Canadian Arctic Expedition,
1913-1918

v.9, pt.fi

REPORT

OF THE

CANADIAN ARCTIC EXPEDITION

1913-18

VOLUME IX
ANNELIDS, PARASITIC WORMS, PROTOZOANS, Etc.

PART A: OLIGOCHAETA

Lumbriculidae Frank Smith

Enchytraeidae . Paul S. Welch

SOUTHERN PARTY, 1913-16

OTTAWA

J. de LABROQUERIE TACHE

PRINTER TO THE KING'S MOST EXCELLENT MAJESTY
1919

Vol. ix— 45455— 1 Issued September 29, 1919

Oligochaeta Collected by the Canadian Arctic Expedition

1913-18.

By FRANK SMITH and PAUL S. WELCH.
(With three figures in text.)

An opportunity to examine the collections of Oligochaeta made in the
Arctic regions of North America by the Canadian Arctic Expedition has been
very welcome, since it has made it possible to materially extend our knowledge
of the oligochaete fauna of the Arctic zone as well as that of North America.

The Lumbriculidae and Enchytraeidae are the only families represented,
and are the ones which our knowledge of their distribution in the Arctic regions
of Eurasia would lead us to expect. A general similarity is found between
the oligochaete faunas of the two regions, although they are not identical.

The Lumbriculidae in the collections have been studied and are here dealt
with by the senior author, while the part dealing with the Enchytraeidae and
including the descriptions of the new species is by the junior author. The
collections which are from quite a wide range of localities and a considerable
variety of habitats were made by Mr. F. Johansen, the marine biologist, ento-
mologist, and botanist of the expedition.

It is interesting to learn from the field notes of the collector, that some of
these worm's were found alive in dried-out creek beds and pond bottoms where
they had taken refuge in moss or other vegetation. Others were found unharmed
by exposure to freezing conditions. In these respects they resemble some of
their relatives of more temperate regions.

I.— THE LUMBRICULIDAE1
BY FRANK SMITH.

This family is represented by the single genus Lumbriculus, and there are
but two species. L. variegatus (Miiller), the well-known Eurasian species is
the more abundant and widely distributed form, and L. inconstans (Smith), was
found associated with it in two localities.

Lumbriculus Grube.

Prostomium rounded, without proboscis-like elongation. Setae paired and
cleft. Spermiducal pores one or two, exceptionally three; paired or asymmetric-
ally placed on one or two somites, more commonly on 82 or 10. Oviducal
pores usually paired in one or two somites next following those with spermiducal
pores, sometimes otherwise arranged. Spermathecal pores laterad of ventral
setae; usually in three to five pairs, frequently fewer or more numerous, often
asymmetrically arranged; commonly posterior to spermiducal pores. Dorsal
and ventral vessels connected in posterior part of each of nearly all somites by
a pair of transverse vessels; dorsal vessel in anterior part of each somite, except
a few anterior ones, with one pair of contractile caecal transverse appendages.
Atria, one, two, or three; paired, or asymmetrically arranged in one or two

iContributions from the Zoological Laboratory of the University of Illinois, No. 139.
2Arabic numerals are used to designate somites, counting from the anterior end.

46455—2

4A

Canadian Arctic Expedition, 1913-18

somites. Ovaries and oviducts usually paired in one or two somites next
posterior to one with atria, sometimes otherwise arranged. Spermathecae
usually three to five pairs, number highly variable; commonly in somites
posterior to those containing the atria.

Lumbriculus fariegatus (0. F. Miiller.)

Length, 40-80 mm. Diameter, 1-1-5 mm. Number of somites, 140-200,
or more. Colour in life, reddish to dark brown, anteriorly greenish. Prpsto-
mium rounded conical, length about one and one-half times the basal width.
Spermiducal pores usually on 8, less frequently on 7, rarely on 6, 9, 10, or 11.
Oviducal pores, one to three pairs, often asymmetrically arranged, more com-
monly one pair on 9. Spermathecal pores highly variable in number and
position, nearly always posterior to spermiducal pores and on three or more
somites. Spermaries and spermiducal funnels variable in number and position,
usually associated with the atria and more commonly a pair of spermaries and
one or a pair of funnels in 8. Ovaries and oviducts variable in number and
position, more commonly a pair of each in somite next posterior to atrial somite.
Spermathecae highly variable in number and position, more commonly three
to five pairs in 9-13.

This species is represented in collections from the following localities,
mentioned in order of distribution from west to east; the accompanying data
are given as supplied by the collector : Konganevik Point (Camden bay) ,
Alaska, June 25-26, 1914, in mud at margin of lake. Collinson point (in Camden
bay), Alaska, June 13, 1914, in mud in tundra pond (margin). Herschel island,
Yukon Territory, July 30-31, 1916, pond in tundra swamp at east- end. Cape
Bathurst, Northwest .Territories, July 26, 1916, waterhole in brook, swamp
(tundra). Bernard harbour, Northwest Territories, June 28, 1915, in mud at
margin of tundra pond; July 13, 1915, from green algae in streaming water
(tundra); September 30, 1915, in bottom mud of big lake, one fathom water;
June 6, 1916, in mud of tundra pond (shallow).

^

.-sy

8

-of
st
12

FIG. 1.

FIG. 2.

Diagrams showing distribution of reproductive organs in two specimens of Lumbriculus
variegatus. Numerals designate the somites; a, atrium; of, oviducal funnel; oy, ovary; sf,
spermiducal funnel; st, spermatheca; sy, spermary.

Seven specimens of L. variegatus, among a total of about twenty, have at
least traces of atria, and in each case they are paired in 8 and spermaries are
paired in the same somite. Spermiducal funnels are found in but four of these
and' are also paired in somite 8. In each of the seven specimens haying atria,
paired ovaries and oviducts are contained in 9; an additional pair of well-

Oligochaeta 5 A

developed ovaries and of oviducts is present in 10 in one specimen; two have
small ovaries and either a pair of oviducts or a single one in 10; one has small
ovaries and no oviducts in 10; and three specimens have neither ovaries nor
oviducts in that somite. But two specimens are in the proper stage of sex
organ development to show spermathecae, and in one of them these organs are
paired in 11 and 12 and there is a single one in the right hand side of 10 (fig. 1).
In the other specimen spermathecae are paired in 9, 11, and 12, there i§ one
on the left side of 10 and two on the right side of that somite (fig. 2). As will
appear in the general discussion of this species, the collections contain an un-
usually large percentage of specimens with partially or well developed sex organs.
Such individuals are ordinarily very rare. The diameters of these specimens
are 1-1 -4 mm.

Lumbriculus inconstans (F. Smith.)

Length, 30-60 mm. Diameter, 0 • 6-0 • 8 mm. Number of somites, 150-200,
or more. Colour in life, reddish, anteriorly greenish. Prostomium rounded,
length one and one-half times the basal width. Spermiducal pores usually
on 10, occasionally on 9 or 11. Oviducal pores usually two pairs in two somites
next following the one with spermiducal pores, occasionally but one pair.
Spermathecal pores paired or asymmetrically distributed on four or five somites
posterior to the atrial somite. Spermaries and spermiducal funnels one to
three pairs, commonly two pairs in 9 and 10. Ovaries and oviducal funnels
one or two pairs, commonly two pairs in 11 and 12. Spermathecae paired or
asymmetrically distributed in four or five of somites 11-16.

A few specimens of this species were found associated with L. variegatus
in the collections from the following localities: Collinson point, Alaska, June 13,
1914, in mud in tundra pond (margin). Herschel island, Yukon Territory,
July 30-31, 1916, pond in tundra swamp at east end.

sy

FIG. 3

Diagram showing distribution of reproductive organs in a specimen of Lumbriculus inconstans.
Numerals and letters as in figs. 1 and 2.

But two of the specimens of L. inconstans have atria. One of them has
paired atria in 10, but no traces of spermathecae. It has also paired spermaries
and spermiducal funnels in 9 and 10, paired ovaries and oviducts in 11, one
well developed ovary and one minute one in 12, but no oviducts are recognizable
in that somite. Small cell aggregates that are apparently similar to rudimentary
ovaries are present on one side of 13—17. Similar bodies were found by Mrazek
(1907:429) in L. variegatus. Another specimen that is more nearly mature
has also paired atria in 10 (fig. 3) and paired spermaries and spermiducal funnels

46455— 2J

GA

Canadian Arctic Expedition, 1913-18

in 9 and 10. Ovaries and oviducts are paired in 11 and 12; spennat hrcac are
present on one side of 12-14, and on the other side there are vestiges of spcr-
mathecal ducts in 11-14. The diameter of these specimens is Inn o-f) -n-.v>
mm. which is but half of that of the mature individuals of /.. rnrnijutus \\\ the
same collections.

DISTRIBUTION AND SYSTEMATIC RELATIONSHIPS.

DISTRIBUTION. — L. variegatus has long been known from various pan- of
Europe. Eisen (1879:11) reported specimens from several localities in western
arctic Siberia as doubtfully belonging to this species. His figures and desci -ipi ion
•of certain " glands " in 9-12 make it probable that he was really dealing with
the spermathecae in L. variegatus. In an earlier paper Eisen (1872:122) also
records the species in Greenland, but this is regarded by some subsequent
writers as doubtful. It is now shown to have an extensive distribution in the
northern fringe of the continental areas of Eurasia and North America and
probably the area occupied by it is fairly continuous.

L. inconstans has previously been known only from the North Central and
Great Lakes regions of the United States, and the new records greatly extend
its range.

SYSTEMATIC. — L. variegatus has had an especially prominent place in the
oligochaetological literature of Europe, and is well known, not only to the
systematists, but also to students of regeneration phenomena and the related
morphological problems. Although it was known to Bonnet in 1745 and was
formally described and named by 0. F. Muller in 1774, yet it was not until
1907 that an adequate treatment of the reproductive organs appeared, when
Mrazek published his paper giving the results of an examination of something
over 200 sexually mature specimens. Such individuals are extremely rare and
the reproductive system is extraordinarily variable, and, in consequence, earlier
writers with not more than one to three specimens each had published more
or less discordant accounts. These discrepancies are easily explained in the
light of Mrazek's results.

The atria are found in one or more of somites 6-11. There is sometimes
a single atrium, sometimes a pair, and sometimes either t'wo or three atria
asymmetrically disposed. In the latter case the atrial somites are sometimes
not consecutive. The paired relation is the most common, and 8 is most fre-
quently the atrial somite. Aside from 8, somite 7 is the one more commonly
involved. The number of spermiducal funnels varies from a single one to three
pairs, with the presence of a single one or a single pair in 8 occurring most
frequently. Asymmetry in the arrangement of the spermiducal funnels is of
frequent occurrence. Similar irregularities in the number and arrangement
of spermaries, ovaries, and oviducts are found; the most common condition is
one pair of spermaries in 8 and one pair each of ovaries and of oviducts in 9.
An additional pair of ovaries and of oviducts in 10 is also of frequent occurrence.
There is most surprising variability in the number and arrangement of the
spermathecae. There were none of somites 7-17 that did not contain sper-
mathecae in some one or more of Mrazek's series of worms. It is very common
to have two or even three spermathecae on one side of a single somite with the
result that as many as five and even six spermathecae in a single somite have
been found. The total number of these organs in one individual has been found
varying from three in one specimen to twenty-seven in another, and the number
of somites containing spermathecae has varied from two to nine. In nearly all
specimens, 10, 11, and 12 are found to contain spermathecae, and also in quite
a large number, 9 or 13 or both have these organs, while they are found in any
of the other somites far less frequently.

Oligochaeta 7 A

L. inconstans, like its congener, is very rarely found sexually mature and
the few specimens thus far obtained in that condition indicate that in this
species also, there is much variability in the number and arrangement of the
various reproductive organs. The most characteristic difference" between the
two species is in the general location of these organs and in the number
of spermaries and spermiducal funnels. Paired atria, spermaries, and
spermiducal funnels in 10, with an additional pair of spermaries and sper-
miducal funnels in 9, is the usual condition in L. inconstans, while a single pair
of atria and of spermaries in 8, and a single spermiducal funnel or a pair in that
somite is the usual condition in L. variegatus. The spermathecae of L. inconstans
are normally about two somites farther posterior than are those of the other
species. Size is not ordinarily very significant as a distinguishing character,
but no specimen of L. inconstans has been found with a diameter much more
than half of that of ordinary mature specimens of L. variegatus, and now a similar
difference is apparent between mature specimens of the two species when found
in the same situation. This is true in each of the two collections of the expedition
in which L. inconstans is contained.

Unfortunately there has been no series of sexually mature specimens of L
inconstans at all comparable in numbers with that of the European specie^
obtained by Mrazek. Including the single specimen to which reference was
made by Moore (1906 : 169), we know the position of the atria in but eleven j
and of the spermathecae in but five specimens of L. inconstans from the Great
Lakes region of North America. The atria are paired in 9 in two specimens;
paired in 10 in eight specimens; and paired in 11 in one. In each of the two
with atria paired in 9 there are spermaries and spermiducal funnels in 9 ; ovaries
and oviducts in 10 and 11; one has spermathecae in 11-15, in part asymmetrical,
and in the other they are undeveloped. In the specimen with the atria paired
in 11, there are paired spermaries and spermiducal funnels in 9, 10, and 11;
a pair of ovaries and oviducts in 12; and spermathecae in 12-16, in part asymme-
trical. In two earlier papers (1895 and 1905) I have referred to this specimen
with atria in 11, and included it with others in a statement that spermaries are
present in 9 and 10 and ovaries in 11 and 12. A re-examination of this specimen,
with greater experience in interpreting vestigial structures in these worms,
has convinced me that in this particular individual, the gonads and ducts of
11 are in reality spermaries and spermiducal rather than ovaries and oviducal
as previously announced. Vestiges of the funnels is about all that is present
to represent the ducts, and their position indicates that they probably are
spermiducal structures. In each of the eight specimens with paired atria in
10 there are paired gonads, presumably spermaries, in 9 and 10; and five of
them have at least traces of spermiducal funnels in 9 and 10. • In the others,
recognizable traces of the funnels are lacking. Ovaries and oviducal funnels
are present in 11 and 12 in each of five of these eight specimens; two others
have gonads, presumably ovaries, in 11 and 12, but no. recognizable funnels;
and one has one pair of ovaries and a pair of oviducal funnels in 11, but none
in 12, except a rudimentary gonad on one side. Spermathecae are recognizable
in but five of the specimens and in some are somewhat asymmetrical, having
in some somites a single spermatheca instead of a pair. One specimen with
atria in 9 has spermathecae in 11-15; one with atria in 11 has spermathecae in
12-16; and of three specimens with atria in 10, one has spermathecae in 11-15,
one has them in 12-15, and one has them in 12-16. The specimen described
from arctic North America has them in 11-14. No two of the six specimens
which have the organs all present have an identical arrangement of them.

In connection with the question of the possible identity of the two species
of Lumbriculus mentioned in this paper, it is interesting to note that in all of
Mrazek's series there were but three that had atria in 10, and each of these
had an asymmetrical arrangement of these organs, there being in each case
one or two atria in a more anterior somite. In at least one of the very small

8 A

Canadian Arctic Expedition, 1913-18

number of specimens in his series having atria in 11, there was evidence of
regeneration of anterior somites. This would be very likely to alter the number
of somites and hence the numerical relations of organ positions.

Lumbriculus, sp.

The collection contains two small specimens having data as follows:
'Brackish pond at Teller (Port Clarence), Alaska. In moss-brink. August 3,
1913." They apparently are sexually immature specimens of some species of
Lumbriculus. Sections of the most promising looking specimen revealed defective
preservation, some evidence of regeneration of a few anterior somites, and a lack
of reproductive organs except apparent traces of gonads. These latter are in
somites 10-13 and possibly in one side of 9. Because of the change in the number
of somites liable to occur in the process of regeneration, the location of the gonads
has little systematic significance, and there is insufficient basis for identification.

The occurrence of representatives of Lurroriculidae in brackish water is at
least unusual.

LITERATURE CITED, PART I.

Eisen, G.

1872. Om nagra arktiska Oligochseter. Ofv. Kongl. Vet. Akad. Forh. 29 : 119—124.

1879. On the Oligochaeta collected during the Swedish Expeditions to the arctic regions in
the years 1870, 1875 and 1876. Kongl. Sv. Vet. Akad. Handl. 15 : 1—49.

Moore, J. P.

1906. Hirudinea and Oligochaeta collected in the Great Lakes Region. Bull. Bur. Fisheries

25 : 153—171.

Mrazek, Al.

1907. Die Geschlechtsverhaltnisse und die Geschlechtsorgane von Lumbriculus variegatus

Gr. Zool. Jahrb. Anat. u. Ont. 23 : 381—462.

Smith, F.

1895. Notes on Species of North American Oligochseta. Bull. III. State Lab. Nat. Hist.
4 : 285—297.

1905. Notes on Species of North American Oligochaeta. V. The Systematic Relationships
of Lumbriculus (Thinodrilus) inconstans (Smith). Bull. 111. State Lab. Nat.
Hist. 7 : 45 — 51.

II.— THE ENCHYTRAEIDAE.1
BY PAUL S. WELCH.

Eight of the collections contained mature enchytraeids representing six
species distributed among four genera. Immature specimens not permitting
specific identification were also present. Serious difficulties encountered in
connection with some of the material necessitated provisional identification of
two species. These difficulties were mainly due to incomplete, original descrip-
tions of arctic Eurasian species made many years ago when the Enchytraeidae
were little known and before the taxonomic importance of several anatomical
features was recognized. Since the original Eurasian specimens are not avail-
able, the descriptions and accompanying illustrations are the sole dependence,
thus preventing positive identifications, at least in some cases. For this reason,
brief anatomical descriptions of the representatives of each species treated in
this report have been included and it will be understood that they are based
exclusively upon the characters of the Canadian specimens. Characteristics of
immature specimens are listed only in case it is unlikely that they are juvenile
individuals of a species already discussed.

Henlea nasuta (Eisen.)

A collection of enchytraeids made July 7, 1915, from wet moss roots on a
tundra, Bernard harbour, Northwest Territories, contained two specimens, both
mature, and the posterior portion of another which exhibit the characters of
Henlea nasuta (Eisen.)

CHARACTERISTICS. — Length, 19.3 mm. Diameter, about 1.0 mm. Somites,
about 58. Colour of alcoholic specimens light brown. Prostomium small;
short; pointed. Setae 4-6 per bundle in ventral rows; 3-5 per bundle in lateral
rows; straight; inner ones of bundle slightly smaller. Clitellum well developed ;
on 12-f 13; continuous around body. Lymphocytes scanty; ovoid to circular.
Peptonephridia present; one dorsal and one ventral strand; both arising from
digestive tract in 5; both in close contact with wall of latter; branches pro-
jecting into coelom in 6; ventral strand longer than dorsal, extending to 8.
Oesophagus expanding abruptly into intestine in 8. Two conspicuous intestinal
diverticula in 8, each opening independently by single opening into latero-
ventral part of the digestive tract; bag-shaped; reflected cephalad; slightly
flattened laterally; large central cavity with many deep folds of lining. Dorsal
blood-vessel arising in 8. Nephridia with small anteseptal and large postseptal
part; efferent duct arising from anterior ventral surface of latter. Spermiducal
funnel about 1J times longer than wide. Sperm duct confined to 12. Penial
bulb of simple lumbricillid type; body of bulb composed of but one set of cells.
Spermathecae with no definite demarcation between duct and ampulla; few,
small, unicellular glands at ectal opening; duct about one-half as long as
ampulla; body of ampulla ovoid, decreasing entad into very fine tube above
digestive tract and uniting with similar tube from corresponding spermatheca
on opposite side of body to form common duct; apparently uniting with alimen-
tary canal in posterior part of 5.

DISTRIBUTION. — Henlea nasuta was originally described by Eisen (1879 : 20-
21) from specimens collected all along the Yenisei river, northern Siberia, at
localities varying from 60° 50' to 72° 40' n. lat. According to the synonymy

Contribution from the Entomological Laboratory, Kansas State Agricultural College,
No. 34.

10 A Canadian Arctic Expedition, 1 913-1 S

given by Michaelsen (1900 : 69), it was later reported under other names from
Solowetski island in the White sea, Denmark, Germany, Bohemia, France, and
Italy.

Henlea tenella (Eisen.)

Two mature enchytraeids belonging to the genus Henlea were collected
under old driftwood logs on the tundra at Collinson point, Alaska, September
27, 1913. These specimens agree in so many particulars with the description
of Henlea (Archienchytraeus) tenella (Eisen) that they are placed provisionally
under that name. Unfortunately, the description of this species, like many of
the older ones, is very brief, indefinite with respect to some features, and entirely
lacking in information concerning certain important anatomical characters. In
order to facilitate future revision, the principal characters of the specimens
studied are given below.

CHARACTERISTICS. — Length, 10 • 8-12 mm. Diameter, in region of clitellum,
about 0-42 mm. Somites, 52-53. Colour of alcoholic specimens uniformly
dark brown. Prostomium short; very blunt; rounded; smooth. Setae almost
straight; inner ones of bundle slightly smaller than outer; 2-5 per bundle in
lateral rows, usually 3-5; 2-6 per bundle in ventral rows, usually 4-6; number
per bundle decreasing at posterior end of body. Clitellum only slightly devel-
oped; inconspicuous; on 12-13; diameter of body slightly or not at all greater
at clitellum than at adjacent somites; continuous around body. Lymphocytes
scanty; elliptical; variable in size. Brain 0-19 mm. long, 0-12 mm. wide;
depressed; lateral margins slightly divergent caudad; posterior margin deeply
emarginate; anterior margin slightly concave; one pair supporting strands from
apices of posterior lobes, another pair from latero-posterior margins-, both pairs
extending caudo-laterad to body-wall. One pair of diminutive peptonephridia
in 5 on ventral side of digestive tract; slightly branching. Chloragog cells in
dense layer about digestive tract for its whole length. Oesophagus expanding
abruptly into intestine in 8. Intestinal diverticulum in 8; entirely surrounding
digestive tract; conspicuous in transverse section; tubular in structure, tubules
uniting with intestine in posterior part of 8. Origin of dorsal blood-vessel in
posterior part of 8 or beginning of 9. Nephridia with small, anteseptal part;
postseptal part large, compressed; efferent duct as long as postseptal part,
arising from ventral surface of latter near septum. Spermiducal funnel small;
somewhat cylindrical; expanded near middle; length about twice maximum
diameter. Sperm duct much contorted; confined to 12. Penial bulb small;
of lumbricillid type; body of organ with but one set of cells. Spermathecae
very small and simple; wall thinned to form ampulla without increasing diameter
externally; no diverticula; inconspicuous glands at ectal opening of duct;
spermathecae uniting dorsad of digestive tract to form small, common tube;
latter connecting with lumen of digestive tract; ental portions of spermathecae
very slender.

DISCUSSION AND DESCRIPTION.

When compared with the original description of Henlea tenella (Eisen,
1879 : 17), differences appear in the number of setae per bundle, the length and
breadth of the brain, and the shape of the spermiducal funnel, but these disagree-
ments are minor and cannot be regarded as having any particular importance.
It is not known whether the original material from which Henlea tenella was
described had an intestinal diverticulum, penial bulbs, and dorsal blood-vessel
of the kinds herein described for the Collinson point specimens or not and an
element of uncertainty thus exists in the identification. For the purpose of this
report none of the anatomical features demand special mention except the
intestinal diverticulum, the penial bulb, and the spermathecea.

Oligochaeta 11 A

INTESTINAL DIVERTICULUM. — Structurally, the intestinal diverticulum in
these Collinson point specimens resembles very closely those in Henlea maderata
Welch and Henlea tubulifera Welch. The writer pointed out in an earlier paper
(Welch, 1914 : 160) that a similar type of structure is reported for the intestinal
diverticulum in the genus Hepatogaster 6ejka. In fact, Cejka (1910) uses this
type of structure as one of the distinguishing characters of Hepatogaster. How-
ever, since the Collinson poinl^ material lacks the canals in the intestinal epithe-
lium and since the peptonephridia and spermathecae differ from those described
for that genus, this species is herein recorded as belonging to Henlea.

This organ arises from the posterior end of the oesophagus in 8 and is
reflected cephalad, closely investing the latter for almost the entire length of
the somite. It occupies approximately one-half of the body-cavity in 8. A
deep, narrow, longitudinal depression on the mid-dorsal side partly encloses the
dorsal blood-vessel. A smaller, longitudinal depression on the mid-ventral
surface partly encloses the ventral blood-vessel. The organ is largely composed
of a series of branching, longitudinal, thick-walled tubules closely crowded
together, which unite near the caudal end of the diverticulum and open into the
lumen of the digestive tract through a few larger tubes.

It might be mentioned that Friend (1914 : 151-153) lists Henlea tenella
under a heading " cesophageal glands absent," but, since the English record is
uncertain, this statement is disregarded.

PENIAL BULB. — The penial bulbs are small, being but little thicker, in
transverse section of the body of the worm, than the adjacent muscle-layer of
the body-wall. They are simple in structure, conforming to the lumbricillid
type. Each is composed of but one set of gland cells and is covered by a well-
developed muscle-layer. The sperm duct extends well into the bulb to unite
with a distinctly differentiated penial lumen. In the description of the species
given by Michaelsen (1900 : 70), the statement is made that the penis is much
smaller than the sperm funnel but it is uncertain what part or how much of the
penial bulb apparatus is referred to. If this statement applies to the whole
penial bulb, it is not true for the American specimens since the penial bulb is
as large as the sperm funnel although but little larger.

SPERMATHECAE. — The original description by Eisen (1879 : 17) contains no
mention of any connection of the spermathecae with the digestive tract, and
his accompanying figures (PL III, 5g; PL IV, 5m) seem to indicate that they
end blindly, being very much narrowed at the ental end. Since the ental
extremities of the spermathecae in the Collinson point specimens are very much
reduced in diameter, the writer is inclined to suspect that Eisen overlooked the
very inconspicuous alimentary connection, or else examined immature speci-
mens, since there is evidence that the connection with the digestive tract is
one of the last features to be completed in the development of the spermatheca.
Eisen also states that "A form with the receptacle bent at the top and some-
what longer could perhaps be separated as a variety: elongatus (PL IV, fig. 5m).
This variety was found in Norway near Tromso, together with Neoenchytraeus
Ratzelii, and may be considered as a local variety or a beginning species." This
" variety " was also probably described from immature specimens or else the
alimentary connection was overlooked. Judging from the above description
and an examination of the original figures, there seems to be no grounds for
establishing such a variety.

DISTRIBUTION. — The specimens studied by Eisen and classified as Henlea
tenella were collected on Solowetski island in the White sea (65° n. lat.), at
Tromso, Norway (about 69° n. lat.) and at " Sapotschnaja Korga, Jenissej,"
Siberia (71° 41' n. lat.). Aside from an uncertain record by Friend (1912 : 585-6)
of its occurrence in England, no other records appear in the literature, and it
thus appears that the species, as known at present, is Arctic in its distribution.
Although widely removed geographically from the known Old World localities,
this species apparently exists under similar conditions in arctic America.

12 A

Canadian Arctic Expedition, 1913-18
Henlea ochracea (Eisen.)

A collection made July 26, 1916, at cape Bathurst, Northwest Territories,
from a " water-hole in brook swamp (tundra) " contained two mature enchy-
traeids which exhibit characters very similar to those described by Eisen
(1879 : 20) for Henlea ochracea. Eisen's description is so meager that other
workers have had difficulty in using it. The writer, in a careful examination
of the original description and its accompanying figures, has found so <•!<>«'
an agreement with the Canadian specimens that they are listed here as Henlea
ochracea, at least provisionally.

CHARACTERISTICS. — Length, 12 mm. Maximum diameter, .0-7-1 mm.
Somites, 47. Colour of alcoholic specimens dark brown. Prostomium prom-
inent; smooth; rounded at tip; extending well beyond mouth. Setae straight :
inner ones of bundle slightly smaller than outer; 3-5 per bundle, fewer at
extremities. Clitellum on 12-13; well developed; continuous around body.
Lymphocytes sparse; ovoid to circular. Brain almost as long as wide; anterior
margin slightly concave; posterior margin emarginate; lateral margins divergent
caudad; almost circular in transverse section. Two sets of peptonephridia
arising from digestive tract in 5; dorsal pair arising as large, tubular, glandular
organs, extending caudad for short distance between epithelium and muscle-
layer of alimentary canal, ending in a few long branches into coelom in 6; ventral
pair arising as two tubular strands, more massive than dorsal peptonephridia,
also few branches into coelom in 6. Oesophagus expanding abruptly into
intestine in 8. Intestinal diverticulum in 8; completely surrounding digestive
tract as single organ; with numerous internal tubules. Origin of dorsal blood-
vessel in 9; vessel exceptionally enlarged above diverticulum. Nephridia with
very small anteseptal and large postseptal part; efferent duct arising from
posterior end of latter. Spermiducal funnel about twice as long as wide; with
thin, reflected collar; sperm duct in 12 only, much coiled. Penial bulb of
simple lumbricillid type; but one set of cells in body of bulb. Spermathecae
spindle-shaped; without distinct separation between ampulla and duct; no
diverticula; two groups of glands at ectal opening; extending caudo-dorsad to
union with corresponding part of opposite spermatheca; common duct leading
to lumen of alimentary canal; diameter of ampulla about twice that of duct.

DISCUSSION. — Slight differences between the Canadian specimens and
those described by Eisen from Nova Zembla exist in one or two sets of organs.
The spermiducal funnel in the former is not as stout as Eisen indicates in his
figure, and the ampulla of the spermatheca is less distinctly set off from the
duct, but these are differences of degree and are of little significance. Eisen
figures several glands at the ectal openings of the spermathecae but the Canadian
specimens have only two.

DISTRIBUTION. — Aside from the original discovery of a few specimens of
Henlea ochracea in Nova Zembla, nothing has hitherto been known concerning
its distribution. The discovery of enchytraeids apparently belonging to this
species in arctic North America is therefore a matter of interest and indicates
for this annelid a wide distribution.

Henlea arctica, n. sp.

A collection made July 2, 1915, from the mud at the margin of a tundra
pond, Bernard harbour, Northwest Territories, contained six enchytraeids,
two of which were mature. These annelids show a combination of characters
which does not seem to agree with descriptions of species already known. In
some respects, they appeared to resemble certain poorly described species and
incomplete descriptions have made the determination very puzzling, but the
lack of complete agreement has led to the final conclusion that these worms
must be regarded as representing a new species.

Oligochaeta 13 A

DEFINITION. — Length, 6-3-13 mm. Diameter, about 0-45 mm. Somites,
40-51. Colour of alcoholic specimens light brown. Prostomium blunt; round;
smooth. Setae 3-5 per bundle in both lateral and ventral rows, fewer at
extremities; all in bundle about same size; distinctly sigmoid, distal points
recurved. Clitellum slightly developed; on 12-13. Lymphocytes scanty;
ovoid. Brain about one and one-half times longer than wide; anterior and
posterior margins slightly concave; lateral margins divergent caudad; two
pairs of supporting strands from posterior part. Peptonephridia present; one
dorsal and one ventral strand; dorsal strand arising from digestive tract in 5
and extending caudad, thickened at origin, in contact with wall of digestive
tract to posterior part of 7 and extending under intestinal diverticulum, giving
off branches into coelom in 6; ventral strand also in contact with walls of digestive
tract in 3, 4, and 5, tubular, rather thick walled, apparently no branching but
merely irregular folding. Oesophagus enlarging abruptly into intestine in
anterior part of 8. Intestinal diverticulum of- peculiar structure; completely
surrounding digestive tract in 7; thin walled; large internal cavity; very few
folds of lining; no evidence of internal tubules; longitudinal, mid-dorsal partition
separating internal space into right and left parts; connection with digestive
tract very obscure and not definitely determined. Dorsal blood-vessel arising
in 8. Nephridia with anteseptal part as mere nephrostome; postseptal part
large; efferent duct arising from ventral surface of latter near septum. Spermi-
ducal funnel but little longer than wide; with thin, reflected collar; attached
end merging gradually into sperm duct for some distance. Sperm duct in 12
only; much coiled. Penial bulb of simple lumbricillid type; small, maximum
diameter only about one and one-half times greater than that of ventral nerve
cord in same transverse section; cells of one kind only in bulb. Spermathecae
with one group of very small glands at ectal opening; duct expanding gradually
into globular ampulla; ental end of ampulla joining corresponding part of
opposite spermatheca, dorsad of digestive tract, to form a common duct leading
into digestive tract in posterior part of 5; diverticula lacking.

Henlea, sp. (a).

Among the specimens collected at Pihumalerksiak island (Cockburn point) ,
Northwest Territories, July 15,' 1916, from between moss roots, one specimen
was found which, though too immature to identify to species, showed clearly
the characters of the genus Henlea. There are some indications that this
specimen may be an immature Henlea nasuta (Eisen).

Henlea, sp. (b).

The collections from Bernard harbour included one bottle containing four
specimens from the mud of a shallow tundra pond, May 31, 1916. Two of the
specimens were hopelessly deteriorated while the other two were immature,
thus making specific identification impossible. The two best preserved speci-
mens were sufficiently developed to make it certain that they belong to the
genus Henlea. There is some evidence that these specimens are immature
examples of the same form listed provisionally on a preceding page as Henlea
tenella (Eisen).

Enchytraeus hyalinus, var.

A collection from Pihumalerksiak island (Cockburn point), Northwest
Territories, July 15, 1916, contained eighty-seven specimens, of which about
one-fifth were mature. These enchytraeids were collected among moss roots,
and described in the field notes as " white worms," referring, no doubt, to the

14 A

Canadian Arctic Expedition, 1913-1$

colour of the living specimens. An inspection of this collection indicated that
more than one species was represented, and this was confirmed by subsequent
examination of cleared specimens and serial sections of selected individuals.
Not all of the material was sectioned and, since specific identification depends
upon such procedure, the identity of many of the worms was judged from t In-
external examination of cleared specimens and a comparison of the results with
those obtained by a study of the sectioned specimens. Most of this collection
belongs to a variety of Enchytraeus hyalinus (Eisen). The other species repre-
sented (Henlea sp. (a) and Mesenchytraeus johanseni n. sp.) are referred to
elsewhere. Unfortunately, the original description of hyalinus is meagre,
certain important structural features being unmentioned.

CHARACTERISTICS. — Length, about 11 mm. Maximum diameter, about
0-5 mm. Somites, about 50. . Colour " white " according to collection record;
alcoholic specimens yellowish brown. Setae straight; those of a bundle equal
in size; 2-3 in lateral bundles, mostly 3; almost inyariably 3 in ventral bundles,
except in immediate vicinity of extremities. Clitellum on 12-13; moderate in
size; incomplete on mid-ventral surface. Prostomium blunt; rounded; smooth.
Lymphocytes sparse; small; ovoid to circular. Brain about 1-8 times longer
than wide; lateral margins divergent caudad; anterior margin very slightly
concave; posterior margin straight or very shallowly concave; nearly circular
in transverse section through middle. One pair peptonephridia arising from
mid-dorsal part of oesophagus in 4; strands arising independently but almost in
contact; unbranched; thick-walled; somewhat contorted; each strand extend-
ing ventro-laterad from point of origin; thin-walled, somewhat spherical expan-
sion in middle of length of each organ, walls differing in structure from adjacent
ones; both organs confined to 4. Dorsal blood-vessel arising in 14. Anteseptal
part of nephridia small, comprising little more than mere nephrostome; post-
septal part large, efferent duct arising from posterior end of postseptal part.
Spermiducal funnel about three times longer than wide; sperm duct short,
confined to 12. Penial bulb of the typical enchytraeid type as defined by Eisen
(1905 :7-8); consisting of separate and distinct sets of glands, each opening
independently on surface of body; mainly composed of two well-developed,
multi-cellular, spherical glands or " cushions " on each side of body, arranged
in longitudinal sequence and close together but not in contact; sperm duct
opening to exterior independently of anterior gland on its ectal side; cells in
glands apparently all of same kind.

DISCUSSION AND DESCRIPTION.

When compared in detail with the original description of Enchytraeus
hyalinus. (Eisen) (1879 : 26-27), it is noted that the correspondence is very close
for all of those features mentioned, with the exception of the form of the sperma-
thecae. It appears that the specimens from Nova Zembla on which the original
description was based had spermathecae which showed an atrium-like enlarge-
ment of the duct. No such enlargement appears in the spermathecae of the Can-
adian specimens. There is some resemblance to the spermathecae in Enchytraeus
marinus as described by Moore (1902 : 80-82).

LYMPHOCYTES. — The original description also states that no " perigastric
cells " (lymphocytes) are present in the Nova Zembla material. If such obser-
vation was correct, another difference appears since lymphocytes do occur in
the specimens from northern Canada.

DEVELOPING OVA. — Masses of developing ova occurred in the usual position
in 12. In addition, ova were observed in the body-cavity scattered at' intervals
from 29 to 42 inclusive. The remoteness from the oviducal pores makes it
doubtful if such ova ever pass to the exterior.

Oligochaeta 15 A

DISTRIBUTION. — Thus far Enchytraeus hyalinus has been definitely known
only from Nova Zembla. The habitat of the original specimens is not given.
Friend (1912 : 222-223) has described, apparently with some hesitation, a variety
of hyalinus under the name densus from specimens existing under inland con-
ditions in England. The brief description of this variety corresponds closely
with that given above for the North American forms, with the exception of the
spermathecae. The structure of the penial bulb is not described and no com-
parison is possible. Until this variety densus is more certainly and more com-
pletely described, the writer- prefers to leave this arctic American variety without
a name.

Mesenchytraeus johanseni, n. sp.

As mentioned before, the collection made July 15, 1916, from moss roots
on Pihumalerksiak island (Cockburn point), Northwest Territories, contained
enchytraeids other than Henlea sp. and Enchytraeus hyalinus var. About ten
of the specimens show characters of the genus Mesenchytraeus. All of these
specimens which were sectioned proved to be representatives of a new species
of Mesenchytraeus, which is named in honour of the collector.

DEFINITION. — Length, 12-14 mm. Diameter, about 0-7 mm. Somites,
55-57. Colour " white " according to field notes accompanying specimens;
light yellowish in alcohol. Prostomium blunt; smooth; rounded. Head pore
near tip of prostomium. Setae sigmoid; similar in size; 3-5 in ventral rows,
2-4 in lateral rows, fewer at extremities. Clitellum well developed; on f 11-13.
Lymphocytes scanty; ovoid. Dorsal blood-vessel arising in 22-23; cardiac
body present. Chloragog cells with heavy load of brownish-black pigment.
Nephridia with typical mesenchytraeid structure; anteseptal part small; post-
septal region large; efferent duct arising from ventral part of latter near septum.
Spermiducal funnel about one and one-half times longer than wide. Sperm ducts
elongate, extending caudad to 16 within ovisac; much contorted. Testes
massive; undivided. Sperm sac present; single; extending caudad to
16; within ovisac; bifurcating at posterior end. Ovisac present; extending
caudad to 16; bifurcating at posterior end. Penial bulb of mesenchytraeid
type; small; atrium present, largely within body of bulb, about five irregular
atrial glands extending into coelom; no accessory glands observed. One pair
spermathecae; few unicellular glands at'ectal opening; no distinctly differen-
tiated ampulla, body of organ and duct of about same diameter; each sperma-
theca joining digestive tract very near mid-ventral line in posterior part of 5;
each with two long, slender diverticula extending dorsad over alimentary canal.

The above description is based upon a careful study of three mature speci-
mens. This species exhibits some resemblances to Mesenchytraeus solifugus
(Emery), but lack of agreement appears in such features as the amount of pig-
mentation, place of origin of the dorsal blood-vessel, number of atrial glands,
and presence or absence of accessory glands.

Lumbricillus lineatus (Mull.)

One specimen belonging to the genus Lumbricillus was collected under
sea-weed on the beach at Bernard harbour, Northwest Territories, June 12, 1916.
The posterior part of the body of the worm was badly deteriorated but fortu-
nately the anterior 19-20 somites were in a fair state of preservation and thus
specific identification was possible. The specimen lacked a little of being
completely sexually mature but the development of the various organs was
sufficiently completed to make it evident that the worm is Lumbricillus lineatus
(Mull.).

16 A

Canadian Arctic Expedition, 1913-18

Another collection made on July 19, 1915, from green algae in a creek
outlet, Barnard harbour, Northwest Territories, contained twenty-five specimens,
of which fourteen were mature. All the specimens of this collection which
were sectioned also proved to be Lumbricillus lineatus (Mull.).

CHARACTERISTICS. — Length, about 10-12 mm. Somites, about 50. Diam-
eter, in region of clitellum approximately 1 mm. Colour " yellow brown "
according to data accompanying the June 12th collection; dark brown in alcohol.
Setae sigmoid; approximately uniform in size; 4-6 in ventral bundles; 3-4 in
lateral bundles. Clitellum slightly developed; on 12-13; incomplete on ventral
surface of body. Anterior and posterior margins of brain slightly concave;
lateral margins divergent caudad; length apparently about equal to maximum
width. Peptonephridia lacking. Dorsal blood-vessel arising in 14. Nephridia
with very small anteseptal part and large postseptal part; efferent duct arising
from ventral surface of posterior part of latter. Testes multilobed. Spermi-
ducal funnel cylindrical; about three times longer than diameter. Sperm duct
confined to 12. Penial bulb of typical lumbricillid form; cells of one kind only
in interior. One pair spermathecae in 5; thick-walled; no diverticula; duct
and ampulla large, without sharply defined line of division; ental region much
narrowed and reflected cephalad before reaching digestive tract; duct shorter
than ampulla, surrounded full length by conspicuous glands; ental portion cf
each spermatheca uniting independently with dorso-lateral surface of digestive
tract; lumen of spermatheca uniting with lumen of latter. Ventral glands in
13 and 14; anterior larger than posterior; similar in shape; surrounding ventral
nerve cord closely on ventral and lateral surfaces only.

DISCUSSION AND DESCRIPTION.

COLOUR. — The data accompanying the June 12, 1916, specimen contained
the statement " yellow brown worm," which the writer interprets as referring
to the colour of the living animal. Since Lumbricillus lineatus has been described
from other regions as being brownish red, or red tinged with yellow, it would
appear that the Bernard harbour form is typical so far as colour is concerned.
l$o data are available on the colour of the living worms in the collection of July
19, 1915.

SPERMATHECAE. — In the specimens collected July 19, 1915, the spermathecae
were as described above, but in one 'specimen collected June 12, 1916, they
lack the connection of the lumen of the ampulla with that of the digestive tract.
The ental portion of each spermatheca is adherent to the digestive tract but
the connection has not yet been established. That this is a condition of imma-
turity is suggested by the fact that there is evidence that such connection is
the last of the parts of the spermatheca to be completed. The total absence
of spermatozoa in the spermathecae also indicated immaturity.

PENIAL BULB. — It is interesting to note that in the single, partially
immature specimen, collected on June 12, 1916, there is a distinct difference
in the development of the penial bulb on the two sides of the animal. The
right penial bulb is apparently completely developed, and all of the parts are
represented in their usual condition in the lumbricillid type. However, on the
left side, the bulb is in a very early stage of development, represented only by
a small mass of developing tissues in the body-wall, and none of the regions
of the fully-developed bulb can be distinguished. Mature specimens in the
other collection showed the penial bulbs with the structure as previously described
(Welch, 1917a : 128).

DISTRIBUTION. — The writer (1917a : 123-125) has recently pointed out
that the enchytraeids formerly regarded as distinct and designated under the
names litoreus, subterraneus, verrucosus, lineatus, and agilis, are all one and the
same. This synonymy must, of course, be taken into account in considering

Oligochaeta 17 A

the matter of distribution. It was. also shown that lineatus is evidently a
cosmopolitan form, having been reported from England, Ireland, Scotland,
Hebrides, North Russia, Denmark, Germany, France, Bohemia, Switzerland,
Terra del Fuego, and the coast of the New England States. It occurs commonly
under sea-weed on the shore in various parts of the globe. It has also been
found existing under fresh-water conditions.

Lumbricillus, sp. (a).

A collection from Pihumalerksiak island (Cockburn point), Northwest
Territories, July 15, 1916, contained thirty-three enchytraeids, of which thirty-
two were of similar size and general external appearance. These worms were
taken from the mud in a " water hole." Unfortunately, none of them are
mature and specific identification is impossible. However, all of the specimens
examined in detail show the testes composed of a number of distinct lobes —
a character which has been regarded as diagnostic of the genus Lumbricillus.
Mention should be made of the fact that these specimens, while possessing the
.divided testes, have straight setae — a feature which departs from the usual
definition of the genus Lumbricillus.

Lumbricillus, sp. (b).

The above mentioned collection from Pihumalerksiak island (Cockburn
point), Northwest Territories, July 15, 1916, also contained one specimen
which shows distinct differences when compared with the other individuals of
the same lot. The significant feature is the possession of sigmoid setae. It also
exhibits the divided condition of the testes and apparently belongs to the genus
Lumbricillus. However, the immaturity of the specimen prevents a specific
identification.

GENEEAL CONSIDERATIONS.

PIGMENTATION. — In an earlier paper, the writer (1916 : 120) discussed the
presence of the pigmentation which occurs in all of the known glacier enchytraeids
and pointed out that it does not seem possible to explain its presence on the
grounds of low temperatures, since certain species in warmer latitudes show
pigmentation while others in frigid conditions show no indication of it. In this
connection, it is interesting to note that none of the specimens collected in
arctic Canada exhibit any marked pigmentation such as occurs in the above
mentioned glacier worms. The specimens of Mesenchytraeus johanseni n. sp.
show a heavy load of brownish-black pigment granules in all the chloragog cells
but none elsewhere. This is also true of the specimens of Lumbricillus sp. (a).
No pigmentation of any sort was observed in any of the other specimens.
Additional evidence is thus presented against the view that heavy pigmentation
of the internal organs in enchytraeids is directly correlated with frigid habitats.

SEASONAL RELATIONS. — According to the field records of the collector,
some of the enchytraeids of this collection were found alive among moss roots
in dried-out stream beds in mid-summer. Possibly this is an indication of their
method of surviving drouth conditions of the summer season. At least it would
appear that these aquatic worms can tolerate a marked reduction of the water
in the surroundings. Another noteworthy feature is the very low temperatures
to which these worms are subjected during the winter months. The remarkable
resistance of certain enchytraeids to frigid conditions has been discussed in
other papers (Welch, 1916; 1917b) and it is evident that the worms living in
arctic Canada are also successfully adjusted to long exposure at freezing
temperatures.

ISA

Canadian Arctic Expedition, 1913-18

DISTRIBUTION OF ENCHYTRAEIDAE. — The enchytraeid collections on which
this report is based are of particular interest because of the fact that nothing
has hitherto been known concerning this group in the region visited by the
Expedition. In fact, the entire arctic portion of the North American continent
is practically unknown territory with ivgnrd to this group of animals, although,
as will be mentioned later, there is reason to believe that it is abundantly
represented there. Eisen (1872 : 122-123) reported Lumbricillus profugus
(Eisen), under the name Enchytraeus pagenstecheri Ratzel, from Godhavn, Green-
land. According to Michaelsen (1900 : 82), 0. Fabricius (1870) reported
Lumbricillus minutus (Mull.), under the name Lumbricus minutus, from Green-
land, but the exact locality was apparently not given. Ude (1896: 27) recorded
the occurrence of Enchytraeus albidus Henle (Enchytraeus vejdovskyi Eisen) at
Karajak, Greenland. Emery (1898), Moore (1899), and Welch (1917b) made
studies on Mesenchytraeus solifugus (Emery) collected in Alaska on and in Ihe
general vicinity of Mount St. Elias. Eisen (1905) studied the collections of
the Harriman Alaska Expedition on the west coast of Alaska, the vicinity of
Port Clarence being the northern limit. No other records occur in the literal u re,
and it thus appears that the Canadian Arctic Expedition collections present the
first indications of the enchytraeid fauna existing in the vast territory between
the west coast of Greenland and Bering strait.

It will be noted that the representatives of the genus Henlea comprise a
considerable part of the collections. Ude (1901 : 19) has already pointed out
that Henlea is especially well represented in the arctic region. The other genera
appearing in the Canadian Arctic Expedition collections are also known from
other "parts of the far north. The common occurrence of Mesenchytraeus in
the high latitudes has been pointed out by Ude (1901 : 23), and on the snow-
fields and glaciers of high altitudes by Emery (1898), Moore (1899), and Welch
(1916 : 1917b).

The arctic Canadian enchytraeid fauna, as indicated by the collections of
the Canadian Arctic Expedition, shows a similarity to that of arctic Europe
and Asia which is worthy of special mention. As nearly as specific identifica-
tions have been possible, all of the species represented by sexually mature
material, exclusive of new forms, are known from arctic Eurasia, four having
been originally described from that region. Two species (Henlea nasuta and
Lumbricillus lineatus) are apparently cosmopolitan.

While the list of enchytraeids taken by this Expedition is not extensive,
and while the collections were not made in an exhaustive manner, the results
confirm previous conclusions concerning the geographical relations of this family,
viz., that the group is preeminently northern in its distribution. Such Arctic
regions as have been examined yield evidence of a rich enchytraeid fauna.
The abundance of some of these worms reported by earlier collectors is an
indication of their successful adjustment to the rigid conditions of the environ-
ment. There is every indication that future collections will show the Enchy-
traeidae to be one of the most prominent and most widely distributed of the
groups of animals occupying the north frigid zone.

Oligochaeta 19 A

LITERATURE CITED, PART II.
Cejka, B.

1910. Die Oligochseten der Russischen in den Jahren 1900-1903 unternommenen Nord-
polarexpedition. I. Ueber eine neue Gattung der Enchytrseiden (Hepatogaster) .
Mem. Acad. Sci. St. Petersb., (8), 29, No. 2, 1-29; 3 pi.

Eisen, G.

1872. Om nagra arktiska Oligochseter. Ofv. Ak. Forh., 29 : 119 — 124.

1879. On the Oligochseta collected during the Swedish Expeditions to the Arctic Regions

in the Years 1870, 1875 and 1876.
Kong. Sv. Vet. Akad. HandL, Bandet 15, N:o 7, 49 pp.; 16 pi.

1905. Enchytraeidae of the West Coast of North America. Harriman Alaska Expedition,
12 : 1—126; 20 pi. New York.

Emery, C.

1898. Diagnosi di un nuovi genere e nuova specie di Anellidi della famiglia degli Enchy-

trseidae.
Atti della R. Accad. dei Lincei, (5), 7 : 110—111.

Fabricius, O.

1780. Fauna Groenlandica. (Original not seen.)

Friend, H.

1912a. British Enchytrseids. IV. The Genus Henka. Journ. Royal Micr. Soc., pp.
577—598.

1912b. New British Oligochsets.

The Zoologist, (4), 16 : 220—226.

1914. British Enchytrseids. VI. New Species and Revised List. Journ. Royal Micr. Soc.,
pp. 128—154.

Michaelsen, W.

1900. Oligochaeta. Das Tierreich, 10 Lief. XXIX +575 pp.; 13 fig. Berlin.

Moore, J. P.

1899. A Snow-inhabiting Enchytraeid (Mesenchytraeus solifugus Emery) collected by Mr.

Henry G. Bryant on the Malaspina Glacier, Alaska.
Proc. Acad. Nat. Sci. Phil., pp. 125—144; 1 pi.

1902. Some Bermuda Oligochaeta, with a Description of a New Species. Proc. Acad. Nat.
Sci. Phil., pp. 80—84; 2 fig.

Ude, H.

1896. Enchytraeid en. Hamburger Magalh. Sammelr., 42 pp.; 1 pi.

1901. Die arktischen Enchytraiden and Lumbriciden, sowie die geographische Verbreitung

dieser Familien.
Fauna Arctica, Band II, Lief. 1, 34 pp.; 2 pi.

Welch, P. S.

1914. A new Henlea (Enchytrseidse) from Northern Michigan. Trans. Am. Micr. Soc.,
33 : 155—164; 1 pi.

1916. Snow-field and Glacier Oligochseta from Mt. Rainier, Washington. Trans. Am.
Micr. Soc., 35 : 85—124; 4 pi.

1917a. The Enchytrseidse (Oligochseta) of the Woods Hole Region, Mass. Trans. Am.
Micr. Soc., 36 : 119—138.

1917b. Alaskan Glacier Worms (Oligochseta). Bionomical Leaflets, McGill University,
Montreal, No. 2.

Q. Canadian Arctic Expedition,
115 1913-1918
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