristic species from
North American fresh-water fishes, though the present records greatly extend
the known geographical range of both. Echinorhynchus gadi, the only species
found in marine fishes by the expedition, is a species common to marine hosts
of both Europe and America. A new species of the genus Filicollis from the
King Eider stands intermediate between the European and the North American
specics of this genus, but in some respects shows much closer relationship with
the previously described American species. Data concerning these four species
are given in the following paragraphs.

Filicollis arcticus, n. sp.
(Plate I, Figs. 1 to 5).
Color: orange (Johansen's notes).

Speciric DiaGnosis.—With the characters of the genus as emended by
Van Cleave, 1916. Body of fully mature individuals preserved in alcohol,
large, thick, approximately cylindrical in cross section, much folded or wrinkled,
especially in posterior region, due to contraction; about 20 mm. long and 5 mm.
in maximum diameter. Anterior portion of body wall covered with delicate
cuticular spines, visible only in cleared portions of the body-wall. Proboscis
ovoid, 0-77 mm. long, and 0-73 mm. in diameter, armed with twenty-two
longitudinal rows of seven or eight hooks each. Four or five hooks nearest
base of each row more delicate than those anterior to them and supplied with
less prominent roots. Anterior hooks provided with prominent posteriorly
projecting roots longer than the hook proper. Hooks range in length from 89
to 118 uw. Heavier hooks of anterior proboscis region are 35 yp thick at point
of emergence from proboscis wall, while those of posterior region range from
18 to 24 » thick. Neck approximately cylindrical, 1-2 mm. long by 0-44 mm.
in diameter. Proboscis receptacle inserted at base of proboscis and extending
through neck and into body cavity, entire length 2-5 mm. Embryos within
body cavity of gravid female 126 to 155 uw long by 30 to 41 » in diameter, most
frequent size about 150 by 41 u.

Type host, King Eider, Somateria spectabilis (Linnzeus), in intestine.

Type locality, Bernard harbour, Northwest Territories, Canada.

Collected hy Mr. F. Johansen, June 16, 1916.

Cotypes are deposited in the Victoria Memorial Museum, Ottawa, Canada,
and in the collection of the writer at Urbana, Illinois.

The above species description is based upon the examination of four speci-
mens. In all of these the proboscis was completely retracted within the body.
The data concerning the proboscis and its associated structures such as the hooks
and the proboscis receptacle are from a single fully mature female from which
the writer dissected out the proboscis. This dissection was stained and cleared
in the usual manner and mounted as a permanent mount in damar. Portions
of the body wall of this same individual were in like manner cleared and mounted
to permit of a study of the body spines which are unrecognizable in surface
view of alcoholic specimens. These spines recall those previously described
by the writer for Filicollis botulus in that in the gravid female they are very
poorly developed, appearing as very slight elevations of the cuticula in which
spines are observable only through close observation (see fig. 3).

Acanthocephala 5E

A comparison of F. arcticus with other known members of the same genus
discloses some interesting facts regarding the geographical distribution of the
members of this genus. F. anatis is the common European representative of
Frilicollis while F. botulus occurs in the Eiders in the United States. Filicollis
arcticus, n. sp:, differs in definite manner from both the previously mentioned
species but shows a distinctly closer relationship to F. botulus. Embryos
within the body cavity of the gravid female of the last named species range
from 71 to 83 y in length while those of F. arcticus vary from 126 to 155 uw. The
two American species also differ in the number of hooks on the proboscis. In
F. botulus there are but sixteen longitudinal rows of hooks while for F. arcticus
the writer has found twenty-two. Both of these American species lack the
spherical enlargement of the proboscis characteristic of the European species.

The King Eider, the host of F. arcticus, though cireumpolar in its distri-
bution, evidently does not carry the same acanthocephalan infestation through-
out its range. From the West Tajmirland peninsula, von Linstow (1905:3)
described Echinorhynchus pupa from this same host species. Unfortunately
his description and his figures of this species fail to give a full enough account
of the structure to enable anyone to place it with certainty in any of the genera
recognized in modern taxonomy of the Acanthocephala. In his figure of a
peculiarly coiled specimen the first loop posterior to the proboscis fails to show
any connection with the rest of the body. No mention is made in the text of
any peculiarity in the structure of the neck or anterior body region so it seems
probable this is not a unique structure but is simply an error in the execution
of the drawing. No evidence is presented, either in his description or in his
figures, which would make it seem probable that his species belongs to the
genus Filicollis. Thus on opposite sides of the arctic circle the King Eider
apparently is parasitized by Acanthocephala representing two distinct genera.

Echinorhynchus salvelini Linkins, 1918 (In Ward and Whipple).
(Plate II, Figs. 6 to 9).
Color: orange or greenish yellow (Johansen’s notes).

Speciric Diacgnosis.—With the characters of the genus. Anterior portion
of body-proper rather conspicuously enlarged, especially in the females. Males
7 to 12 mm. long and 0-8 to 2 mm. in maximum diameter. Females 10 to 20
mm. long and 1-2 to 2-25 mm. in maximum diameter. Proboscis cylindrical,
armed with sixteen to eighteen longitudinal rows of about thirteen hooks each.
Basal hooks about 39 to 50 » long, hooks on middle and anterior regions of
proboscis 44 to 68 uw long, some with a root about 83 uw long. Lemnisci short.
little longer than proboscis receptacle. Embryos within body cavity of gravid
female 115 to 165 uw long by 20 to 25 » broad, middle membrane forming a pro-
longation at each pole more than twice as long as it is broad, frequently charac-
teristically curved.

Arctic hosts: Cristivomer namaycush (Walbaum) and Salvelinus malma
(Walbaum). More detailed data of the occurrence of this species is given in
the following table:—

RECORDS OF THE OCCURRENCE OF £E. SALVELINI IN HOSTS Or
ARCTIC AMERICA.

Host. Locality. Date. Remarks.
Cristivomer namaycush....|Lake at Bernard harbour. .|October, 1915........ E. sulvelini only.
= ee es ../June 15, 22, 26, 1915..|#. coregoni also.
o Me re ctApprils UOC: i iysdau ara. E. coregoni also.
Salvelinus malma.........|Creek at Bernard harbour. |July 13, 1915.........|E. coregoni also

OE Canadian Arctic Expedition, 1913-18

Individuals of this species from the arctic are of considerably larger size
than those examined by the writer from hosts of The Great Lakes. The fully
mature females frequently reach a length of 20 mm. and show a very prominent
enlargement of the anterior portion of the body (fig. 6). This enlargement
is not of the type commonly occurring in gravid females of other species, involving
a practically uniform distention of the body, but is restricted chiefly to the
dorsal surface thereby giving the individuals a distinctly humped appearance.
In males this anterior enlargement occurs only rarely (fig. 8). The males of
this species bear a slight resemblance to those of E. gadi, in that the cement.
glands are arranged in approximately linear order in the posterior region of
the body, though frequently those near the end of the chain in E. salvelina
overlap one another.

In the original description of this species the number of proboscis hooks
was limited to sixteen longitudinal rows. The present writer has found indivi-
duals in the arctic collections possessing eighteen longitudinal rows of hooks
but in all other details agreeing with the description of this species.

Echinorhynchus coregoni Linkins (in Van Cleave, 1919).
(Plate II, Figs. 10 and 11).
Color: as the preceding species (Johansen’s notes).

Speciric DiaGnosis.—With the characters of the genus. Males 3 to 5
mm. long, with a maximum diameter at about the anterior fourth of the body,
0-8 to 1-05 mm. Females 3 to 10 mm. long, with maximum diameter of 0-6
to 1-7 mm. Proboscis cylindrical, carrying twelve to fifteen longitudinal rows
of hooks, each with ten or eleven hooks. Hooks not crowded on proboscis.
Basal hooks 28 to 53 u long; those on middle of proboscis 65 to 80 » long; those
near anterior tip smaller and weaker than those on middle of proboscis. Ventral
hooks slightly larger and stronger than dorsal. Lemnisci not longer than
proboscis receptacle. Cement glands of male in a compact mass. Embryos
within body cavity of gravid female 51 to 91 » long by 17 to 20 uw wide, with a
slight, approximately globular, prolongation of the middle membrane at each
pole.

RECORDS OF THE OCCURRENCE OF E. COREGONI IN HOSTS OF
ARCTIC AMERICA.

Host. Locality. Date. Remarks.
Cristivomer namaycush....|Lake at Bernard harbour. .|June 15, 22, 26, 1915. .|\also infested with
§ a & ..jApril, 1916...........|f BE. salvelini.
Salvelinus malma......... Creek at Bernard harbour. |July 18, 1915......... E. salvelini also.
e - SS ../June 28, 1916........ E. coregoni only.
Ss oquassa marstont....... Lake at Bernard harbour. .|December, 1915...... E. coregoni only.
; if oe . {October 1 and 2, 1915.|E. coregoni only.

In this species the nuclei of the subcuticula are very numerous and are
distributed almost uniformly over the extent of the body wall. Each mass of
nuclear material is surrounded by a comparatively distinct clear region, many
of which have the appearance of belonging to the lacunar system (fig. 11).
The writer has found the individuals of this species from arctic hosts to be
considerably larger than individuals from the region of The Great Lakes.

Acanthocephala 7E
Echinorhynchus gadi Miiller, 1776.

Over fifty specimens of this species were obtained from fish specimens
collected by Dr. R. M. Anderson at Banks peninsula, Bathurst inlet, Northwest
Territories, May 18, 1916. A single large specimen was taken from the stomach
of the Cod, Gadus callarias Linnaeus, at Station 48c (Banks peninsula). The
remaining worms came from the California Tomcod, Microgadus proximus
(Girard), at Station 48b (Banks peninsula, same date and place as above).
As is customary in this species, the specimens vary greatly in size. Those of
the present collection range from about 8 mm. to over 50 mm. in length.

This is one of the few species of Acanthocephala common to hosts both of
Europe and of America. Under the name EF. acuws Rudolphi, this species has been
reported frequently by Linton from numerous species of fishes from the Atlantic
coast of North America, including Gadus callarias, and also from Microgadus
tomcod which is closely related to M. proximus. The writer has also examined
specimens from both of these last-named hosts and has compared them with
individuals from European hosts and with the arctic representatives. Speci-
mens from all three localities undoubtedly belong to the same species.

CONCLUSIONS.

The Acanthocephala collected by the Canadian Arctic Expedition comprises
two typically North American species of the genus Hchinorhynchus parasitic
in fresh-water fishes, one species of Echinorhynchus common to marine fishes
of both Europe and America, and a fourth species of the genus Filicollis from the
King Eider which shows distinctly closer affinities to the previously described
North American representative of the genus than tothe European. There is no
evidence indicating any tendency toward a mixing of American and European
species of Acanthocephala in the arctic regions. The evidence is too frag-
mentary to permit of any statements of the general distribution of members of
this group within the arctic circle.

LITERATURE CITED.

Goeze, J. A. E. 1782. Versuch einer Naturgeschichte der Kingeweide-wirmer thierischer
Kérper. Blankenburg. ;
von Linstow, O. 1905. Helminthen der Russischen Polar-Expedition 1900-1903. Mem.
Acad. Imp. Sc. St. Petersbourge, Serie 8, Class Physico-Math., 18:1-17._
Lithe, M. 1904-05. Geschichte und Ergebnisse der Echinorhynchen-Forschung bis auf West-
rumb (1821). Zool. Annal., 1:139-250; 251-353.

; 1911. Die Acanthocephalen. Die Stisswasserfauna Deutschlands, Heft 16. Jena.
Van Cleave, H. J. 1916. Fulicollis botulus, u. sp., with notes on the characteristics of the genus.
Trans. Amer. Micros. Soc. 35:131-134. ;
———1918. The Acanthocephala of North American Birds. Trans. Amer. Micros. Soc.,

37:19-48.
——1919. Acanthocephala from the Illinois River, with descriptions of species and a synopsis
of the Family Neoechinorhynchide. Bull. Ill. Nat. Hist. Survey, vol. 13, No. 8.
Ward, H. B. and Whipple, G. C. 1918. Fresh-water Biology. John Wiley and Sons, New
York.

Canadian Arctic Expedition, 1913-18

EXPLANATION OF PLATES.

All figures were drawn with the aid of a camera lucida from hexatoxylin stained mounts
in damar, except figure 1, which was from an alcoholic specimen. Magnification is indicated
by a projected portion of a stage micrometer accompanying each drawing, values of these scales
are given in a footnote.

Fig. 1.

Piatr I.*
Filicollis arcticus, n. sp.

Female, showing general body form. It was from this individual that the proboscis was
later removed for study.

2. Profile of proboscis of fully mature female.

3. Surface view of a portion of the body wall from mature female, showing cuticular spines.
4,

5. An embryo from body cavity of gravid female.

Proboscis of same individual shown in fig. 1.

_ *The scale indicating magnification accompanying Fig. 1 has the value of 1 mm.; scales accompany-
ing all other figures of this place have the value of 0.1 mm.

9E

Puate I.

Acanthocephala

108

Tig. 6.
We
8.
9.
10.
11.

Canadian Arctic Expedition, 1913-18

Puate IT.**

Echinorhynchus salvclint. Outline drawing showing general body form of mature female.
Embryo from body cavity of gravid female of #. salvelint.

Male of F. salvelini in optical vertical section.

Selective apparatus from genital tract of mature female of EZ. salvelini.

Outline drawing showing general body form of Echinorhynchus coregoni, female.
Portion of body wall of #. coregoni, showing numerous small subcuticular nuclei.

** Scales indicating magnification accompanying figures 6, 8, and 10 have the value of 2 mm.; those
accompanying figures 9 and 11, 0.1 mm. and figure 7, 0.05 mm.

Acanthocephala Prats Il. 118

Report of the Canadian Arctic Expedition, 1913-18.
_ CONTENTS OF PARTS IN PREPARATION.

: : "Felume I: General Introduction, Narrative, Bte.
Part A: Worthen Party, 1913-18.
Part B: Southern Party, 1913-16. By Rudolph Martin iatteraon.” “@ n: preparation).

. Volume I: Mammals and Birds.
‘ Part A: a “ay Rudolph Martin Anderson. (In preparation).
Part B: Birds. By .R.M. Anderson and P. A. Taverner. (In preparation).

Volume IM: Insects.
Part A: Collembola. By Justus W. Folsom. (Issued).
Part B: Neuropteroid Insects. By Nathan Banks. . (I ssued).
Part C: Diptera. By Chas. W. Alexander, Harrison G. yar, and J. R. Malloch. (Issued).
Part D: Mallophaga and Anoplura. By A. W. Baker, ee . Ferris, and-G. H. F. Nuttall. (Issued).
_Part E: Coleoptera. By J.M. Swaine, H.C. Fall, C. W . Leng, and J. D DicGberetiny Jr. (Issued).
Part F: Hemiptera. By E, P. Van Duzee. {ssued), si
Part:G: Hymenoptera and Plant _Galls.. By Alex. D - MacGillivray, Charles T. Brues, F. Ww. L. Sladen,
and E. Porter Felt. (Issued). %
Hart Hs oe Mites, and Myriapods. By J. H. Hurtin, Nathan Banks, and Ralph, Vis Chamberlin.
sou
Part I: Lepidoptera. By Arthur Gibson. Ussued).
Part J: Orthoptera. By E. M. Walker.
Part K: General Observations on Insect Life in the Arctic. By Frits shun (In preparation). °

‘Volume IV: Botany. : ;

‘ Part A: _ Freshwater Algae and Freshwater Diatoms. - ’ By Charles W. Lowe. (In preparation).
_ Part B: Marine Algae.~ By F.Collins. (In preparation). ?

Part C: Fungi. By John Dearness., (In preparation).

Part D: Lichens. By K, L. Merrill. (In preparation).

Part ‘E: Mosses. By R. Ss. Williams. (In preparation).

a Volume V: Botany.

Part A: Flowering Plants and Ferns.,, By James M. Macoun, and others. (In preparation).
Part B: General Notes on Arctic Vegetation. y Frits Johansen. (In preparation).

Volume VI: Fishes, Tunicates, Ete.

Part A: Fishes... By F. Johansen. (In preparation).
Part B: Ascidians, ete. Byes G. Huntsman. (In preparation).

Volume VII: Crustacea.
Part A: — iristanaiiniss By Mary J. Rathbun, (Issued).
Part B: Schizopod. Crustaceans. By Waldo L. Schmitt. (Issued)
Part C: Cumacea. By W.T. Calman. (Jn preparation).
Part D: Isopoda. By Miss P. L. Boone. (In preparation). j
-Part E: Amphipoda. By Clarence R. Shoemaker. (In press. t
Part F: Pantopoda, Leon J. Cole. (In preparation).
Part G: Euphyllopoda. By F. Johansen. (In preparation).
Part H: Cladocera. By Chancey Juday. - In press).
Part I: Ostracoda. By R.W. Sharpe. (In preparation).
Part J: Freshwater Copepoda. By C. Dwight oe (In press).
Part K: Marine Copepoda. By A. Willey. (In press).
Part L: Parasitic Copepoda. ‘ By Chas. B. Wilson. ti n press).
Part M; Cirripedia. By H. A. Pilsbry. (In preparation).

Volume VII: Molluscs, Echinoderms, Coelenterates, Ete.

Part A: Mollusks, Recent and Pleistocene. By Wm. H. Dall. (Issued.)
Part B: Cephalopoda and Pteropoda. By S.S. Berry and W. F. Eiapie (In preparation).
Part C: Echinoderms. By Austin'H. Clark. (In press). ;
Part D: Bryozoa. By R.C. Osburn. (In 2 A ata
Part E. Rotatoria. By H. K. Harring. preparation).
Part F: Chaetognatha. By A. G. Tidisnak (In preparation).
Part H: Medusae and Ctenophora. By H.B..Bigelow. (In press).
Part I: Hydroids. By McLean Fraser. (In preparation).
(Porifera, Actinozoa, and Alcyonaria:* material small i in amount, and no specialists selected).

Volume IX: Annelids, Parasitic Worms, Protozoans, Etc.

Part A: Oligochaeta. By Frank Smith and Paul S. Welch. (Issued).

Part'B: Polychaeta. By Ralph V. Chamberlin. (In press).:

Part C: Hirudinea. By J. P. Moore. (In preparation).

Part D: Gephyrea. By Ralph V. Chamberlin. (In press).

Part E: Acanthocephala. By H. J. Van Cleave. (In press).

Part F: Nematoda. By N. . Cobb. (In preparation).

‘Part G: Trematoda. By eee . Cooper. (In preparation).
oe .H; Cestoda. By A.R ’ Cooper. (In preparation).

Part I: "Turbellaria. By A. “Hassell, (In preparation).

Part J: Gordiacea. ,

‘Part K. Sporozoa. By ww. ‘Mavor. (In preparation).

Part M: Foraminifera. By J. A. Cushman. (Issued).

Volume X: Plankton, Hydrography, Tides, Etc,

‘Part A: Plankton. Marine Diatoms..
Part B: Tidal Observations. By -W. Bell Dawson. (In press).
Hydrography. ‘